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INSTITUTION NOILN. :* ee ee eo a | > =a = =) 4 =) = > f= *, > kK S E = ae = = oO z z Zz iS SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS Saluvugi7a_LIBRA NVINOSHLIWS SMITHSONIAN NVINOSHLINS XS AW NS SMITHSONIAN NVINOSHLINS SMITHSONIAN LIBRARIES SMITHSONIAN INSTITUTION NOILN. N ) -S SMITHSONIAN N NVINOSHLIWS NOILNLILSNI LIBRARIES NOILNLILSNI NOILNLILSNI INSTITUTION NOILALILSNI NVINOSHLINS S3IYVYdIT LIBR. 7 INSTITUTION S3INVUSIT LIBRARIES INSTITUTION INSTITUTION S sazI¥vVudIT LIBRARIES S S31u¥Vvud =z < =a «ss aN x BSA = = S SAIYVYaIT_ LIBRARIES SMITHSONIAN _ INSTITUTION NOILN N S) N ISSN: 0098-4590 cilentist Volume 52 Winter 1989 Number 1 CONTENTS Response of Laying Hens to Various Grains and PeePeAMe SUP PICTMENtAatiON ooo. kei ee ee eis ee ee ee os R. D. Miles, J. E. Marion, R. D. Barnett, N. Ruiz, and R. H. Harms i First Record of Pawpaw Consumption by the Florida Mouse ....... Cheri A. Jones a Atlantic White Cedar (Chamaecyparis thyoides) in the NMR TIALS we See ees Sees chery ese eo wa 24m es elec bor dd cid’ oe Daniel B. Ward and Andre F. Clewell 8 Water Quality Efficiency of an Urban Commercial Wet Detention Stormwater Management System at the Boynton Beach Mall in South Palm Beach County, ORAM cst satis Me OOS eM bee ble POL Orme i eae ast) Jeffrey Dee Holler 48 Allocation of Energy Resources in the Freshwater Angiosperms Vallisneria americana Michx. and Potamogeton pectinatus cs TL ELCTE CIS 0 0a aa ees Ota Nee Clinton J. Dawes and John M. Lawrence 58 Puma ICAPIHETILOL REVIEWEIS 2. 0. 1k ki ct cee cs ete ee bes 64 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1988 Editor: Dr. DEAN F. MarTIN _Co-Editor: Mrs. BARBARA B. MARTIN Chemical and Environmental Management Services (CHEMS) Center Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoriDA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. Applica- tions may be obtained from the Executive Secretary. Both individual and institutional members receive a subscription to the FLoripa ScrENTIsST. Direct subscription is avail- able at $20.00 per calendar year. Original articles containing new knowledge, or new interpretation of knowledge, are welcomed in any field of Science as represented by the sections of the Academy, viz., Biological Sciences, Conservation, Earth and Planetary Sciences, Medical Sci- ences, Physical Sciences, Science Teaching, and Social Sciences. Also, contributions will be considered which present new applications of scientific knowledge to practical problems within fields of interest to the Academy. Articles must not duplicate in any substantial way material that is published elsewhere. Contributions are accepted only from members of the Academy and so papers submitted by non-members will be accepted only after the authors join the Academy. Instructions for preparation of manuscripts are inside the back cover. Officers for 1987-88 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. LESLIE SUE LIEBERMAN Treasurer: Dr. ANTHONY F. WALSH Department of Anthropology 5636 Satel Drive University of Florida Orlando, Florida 32810 Gainesville, Florida 32611 Executive Secretary: President-Elect: Dr. MARVIN L. Ivey Dr. ALEXANDER DICKISON Department of Natural Sciences Department of Physical Sciences St. Petersburg Junior College Seminole Community College P.O. Box 13489 Sanford, FL 32771 St. Petersburg, FL 33733 Program Chairs: Dr. GEorcE M. Dooris Secretary: Dr. PATRICK J. GLEASON Dr. Patricia M. Dooris 1131 North Palmway P.O. Box 2378 Lake Worth, Florida 33460 St. Leo, Florida 33574 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist ~~ - QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BARBARA B. MarrTIN, Associate Editor Volume 52 Winter 1989 Number 1 Agricultural Sciences RESPONSE OF LAYING HENS TO VARIOUS GRAINS AND ENZYME SUPPLEMENTATION R. D. Mixes”, J. E. Marton®, R.D. BARNETT”, N. Ruiz” anp R. H. Harms” (Department of Poultry Science, University of Florida, Gainesville, FL 32611, Department of Poultry Science, North Carolina State University, Raleigh, NC 27695, “North Florida Research and Education Center, University of Florida, Quincy, FL 32351-9529, “)Nutrena Feed Division #4, P.O. Box 9300, Minneapolis, MN 55400. ABSTRACT: Two experiments were conducted to determine if an enzyme preparation contain- ing alpha amylase and bacterial proteases would result in improved laying hen performance when supplemented to diets containing various grains. In Experiment 1, the enzyme preparation was added at 0, .05 and .15 % to a corn soybean meal basal diet. Experimental diets were mixed every two weeks. Fourteen replicates of 10 hens each were fed the experimental diets ad-libitum during five 28-day periods. Results indicated no significant beneficial or detrimental effect on egg production, feed conversion or egg weights due to the addition of the enzyme. Grains used for feeding comparisons in experiment 2 were yellow corn, milo, Florida 301 variety of wheat and Beagle 82 variety of triticale. Each grain was substituted for 50% of the yellow corn in the basal diet. Diets were balanced to maintain equal levels of sulfur amino acids and lysine while protein and energy were allowed to vary among treatments. Treatments consisted of eight replicate groups of 5 hens and were fed for four 28-day periods. Results indicated that grain source but not enzyme, influenced feed consumption and conversion. Neither grain source nor enzyme statisti- cally influenced egg production, egg weight or egg specific gravity. New cereal grain varieties are constantly being developed for use in cer- tain geographical locations in the U.S. These varieties merit testing for their potential as a partial or total replacement for corn in poultry diets. The use of many of these varieties would give Florida farmers an opportunity to double crop cultivated land and might also lower the cost of poultry feeds in the Southeast. The nutritive value of triticale, a grain developed from the intergeneric cross of durum wheat and rye, has been evaluated for lambs (Sherrod, 1972; 1Florida Agricultural Experiment Stations Journal Series No. 8854 2 FLORIDA SCIENTIST [Vol. 52 Jordan and Hanke, 1972), swine (Erickson, et al., 1979; Hale, 1983) and broilers (Sell, et al. 1962; Bragg and Sharby, 1970; Wilson and McNab, 1975). Ruiz et al. (1987) reported that Florida 301 wheat is an excellent cereal grain for broiler chicks as measured by feed efficiency and growth response. However, the growth response to broiler chicks was inconsistent when Beagle 82 triticale was fed in three different experiments. Birds fed the triticale-based diets had better feed efficiency than birds fed corn-based diets. Kim and co-workers (1976) fed wheat-based diets to laying hens and found that egg production and body weights were equal to hens receiving corn- based diets. Quart and co-workers (1986) fed 68 week old, laying hens, diets containing 50% wheat of varying grain sizes (fine to whole). These authors reported no differences in hen performance when compared to hens fed 100% finely ground corn. Enzyme preparations have been used to enhance feed utilization by tur- keys (Moran and McGinnis, 1965, 1966, 1968; Fry et al., 1958), broilers (Reese et al., 1983; Willingham et al., 1959; Rexen, 1981; Potter et al., 1965; Leong et al., 1961) and laying hens (Gleaves and Dewan, 1970; Gohl, e¢ al., 1978). The objective of the two experiments reported herein was to evaluate laying hen performance when fed different grains and also evaluate the effect of adding a commercially available enzyme preparation to these grain based diets. MATERIALS AND METHOps—Enzyme supplement and grains. The enzyme supplement KEMZYME® was obtained from Kemin Industries, Des Moines, Iowa. The product was a free flowing, dry preparation and contained alpha-amylase activity from Bacillus licheniformis, Fun- gal and bacterial protease activity from Aspergillus oryzae and Bacillus subtilis, respectively. The enzyme preparation was stored refrigerated at 2-5C until being incorporated into diets at time of mixing. The yellow corn in experiment 1 and the yellow corn and milo in experiment 2 were obtained from a commercial source. The Beagle 82 triticale was selected, developed and tested at the Georgia Coastal Plain Experiment Station, Tifton, GA and the North Florida Research and Education Center (NFREC) Quincy, FL (Hale, 1983; Barnett et al., 1982). The Florida 301 wheat was developed by the University of Florida at the NFREC in cooperation with the Science and Education Administration, US Department of Agriculture (Barnett and Luke, 1980). The proximate analysis, calcium and phosphorus contents of Beagle 82 triticale and Florida 301 wheat were the same as reported by Ruiz et al. (1987). Experiment 1: A total of 420 individually caged 36 weeks old Single Comb White Leghorn (SCWL) hens were fed a yellow corn-soybean meal basal diet or the same basal diet supple- mented with .05 or .15% KEMZYME® (Table 1). Each dietary treatment was fed ad libitum to 14 replicate pens of 10 individually caged hens each for five 28-day periods. Water was furnished by a flow-through trough system for 15 min. every two hr. Experimental diets were mixed at two week intervals. Daily egg production records were maintained and were summarized at the end of each 28-day experimental period. Egg weights and egg specific gravity were determined on all eggs collected on day 21 of each 28-day experimental period. The experiment was conducted from March through July. Experiment 2: The grains tested in this study were substituted for 50% of the yellow corn in the basal diet (Table 1). Diets were formulated so that sulfur amino acids and lysine levels were kept constant. The levels of other amino acids, protein and energy were allowed to vary among treatments. Experimental diets were fed according to the recommendation of Harms (1981) and nutrient levels of the diets were adjusted based on daily feed intake. A total of 320 individually caged 50 weeks old SCWL hens were housed in an open-type house. Each dietary treatment was No. 1, 1989] MILES, ET AL.—RESPONSE OF LAYING HENS 3 TaBLE 1. Composition and calculated analysis of diets used in Experiment 1 and 2. Experiment il Y) Treatment Ingredient 1 2 3 4 %o Yellow Corn COPA 72.02 36.00 36.00 36.00 Milo 36.00 Beagle 82 triticale 38.19 Florida 301 wheat SM o74 Soybean meal (48.5% ) 18.74 18.08 17.86 15.90 16.92 Iodized salt 0.40 0.42 0.42 0.42 0.42 Limestone 8.39 (236 7.35 7.35 7.36 Dicalcium phosphate 1.65 on 1.57 1.54 1.53 (24% Ca, 18.5% P) Microingredients* 0.50 0.50 0.50 0.50 0.50 DL-methionine 0.10 0.06 0.11 0.09 0.06 Calculated analysis Crude protein, % [555 15.10 15.02 15:27 15.80 Arginine, % 1.04 0.91 0.91 0.98 0.91 Sulfur amino acids, % 0.62 0.60 0.60 0.60 0.60 Lysine, % 0.76 0.72 0.74 0.72 0.76 Tryptophan, % 0.19 0.18 0.18 0.18 0.19 Calcium, % 3.40 3.20 3220 3.20 3.20 Total phosphorus, % 0.45 0.60 0.60 0.60 0.60 Metabolizable energy, kcal/kg 2899 2911 2883 2799 2809 aSupplied per kilogram of diet: 6600 IU vitamin A; 2,200 IU vitamin D,; 2.2 mg menadione dimethylpyrimi- dinol bisulfite; 4.4 mg riboflavin; 13.2 mg pantothenic acid; 39.6 mg niacin; 499 mg choline chloride; 0.022 mg vitamin B12; 125 mg ethoxyquin; 60 mg manganese; 50 mg iron; 6 mg copper; 0.198 mg cobalt; 1.1 mg iodine; 35 mg zinc. TABLE 2. Response of laying hens to enzyme supplementation of a yellow corn-soybean meal based diet in Experiment 1. Feed Feed Egg Egg Dietary Hen-Day Egg Consumption Conversion Weight Specific Treatment Production (%) (gm/b/day) (Kg/doz) (g) Gravity Control 80.392 iene 1.462 63:62 07083 KEMZYME (.05%) 81.054 98.38 1.462 63.44 1.07798 KEMZYME (.15%) 79.692 97.48 1.482 63.34 1.0783 aMeans within a column with different superscripts differ significantly (p <.05). fed ad libitum to 8 replicate groups of 5 hens each for four 28-day periods. The grain based experimental diets were fed with and without .05% KEMZYME. The experiment was conducted from late September through January. RESULTS AND Discuss1on— Hens fed enzyme preparation at two different levels in the all corn-soybean meal based diet had no better performance than those hens fed the control diet (Table 2). Hen day egg production, feed con- sumption, feed conversion, egg weight and egg specific gravity were similar among treatments. In Experiment 2, feed consumption and feed conversions (Table 3) were significantly influenced (Table 4) by grain source but not by enzyme supple- 4 FLORIDA SCIENTIST [Vol. 52 TABLE 3. Average mean performance criteria of laying hens to enzyme supplementation of a diet containing various grains (Experiment 2). Feed Feed Egg Egg Dietary Hen-day Egg Consumption Conversion Weight Specific Grain Production (% ) (gm/b/day) (Kg/doz) (g) Gravity No enzyme supplementation Corn 16.1 100.8 Gh 63.0 1.0807 Milo ies 96.4 1.50 62.7 1.0799 Wheat 78.3 102.3 1.59 62.8 1.0816 Triticale loa 105.6 1.62 63.0 1.0796 + Enzyme supplementation Corn 77.6 O9e 1.55 64.9 1.0816 Milo 79.6 98.8 1.50 63.6 1.0812 Wheat 78.8 104.3 1.59 63.3 1.0810 Triticale (shar 109.0 1.68 64.3 1.0831 a See Table 4 for statistical analysis. TABLE 4. Statistical analysis of production of hens fed different dietary grains with and with- out enzyme supplementation?. Probability F Measure Grain (G) Enzyme (E) GxE Egg production 0.5726 0.2673 0.9515 Feed consumption 0.0001 0.2099 0.3513 Feed conversion 0.0001 0.7349 0.3495 Egg weights O17 0.0699 0.8896 Egg specific gravity 0.7747 0.0615 0.0877 aSee Table 3. mentation. The hens fed the diet containing milo consumed the least amount of feed and exhibited the best feed conversions. Hens fed diets with Triticale consumed the greatest amount of feed and had the poorest feed conversion. Hens fed the wheat based diets were intermediate to the hens fed the corn and Triticale diets in feed conversion. There were no significant differences in egg production due to diet among any treatments. | The type of grain did not affect egg weights or specific gravity. However, both measures were approaching significance due to enzyme supplementa- tion. Even though not statistically significant the egg weights were numeri- cally heavier when the enzyme was supplemented to the diet. This may have been attributed to the increased energy intake due to the increased feed con- sumption observed when the enzyme was supplemented. Normally, when egg weight is less, specific gravity is greater. However, such was not the case in this experiment. As egg weight increased due to enzyme supplementation the specific gravity increased. It is unlikely that the small increase in calcium consumption due to a 2-4 gram increase in feed consumption resulted in the increase in egg specific gravity. The hens fed wheat and triticale consumed more feed and this resulted in poorer conversion. This could have resulted from a combination of two rea- No. 1, 1989] MILES, ET AL.—RESPONSE OF LAYING HENS 5 sons. Both, the energy value and density of these ingredients were less than those of corn and milo. With current feed prices and calculated energy values from these data collected in this experiment triticale had approximately 85 % the feeding value of corn and wheat had a value of 88%. LITERATURE CITED BarNETT, R.D., AND H.H. Luke. 1980. Florida 301 a new wheat for multiple cropping systems in North Florida. Circ. $-273, Agric. Exp. Sta. Inst. Food Agric. Sci. Univ. Florida, Gaines- ville, FL. BarNETT, R.D., D.D. Morey, H.H. Luke, ann P.L. PrAHuer. 1982. Beagle 82 triticale a new winter feed grain for multiple cropping systems in the coastal plains regions of South Georgia and North Florida. Circ. S-297, Agric. Exp. Sta. Inst. Food Agric. Sci., Univ. Florida. Bracc, D.B., AND T.F. SHArBy. 1970. Nutritive value of triticale for broiler chick diets. Poultry Sci. 49:1022-1027. Erickson, J.P., E.R. Mruuer, F.C. Exyior, PK. Ku, anp D.E. ULtrey. 1979. Nutritional evalua- tion of triticale in swine starter and grower diets. J. Anim. Sci. 48:547-553. Fry, R.E., H.B. ALLRED, L.S. JENSEN, AND J. McGinnis. 1958. Influence of enzyme supplemen- tation and water treatment on the nutritional value of different grains for poults. Poultry Sel. Of 731 2-370. G.eaves, E.W., AND S. Dewan. 1970. Influence of a fungal enzyme in corn and milo layer rations. Poultry Sci. 49:596-598. Gout, B., S. ALDEN, K. ELWINGER, AND S. THOMKE. 1978. Influence of beta-glucanase on feeding value of barley for poultry and moisture content of excreta. Br. Poult. Sci. 19:41-47. Hater, O.M. 1983. Triticale, peanut meal, sunflower seed meal and other by-products for swine. 135-146. In: Proc. Georgia Nutr. Conf. Hate, O.M. 1984. Performance of swine fed grain sorghum and Beagle 82 triticale diets. Pp 53- 61. In: Proc. Georgia Nutr. Conf. Harms, R.H. 1981. Specifications for feeding commercial layers based on daily feed intake. Feedstuffs 53(47):10,40-41. JorpANn, R.M., anp H.E. Hanke. 1972. Finishing lambs with triticale, barley or corn. Feedstuffs 44(19):30. Kim, S.M., M.B. Parte, S.J. REEpy, AND J. McGinnis. 1976. Effects of different cereal grains in diets for laying hens on production parameters and liver fat content. Poult. Sci. 55:520- 530. Leone, K.C., L.S. JENSEN, AND J. McGinnis. 1961. Influence of fiber content of diet on the chick growth response to enzyme supplements. Poultry Sci. 40:615-619. Moran, E.T., anp J. McGinnis. 1965. The effect of cereal grain and energy level of the diet on the response of turkey poults to enzyme and antibiotic supplements. Poultry. Sci. 44:1253- 1261. Moran, E.T., anp J. McGinnis. 1966. A comparison of corn and barley for the developing turkey and the effect of antibiotic and enzyme supplementation. Poultry Sci. 45:636-639. Moran, E.T., ano J. McGinnis. 1968. Growth of chicks and turkey poults fed western barley corn grain-based rations: Effect of autoclaving on supplemental enzyme requirement and - asymmetry of antibiotic response between grains. Poultry Sci. 47:152-158. Ouart, M.D., J.E. Marion, AND R.H. Harms. 1986. Influence of wheat particle size in diets of laying hens. Poultry Sci. 65:1015-1017. Potter, L.M., M.W. SHutz, AnD L.D. Matrterson. 1965. Metabolizable energy and digestibility coefficients of barley for chicks as influenced by water treatment or by presence of fungal enzyme. Poultry Sci. 44:565-573. Reese, G.L., R.S. Karc, B.I. FANCHER, AND P.W. Wa.proup. 1983. The use of supplemental enzymes in the diet of broiler chickens. Nutr. Rep. Int. 28(4):919-929. REXEN, B. 1981. Use of enzymes for improvement of feed. Anim. Feed Sci. Technol. 6:105-114. Ruiz, N., J.E. Marion, R.D. Mites, AnD R.D. Barnett. 1987. Nutritive value of new cultivars of triticale and wheat for broiler chick diets. Poultry Sci. 66:90-97. SELL, J.L., G.C. Hopcson, ano J.H. SHEBEsxI. 1962. Triticale as a potential component of chick ration. Can. J. Animal Sci. 42:158-166. 6 FLORIDA SCIENTIST [Vol. 52 SHERROD, L.B. 1972. Nutritive value of selected triticale and wheats. J. Anim. Sci. 34:909 (Ab- str). WILLINGHAM, H.E., L.S. JENSEN, AND J. McGinnis. 1959. Studies on the role of enzyme supple- ments and water treatment for improving the nutritional value of barley. Poultry Sci. 38:539-544. Witson, B.J., AND J.M. McNas. 1975. The nutritive value of triticale and rye in broiler diets containing field beans (vicia Faba L.) Br. Poult. Sci. 16:17-22. Florida Sci. 52(1): 1-6. 1989. Accepted: May 4, 1988. No. 1, 1989] JONES—PAWPAW CONSUMPTION ff FIRST RECORD OF PAWPAW CONSUMPTION BY THE FLORIDA MOUSE—Cheri A. Jones, Department of Zoology, University of Florida, Gainesville, Florida 32611 Asstract: A Florida mouse was observed collecting and transporting a fruit of the flag pawpaw (Asimina incarna). This is the first record of pawpaw in the diet of this species. THE Florida mouse (Podomys floridanus) is known to eat insects, seeds, nuts, fungi, and other plant material (Layne, 1978), but details of its natural diet have not been reported. On 20 July 1987 on the Katharine Ordway preserve in Putnam County, Florida, I watched a Florida mouse collect and transport a fruit of the flag pawpaw (Annonaceae: Asimina incarna). The mouse, an adult male weighing 29 g, was captured in a Sherman trap at the entrance of an inactive burrow of the gopher tortoise (Gopherus polyphe- mus). After the mouse was released, at approximately 0600 EST, he wan- dered through the vegetation surrounding the apron of the burrow. Then he climbed a small pawpaw plant to a height of 30 cm, knocking off a ripe fruit as he did so. After looking through the leaves, the mouse appeared to see the pawpaw fruit on the ground. He ran down the stem, picked up the fruit in his mouth, and carried it to another tortoise burrow 11.4 m away, dropping it once enroute. Ripe pawpaws subsequently presented to captive Podomys were entirely consumed except for the seeds, which were gnawed upon. The flag pawpaw is abundant in the longleaf pine-turkey oak sandhills on the Ordway Preserve and might be a significant food source for Florida mice and other mammals. Garner and Landers (1981) and Norman and Clayton (1986) reported that the gopher tortoise eats pawpaws, and Norman and Clayton (1986) also suggested that the fruits might be dispersed by small mammals. This apparently is the first record of a small rodent taking the fruit of A. incarna. ACKNOWLEDGEMENTS—I thank R. A. Kiltie, J. F. Eisenberg, and two anonymous reviewers for their comments on the manuscript. LITERATURE CITED Garne_r, J. A., AND J. L. LANpErs. 1981. Foods and habitat of the gopher tortoise in southwestern Georgia. Proc. Ann. Conf. S.E. Assoc. Fish and Wildl. Agencies 35:120-134. Layne, J. N. 1978. Florida mouse. Pp. 21-22. In: Layne, J. N. (ed.). Rare and Endangered Biota of Florida. Vol. 1: Mammals. Univ. Presses of Florida, Gainesville. Norman, E. M., anno D. Crayton. 1986. Reproductive biology of two Florida pawpaws: Asimina obovata and A. pygmaea (Annonaceae). Bull. Torrey Bot. Club. 113:16-22. Florida Sci. 52(1): 7. 1989. Accepted: April 19, 1988. Biological Sciences ATLANTIC WHITE CEDAR (CHAMAECYPARIS THYOIDES) IN THE SOUTHERN STATES DANIEL B. Warp” AND ANDRE F. CLEWELL” ()Department of Botany, University of Florida, Gainesville, FL 32611; ®A. F. Clewell, Inc., 1345 University Parkway, Sarasota, Florida 34243 Asstract: Atlantic white cedar (Chamaecyparis thyoides (L.) BSP.), an often dominant swamp-forest tree of the eastern seaboard, is represented in the southern part of its range by two isolated populations in north peninsular Florida and more extensive but discontinuous stands along the northern Gulf coast from central panhandle Florida to Mississippi. These southern stands have been surveyed from the standpoint of distribution, autecology, habitats, taxonomy, and protection. Selected populations are described in detail, including listing of associated spe- cies. Most populations occupy valleys of small streams through deep sandhills, where soils are perennially moist or wet from constant seepage of groundwater but are only briefly, if at all, inundated. White cedars have also colonized boggy pine flatwoods near the coast in panhandle Florida, following the suppression of fire in recent decades. Soils may be peaty or sandy, and are circumneutral where charged by spring water. Fires are less frequent or at least less destructive than in the northern range of the species, due to the incised topography, the constantly moist soil and leaf litter, and the intermixture of relatively poorly burning vegetation of other species. The infrequency of fires favors a mixed forest of white cedars, dicotyledonous hardwoods, and some- times palms, rather than the uniform stands typical of white cedar in northern stands. Herba- ceous species are numerous. Reproduction of white cedar typically occurs during gap succession, rather than after crown fires. Stand age is therefore uneven, in contrast to northern stands. White cedars were found to be essentially unharmed by insect or fungal pests, with lightning appearing to be the major cause of death of mature trees. Utilizing lightning ground-flash frequency data derived from engineering studies, a scale of probabilities is derived for estimating lightning-strike intervals for trees protruding above the forest canopy. White cedars in mixed-growth southern stands commonly show suppression in rate of growth that may continue for 75 years or more. As the tree attains a position in the canopy, the growth rate increases. Canopy trees in a Florida stand, measured over a 22-year span, indicate an annual diameter increase of 0.68 cm; this value is greater than recorded elsewhere for the species. Trunk diameters were observed to approach, but not surpass, 100 cm (except for two Alabama patriarchs, which are about 150 cm. dia.). Height was recorded to 27 m. Age probably does not exceed 190 years (except the Alabama giants, which may reach 275 years). White cedars were observed or reported in association with Parnassia grandifolia, Salix floridana, Taxus floridana, and other endemic or rare species of the Gulf coastal plain. Pieris phillyreifolia, an ericaceous vine, was found to grow beneath the bark of white cedar and produce its shoots at heights to 13 m. In peninsular Florida, a large white cedar has been used by successive generations of black bears as a territorial marker, by claw lacerations of the bark. A variety of white cedar, C. thyoides var. henryae (Li) Little, differenti- ated largely on the absence of foliar glands, is recognized from five counties in western Florida and adjacent Alabama. “The third kind of swamps are those spongy tracts, where waters continually ooze through the soil, and finally collect in streams and pass off. These are properly termed galls, sometimes sour, sometimes bitter lands. They are the coldest soils we have...When their foundation is alluvial matter, it is usually very thin, like quagmire... When the base is sand, it is always a lively quicksand, very dangerous to cattle. These galls are usually covered with titi and other androm- edas, loblolly and other laurels, vaccinium and ...coast cypress, Cupressus thyoides [ = Chamae- cyparis thyoides].” —John Lee Williams, A View of West Florida. Philadelphia. 1827. No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 9 ATLANTIC white cedar, Chamaecyparis thyoides (L.) BSP., is an uncom- mon yet important forest tree in northern Florida and near the Gulf coast in Alabama and Mississippi. Though a species long familiar to botanists, ecolo- gists, and wetland managers, nearly all information regarding the tree is derived from observations in its northern range, with little recorded about the requirements and behavior of white cedar in these southern states. What is the exact range of white cedar? How large does it grow, and how long does it live? Why is it so often the dominant tree in its specialized habi- tat? What is there about the seepy woodlands where it is found that is differ- ent from the much more extensive wetlands where it does not occur? What physical or environmental factor limits the size to which it can grow? Is it important that occasional trees are struck by lightning. Although easily de- stroyed by fire, how does it survive the conflagrations that periodically sweep the adjacent uplands? What other plant species are characteristically associ- ated with white cedar? And what taxonomic rank, if any, should be given to the second entity, C. henryae Li, described from the Gulf coast? The present study, in which an attempt is made to answer these questions in southern states, is part of a broad examination by a number of investigators of Atlantic white cedar and its habitat throughout the range of the species, and is the outgrowth of the Atlantic White Cedar Wetlands Symposium spon- sored by the Marine Biological Laboratories, Woods Hole, Massachusetts, in October 1984. Thirty-two of the papers presented at that symposium have been assembled (Laderman, 1987), and summarize these related but inde- pendent studies. A synopsis of the observations documented in the present study has been included in the symposium report (Clewell and Ward, 1987). Most of the white cedar stands mentioned in the text are shown on an outline map of the southern states (Fig. 1). DistRIBUTION— The distribution of Atlantic white cedar, or “juniper” as it is called locally, is distinctive in Florida, Alabama, and Mississippi, and is not closely paralleled by that of any other vascular plant species. Two stations occur in northern peninsular Florida, while the population centers are in the central and western parts of the Florida panhandle, southern Alabama, and southern Mississippi. Except for a cluster of small populations in west-central Georgia, a large hiatus separates the Florida-to-Mississippi stands from those extending from the Carolinas northward. The Gulf coastal distribution of white cedar (Fig. 2) may be divided into regions separated both for convenience and for reasons of possible biogeo- graphic significance. North peninsular Florida: A stand along Juniper Creek and Morman Branch, Ocala National Forest, at the eastern edge of Marion County; and a stand 45 km to the north-northwest, 6 km south of Interlachen, along Deep Creek, which discharges into the Oklawaha River. The drainage of both of these stands is eastward to the St. John’s River and thence to the Atlantic Ocean. 10 FLORIDA SCIENTIST [Vol. 52 Fic. 1. Outline map of the southeastern United States with major streams of the Gulf coastal plain, showing location of white cedar stands mentioned in the text. A. Deep Creek stand, Putnam Co., Fla.; B. Ocala stand, Marion Co., Fla.; C. Tates Hell, Franklin Co., Fla,; D. Johnson Juniper Swamp, Liberty Co., Fla.; E. Yellow River Tidal Swamp, Santa Rosa Co., Fla.; F. Blackwater River, Okaloosa Co., Fla.; G. Sweetwater Creek, Santa Rosa Co., Fla.; H. Escambia River and Perdido River, Escambia Co., Fla.; I. Bluff Creek, Jackson Co., Miss.; J. Westernmost station. Catahoula River, Pearl River Co., Miss.; K. Largest tree. Brewton, Es- cambia Co., Ala. East side of Apalachicola River, Florida Panhandle: Telogia Creek and other eastward-draining tributaries of the Ochlockonee River, Gadsden and Liberty counties; New River of Liberty and Franklin counties, which drains into the Gulf at St. George Sound; and various westward-draining tertiary streams entering the Apalachicola River. The easternmost of these stations is separated by nearly 300 km from the stands of north peninsular Florida. West side of Apalachicola River: Tertiary eastward-draining tributaries of the Apalachicoia River, Calhoun and Gulf counties; streams directly entering the Gulf at St. Vincent Sound and St. Joseph Bay, Gulf County; and an outlying station in the Black Creek drainage, east of Choctawatchee Bay, southern Walton County. This outlying station, though physically somewhat closer to the region next described, appears biologically more similar (in pres- ence of glands on facial leaves) to the stands to the east. Western panhandle Florida and adjacent Alabama: Numerous stations along the Yeliow River and its tributaries, north and central Okaloosa County and southeastern Santa Rosa County; the Blackwater River, Escambia County, Perdido River and secondary streams, Santa Rosa and Escambia counties; headwater extensions of the Escambia River and Perdido River into No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR al Fic. 2. Outline map of the southeastern United States, showing location of known white cedar stands. Each stand has been documented by one or more herbarium specimens collected or examined by the authors. Open circles represent typical Chamaecyparis thyoides. Solid circles represent Chamaecyparis thyoides var. henryae. Escambia County, Alabama; and tributaries entering the Perdido River, Per- dido Bay, and Bon Secour Bay, Baldwin County, Alabama. Southern Mississippi: Tributaries of the Escatawpa River and Pascagoula River, Jackson County; and branches of the Catahoula River, Pearl River County. West-central Georgia: A small stand on a tributary of Upatoi Creek, be- tween Talbot and Marion Counties; and a stand on Whitewater Creek, Tay- lor County. These two stations, although separated by less than 30 km, are in different drainages, Upatoi Creek in the drainage of the Chattahoochee River, and Whitewater Creek in that of the Flint River. These rivers converge at the Georgia-Florida state line to form the Apalachicola River. The Florida peninsular and panhandle populations are separated geologi- cally by the Suwannee Straits (Puri and Vernon, 1964), which affected depo- sition of both Mesozoic and Cenozoic sediments. Many plant species are re- stricted to one side or the other of this saddle-like geological feature, which evidently has exercised considerable influence over plant distribution within the state (Clewell, 1985). Older distribution maps have shown white cedar occurring throughout northern Florida and extending half-way down eastern peninsular Florida (Korstian, 1931; Munns, 1938; Brush, 1947; James, 1961). These range maps 12 FLORIDA SCIENTIST [Vol. 52 have been found (Ward, 1963) to be based upon an unpublished map pre- pared by G. B. Sudworth, which in turn was documented by his file of local- ity records compiled from various sources. Sudworth’s records for northeast- ern Florida were based upon R. M. Harper’s Geography and Vegetation of Northern Florida (1914: 324, 334, 342). But Harper is not to be discredited; when he failed to observe a major forest tree in a particular area, and yet could not claim that no individuals were present, he customarily listed it under the conservative heading, “rare or absent.” Sudworth chose to place the emphasis on the first possibility, thereby initiating a prolonged series of over-estimations of the abundance of white cedar in northeastern Florida. Only in recent years have maps been published (Little, 1971; Taras, 1971) correcting this half-century misunderstanding. GroLocy— White cedars grow primarily on siliceous sands of Plio-Pleisto- cene deposition in coastal lowlands, including extensions of these lowlands northward along streams that dissect adjacent highlands (Puri and Vernon, 1964; White, 1970). The large majority of white cedar stands occur from near mean sea level (msl) to about 23 m above msl. Both of the two stands in peninsular Florida occur at the edge of the physiographic province of the coastal lowlands called the St. John’s River Offset. Elevations are 15 m higher within 0.8 km of the Marion County stand and within 2.4 km of the Putnam County stand; in peninsular Florida these are substantial changes in relief. In the panhandle, white cedar stands are commonly at the base of areas of higher elevation, although in the boggy flatwoods in the southern panhandle large differences in terrain are lacking. The most characteristic geomorphological feature associated with white cedar stands is the presence of perennial or near-perennial seepages. These seeps often represent base flow from sand ridges or hills flanking stream drainages. Where hills are not closely adjacent, the seepage areas occupy broad basins near sea level and receive base flow from more distant physio- graphic provinces to the north. PENINSULAR FLORIDA STANDS— Two stands of Atlantic white cedar occur in north-central peninsular Florida. They are discussed here in order of date of discovery and descending size. Deep Creek Stand: Along Deep Creek, approximately 6 km south of In- terlachen, Putnam County, is an extensive stand of “juniper” (Fig. 3). This stand may not have been known to botanists prior to 1942, the date of the earliest herbarium collections encountered (W. B. DeVall, FLAS); it was not mentioned in print until 1946 (West and Arnold, 1946). In 1963 it was briefly examined by students under the direction of C. D. Monk (Collins et al., 1964). The stream south of Interlachen along which the white cedar grows has only recently been known as Deep Creek. Herbarium labels dating from the 1940’s refer to it as Cabbage Creek. Ward (1963) referred to the white cedars as occurring for several miles along “a small clear stream known as Cabbage No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 13 To Interlachen Cousintown Road ee / = eat 2 oo fa. Power Line Teste) | Sra ; ba eee WA car At Se Vi / To Okiawaha River y White Cedar Stand Fic. 3. Map of white cedar stands along Deep Creek, Putnam County, Florida. Flow of the stream is from west to east. The densest portion of the stand, and the only area in which a continuous canopy is present, is east of State Road 315, both north and south of the power line right-of-way. Creek almost to its mouth on the Oklawaha River.’ Collins and co-workers (1964) repeated the designation of the stream as Cabbage Creek, defining it as arising in a headwater swamp (known as Miller Hammock) and flowing southeast for about four miles, then joining Deep Creek, a tributary of the Oklawaha River. Residents of the area consistently refer to the stream as Cabbage Creek, a name surely derived from an extensive cabbage palm (Sa- bal palmetto) savannah along both sides of the stream below (east of) the area occupied by white cedar; this savannah at present is shallowly covered by waters backed up by the Rodman Reservoir, a now-flooded portion of the Oklawaha River. By cartographic license, however, the name Cabbage Creek has been transferred to a stream that enters the Oklawaha River by a different route (U.S.G.S., Rodman and Keuka quadrangles, 1949; Florida Department of Transportation, Putnam Co., 1968). This second stream, apparently previ- ously unnamed and without white cedars along its course, flows west and then south from the headwater swamp that also supplies the east-flowing stream along which the white cedars grow. The name Deep Creek, though originally referring only to what is now an arm of the Rodman Reservoir, was then extended upstream so that it applied to the entire east-flowing drainage. Though standard maps of the area are clearly at variance with local us- age, the ambiguity that now accompanies the name Cabbage Creek suggests that greater clarity is achieved by acceptance of the cartographic designation 14 FLORIDA SCIENTIST [Vol. 52 of the cedar-bordered stream as Deep Creek. Both the early observations by Ward (1963) and the study by Collins and co-workers (1964) were entirely within the drainage of this east-flowing stream. Deep Creek, as thus defined, is a clear-flowing, sand-bottomed stream approximately 7 km in length. It is heavily supplied by seepages from the adjacent sandy slopes, as well as from its headwater swamp and, in its lower portion, from Gum Creek, a major tributary entering from the north. There is essentially no surface supply even in heavy rain. At mid-point, it is approxi- mately 5 m wide and 0.3 m deep, although these dimensions vary greatly within short distances. Its elevation is 17 m (55 feet) above msl at headwaters, and 8 m (25 feet) at its mouth on Rodman Reservoir. This slope and constant supply produces a brisk and stable flow. The banks of Deep Creek are well-vegetated and scarcely at all eroded by the flow. The surface beyond the bank, on both sides, is a level riverbottom, densely forested with a large variety of mesic to hydric woodland trees and other species (Table 1). The surface is covered with black, organic peat that overlies sand. The soil is a seepage surface and is constantly wet, though irregularities associated with hummocks at the base of trees and fallen logs provide moisture diversity. TABLE 1. Flora of the Deep Creek stand, Putnam County, Florida. (Potential) Overstory Trees Acer rubrum Chamaecyparis thyoides Fraxinus caroliniana Fraxinus profunda Gordonia lasianthus Agarista populifolia (= Leucothoe populifolia) Alnus serrulata Baccharis glomeruliflora Bumelia aff. lycioides Callicarpa americana Carpinus caroliniana Cephalanthus occidentalis Amphicarpa bracteata Berchemia scandens Decumaria barbara Gelsemium sempervirens Liquidambar styraciflua Liriodendron tulipifera Magnolia virginiana Nyssa biflora Persea palustris Understory Trees and Shrubs Cornus foemina Euonymus americanus Ilex coriacea Itea virginica Leucothoe axillaris Lyonia lucida Myrica cerifera Ostrya virginiana Woody Vines Matelea floridana Parthenocissus quinquefolia Rhus radicans Smilax glauca Pinus elliottii Quercus laurifolia Quercus nigra Sabal palmetto Ulmus americana var. floridana Rhapidophyllum hystrix Rhododendron serrulatum Rhus vernix Rubus argutus Salix floridana Serenoa repens Vaccinium fuscatum Viburnum obovatum Smilax hispida Smilax laurifolia Smilax walteri Vitis rotundifolia No. 1, 1989] TABLE 1. Continued. Dryopteris ludoviciana Isoetes flaccida Lorinseria areolata Carex chapmanii Carex leptalea Chasmanthium ornithorhynchum Apteria aphylla Arisaema acuminatum Chamaelirium luteum Dioscorea floridana Habenaria odontopetala Aristolochia serpentaria Aster carolinianus Bidens mitis Boehmeria cylindrica Cacalia diversifolia Cadamine bulbosa Cirsium aff. muticum Ferns and Allies Osmunda cinnamomea Osmunda regalis Phlebodium aureum Graminaceous Species Cladium jamaicense Leersia virginica Oplismenus setaceus Forbs (Monocots) Hymenocallis rotata Malaxis spicata Orontium aquaticum Pontederia cordata Forbs (Dicots) Eupatorium fistulosum Hydrocotyle umbellata Lobelia amoena var. glandulifera Mikania cordifolia Mitchella repens Nasturtium microphyllum WARD AND CLEWELL—ATLANTIC WHITE CEDAR 15 Polypodium polypodioides Selaginella apoda Thelypteris palustris Rhynchospora inundata Rhynchospora miliacea Pothieva racemosa Sagittaria lancifolia Tillandsia bartramii Tillandsia usneoides Pluchea longifolia Polygonum punctatum Ruellia caroliniensis Sabatia calycina Samolus parviflorus Solidago aff. sempervirens Viola floridana Clematis ?crispa Parnassia grandifolia Atlantic white cedar grows only along part of Deep Creek. To the west, in the headwater swamp, the trees are increasingly scattered, and a final range determination has not been established. To the east, downstream, the density of white cedar increases, in places comprising the entire canopy. The stand ends rather abruptly between the confluence with Gum Creek and the de- bouchure at Rodman Reservoir. The area in which white cedar occurs is approximately 5.5 km (3.4 miles) in length; the width is greatly variable, governed by the width of the creek bottom—a total of 200 ha (495 acres) may be included. Within this area the number of white cedars above 10 cm diam- eter-breast-height (dbh) may lie between 2,000 and 5,000 individuals. The Deep Creek stand has been divided in years past by two north-south roads—to the west the Cousintown Road, an unpaved county road, and to the east State Road 315, a paved highway. In 1981, the stand was further dissected by a Seminole Electric transmission line with an east-to-west 50 m- wide easement clearing. Within the past five years, a private land owner constructed a farm pond on a small tributary entering Deep Creek from the north, clearing the adjacent land and removing a dense growth of old-growth white cedars. Other land owners have occasionally cut portions of the stand 16 FLORIDA SCIENTIST [Vol. 52 for fenceposts or sawtimber, or have developed improved pastures intruding on the wet bottomlands. These introgressions, however, have not yet changed the character of the stand to such an extent that its natural features and great beauty are obscured. Collins and co-workers (1964) sampled three sites in the Deep Creek stand. The first site was dominated by white cedar, the second site included white cedar among the more important species, and the third site contained little white cedar. Importance values were determined for all trees that were at least 10 cm dbh. In the first site sampled by Collins and co-workers (1964), white cedar had an importance value of 167 (of a possible 300). Values for the next three trees were Magnolia virginiana - 41, Persea palustris - 33, and Pinus taeda (P. elliottii?) - 26. Six less important tree species were also present. In the second site, white cedar had an importance value of 86. Other trees with high values were Persea palustris - 45, Acer rubrum - 36, and Fraxinus caroliniana (F. profunda?) - 32. Four other tree species were also present. The third site was dominated by Sabal palmetto - 79, Quercus laurifolia - 42, Nyssa biflora - 39, and Magnolia virginiana - 38. Among the eight less important tree species was white cedar with a value of 12. White cedar saplings were common at the first site and uncommon at the other stations. As reported by Collins and co-workers (1964), the soil in the Deep Creek stand consisted of coarse sand which graded into gravel. Overlying the sand was an organic layer 0-46 cm deep. The pH was 6.6-7.5. Calcium was 791- 2144 ppm, magnesium 115-404 ppm, potassium 12-38 ppm, and phosphorus 2-7 ppm. Fire, which is recognized as a powerful force of both destruction and regeneration of white cedar stands in the northern Atlantic states (Korstian, 1931; S. Little, 1979), seems not to be importantly influential in the Deep Creek station (see “Relationship with Fire’). Soon after its discovery, the Deep Creek white cedar stand was recognized as a location for other cool-temperate plant species (Table 1). Some of these species are unknown from peninsular Florida except in this stand and some- times in the Ocala white cedar stand of Marion County. Among these dis- juncts are fairy-wand (Chamaelirium luteum), not otherwise known south- east of Leon County in the panhandle; a swamp thistle (Cirsium aff. muticum) differing in size and flower color from those few plants known in the panhandle; and grass-of-Parnassus (Parnassia grandifolia), found also abundantly with white cedar in the Ocala stand but otherwise known on the Gulf coast except for a few stations just east of the Apalachicola River. Other species, as tulip tree (Liriodendron tulipifera), are near the southern limit of their range here, while alder (Alnus serrulata) terminates its southern range at this station. Deep Creek is one of the few stations for the Florida willow (Salix floridana), a small tree endemic to peninsular Florida and adjacent Georgia; Florida willow also occurs in the Ocala stand. No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR iLaf San, = Lake George i To Y Salt | Springs | I 2 7 : 5 ae 3 \ ig N \ | x | Io Fea | 5 | 1 KM FAS ST TF | | | | Sal | --{ White Cedar Stand | 12 | es F, oe eS S7est Service | Oe oe es | Fic. 4. Map of white cedar stand along Juniper Creek and Morman Branch, Ocala National Forest, Marion County, Florida. Flow of the stream is from west to east. The densest portion of the stand is along Morman Branch immediately south of the Forest Service access road. Most of the associated plants of restricted distribution, such as Pieris phillyreifolia and Parnassia grandi- folia, occur only in this area. Ocala Stand: A second peninsular Florida stand of white cedar is present along a portion of Juniper Creek and its tributary, Morman Branch, in the eastern part of the Ocala National Forest, Marion County (Fig. 4). Juniper Creek arises as a major spring approximately 40 km (25 miles) east of Ocala. The exceptionally clear water has a pH of 8.5 at the head-spring (Rosenau et al., 1977). It flows generally eastward for about 16 km and enters the south- western corner of Lake George, a widened portion of the St. John’s River. Along the way it is supplemented by secondary springs, some of appreciable size. About 11 km below the head-spring it is bridged by State Road 19. The stream is heavily trafficked by canoeists between the head-spring and take- out landing at the bridge, but is very little visited downstream of the bridge. To the south of Juniper Creek and largely to the east of State Road 19 is a 250 ha swamp fed by seepages and small springs from the surrounding stabi- lized or “fossil” dunes that constitute much of the Ocala National Forest. These seeps and springs coalesce to form a small stream, Morman Branch, about 3 km in length, which enters Juniper Creek from the south about 1 km below the State Road 19 bridge. The branch is crossed about midpoint by a rudimentary Forest Service road (No. 71), the only practical means of access. 18 FLORIDA SCIENTIST [Vol. 52 Juniper Creek at its head-spring lies at an elevation of 9 m (30 feet) above msl, while its mouth on Lake George is near 1.5 m (5 feet). Nearly all white cedar along both Juniper Creek and Morman Branch are within 2 to 4 m of sea level. Atlantic white cedars grow along both sides of Juniper Creek from just below the State Road 19 bridge, to a point shortly beyond the entrance of Morman Branch, a distance of about 1 km. These white cedars are conspicu- ously visible as the dominant tree along the streamcourse—and are undoubt- edly the source of the name, “Juniper Creek.” The trees also occur along the middle and lower third of Morman Branch, both above and below the Forest Service road and extending to the joining with Juniper Creek. (Maps suitable for an overall view of these stands are: U.S.G.S., Astor and Juniper Springs quadrangles, 1972; Florida Department of Transportation, Marion Co. (sheet 1), 1977; and U.S.D.A. Forest Service, Ocala National Forest, 1965.) The Ocala stand of white cedar was unknown to botanists until 1962, when it was called to the attention of one of the present authors who then described it briefly (Ward, 1963). At that time, the extent of the stand was not fully appreciated; it was described as not extending beyond 4 ha (10 acres). In later investigation by canoe and overland through the headwater swamp of Morman Branch, the stand has been observed to be much larger, perhaps 50 ha (124 acres). An exact determination of the size of the stand has not yet been undertaken because of logistical considerations, among them being nearly impenetrable vegetation, frequent water moccasins, signs of black bears (see “Bear Damage’), and inconceivable numbers of minute “seed” ticks. (One investigator removed 450 of the latter from his body fol- lowing an August survey. ) The number of white cedars in the Ocala stand is fewer than along Deep Creek in Putnam County. Simultaneous on-site estimates by three different observers gave mean values of between 1,000 and 1,500 trees of at least 10 cm dbh in the Morman Branch drainage and about 550 trees along Juniper Creek. Of those along Juniper Creek, nearly half grow on a small complex of islands on the south side just below the State Road 19 bridge, with the others distributed almost evenly along both banks of the stream. In all parts of the white cedar stand along Juniper Creek and Morman Branch, soil moisture within the root zone is high and remains constant in all seasons. This constancy of moisture is in part attributable to the extraordi- nary stability of the spring-fed flows of both Juniper Creek and Morman Branch. This stability is apparent not only by observation at different seasons and by lack of erosion along the stream banks, but by the luxuriant growth on sand bars 3 cm below the surface of Juniper Creek, of a pondweed (Potamo- geton pectinatus), a species unable to withstand the desiccation accompany- ing any lowering of the water flow. The pH of small streams within the stand, as measured by Dunn (1985), varied only between 6.9 to 7.1. Such circumneutral pH values appear to reflect the neutral to mildly alkaline waters of the perpetual springs and seeps No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 19 that supply the stand, relatively uninfluenced by the probably more acidic peat of the soil surfaces. Within most of the Morman Branch portion of the stand, the exposed soil is wet, nearly black peat, with fallen logs and stumps of long-dead trees providing a degree of microtopographic relief. The surface is consistently above the level of the stream, which is edged by a 10-30 cm bank. In most areas the white cedars are conspicuous, often being the largest trees of the association, but usually are not densely spaced or exclusive of other tree spe- cies. South of the Forest Service crossroad are a number of open glades— grassy avenues among the denser vegetation—each with a shallow central channel of flowing surface water from adjacent springs, and edged by species of Parnassia, Pinguicula, Aletris, and other herbs. The curious primitive quillwort (Isoetes flaccida) is frequent, its leaves spreading on the surface of mucky openings. In contrast to Morman Branch, the banks above Juniper Creek are higher (to 1.5 m), and the vegetation is less dense. An area of exceptional interest lies just downstream of the State Road 19 bridge on the south side of the creek. There, a labyrinth of steep-sided, flat-topped islets, varying in length from 2 m to about 30 m, are separated by open channels supplied by Juniper Creek. In this complex of islets white cedar is the dominant tree species, favored by the multiplicity of stream edges suitable for seed germination and protected by the channels from low-intensity fires that might otherwise creep into the stand from adjoining pinelands. Two white cedars in the Ocala stand are very close in size to the largest individuals of the species recorded in Florida. An ancient tree, its trunk much damaged by bear-scratching, in the upper reaches of Morman Branch, has a trunk diameter of 91 cm (36 inches), while a tree on Juniper Creek across from the mouth of Morman Branch reaches 88 cm (35 inches). In 1962 and again in 1984 all trees above 30 cm dbh were measured in a small part of the stand (see “Age, Size, and Rate of Growth’). Much of the fascination with the Ocala stand of white cedar lies in the associated species of vascular plants (Table 2). Among the most curious is the climbing pieris (Pieris phillyreifolia), a common plant of the stand. This eri- caceous evergreen frequently assumes the growth habit of a low shrub, with stems arising from atop stumps and fallen logs. But under suitable conditions its rhizome penetrates the outer bark of certain associated trees and burrows its way upward, becoming visible only where its lateral branches break through the host tree’s bark to appear as green sprays of foliage and axillary racemes of white flowers. Climbing pieris nearly always selects white cedar or pond cypress (Tax- odium ascendens) as its host (Judd, 1982). It has been reported as emerging from white cedar trunks to a height of 7 m (Ward, 1963) and pond cypress trunks to 10 m (Judd, 1982). During the present study Pieris branches emer- gent from white cedar trunks were observed frequently at heights of 6-8 m throughout the Morman Branch Station. Those that ascended even higher 20 FLORIDA SCIENTIST TABLE 2. Flora of the Ocala stand, Marion County, Florida. Acer rubrum Chamaecyparis thyoides Fraxinus profunda Gordonia lasianthus Magnolia virginiana Agarista populifolia (= Leucothoe populifolia) Baccharis glomeruliflora Bumelia aff. lycioides Carpinus caroliniana Cephalanthus occidentalis Cornus foemina Berchemia scandens Decumaria barbara Gelsemium sempervirens Lonicera sempervirens Dryopteris ludoviciana Isoetes flaccida Lorinseria areolata Osmunda cinnamomea Carex chapmanii Carex leptalea Aletris Plutea Apteria aphylla Arisaema acuminatum Dioscorea floridana Bidens mitis Cirsium aff. muticum Lobelia amoena var. glandulifera (Potential) Overstory Trees Morus rubra Nyssa biflora Persea palustris Pinus elliottii Quercus laurifolia Understory Trees and Shrubs Euonymus americanus Ilex coriacea Illicium parviflorum Itea virginica Leucothoe axillaris Lindera aff. benzoin Lyonia lucida Woody Vines Parthenocissus quinquefolia Pieris phillyreifolia Rhus radicans Ferns and Allies Osmunda regalis Polypodium polypodioides Pteridium aquilinum Graminaceous Species Cladium jamaicense Fimbristylis castanea Forbs (Monocots) Epidendrum conopseum Habenaria odontopetala Hypoxis leptocarpa Forbs (Dicots) Ludwigia Ppalustris Mikania cordifolia Parnassia grandifolia Pinguicula caerulea [Vol. 52 Quercus nigra Quercus virginiana Sabal palmetto Ulmus americana var. floridana Myrica cerifera Rhapidophyllum hystrix Rhododendron serrulatum Rubus argutus Salix floridana Vaccinium fuscatum Schrankia uncinata Smilax laurifolia Vitis rotundifolia Selaginella apoda Thelypteris palustris Vittaria lineata Rhynchospora inundata Rhynchospora miliacea Malaxis spicata Ponthieva racemosa Tillandsia bartramii Saururus cernuus Solidago aff. sempervirens Viola floridana No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 74, | were measured with a forester’s clinometer; three were recorded above 10 m, with the most lofty emerging at 13 m. As Pieris continues to ascend, new branches protrude through the bark of the host tree, and older branches die and break off. Thus at any given time, only a short vertical array of Pieris branches extends from the cedar trunk. On those trees where recent branches are at a considerable height, ablation of the cedar bark on the lower trunk, as is normal with time, exposes the flaky red bark of the previously hidden Pieris stems. This exposure seems not to affect the continued growth of the climbing vine. Other plants associated with white cedar in the Ocala stand, as with the Deep Creek stand, reflect either extremes of range or disjunctions with popu- lations to the north. At least one species (Lindera aff. benzoin) appears not to occur elsewhere southeast of the central panhandle. Others (Cirsium aff. muticum, Lobelia amoena, Parnassia grandifolia) are similarly disjunct but share peninsular stations with the Deep Creek stand. Pieris, though absent at the Deep Creek station, occurs apart from white cedar along another spring run in adjacent Lake County, but is otherwise unknown south of northern and western Florida. An array of northern species found in the Deep Creek stand do not reach this more southern station (Alnus serrulata, Cacalia diver- sifolia, Chamaelirium luteum, Eupatorium fistulosum, Liriodendron tulipi- fera), a possible indication of progressive species loss during the Pleiocene- Pleistocene southward migrations of the white cedar forest and its northern associates. Along Juniper Creek the number of species directly associated with white cedar is appreciably less than in the Morman Branch drainage (28 vs. 63 in an August 1984 survey). No species found directly associated with cedar along Juniper Creek was not also seen at Morman Branch, a suggestion that the flora of the main creek may be a depauperate derivative from a refugial area along the small tributary. In and along the edges of the open waters of Juni- per Creek itself, however, additional species do occur. Several of these proxi- mal species (Acrostichum danaeifolium, Cyperus articulatus, Fimbristylis castanea, Kosteletzkya virginica, Solidago aff. sempervirens) are elsewhere characteristic of brackish habitats, and suggest a significant mineral content in the springfed water of Juniper Creek. The rare endemic vetch, Vicia oca- lensis, grows abundantly among the sawgrass and other emergent vegetation of the stream, very close to but not directly associated with the white cedar. A number of species that occur only in the channel of Morman Branch have arbitrarily been excluded as not growing within the white cedar associa- tion. These species include: Ceratophyllum demersum, Cicuta mexicana, Lo- belia cardinalis var. meridionalis, Pontederia cordata, Vallisneria neotropica- lis. PANHANDLE FLoripA STANDS— Many stands of Atlantic white cedar occur in panhandle Florida; indeed, along many streams in the central panhandle and again in the western panhandle, the stands in places, though differing 22 FLORIDA SCIENTIST [Vol. 52 greatly in density, are so frequent as to be considered contiguous. The follow- ing are selected as representative. Tates Hell: Much of Franklin County and adjacent southern Liberty County is a vast complex of boggy pine flatwoods, grass-sedge bogs, and shrub bogs. These wetlands are slowly drained by shallow sloughs dominated by pond cypress (Taxodium ascendens), sweetbay (Magnolia virginiana), black gum (Nyssa biflora), and sometimes white cedar. This inhospitable mo- saic is known locally as Tates Hell. White cedar is abundant from the New River drainage westward to the Kennedy Creek drainage, but is only locally common nearer the Ochlockonee and Apalachicola rivers and in central and northern Liberty County (Clewell, 1971). The vegetation of Tates Hell (Table 3) and the changes associated with reforestation activities have been surveyed by Conde and co-workers (1977). Two sites were described, one undisturbed and the other prepared and planted to seedling slash pines. The area has also been examined by one of the present authors (A. F. C.). TABLE 3. Flora of the Tates Hell stand, Franklin Countys. Chamaecyparis thyoides Magnolia virginiana Aronia arbutifolia Clethra alnifolia Cliftonia monophylla Cyrilla racemiflora Gaylussacia mosieri Hypericum brachyphyllum Gelsemium Prankinii Pieris phillyreifolia Lycopodium alopecuroides Andropogon glomeratus Andropogon tenarius Andropogon virginicus Aristida stricta Aristida virgata Carex joorii Ctenium aromaticum Dichromena latifolia Fuirena squarrosa (Potential) Overstory Trees Nyssa biflora Persea palustris Understory Trees and Shrubs Hypericum fasciculatum Ilex coriacea Ilex glabra Ilex myrtifolia Lyonia fruticosa Lyonia lucida Woody Vines Smilax laurifolia Ferns and Allies Lycopodium carolinianum Graminaceous Species Juncus marginatus Juncus polycephalus Panicum acuminatum Panicum dichotomiflorum Panicum rigidulum Rhynchospora baldwinii Rhynchospora cephalantha Rhynchospora chapmanii Pinus elliottii Taxodium ascendens Myrica cerifera Myrica inodora Osmanthus americanus Rhododendron serrulatum Styrax americana Vaccinium corymbosum Vitis sp. Woodwardia virginica Rhynchospora corniculata Rhynchospora filifolia Rhynchospora gracilenta Rhynchospora plumosa Rhynchospora rariflora Scirpus cyperinus Scleria baldwinii Scleria reticularis No. 1, 1989] TABLE 3. Continued. Aletris lutea Calopogon pallidus Eriocaulon decangulare Hypoxis leptocarpa Lachnanthes caroliniana Lachnocaulon anceps Agalinis linifolia Agalinis purpurea Aster chapmanii Balduina uniflora Bartonia paniculata Carphephorus pseudoliatris Chondrophora nudata Coreopsis aff. leavenworthii Cuscuta compacta Drosera capillaris Drosera tracyi Eupatorium recurvans Forbs (Monocots) Lilium catesbaei Lophiola americana Platanthera ciliaris Platanthera nivea Pleeia tenuifolia Sagittaria graminea Forbs (Dicots) Eupatorium semiserratum Euthamia minor Helianthus floridanus Justicia crassifolia Liatris spicata Lobelia floridana Ludwigia linearis Ludwigia pilosa Oxypolis filiformis Pluchea camphorata Pluchea foetida Pluchea odorata WARD AND CLEWELL—ATLANTIC WHITE CEDAR Syngonanthus flavidulus Xyris ambigua Xyris baldwiniana Xyris elliottii Xyris stricta Zygadenus glaberrimus Pluchea rosea Polygala cruciata Polygala cymosa Polygala lutea Proserpinaca pectinata Rhexia alifanus Rhexia virginica Sabatia quadrangula Sarracenia flava Sarracenia psittacina Utricularia cornuta Utricularia juncea 23 2As modified from Conde et al. (1977). Elsewhere, boggy pine flatwoods typically experience surface fires every several years (Clewell, 1981). The Tates Hell survey areas described by Conde and co-workers (1977), however, evidently have not been burned for many years. The overstory includes fire-intolerant white cedars and pond cypress, as well as 20 m.-tall slash pines (Pinus elliottii). The understory consists of a dense thicket 3 to 6 m. tall, primarily of Cliftonia monophylla, Cyrilla ra- cemiflora, Lyonia lucida, Ilex coriacea, and Ilex myrtifolia. Neither the over- story white cedars nor the undergrowth could have developed except with fire suppression. The larger pines, however, bear turpentine “faces.” It was customary for naval-stores operators to burn pine flatwoods frequently to suppress undergrowth and permit access. The presence of these turpentine faces serves as evidence that in all probability the survey areas have been burned repeatedly in the past, and that the white cedars have colonized the sites following fire suppression. Ground cover in the undisturbed survey area is limited to a few grassy openings. Site preparation in the planted survey area has released the many suppressed plants of the ground cover from competition. Where seed trees have survived in the cypress sloughs that interrupt the pine flatwoods, nearby planted areas are being readily colonized by white cedar seedlings. Wiregrass (Aristida stricta) dominates the species-rich herbaceous ground cover which contains many insectivorous plants (species of Drosera, Sarracenia, and ter- restrial species of Utricularia). Johnson Juniper Swamp: Kurz (1927) noted white cedar in Johnson Juni- per Swamp in Liberty County, 13 km south of Bristol. He listed the following 24 FLORIDA SCIENTIST [Vol. 52 species associated with white cedar: Magnolia virginiana, Cliftonia monophylla, Pinus taeda, Taxodium ascendens (= T. “imbricatum”), Nyssa biflora, Persea palustris (=P. “pubescens”), Taxus floridana, and on the higher accumulated peat, Magnolia grandiflora (=M. “foetida”). The stand was remarkable for the presence of Florida yew (Taxus floridana), which had previously been known only from ravines between Bristol and the Georgia- Florida state line on the east side of the Apalachicola River. White cedar and Florida yew are not known elsewhere to occur in association. Yellow River Tidal Swamp: Duever (1984) has made available the file of a white cedar swamp at the mouth of the Yellow River, Santa Rosa County. Water levels within the swamp were held relatively constant by the Gulf of Mexico, which acted much like the dam of a reservoir in retaining the fresh waters of the river. The riverine water held back by this “dam” was under gentle tidal influence but was not saline. The swamp was dominated by white cedar, Magnolia virginiana, Ilex cassine, Ilex myrtifolia, Cyrilla ra- cemiflora, and Myrica spp. Blackwater River: A 29-km reach of the Blackwater River, Okaloosa County, was surveyed by canoe from a starting point just south of the Ala- bama state line, with an inventory taken of the woody species (Table 4). The forest on the low levee of the river was dominated by white cedar for its entire length. Ilex opaca was dominant in the understory. The habitat was generally mesic, but the water table appeared to be continuously near the soil surface. There was no evidence of prolonged flooding, deep scouring, or recent allu- vial deposition. The soil consisted of very fine sand and silt with a clayey subsoil that may impede percolation. TABLE 4. Partial flora of selected locations. Blackwater River stand, Okaloosa County. (Potential) Overstory Trees Acer rubrum Magnolia grandiflora Quercus laurifolia Chamaecyparis thyoides var. Nyssa biflora Quercus nigra henryae Persea palustris Quercus virginiana Liquidambar styraciflua Pinus taeda Taxodium distichum Understory Trees and Shrubs Cyrilla racemiflora Kalmia latifolia Osmanthus americanus Cliftonia monophylla Leucothoe sp. Symplocos tinctoria Ilex coriacea Lyonia lucida Vaccinium elliottii Ilex vomitoria No. 1, 1989] TABLE 4. Continued. WARD AND CLEWELL— ATLANTIC WHITE CEDAR ae Sweetwater Creek stand, Santa Rosa County Acer rubrum Chamaecyparis thyoides var. henryae Liriodendron tulipifera Callicarpa americana Cliftonia monophylla Cornus foemina Cyrilla racemiflora Euonymus americanus Smilax glauca Lorinseria areolata Mitchella repens (Potential) Overstory Trees Magnolia grandiflora Magnolia virginiana Nyssa biflora Understory Trees and Shrubs Ilex coriacea Ilex glabra Leucothoe axillaris Myrica heterophylla Woody Vines Vitis rotundifolia Ferns and Allies Osmunda regalis Herbs Pinus taeda Quercus laurifolia Quercus nigra Osmanthus americanus Persea palustris Styrax sp. Symplocos tinctoria Woodwardia virginica Bluff Creek stand, Jackson County, Mississippi.# Acer rubrum Carya sp. Chamaecyparis thyoides var. henryae Ilex opaca Alnus serrulata Calycanthus floridus Castanea pumila Clethra alnifolia Cliftonia monophylla Cyrilla racemiflora Hamamelis virginiana Arundinaria gigantea (Potential) Overstory Trees Liquidambar styraciflua Liriodendron tulipifera Magnolia grandiflora Magnolia virginiana Nyssa biflora Understory Trees and Shrubs Ilex coriacea Ilex glabra Ilex vomitoria Itea virginica Kalmia latifolia Lyonia lucida Myrica cerifera Graminaceous Species 4As modified from Eleuterius and Jones (1972). Quercus laurifolia Quercus nigra Pinus elliottii Sabal palmetto Taxodium distichum Osmanthus americanus Persea borbonia Rhododendron serrulatum Styrax americana Vaccinium arboreum Vaccinium elliottii 26 FLORIDA SCIENTIST [Vol. 52 Sweetwater Creek: A white cedar swamp was inspected along Sweetwa- ter Creek at State Road 4, 1.5 km east of Munson, Santa Rosa County (Table 4). Sweetwater Creek is a tributary of the Blackwater River. The adjacent uplands, forested with longleaf pine (Pinus palustris), slope downward to a broad floodplain. The soil of this floodplain is clayey, and in places is covered with sand apparently eroded from the uplands. Surficial soil of the floodplain is darkened with organic matter and is moist with ground water seeping from the uplands. At the base of the slope the seepage area is covered with a swamp of titi (Cyrilla racemiflora). Although there was no evidence of recent fire, an old stump was charred, and several trees of Magnolia grandiflora had multiple trunks, often a consequence of coppicing after fire. White cedar is among the dominant trees, particularly near the creek, with the largest about 40 cm dbh. Other dominants are Acer rubrum, Quer- cus laurifolia, and Quercus nigra. None of the oaks are of large diameter, although large trees of Pinus taeda are common. A few large trees of Nyssa biflora are present. In the understory Ilex opaca is common, while Leucothoe axillaris, Mitchella repens, and Smilax glauca are common to abundant in the ground cover. Escambia County sites: White cedar stands were observed near State Road 184 in bayheads of streams that drain eastward into the Escambia River. The bayheads are dominated by white cedar, Pinus elliottii, Magnolia virginiana, and Cliftonia monophylla. Flats near the streams bear stands of longleaf pine (Pinus palustris) with a discontinuous growth of white cedar and Cliftonia. These flats lack saw palmetto (Serenoa repens), a characteris- tic species of most longleaf pinelands. Burkhalter (1984) has reported white cedars as common along the Per- dido River west of Barrineau Park, with the infrequent ericaceous shrub, Leucothoe racemosa, growing abundantly in association, particularly on the Alabama side of the river. Mississipp1 STANDS—FEleuterius and Jones (1972) carefully examined a stand of white cedar along Bluff Creek, a tributary of the Pascagoula River, near Vancleave, Jackson County, southeastern Mississippi. The Bluff Creek stand has also been surveyed (by D. B. W.) during the present study. Other than for two very limited stations along branches of the Catahoula River, approximately 80 km to the west, this Bluff Creek population is the western- most station for the species. Eleuterius and Jones found white cedar to extend along Bluff Creek for 11 km (7 miles). Most of the stand was within 3 m (10 feet) in elevation above the stream, although some trees extended to twice this distance where they intergraded with pine and hardwood forest. The stand had a maximum width of about 0.8 km. Along part of the stand steep bluffs bordered the stream; in other areas the stream was flanked by a gently sloping floodplain. White cedars grew on the bluffs, on the levees along the stream, in bogs behind the levees, and on gentle slopes that terminated in sand bars along the No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR Aah stream. The soil was of sandy texture, gray in color, non-stratified, and with a pH of 4.8 to 5.0. Areas along the creek upon which water stood for long periods did not have white cedar, but instead supported cypress (Taxodium distichum) and black gum (Nyssa biflora). Some white cedars in the stand were affected by a witches’-broom (Gym- nosporangium sp.) that produced oblong or spherical swellings on trunks and branches. Although the authors were aware that elsewhere white cedar is fire-maintained (Korstian, 1931), they did not find evidence in this Missis- sippi stand that past fires had occurred. Eleuterius an Jones selected 50 stations within the stand for sampling with the quarter method of Cottam and Curtis (1956), to produce a total of 200 quarters. White cedar was dominant in all areas sampled, although other conifers and both deciduous and evergreen hardwoods were conspicuously present. Slash pine (Pinus elliottii), black gum (Nyssa biflora), bald cypress (Taxodium distichum), American holly (Ilex opaca) and red maple (Acer ru- brum) were the major associated species (Table 4). Though of less relative density and less relative frequency than the mature trees, seedlings and sap- lings of white cedar were abundant, and received higher values in the sam- ples than did those of other tree species. AuTECOLOoGy—Brief reviews of the physiological and growth require- ments to be expected for white cedar in the Florida panhandle have been prepared by Clewell (1971, 1981), based on studies made in North Carolina and elsewhere (Korstian, 1931; Wells, 1942; Little, 1950; Fowells, 1965). The general assumption was made that white cedars in Florida are similar in behavior and requirements to those of the Atlantic seaboard provinces. In most respects the present study has shown this assumption to be justified, although in many details, and particularly with regard to fire, the white cedar stands of Florida and the Gulf coast are different and characteristic of the special conditions under which they grow. Reproduction: Atlantic white cedar in peninsular Florida and west along the Gulf coast, although often overtopping its associated hardwoods and fre- quently a dominant component of the canopy, is almost never found in the exclusive, uniform-aged stands characteristic of white cedar on the northern Atlantic coastal plain. The uneven-aged, mixed-species stands typical of the southern white cedar forests are a consequence of gap succession in the ab- sence of fire. The seeds of white cedar are produced in appreciable numbers; by use of seed traps at ground level, a mature stand in New Jersey was found to pro- duce up to 22 million seeds per hectare per year (Little, 1950). Viability in the soil may be brief; an earlier New Jersey study found numbers ranging down from 2.7 million viable seeds per hectare in the uppermost 7.6 cm soil layers (Moore and Waldron, 1940). Seed production in Florida and the Gulf coast may be much less on an area basis, for the tree density is comparatively low. However, beneath or adjacent to a given tree, the number of viable seed may be fully equal to the levels recorded in New Jersey. A single favorable 28 FLORIDA SCIENTIST [Vol. 52 location under optimal conditions may temporarily support five or six seed- lings per dm’, a number that would require 20-24 million seeds per hectare, assuming a single- year supply and a 25% rate of germination. Whenever a gap occurs in the canopy, either by limb-fall or more major disruption of one of the larger trees, and light intensity at the ground level is increased, large numbers of white cedar seedlings appear either from seeds dormant in the soil or newly shed by adjacent trees. The seedlings have little or no capability of penetrating a litter of undecomposed leaves, but are able to germinate on exposed peat, rotting logs or stumps, mossy hummocks, ex- posed sandy seepage areas, or the banks of spring runs. Seedlings at a given spot tend to be uniform in size and (presumably) age, reflecting a single opportunity when light, seed availability, moisture, and season were favor- able. Korstian (1931) has provided illustrations of seedlings of known ages that help in the estimation of the age of the young plants. Nearly all white cedar seedlings die following re-growth of the canopy and closure of the light-transmitting gap. Although the seedlings, once they reach an age of three or four years, are able to persist for a time under conditions of low light intensity, they eventually succumb. Commonly an entire cohort of seedlings will perish within the same season. Thus when a tree- or limb-fall later reopens the canopy, there are no suppressed seedlings to exploit the high-light opportunity, and the cycle of seed germination, seed- ling suppression, and death, begins again. Infrequently a second gap in the canopy develops before all of a cohort of seedlings have expired, and the survivors are able to continue growth. Se- quential, suitably spaced breaks in the canopy permit a seedling to develop into a small tree which, although still suppressed by the overstory, is increas- ingly capable of enduring periods of reduced light. Thus the longer the sap- ling persists and the larger its size, the greater the probability of its eventually attaining a place in the canopy. Once a tree enters the canopy and is no longer subjected to a regime of reduced light, its growth accelerates (see “Age, Size, and Rate of Growth”). With time, and with the limited life span of the associated tree species, the few white cedars that survive to reach the canopy become the dominant members of the forest association. Seed production is essentially limited to white cedars that have attained the canopy. Fire is rare and usually of low intensity in white cedar stands on perpetu- ally wet seepage slopes (see “Relationship with Fire”). Thus the periodic de- struction by fire of all mature trees, followed by re-growth from soil-pro- tected seeds, to produce a uniform-aged single-species stand (Korstian, 1931; Little, 1979), is not characteristic of the Florida and Gulf coast white cedar forests. Only in limited areas, most often at the upper edges of the stands, are there small, dense growths of uniform-aged trees, suggesting fire incursion from adjacent uplands. On most sites, the admixture of other species, primar- ily hardwoods with green foliage that retards fire, and with an irregular but predictable sequence of openings in the canopy, generates a white cedar stand No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 29 of long-lived trees with the addition of new individuals only fortuitously and in small numbers. Age, Size, and Rate of Growth: The growth rate of Atlantic white cedar in Florida and the Gulf coast, and the age and size that the trees may eventu- ally attain, has not been recorded, except in the most general of terms. There is now no way to determine the size to which white cedars grew in pre-Columbian times, for the easily worked wood, highly valued in pioneer days for shingles, boat timber, tubs and buckets, and uncounted other uses, led to the rapid exploitation of the larger trees in all accessible stands. But a very few surviving trees of exceptional size suggest the dimensions that may have been frequent in the primordial white cedar forest. The tree presently listed in the “National Register of Big Trees” as the largest Atlantic white cedar is located near Brewton, Escambia County, Ala- bama (Hartman, 1982); it has a diameter of 150 cm (circumference: 15 ft. 6 in.) and a height of 26.5 m (87 ft.). A second tree of even more imposing size was located in 1962 northwest of Flomaton, also in Escambia County, Ala- bama (Li, 1962); its trunk measured 155 cm dbh, its height 27m, its trunk to the first limb 9.75 m, and its crownspread 10 m. Both trees are on upper tributaries of the Escambia River and are relics of a stand that once extended south along the Escambia to its mouth on Pensacola Bay in western panhan- dle Florida. The Escambia River record-sized trees are not convincingly surpassed in earlier literature, nor in records from the more northern portions of the range of the species. Diameters of 152 cm and heights of 36.5 m have been men- tioned (Korstian, 1931; Fowells, 1965; Taras, 1971), although without loca- tion or other authentication. The classic treatise on North American trees, C. S. Sargent’s Sylva of North America (1896) reported white cedars “seventy or eighty feet [21-24 m] in height,’ with trunks “usually about two [60 cm], but occasionally three or four feet [90-120 cm] in diameter.” Other late nineteenth century accounts, written after the prime stands had been cut, cited an aver- age height of 12.2 m and an average diameter of 61 cm in North Carolina (Pinchot and Ashe, 1897) and maximum heights of 24 to 27 m and diameters seldom exceeding 60 cm in Alabama (Mayr, 1890; Mohr, 1901). The largest extant Atlantic white cedars in the northern states are appre- ciably smaller than the largest of those in southern Alabama. The New Jersey record white cedar, in Ocean County, has a diameter of 89 cm (Porcella, 1984). The three listed champions in North Carolina, in Camden, Chowan, and Beauford counties, have diameters of 71, 63.5, and 61 cm and heights of 22.5, 25.6, and 23.8 m (Anonymous, 1984). Other extant Gulf coast trees are also appreciably smaller than the two record-sized survivors on the Escambia River. The tree previously accepted as Florida’s record white cedar, in the Blackwater River State Park, Santa Rosa County (Will and Simmons, 1984), has a diameter of 97 cm, a height of 16 m, and a spread of 10.4 m; it is a “wolf,” a tree spared during turn-of-century logging because of its forked, defective trunk. Two trees encountered during 30 FLORIDA SCIENTIST [Vol. 52 the present study in the Ocala National Forest stand, Marion County, Flor- ida, are of similar dimensions. One, along Juniper Creek, has a trunk 88 cm dbh. A specimen along Morman Branch, even though its trunk has been deeply and repeatedly clawed by bears, has a diameter of 91 cm, a height of 26.5 m, and a crown-spread of 9.5 m.; it is the currently reigning Florida champion white cedar (Simmons, 1986). In Mississippi the largest white ce- dars have trunks 71 cm dbh and heights estimated at 24 to 30.5 m (Eleuterius and Jones, 1972). It is worth noting that the cross sectional areas of the trunks of the two large trees on the Escambia River are 2.4 or more times greater than the cross sectional area of these second-rank large trees. The absence of trees approach- ing the size of the largest specimens, as would be anticipated if a normal size distribution were still present, is an indication of the extent to which nine- teenth and early twentieth century logging has depleted the white cedar stands of the Gulf coast. The rate of growth of Atlantic white cedar, as measured in North Caro- lina, Virginia, and New Jersey (Korstian, 1931), is tabulated as remaining rather constant for up to 100 years. The trees in these more northern swamps are usually of uniform age, resulting from simultaneous germination follow- ing fire destruction or logging of the previous stand, and show only slight suppression during the juvenile years. The tabulated values (adjusted to met- ric scale) are annual increases in diameter of 0.32 to 0.43 cm for trees 20 years of age, and 0.35 to 0.48 cm for trees 80 years of age, in dense and open stands, respectively. Peninsular Florida and Gulf coast stands, in contrast, are usually of mixed composition, with the cedars sometimes sparsely distributed and seldom of uniform age. The annual rings commonly show evidence of suppression of the young trees, generally for several decades, before the trees attain a position within the mixed-species canopy and begin to grow at an accelerated rate. Thus sections of young trees reveal an age that if extended proportionately to older individuals is in excess of their true age, while shallow increment cor- ings of older trees understate the years necessary for the individuals to reach their mature size. Very few data are available to support estimates of rates of growth of white cedar on the Gulf coast. Fifty-year-old stands in Pearl River County, Mississippi, and Escambia County, Alabama, have been recorded (adjusted to metric) as having diameters of 18.8 and 19.8 cm (Korstian, 1931); these dimensions give annual rates of diameter increase of 0.38 and 0.40 cm. Two small trees cut in the Ocala National Forest with trunk diameters of approxi- mately 10 and 13 cm, showed annual rates of increase of 0.16 and 0.27 cm (Ward, 1963), the relatively slow rate indicating suppression characteristic of such juvenile trees. Three trees in the Deep Creek stand, Putnam County, Florida, cut in 1981 for a power line right-of-way, with trunk diameters of 30, 46, and 48 cm, gave annual rates of increase of 0.27, 0.49, and 0.44 cm. No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR on In 1962 one of the present authors surveyed a small part of the Morman Branch stand in the Ocala National Forest and measured the diameter of 39 of the larger trees (Ward, 1963). The trees selected for measurement were individuals with diameters of greater than 30 cm. Although the selected trees were not marked, notes were taken indicating the physical status of each tree and its location in relation to stream channels and to other measured trees. In 1984 the same area of Morman Branch was re-surveyed. As in 1962, all trees with diameters greater than 30 cm were measured. Some of the earlier samplings, marked as “top gone” or otherwise defective, had disappeared or were reduced to moss and Pieris-covered stumps, while several trees too small to have been tabulated in the earlier study had now attained sufficient size for inclusion. | The measured portion of the stand, consisting of 36 trees in 1962 (with 3 trees excluded as outside the contiguous perimeter of the tabulated area), contained 34 trees in 1984. The average diameter of 45.7 cm in 1962 had increased to 52.5 cm by 1984, an annual increase in diameter of 0.31 cm. This increase, representing the increase observed in the typical larger tree of the stand, rather than the increase of individual trees, suggests a recovery by the stand from some disturbance. What this disturhance may have been— possibly severe windthrow, or conceivably fire—must have been distant in years, and has not been identified. Only 9 of the trees in the 1962 sampling could be identified with certainty in 1984. These trees, which in 1962 had had an average diameter of 43.4 cm, by 1984 had increased to an average diameter of 58.3 cm, a mean increase of 14.9 cm in 22 years, or an annual increase in diameter of 0.68 cm. This rate of annual increase in diameter, as recorded in the Ocala National Forest stand, is thus appreciably greater than other values of the Gulf coast or for the more northern white cedar stands. With these values for rates of growth in diameter, it is possible to make estimates of the age of the largest Atlantic white cedars of Florida and the Gulf coast. Under most conditions a period of at least 50 years, and at times as much as 100 years, is required for trees to attain sufficient stature to enter the canopy and to overcome the shade suppression of their juvenile years; for this period a rate of diameter increase of 0.25 cm per year is appropriate. Beyond 50 to 100 years and continuing to senescence, a significantly higher rate of increase is attained; the observed Ocala Forest rate of diameter in- crease of 0.68 cm may be used. By inverting these values, and using a median age of 75 years for the release from suppression, a simple table may be de- rived, suitable for estimating the age of Florida and Gulf coast Atlantic white cedars (Table 5). If the values from which this table is derived are extended to the two exceptionally large Escambia River white cedars, age estimates may be ob- tained of 268 and 275 years. If the value of 0.40 cm increase per year for a 50- year-old Escambia County, Alabama, stand (Korstian 1931) is used without adjustment for juvenile suppression, the appreciably greater ages of 375 and 32 FLORIDA SCIENTIST [Vol. 52 TABLE 5. Relationship of trunk diameter to age, in white cedar. Size (cm dbh): Age (years): Size (cm dbh): Age (years): 10 40 70 150 20 idl 80 165 30 92 90 180 40 105 100 194 50 P21 110 209 60 136 120 224 387 years are deduced. Other than destructive measurement, there is pres- ently no way the true age of these largest trees may be determined. However, since the value for the 50-year-old stand was most likely reflective of juvenile suppression throughout the entire period of measurement, these latter values may well be in excess of the true ages of the trees. Two earlier estimates for greater maximum ages of white cedars are defec- tive. In the 1962 study of the Ocala Forest stand (Ward, 1963), ring counts from the stumps of small trees were extrapolated to suggest the largest trees of the stand were 250 to 350 years of age. The present study, utilizing actual rates of growth of larger trees as tabulated above, gives values for the two largest trees of the Ocala stand as 181 and 190 years. An 1868 report on the geology of New Jersey (G. H. Cook, in Fowells, 1965) indicated some trees have reached 1,000 years of age. Such an age, with the concomitant enor- mous size, seems outside the physical limits that elsewhere govern the size of the white cedar, and is best dismissed as allegorical. Flooding: Simons (1984) has made observations that demonstrate the sen- sitivity of white cedar to flooding. Simons wrote: “T grew in pots about 20 Atlantic white cedars derived from the Morman Branch [Ocala] population. I planted them out in several spots around the pond cypress and black gum swamps bordering the relatively stable pond known as Cypress Lake [in eastern Suwannee County, Florida]. I planted the trees, which were about 6 inches tall, in places above the average high water level that I thought would remain moist but would not flood very often or for very long. They were in association with Pinus elliottii, Quercus geminata, Ilex glabra, Lyonia lucida, and Leucothoe racemosa. They grew quite well for the first two years and were about two feet tall, vigorous and healthy. Then we had a wet summer during which they were flooded with perhaps 6 inches of water for a month or two. They all died. None of the associated plants died, including pine seedlings of similar and smaller sizes.” Simons’ observations confirm the intolerance of white cedars to flooding during the growing season. Of particular note is that white cedars were less tolerant even of shallow flooding than were plants of associated species. Sen- sitivity to flooding thus precludes white cedar from occupying floodplains of the larger rivers (except the back swamps), such as the Ochlockonee and Apalachicola. The water levels of the larger rivers rise and fall considerably with the season and with headwater rainfall. Sediment erosion and deposi- tion characteristic of such floodplains may also be deleterious to these shal- lowly rooted trees. White cedars, instead, tend to occupy banks of small streams, where gradual colluvial transport of sediments (i.e., erosion from adjacent uplands) is relatively important, compared to alluvial transport. No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 33 Relationship with Fire: Fire is very seldom observed in white cedar stands of Florida and the Gulf coast. The spring-fed saturated soils and lush, broad- leaved understory vegetation serve to suppress fire initiated by lightning strikes and other chance sources. In many white cedar stands superficial ob- servation reveals no signs that fire has been present. There is no blackening on the bases of the trees nor charred hollows in the trunks, and no carbon parti- cles are found in the soil. But where cabbage palms (Sabal palmetto) are associated with the white cedar, as is commonly the case in the disjunct stands in north-peninsular Florida, a distinctive long-lasting fire marker is often present. Trunks of Sabal, as is true of all palms, are fully developed by an apical- thickening meristem within the crown of the tree, and do not expand appre- ciably as the tree ages. Although over a period of years there is some ablation of the surface of the trunk, the rate of tissue disappearance is very slow. The sub-surface cork cambium and ever-renewed bark characteristic of gymno- spermous and dicotyledonous trees is entirely absent. Thus fire-charring of the trunk of Sabal persists as visible blackened tissue and protruding vascular strands long after similar superficial damage to the trunks of other trees has been lost by shedding of the outer layers of bark. In nearly all places in the north-central peninsular stands where careful search was made, at least some of the Sabal trunks were found to be black- ened. The carbonized discoloration invariably extended only part way up the clear bole of the tree, stopping abruptly some distance below the bases of the present oldest leaves. As an example, an 8 m Sabal growing immediately adjacent to white cedars along the south bank of Juniper Creek in the Ocala stand was blackened for only the basal 2 m, while the next 3 m of bare trunk bore no markings. Other Sabal along nearby Morman Branch were black- ened to 3 m, but with the trunks above that level unburned. None of the adjacent white cedars shown any blackening of the bark or other traces of fire. Although blackening of the Sabal trunks seems an unfailing indication that fire has been present, the proportion of the trunk that is blackened is an unreliable measure of the time that has elapsed since the burn. The leaves of Sabal fall from the crown by rupture of the petiole several decimeters from the point of attachment to the trunk. The resultant petiolar “boots” persist for many years, depending on moisture and perhaps on genetic factors, clothing the upper trunk with a fire-resistant sheath. Thus when the charred “boots” later fall, the underlying trunk is unblackened and deceptively suggests, by its apparent growth since the fire, of a burn more distant in time than is actually the case. None of the persistent petiole “boots” were observed to be fire-blackened. The absence of fire-sign on these structures, together with the absence of charring on the trunks of white cedar and its other associated trees, points strongly to a lapse of an appreciable number of years since the passage of fire through the stands. 34 FLORIDA SCIENTIST [Vol. 52 For the Ocala stand, an exact date is known when fire last swept through the surrounding scrub. Scrub vegetation, which covers the largest part of the Ocala National Forest, is characterized by sand pine (Pinus clausa) and other species that are both intolerant of fire and highly flammable. Since scrub vegetation is relatively open and produces a slow build-up of litter, fire is infrequent. But fire, when it does come, runs fast and hot. Fire kills the individual plants and simultaneously triggers the subsequent release of pro- tected seed onto the fire-cleared seedbed. This response of scrub to fire is quite unlike that of the central Florida “high pine,’ a longleaf pine—turkey oak (Pinus palustris— (Quercus laevis) association that is swept by fire every two or three years without killing the individual plants of the association. On 12 March 1935, a fire began on private land just southwest of the Ocala National Forest and, leaping across six 200-foot firebreaks under the impetus of winds averaging 42 mph and gusting to 65 mph, swept northeast- ward to Lake George, blackening 35,000 acres in four hours (Cooper, 1935; Folweiler and Brown, 1946). Soon after it began, the fire “crowned” in the mature sand pine. Although burning brands carried by the wind made the advance uneven, the fire moved at an average pace of 6 mph, in what has been termed the fastest-spreading forest fire in the history of the U.S. Forest Service (Cooper, 1935). Contemporary maps delimiting the extent of the fire (Folweiler and Brown, 1946) do not disclose to what extent this fire was responsible for the fire indications present in the white cedar stands along Juniper Creek and Morman Branch. But since the 1935 fire did sweep over all the adjacent scrub-covered uplands and no fire has occurred in those areas in more recent years, it is most likely that the 1935 fire also penetrated into the white cedar stands and that all fire indications date from that holocaust. The 1935 fire apparently did not reach the white cedar stands along the north side of Juniper Creek. Sabal trunks among the white cedars on the north bank show no trace of fire-blackening, in marked contrast to those trees 100 m to the south, across the freshwater marsh and open stream of Juniper Creek. Although the 1935 fire may have crossed Juniper Creek at other points, the portion of the north bank on which white cedars grow seemingly has been unburned for an interval appreciably longer than the half century since fire has passed through the white cedar stands on the south bank and along Morman Branch. Determination of fire events in the white cedar stand along Deep Creek in Putnam County is less certain. Since that stand is divided by highways into three blocks, fires may have occurred in some areas but not have spread through the stand. Clearly fire has been present, as is shown by blackened Sabal trunks. Burn scars in this stand have been reported as having been seen up to 3.7 m on the boles (Collins et al., 1964), although no such damaged trees were observed in the present survey. That fire has been a recurrent factor over very many years is indicated by the presence of charcoal in each of three samples at all levels to 60 cm (Collins et al., 1964). But no evidence of No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 35 recent fire was encountered in the present study, nor indication of its past influence on the white cedars or other trees. In panhandle Florida, white cedar and Sabal rarely occur together, thus preventing use of the cabbage palm as a convenient marker of recent fire history. Occasionally a charred stump is found, confirming a conflagration in the distant past. More frequently, but less decisively, hardwood trees are en- countered with multiple trunks from the base. With few exceptions (as pop ash, Fraxinus caroliniana), the trees associated with white cedar grow from a single trunk if protected from fire, while multiple trunks commonly develop from coppice sprouts following partial destruction of the tree by fire. A consistent observation in panhandle white cedar stands is that there is considerable difference in tree sizes, and thus in tree ages. This size distribu- tion, together with the scant evidence provided by direct observation or indi- cations given by the occasional coppiced hardwoods, suggests that fire history in most panhandle white cedar stands approximates that described for the Florida peninsula. Where fire has been present it must not have been intense or at least was not uniformly destructive. The Tates Hell region of Franklin County, Florida, merits special men- tion. Although any stand may have individuals thriving in somewhat drier soil beyond and often above the periphery of the seepage surface, these trees are in associations that burn with greater frequency and intensity than the cedar swamp itself, and are thus to be seen as temporary, unsustainable inva- sions. In Tates Hell, large numbers of white cedars are thriving in boggy flatwoods dominated by slash pine (Pinus elliottii), even though such an asso- ciation elsewhere would normally burn every few years. White cedars also occur, although in smaller numbers, in the shallow pond cypress—sweet bay—black gum sloughs that drain the flatwoods. The tree distribution thus superficially suggests a white cedar forest of long duration in the flatwoods, with occasional trees becoming established in the sloughs. However, in pre- settlement days fire in the pine flatwoods of Tates Hell should have been no less frequent than in similar flatwoods elsewhere, and must certainly have been recurrent in the decades of need for access to the pines from which turpentine was being taken. Only within the present century has fire been suppressed, and in the areas of white cedar the dense understory points clearly to the success of fire control. It is highly unlikely that white cedars were common or even present in boggy pine flatwoods until this century, when intentional fire suppression became practiced. In earlier times the ce- dars must have been restricted to the shallow sloughs, and only in recent decades have they expanded into the adjacent flatwoods. During the present study, no mature white cedar stand in Florida or along the Gulf coast was observed to burn or to have recently been burned. Neither were white cedar fires observed by others and reported to the present au- thors. All evidence of fire in white cedar stands was indirect, usually ambigu- ous to some extent, and often distant in time. Clearly fire does occur in many of the stands, as evidenced by carbon in the soil, long-persistent blackening of 36 FLORIDA SCIENTIST [Vol. 52 Sabal trunks, and coppice behavior of associated hardwoods. Such fires may occur at intervals—even though of 50 years or greater—that are much less than the lifespan of the trees, an indication that the fire intensity is not suffic- ient to harm the larger trees. Seedlings are surely destroyed, and perhaps even saplings, but reproduction is not importantly affected, and the stand quickly replaces the small plants that may have been lost. Crown fires appar- ently do not occur, even under the impetus of strong winds and fires that have crowned in adjacent associations. White cedars in Florida and the Gulf coast are surely no less susceptible to fire, as individuals, than are trees in the dense, fire-sensitive stands along the Atlantic coastal plain. This is evidenced by the inability of the southern trees to persist indefinitely in associations where fire is hot and frequent, even though they may repeatedly and with temporary success invade such habi- tats. But in the topographically anomalous associations where they most com- monly occur, the soil is perpetually wet from seepage or surface flow, the cedars are often interspaced with less-flammable hardwoods, and the accom- panying herbaceous undergrowth is often dense and green. All of these fac- tors serve to suppress or to cool fires entering the stand, and to protect the mature white cedars from destruction. Lightning: Since in Florida and on the Gulf coast, fire has only inconse- quential impact upon mature white cedar on perpetually wet soils, and since the species is effectively immune to pathogens or insect predation, those trees that attain a position within the canopy may be expected to experience a lifespan of indefinite duration. Yet even within uncut forests, as in large por- tions of the stands in peninsular Florida, there are no indications of cedars that have attained immense size, greatly disproportionate to the dimensions of associated tree species. The influence of lightning as a force terminating the life of white cedars has not been fully appreciated. Lightning is either not noted as a direct cause of tree damage, or is mentioned only incidentally as a fire initiator (Korstian, 1931; Fowells, 1965). Yet in the mixed-growth Gulf forests, and particularly in the stands in Marion and Putnam counties, north-peninsular Florida, lightning appears to be almost the sole natural factor determining the upper age and size limit of white cedar. Occasional white cedars in all parts of the stands showed prominent signs of lightning strikes. Most trees so affected were of large size and either ex- tended above the canopy or may have done so before truncation. The degree of damage was usually massive and unmistakable, with slabs of wood to 6 m in length and 3 dm in width blasted from the trunk and tumbled to the ground beneath. Numerous fragments of limbs and bark were scattered for considerable distances from the stricken tree. No trees were seen in which a discrete furrow was channeled down the trunk, removing bark only from a narrow strip, as is so commonly observed in lightning-struck pines. The impression was gained that the force of the No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR ii discharge had passed more centrally through the trunk, exploding it perhaps by the instantaneous heating of its contained water. Trees hit by lightning appeared to have no capability of recovery. No tree was observed that showed healed scars attributable to lightning. Two large trees along Morman Branch, in the Ocala stand, noted in a 1962 survey (by D. B. W.) as having apparent lightning damage, were present only as low Pieris-covered stumps in a 1984 re-survey. The importance of lightning in the life history of white cedar in Florida and the Gulf coast is made more pronounced by contrast with its associated canopy species. Most of the other swamp forest trees have little of the cedar’s resistance to insects or fungal attack. Older individuals of red maple (Acer rubrum), pumpkin ash (Fraxinus profunda), sweetbay (Magnolia virgi- niana), and swamp bay (Persea palustris) commonly show cavities in the trunk and other harbingers of eventual collapse; only cypress (Taxodium spp.) and, to a lesser extent, loblolly bay (Gordonia lasianthus), show the white cedar’s immunities to the organisms that terminate the life of most forest trees. Thus with time, with the inexorable disappearance from the canopy of individuals of other species, the surviving white cedars come pro- gressively to protrude above the canopy. With sufficient years the difference in height of the cedars as compared with other canopy species becomes quite marked. Lightning has long been recognized as the most important natural cause of forest fire ignition, as well as a destroyer of individual trees (Komarek, 1964). The level of lightning activity differs very greatly both seasonally and geographically. Recent studies, employing electronic detection and location equipment, have made possible the estimation with a useful degree of preci- sion of the actual frequency of lightning impacts, or ground flashes, in a given area (Darveniza and Uman, 1984; MacGorman et al., 1984). By these studies, Florida has been demonstrated to be almost unique in the United States in its high level of lightning activity, being approached only by portions of Arizona and New Mexico. Measurements obtained in the Tampa Bay area indicate a ground flash density of 12.9+5/km/?/year (Dar- veniza and Uman, 1984), and in the interior of the peninsula a density of 18/ km’/year (MacGorman et al., 1984). Ground flash activity in the Florida panhandle, and in southern Georgia and Alabama, is very nearly of equal magnitude. In contrast, large areas of the United States show less than 6 ground flashes/km/?/year. In hectare units, these data indicate lightning strike frequency in peninsu- lar Florida of 0.129 to 0.18/ha/year (or 0.052 to 0.073 ground flashes/acre/ year). Stated in terms of frequency per unit area, a hectare is predicted to be struck once each 5.6 to 7.7 years (and an acre, once each 14 to 19 years). These data on lightning strike frequency may be integrated with known behavior of lightning discharges, to estimate the probability of a given white cedar being struck. Standard lightning-protection code specifications estab- lish estimates of zones of lightning protection provided by masts of conduct- 38 FLORIDA SCIENTIST [Vol. 52 ing material extending above a base (N.F.P.A., 1980). For masts not exceed- ing 15 m in height, the zone of protection is conventionally taken as the space enclosed by a cone whose apex is the tip of the mast and whose radius at the base is equal to its height. As applied to a forest tree extending above the mean level of its associated canopy species, the protruding tree would be expected to receive all strikes that otherwise would fall within the circle atop the canopy centered on the protruding tree and whose radius was equal to the distance of protrusion. Because of the high-voltage properties of the discharge in creating its own ionized conduit, neither the relative conductivities of the trees nor the properties of the underlying soil would have a material influence in deflecting the strike. TABLE 6. Frequency of lightning strikes on protruding trees. Extension of Tree Protected Strike Frequency Mean Years Above Canopy (m) Area (m7?) Per Year@ Between Strikes 5) ae) 0.0010-0.0014 707-987 10 314 0.0041-0.0057 177-247 15 707 0.0091-0.0127 79-110 aDerived from estimates of ground strikes per km2 provided by Darveniza and Uman (1984) and MacGorman et al. (1984). From these measurements and observations, a table may be constructed to provide an estimate of the probability of a lightning strike upon a protrud- ing tree (Table 6). These values establish that a tree that extends only a few meters above its surrounding canopy does not incur a risk of lightning strike in excess of that risk proportionately shared by all members of the canopy. A tree protruding about 10 m or more incurs an appreciably increased level of risk. A tree extending 15 m above its canopy will be struck with a frequency of once every 100 years or so. If a single strike is lethal, the mean life expectancy of a tree, once its crown extends above the surrounding canopy to a height of 15 m, is approximately one century. These estimates are consistent with the observed age of white cedar trees in Florida and westward along the Gulf coast, and with their size both in absolute terms and relative to that of their associated forest trees. These val- ues are, of course, conditional upon other factors, some of which change constantly with season or with progressive growth of the target and associ- ated trees. Yet this analysis may point to fuller understanding of the life his- tory of the white cedar and its relationships both with its associated species and with the overlying meteorological factors that may govern its maximum age and size. Pathology: The Atlantic white cedar is justly known as a disease-free tree with decay-resistant wood. It is correctly reported to support few fungi, and to have no serious insect enemies (Fowells, 1965). Witches’--broom (Gymnosporangium sp.) has been observed to heavily infect some trees of a Mississippi stand of white cedar (Eleuterius and Jones, No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 39 1972). The report did not suggest the lifespan of the trees was reduced by the infestation. This fungus has not been observed to be significant elsewhere in southeastern stands. No standing tree observed in the present study was judged to be importantly affected by any insect or other pathogen. Cut or fallen timber of white cedar loses the standing tree’s near-immu- nity to fungal or insect attack. In the Deep Creek stand of Putnam County, Florida, trees cut in 1981 and left as logs half-submerged at the edge of a power line right-of-way, showed deterioration by 1984. Exposed sapwood had been extensively attacked by fungi at and above the waterline, and had lost its structural strength. The heartwood, where not excavated by carpenter ants, seemed unchanged from freshly cut wood. Carpenter ants (Camponotus sp.\, however, were found in the heartwood of a majority of logs examined. These insects feed on fruits and other plant materials found within their foraging area, and do not consume the wood of white cedar. But with their powerful mandibles they burrow within the heartwood of the logs where exposed above water, excavating elongate chan- nels which they use for their nests and into which they bring detritus which soon decays. The damage they thus do to cut or fallen logs is considerable, both by weakening the wood structurally and by exposing additional surfaces to weathering and fungal attack. This insect damage appears restricted to fallen timber in wet situations; no instances were observed of carpenter ants within a standing tree. Bear Damage: Damage to Atlantic white cedar by the action of bears has been observed in one Florida stand. The Florida black bear (Ursus americanus floridanus) once was found throughout Florida. It still occurs in limited numbers both in the panhandle and in the less-developed areas of the peninsula. The black bear in its range to the north has repeatedly been noted to scratch the trunk of large trees as a territorial marker and a presumed warning to other bears. The height of the scratching appears to be a signal of the size and social importance of the animal. Only male bears are believed to make these indicators (Maehr, 1984). In Florida, trees marked by bears as social signals are uncommon. They are occasionally observed in the panhandle, but apparently have not previ- ously been seen in the central peninsula. During the present study, several white cedars with greater or lesser amounts of such marking were located in the Morman Branch swamp of the Ocala National Forest. One tree of impressive size, with a trunk diameter of 91 cm, was mas- sively marked. The bark over half the circumference of the lower trunk was scarified or even ripped from the tree to a height just under 2 m. In places the damage had been deep enough to expose the wood, and the tree had formed new tissue to overgrow the destroyed cambium; then subsequent scratching had destroyed this scar tissue in turn. Long grooves, the marks of the bear’s claws, were inscribed in the bark, and the animal’s coarse black hairs were embedded in droplets of resin released by the ruptured tissues. 40 FLORIDA SCIENTIST [Vol. 52 A zone of about 5 m radius immediately surrounding the tree was clear of shrubs or other trees. At the edge of this opening a sapling sweetbay (Magno- lia virginiana) had been bent over and broken off at a height of 3 m. Within a short distance of this marked tree, several cabbage palms (Sabal palmetto) that had been carried down by falling trees so that their crowns were within a meter or two of the forest floor, had been ripped apart by bears who had removed, and presumably consumed, the large succulent bud of the palm and left their claw scratches on the remaining leaf bases. There are no reported occasions in Florida where bears have been ob- served as they marked trees by scratching; only by the few reports from areas to the north and by the evidence left behind can their actions be inferred. It is clear that the Morman Branch tree has been used as a marker by generations of bears. The depth of the damaged tissue and the repeated layers of scar tissue show that many years of similar behavior are involved. The creation of an apparent clearing around the marked tree may be only a consequence of trampling by the animals, or it may be an intentional act; the damage to the young sweetbay on the periphery of the cleared zone suggests the latter. Although the damage to this particular white cedar and, to a lesser ex- tent, to several others in the Morman Branch drainage is exceptional in Flor- ida, it is not unique within the range of the tree. White cedars in the Dismal Swamp, Pasquotank County, North Carolina, have been reported and photo- graphed with the bark stripped from the trunks by bears (Korstian, 1931). It has been suggested (Maehr, 1984) that bear behavior in marking trees is a function of density of the animal’s population. Bears in an area of low population density and hunting stress have no incentive to mark territory. Nor does the behavior appear where large trees are absent. Only where suit- ably dense bear populations co-exist with trees of appropriate dimension, as in the Ocala National Forest where bears are protected from hunting, may one expect to find such distinctively marked trees. Habitats: White cedars in Florida, Alabama, and Mississippi grow in moist woods along streams where alluvial deposition is limited, in bayheads with braided stream flow, in boggy pine flatwoods, in forested sloughs that drain open grass-sedge bogs and savannahs, and in back swamps of larger floodplains where flooding is nominal. (For brief descriptions of these and other communities and habitats, see “Glossary.”) All of these habitats are characterized by perennially moist or wet soils that are supplied by spring-fed streams, hillside seepages, rainfall that col- lects in low, boggy basins. These soils are only shallowly and briefly, if at all, flooded by river overflow. Fires are infrequent and commonly of low inten- sity. In different parts of the Gulf coast, white cedar habitats differ to some extent. The stations in north-peninsular Florida are in wet woods along streams that are primarily spring-fed. In the southern half of the panhandle, white cedar occurs not only on floodplains but also in open bogs and savan- No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 4] nahs, in low areas of pine flatwoods, and in pond cypress strands that slowly drain these habitats. Environments: Two general environments support white cedar. One con- sists of small valleys that dissect deep sandhills. Sandhills absorb nearly all rainwater that falls upon them and release it through perennial seeps to form the streams in the bottom of the valleys. From the Florida panhandle west- ward to Mississippi, soil moisture in white cedar swamps is derived primarily from seepage with a lesser amount from rainfall and surface runoff. In penin- sular Florida, spring-fed streams combine with seepage to maintain perenni- ally moist soils. The Citronelle Formation of the Western Highlands, a prov- ince comprising much of panhandle Florida, consists primarily of deep sands with occasional clay lenses or other impervious strata that impede percola- tion. These impediments force groundwater to move laterally until it appears as seepage in valleys (Puri and Vernon, 1964). Where soils are heavier, as in the loam soils of the Tallahassee Hills in the eastern panhandle, white cedars are absent. The other general white cedar environment consists of lowlands near the western panhandle Gulf coast and in adjacent Alabama that are charged by seepage from adjacent highlands to the north. Percolation of groundwater is prevented by proximity to sea level or by clayey soils. In these areas, white cedars occur in bay swamps and forested sloughs, especially cypress “strands,” which are sloughs containing much pond cypress (Taxodium ascen- dens). Bay swamps rarely burn. Light surface fires may occur in sloughs, but these fires carry poorly because of the scarcity of flammable materials. In response to fire suppression in the present century, white cedars have second- arily invaded boggy pine flatwoods adjacent to bay swamps and sloughs. Paleoecology: Not all terrain on the Gulf coastal plain with the above environments supports white cedar. A notable exception is a 32 km wide expanse of lowlands in Wakulla County, Florida, that begins 3 km east of the Ochlockonee River. Much of this area is occupied by the Bradwell Bay Wil- derness of the Apalachicola National Forest, and consists of boggy flatwoods, bay swamps, and cypress strands that would seem ideal as white cedar habi- tats. Paleoecological data shcw that these wetlands have been in continuous existence for what would seem more than enough time for white cedar migra- tion and establishment (Cameron and Mory, 1976). Since white cedar is abundant in a broad region that begins 2 to 3 km west of the Ochlockonee River, it is difficult to visualize why the cedar has not crossed the river and moved eastward into Bradwell Bay. The absence of white cedar in the Bradwell Bay area may be explained by the drainage pattern. First, the alluvial floodplain of the river would not be hospitable to white cedars, even though they would have access to this flood- plain from the west by way of Telogia Creek, a tributary abundantly flanked by white cedar forests. Second and probably of greater importance in restric- tion of migration, a high sandy bluff separates Bradwell Bay from the Och- 49 FLORIDA SCIENTIST [Velez lockonee River. White cedars could not survive in the dry sandhill vegetation just behind this bluff, which is swept by fire at frequent intervals. If there were stream-cut ravines transecting this bluff, white cedar could perhaps have found a habitat suitable for crossing the dry terrain, but such streams and ravines are lacking. The entire Bradwell Bay region drains toward the south and east, away from the Ochlockonee River, and no migration routes exist from areas where white cedar occurs. The absence of white cedar in Bradwell Bay demonstrates that habitat is not the only determinant of range of white cedar in Florida and the Gulf coast. Paleoecological factors also contribute to present distribution. The ex- istence of several centers of white cedar distribution in the peninsula and the panhandle (Fig. 2) is not explainable solely on the basis of current environ- mental conditions. The lack of suitable avenues for migration, as at Bradwell Bay, may also serve elsewhere to explain white cedar distribution. Another possibility may be considered in explaining why white cedar has its present distribution. It seems reasonable to suppose that the primary habi- tat of white cedar along the Gulf coast since the Pleistocene has been along small steams that drain deep sandhills, as is the present situation. During interglacial periods of high sea levels and correspondingly high water tables, white cedars moved up the slopes of the sandhills. During glacial periods of low sea levels and low water tables, the streams would have been fed by seeps at lower levels, and the white cedars moved down slope. In times of relatively stable sea level, as at present, white cedars have had sufficient time to colo- nize other habitats with appropriate moisture conditions, including bayheads and forested sloughs. Although competition from established vegetation may have prevented white cedars from entering some habitats, perhaps fire was more influential in limiting its distribution. Whatever the constraints in a given era, the fluctuations of sea level with the recurrent glacial cycles may have been the major determinant in permitting or restricting the movement of white cedar in the establishment of its present range. TaxonomMy—Until recently, all Atlantic white cedars of the eastern United States were identified as a single species, Chamaecyparis thyoides (L.) BSP. In 1962 Li (1962) described a new species, Chamaecyparis henryae Li, which included white cedars in western panhandle Florida, southern Ala- bama, and southeastern Mississippi. The new species was named for Mary G. Henry, Gladwyne, Pennsylvania, who had taken the field observations and provided Li with specimens. Chamaecyparis thyoides and C. henryae were distinguished on the basis of several, often overlapping characters, including the twisting of the bark, compression of the branchlets, color of the foliage, several leaf attributes (size, glandularity, keel shape, glaucousness of juvenile leaves), color of the staminate cones, size and color of the ovulate cones, and season of maturity of both staminate and ovulate cones. Botanists in general reacted with indifference to Li’s new species. Little (1966) recognized the entity as distinct, but reduced it to a variety, as C. thyoides var. henryae (Li) Little. Little expanded the range of var. henryae No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 43 to encompass white cedar stands throughout Florida, including the disjunct stations in Putnam and Marion counties as well as the isolated stands in Tay- lor, Talbot and Marion counties, Georgia. Independent botanists, however, did not accept henryae even in varietal status, and Little (1979), in his au- thoritative Checklist of United States Trees, withdrew recognition of Li’s en- tity, retaining it only as asynonym of typical C. thyoides. Angus K. Gholson, Chattahoochee, Florida, has made careful field exam- ination of white cedar within its Florida and Alabama range, with the objec- tive of understanding the differences reported by Li. Gholson has noted (1984) that there is intra-stand variation and that some characters vary within the same tree. Nonetheless, though he stresses that his observations are preliminary and require verification, Gholson is able to see regional patterns of characters that suggest to him that Li (1962) indeed described a valid taxon. Specimens from throughout the southern range of white cedar have been examined (by D. B. W.), with the initial objective of associating the field differences observed by Henry (as recorded by Li) and by Gholson with known stations of the species. Certain of these characters (i.e., color of cones, glaucousness of leaves) were found not to be reliably recorded by preserved specimens, while others (season of cone maturity, twisting of the bark) were largely unavailable. The presence of a small median gland on the facial leaves of typical white cedar, and its absence on the leaves of henryae, was found to have geographic constancy, and is the basis for the mapped distribution of the two variants (Fig. 2). The present authors recognize that a species distributed over a large geo- graphic range, and particularly one with many isolated populations, may be expected to vary morphologically but without necessary taxonomic signifi- cance. However, the correlation of at least the one readily observed differen- tial character (the presence or absence of the small foliar gland) with a dis- crete geographic range, and apparently correlated with at-present less-well-documented ancillary characters as observed by Gholson, appears to be justification for recognition of C. thyoides var. henryae. PROTECTION— White cedar in Florida and along the northern Gulf coast, as the dominant species of a distinctive plant association, is given protection more because of the recognized importance of wetlands than because of the regional rarity of the species. The State of Florida regulates the development or conversion of wetland areas. These areas are defined by the presence of one or more indicator spe- cies (Florida Administrative Code 17-4.02; Florida Statutes 403.901- 403.913). Wetlands contiguous to streams and lakes that are dominated by one of more indicator species are considered as waters of the state. As such, any dredge and fill activities require a permit from the Florida Department of Environmental Regulation or, in the case of agricultural activities, from the appropriate regional Water Management District. Additional state pro- tection is given to those streams classified as Outstanding Florida Waters 44 FLORIDA SCIENTIST [Vol. 52 (Florida Administrative Code 17-3.041). These include all streams in state and federal lands, as well as the Blackwater River, Perdido River, Shoal River, and Apalachicola River, all of which harbor white cedar in their flood- plains. In 1984, white cedar was added to the list of wetland indicator species. This listing, while not protecting individual trees from removal in timber harvest, may be expected to retard changes in land use that would destroy both the white cedar forest and its associated species. If white cedars were to be found to grow with endangered or otherwise protected species, the stands so located may be given additional protection. Nineteen plant species federally designated as “endangered” (and one desig- nated as “threatened”) occur in Florida. Of these, only four occur in the same areas or counties as does white cedar. A trailing morning-glory relative, Flor- ida bonamia (“Bonamia grandiflora”), though occurring in the Ocala Na- tional Forest within short distances of the Morman Branch white cedar stand, is restricted to the sand-pine scrub habitat of the Forest and is never found on the wet soils required by the white cedar. Harper’s beauty (Harperocallis flava), a perennial herbaceous monocot known only from two small areas within the Apalachicola National Forest, Franklin County, grows within a few kilometers of stands of white cedar, and in similar habitats, but the two species have not been observed to be sympatric. Chapman’s rhododendron (Rhododendron chapmanii) is very nearly restricted in its range to panhandle Florida, often occurring in the boggy pine flatwoods sometimes frequented by white cedar, but has only been found sympatric in southeastern Gulf County, southeast of Port St. Joe. Florida yew (Taxus floridana) has been reported as growing with white cedar in the Johnson Juniper Swamp of Lib- erty County in the central panhandle (Kurz, 1927). Other species that may occur with white cedar have been or currently are being considered for possible classification as “endangered” or “threatened” under the federal guidelines and laws. Florida willow (Salix floridana) grows with white cedar in both of the peninsular Florida stations. In the panhandle, species that may attain classified status include Euphorbia telephioides, Gen- tiana pennelliana, Oxypolis greenmanii, Parnassia caroliniana, and Pin- guicula ionantha. If any of these species become designated at the federal level as endangered or threatened, white cedars growing with them will re- ceive almost certain preservation when on federally owned land and an en- hanced level of protection even on privately owned property. The State of Florida requires a permit for the collection of certain desig- nated endangered and threatened plant species (Florida Statutes 581.185). The present statute (as amended October 1987) designates 129 species as en- dangered and 78 species or groups of species as threatened. Of these species, the following are known to occur with white cedar: Illicium parviflorum, Kalmia latifolia, Parnassia grandifolia, Rhododendron chapmanii, Salix floridana, Taxus floridana, and certain ferns and palms (including Rhapi- dophyllum hystrix). Although restrictions on the collection of an associated No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 45 species does not provide direct protection for white cedar, the presence of designated endangered or threatened species is used in justification for the purchase or other protection not only of the designated species but the entire white cedar association. A second Florida listing of endangered and threatened species has been prepared by the Florida Committee on Rare and Endangered Plants and Animals (Ward, 1979). This listing, although without legal sanctions, has been accepted and widely used in the selection of lands for state and county purchase for preservation of natural areas. Of the several included species that occur with white cedar, most have been incorporated into the Florida statute. Each species name is accompanied by a description, a range map, and by recommendations as to appropriate means for its preservation. No area within Florida, Georgia, Alabama, or Mississippi in which Atlan- tic white cedar is the dominant tree species, has been selected for preservation in perpetuity, to the authors’ knowledge. Those areas that are currently under protection have been assigned to that status because of other factors and may be withdrawn with changes in circumstances. No white cedars are presently being cut in the Ocala National Forest, the Apalachicola National Forest, the Blackwater River State Forest, nor the Eglin Air Force Reservation. These areas contain significant stands of white cedar, some virgin (as in the Ocala Forest), other logged in the past but now recovering. But none, either in these federal and state owned properties or elsewhere, has been set aside for perpet- ual protection. A change in economic demand for the timber, a change in policy or in management, and these areas of white cedar dominance may be lost to future study and appreciation. ACKNOWLEDGMENTS— The authors are grateful to the many persons who have assisted us in the assembly of this body of observations. Many of the topics we have touched upon have carried us well beyond classical botany. We wish to thank James R. Burkhalter, Linda C. Duever, and William J. Dunn for providing us with their unpublished field observations, to David S. Maehr for speculation as to the behavior of bears, to Thomas J. Walker for describing the practices of ants, to Ewen M. Thomson for data on lightning and the means of its detection, to Robert W. Simons for his records on the effects of flooding, and to Robert K. Godfrey and Angus K. Gholson, Jr., for observations on the morphology and distribution of white cedars. Our most special gratitude goes to those persons who have accompanied us in the field: Angus K. Gholson, Jr., Robert K. Godfrey, Thomas L. Morris, Robert W. Simons, James Dan Skean, Douglas H. Ward, Gordon C. Ward, Charles H. Wharton, Albert A. Will. This paper is Florida Agricultural Experiment Station Journal Series No. 6155. GLOSSARY Back Swamp: That portion of an alluvial river swamp distant from the main channel that is subject to relatively little overbank flooding and alluvial deposition at high water stage. Bayhead: Evergreen, hardwood swamp dominated by Magnolia virginiana (sweetbay) and sometimes by Persea palustris (swamp bay) or Gordonia lasianthus (loblolly bay), that occupies a headwater stream, often where flow is braided. Soils are peaty and strongly acid. Grass-sedge Bog: Seepage slope or flat dominated by low grasses and sedges (often Aristida stricta, Panicum acuminatum, Rhynchospora spp.) and containing many species of forbs, includ- ing insectivors. Hardwoods are suppressed by frequent surface fires. Pine Flatwoods: Open stands of Pinus palustrus and/or P. elliottii with a low, dense ground cover of Serenoa repens (saw palmetto), Aristida stricta (wiregrass), and many other shrubs and herbs. Kept open and parklike by frequent surface fires. Site conditions are broadly mesic. Stands where the water table is near the soil surface much of the year are called Boggy Flatwoods. 46 FLORIDA SCIENTIST [Vol. 52 Savannah: Grass-sedge bog essentially devoid of shrubs, except Hypericum fasciculatum. Best exemplified in the watershed of the lower Apalachicola River, where clayey soils impede the percolation of rain water on low-lying flats. Shrub Bog: Non-alluvial wetland dominated by mainly evergreen, broadleaved shrubs and small trees, including species of Cliftonia, Cyrilla, Ilex, Leucothoe, Lyonia, and Myrica. Shrub- bogs generally separate pine flatwoods, grass-sedge bogs, or savannahs on higher ground from bayheads or cypress swamps along streams. Titi Swamp: A shrub-bog dominated by Cyrilla racemiflora (titi) and/or Cliftonia monophy- lla (black titi). LITERATURE CITED Anonymous. 1984. Champion trees of North Carolina. North Carolina Div. of Forest Resources, Raleigh. 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Confr., Tallahassee, FL. 3:139-183. KorsTIAN, C. F. 1931. Characteristics, growth, and management of the forest. Pp. 1-35. In Korstian, C. F., and W. D. Brush, Southern White Cedar, U.S.D.A. Tech. Bull. No. 251. No. 1, 1989] WARD AND CLEWELL— ATLANTIC WHITE CEDAR 47 Kurz, H. 1927. A new remarkable habitat for the endemic Florida yew. Torreya. 27:90-92. LADERMAN, A. (ed.) 1987. Atlantic White Cedar Wetlands. Westview Press, Boulder and Lon- don, 401 pp. Li, H. 1962. A new species of Chamaecyparis. Bull. Morris Arb. 13:43-46. Litt te, E. L. 1966. Varietal transfers in Cupressus and Chamaecyparis. Madrono 18:161-167. . 1971. Atlas of United States Trees. Vol. 1: Conifers and important hardwoods. U.S.D.A., Forest Serv. Misc. Publ. No. 1146. . 1979. Checklist of United States Trees (Native and Naturalized). U.S.D.A. Handbook No. 541. 375 pp. Litt.e, S. 1950. Ecology and silviculture of white cedar and associated hard-woods in southern New Jersey. Yale Univ. Sch. For. Bull. No. 56. 103 pp. . 1979. Fire and plant succession in the New Jersey pine barrens. Pp. 297-314. In Forman, R. T. T. (ed.), Pine Barrens: Ecosystem and Landscape, Academic Press. MacGorman, D. R., M. W. Mater, anp W. D. Rust. 1984. Lightning strike density for the contiguous United States from thunderstorm duration records. Ms. prepared for: Office Nuclear Reg. Res., U.S. Nucl. Reg. Comm., Washington, D.C. 44 pp. Maenp, D. S. 1984. Fl. Game and Fresh Water Fish Comm., Naples. Pers. comm. Mayr, H. 1890. Die Waldungen von Nordamerika. Munchen. 448 pp. Monr, C. 1901. Plant Life of Alabama. Contr. U.S. Nat. Herb. No. 6. 921 pp. Moore, E. B., AND A. F. WaLpron. 1940. Growth studies of southern white cedar in New Jersey. J. Forestry 38:568-572. Munns, E. N. 1938. Distribution of important forest trees of the United States. U.S.D.A., Forest Serv. Misc. Publ. No. 287. N.F.P.A. 1980. Lightning protection code. National Fire Protection Assoc., Boston, MA. PiIncHoT, G., AND W. W. AsHE. 1897. Timber trees and forests of North Carolina. N. C. Geol. Surv. Bull. No. 6. M.1I. & J. C. Stewart, Winston, N. C. Porce ia, S. 1984. New Jersey’s record trees. New Jersey Outdoors 11(5):17-22. Puri, H. A., AND R. O. Vernon. 1964. Summary of the geology of Florida and a guidebook to the classic exposures. Fl. Geol. Surv. Tallahassee. Spec. Publ. No. 5, revised. 312 pp. RosENAu, J. C., G. L. FaAukner, C. W. HENpry, JR. AND R. W. Huu. 1977. Springs of Florida. Fl. Geol. Surv. Bull. 31, revised. 461 pp. SARGENT, C. S. 1896. The Sylva of North America. 10:111-114. Simmons, A. P. 1986. Champion Trees of Florida. Pamph., Div. Forestry, Dept. Agric. and Cons. Serv., Tallahassee. 33 pp. Stmons, R. W. 1984. 1122 S.W. 11th Ave., Gainesville, FL. Pers. comm. Taras, M. A. 1971. Atlantic white cedar. In American Woods, U.S.D.A., Forest Serv. FS-225. 8 pp. Warp, D.B. 1963. Southeastern limit of Chamaecyparis thyoides. Rhodora 65:359-363. ___. (ed.) 1979. Rare and Endangered Biota of Florida. Vol. 5: Plants. Univ. Presses of Fl., Gainesville. 175 pp. WeLts, B. W. 1942. Ecological problems of the southeastern United States coastal plain. Bot. Rev. 8:533-561. West, E. AND L. ARNOLD. 1946. The Native Trees of Florida. Univ. Presses of Fl., Gainesville. 212 pp. White, W. A. 1970. The geomorphology of the Florida peninsula. Fl. Geol. Surv. Bull. No. 51. 164 pp. Wits, P. C., anp A. P. Simmons. 1984. Florida trees of record size. Ms. prepared for: Office of Forest Education, Fl. Dept. Agric. and Cons. Serv., Tallahassee. 13 pp. Florida Sci. 52(1): 8-47. 1989. Accepted: May 10, 1988. Environmental Chemistry WATER QUALITY EFFICIENCY OF AN URBAN COMMERCIAL WET DETENTION STORMWATER MANAGEMENT SYSTEM AT THE BOYNTON BEACH MALL IN SOUTH PALM BEACH COUNTY, FLORIDA JEFFREY DEE HOLLER Smith and Gillespie Engineers, Inc., Sarasota, Florida 34243 ApsTRACT: Urban wet detention system investigations were conducted at a commercial shopping mall in Boynton Beach, FL. This study site possessed a permitted drainage area of 25.4 ha (62.8 ac), which was approximately 90 percent impervious. The water management area consisted of three interconnected ponds, each approximately 1.2 ha (3 ac), totaling 3.5 ha (8.7 ac). This site was instrumented with automatic water quality samplers to collect storm-generated runoff samples. In addition, digital stage measurement equipment continually monitored and recorded both surface and ground water elevation. Storm event sampling was initiated upon advent of rainfall in excess of - 1.3 cm (0.5 in) of precipitation over the watershed boundaries. Surface runoff inflow automatic samplers were flow proportionally activated by an electromagnetic flow sensor device. Surface water results indicate the following treatment efficiencies for select nutrients: NO,-N = 87 percent, NH,-N=55 percent, TKN=58 percent, 0o-PO,-P=69 percent, T-PO,-P=76 percent, and TSS = 91 percent. These results were collated with previous results in the literature and compari- sons indicated superior treatment efficiencies. THE Florida Department of Environmental Regulation (FDER) delegated the responsibility of managing stormwater runoff in South Florida to the South Florida Water Management District (SFWMD), pursuant to Chapter 17-25 of the Florida Administrative Code. The intent of this Chapter is “...to regulate potential sources of pollution to Waters of the State from surface water runoff.” SFWMD surface water permitting regulations intended to accomplish the goals of Chapter 17-25 require that stormwater management systems meet specified design criteria to reduce peak flows and enhance water quality pro- tection (SEFWMD, 1987). The Boynton Beach Mall stormwater management system was permitted according to criteria mandating wet detention storage of the first one inch of surface water runoff from the developed project or the total runoff of 2.5 inches times the percentage of impervious area, whichever is greater. The per- mitted stormwater runoff detention storage volume requirement was 2.2 inches. The greater the percentage of impervious area delegated to a project, the greater the required storage volume necessary under SFWMD permitting criteria. This simple concept to reduce runoff peak flow results in additional surface water quality improvements. Water quality benefits associated with wet detention are related to natural physical, chemical, and biological activi- ties which reduce runoff nutrient loadings prior to discharge into the receiving waters. No. 1, 1989] HOLLER—EFFICIENCY OF STORMWATER MANAGEMENT 49 A significant amount of scientific study has been conducted for the purpose of monitoring, controlling, and evaluating nonpoint source pollution (USEPA, 1983; Martin and Smoot, 1983; Wanielista, et al., 1981; and Mattraw, et al., 1978). A major source of nutrient enrichment in surface waters is from non- point source stormwater runoff. Researchers in Central Florida found that the major source of phosphorus entering Lake Eola was from stormwater runoff (Wanielista, et al., 1982). Principal sources of nonpoint pollution are from agricultural and riparian runoff and from unsound construction practices. Ru- ral runoff (a combination of agricultural, pasture and forest) has been reported to contain orthophosphorus concentrations as high as 0.1 mg/L (Schreiber, et al., 1980; Kunishi, et al., 1972). Research conducted in the State of Florida has shown that highway runoff can contain phosphorus concentrations as high as 0.5 mg/L (Yousef, et al., 1985). Sources of urban runoff pollution result from wet and dry atmospheric deposition, street refuse deposition (litter, street dirt, vegetation, and organic residues), traffic emissions and impact, and urban erosion (Novotny, et al., 1985). Phosphorus is a conservative element and is retained in the bottom sedi- ments (benthos) of lakes and ponds (under aerobic conditions). The major phosphorus sink component is the sediments, which exceed the quantity stored in the water column at any time. At low flows, sedimentation is the most efficient mechanism for removing phosphorus from the water column (Chapra and Tarapchak, 1976). Nitrogen may be lost from a surface water body by nitrification-denitrifi- cation processes. Nitrogen associated interactions have been thoroughly in- vestigated by Ritter and Ross (1984). These investigations have shown that a greater percentage of nitrogen is transported in baseflow (ground water) than surface flow, and that soil type has an influence on nitrogen loading rates. Excessively well drained soils had higher nitrogen loading rates than poorly drained soils. These findings have implications on the basic design and opera- tion of wet detention systems with respect to nitrogen control. In addition to nonpoint source stormwater runoff pollutional constituent contributions to wet detention systems, total nitrogen and ammonium con- centrations present in surface water bodies are associated with aquatic ani- mal activity living therein. Aquatic animals, in contrast to most terrestrial forms, commonly excrete ammonium as a waste product of metabolism (Cole, 1979). However, most dissolved ammonium in surface waters occurs as a result of bacterial mineralization of dead plant and animal matter. Wet detention stormwater management systems have been shown to be an effective means of reducing stormwater runoff nutrient loads to surface water bodies. This paper is concerned with investigations and analyses aimed at providing additional information necessary to evaluate surface water qual- ity improvements resulting from the present surface water management per- mitting practices in South Florida. Although a substantial amount of know]- edge and information currently exists regarding nutrient removal aspects of 50 FLORIDA SCIENTIST [Vol. 52 wet detention systems, the need for specific additional research regarding system component interaction and ultimate constituent fate is apparent. Purpose and scope of study—This study investigated water quality and quantity interactions associated with a wet detention stormwater manage- ment system. The site selected for these investigations was an urban commer- cial development known as the Boynton Beach Mall. This study is the third in a series of SFWMD investigations of surface water management systems, pre- ceded by Timbercreek (1984) and Springhill (1987), which were both resi- dential developments. Additional research is presently being conducted by SFWMD to evaluate an ex-filtration stormwater management system receiv- ing roadway drainage, and plans are being finalized to conduct similar re- search on a permitted agricultural citrus operation stormwater management system. Overview of SFWMD stormwater management practices - Individual Permit criteria state that any projects greater than 16 ha (40 ac) are subject to regulation. Projects less than 16 ha (40 ac) total area and/or 130 ha (320 ac) for projects in Dade County are subject to the General Permit criteria. Pro- jects with less than 4 ha (10 ac) total drainage area and less than 0.8 ha (2.0 ac) impervious coverage are exempt from the SFWMD stormwater regula- tory process. Activities in wetlands or existing water bodies prevent exemp- tion status for a project. The quantity performance standard enforced by SFWMD stormwater management permitting criteria dictate that off-site discharge from any de- veloped project area not exceed the 25-year, 3-day storm event, based upon historic discharges, or amounts determined in previous SFWMD permit actions, or amounts specified by SFWMD Allowable Discharge Formulae for specific sub-basins throughout the District. The quality performance stand- ards require detention and/or retention (D/R) in the overall system be pro- vided by either of the following methods, or combinations thereof: (1) Wet detention for the first inch (2.5 cm) of runoff from the developed project or the total runoff of 2.5 inches (6.4 cm) times the percentage imper- viousness, whichever is greater; (2) Dry detention, equal to 75 percent of wet detention volume; (3) Dry retention, equal to 50 percent of wet detention volume. Pre-treatment of one-half inch (1.3 cm) in a dry system is required for either commercial/industrial zoned projects, or projects discharging into “sensitive receiving waters.” Treatment credits are available for projects pro- viding inlets in grassed areas and additional surface water and/or roofed ar- eas. Dimensional design criteria are required to insure optimal system opera- tional efficiency and performance. For example, detention pond side slopes must be a minimum of 4:1, horizontal to vertical. Minimum area and depth requirements are necessary design components stated in the SFWMD regula- tory criteria. Other physical design constraints are intended to promote the development of well-defined littoral zones in the pond systems. Mitigation for No. 1, 1989] HOLLER— EFFICIENCY OF STORMWATER MANAGEMENT 51 BOY4 p 7 y SLTTITTITLTIT TL ISAS, YY V]V]V/US LOT CON GRESS AVENUE. IMPER VIOUS PARKING BON S GW-11@ GW-2 (UNDEVELOPED) OLD BOYNTON ROAD BOYNTON BEACH MALL URBAN RUNOFF STUDY SITE @ ROUTINE SAMPLING STATION Mi RAIN GAGE % SURFACE WATER INFLOW AUTOSAMPLER @ GROUND WATER SAMPLING WELL # SURFACE WATER OUTFLOW AUTOSAMPLER @ RECORDING WELL + SURFACE WATER STAGE RECORDER wee} COMMERCIAL BUILDING 400 FT WZ SURFACE WATER 125 M Fic. 1. Boynton Beach Mall study site. such undesirable designs as the use of bulkheads require the compensation of additional littoral zone coverage. Study area—The Boynton Beach Mall is an urban commercial shopping and service development located in the City of Boynton Beach, Florida, in southern Palm Beach County (S.19, T.45, R.45 E). The entire Mall property consists of 56 ha (139 ac), but it is divided into east (42.7 ha) [105.7 ac] and 52 FLORIDA SCIENTIST [Vol. 52 west (13.3 ha) [33.3 ac] drainage areas. The drainage for these areas is totally separated and the water quality monitoring program considered only the east sub-basin. The east sub-basin surface water management area was approximately 3.5 ha (8.7 ac) of stormwater detention ponds and 3.8 ha (9.5 ac) of green areas (parking medians and lake perimeter). Approximately 10.8 ha (26.8 ac) of the east drainage basin was undeveloped when the monitoring program began, leaving a developed area (parking lots and buildings) of 24.1 ha (59.7 ac). In July of 1987, an additional 1.3 ha (3.1 ac) became developed, result- ing in a developed area of 25.4 ha (62.8 ac). A total parking lot capacity of 5,032 spaces was available (Fig. 1). The stormwater management system consisted of three interconnected ponds which discharged into a local drainage canal (C-16). The detention pond elevation control structure was connected to the C-16 canal by 53 linear meters (175 ft) of 122 cm (48 in) diameter round concrete pipe. The C-16 canal flowed easterly, approximately three km (2 mi) from the detention pond discharge pipe before discharging into the Atlantic Ocean. The wet detention pond effluent was discharged through a single struc- ture, which was a 0.8 m (2.2 ft) wide concrete weir with a crest elevation of 3.0 m (10.0 ft) NGVD, and a 44.5 cm (17.5 in) wide, 60 degree V-notch orifice. The orifice possessed an invert elevation of 2.5 m (8.2 ft) NGVD, and served as a bleeder mechanism. According to SFWMD criteria, the invert elevation of the orifice represents the detention storage volume necessary for this particular system to provide adequate water quality improvements to the raw stormwater runoff entering the ponds. The drainage basin soils belong to the Pompano Fine Sand (Po) and Terra Ceia Muck (Tc) classifications (USDA-SCS, 1978). The Po soil series consist of nearly level, poorly drained, deep, sandy soils, formed in thick beds of sandy marine sediment. The Tc series consists of nearly level, very poorly drained, organic soils. They formed in thick deposits of well-decomposed hydrophytic plant remains. The water table is generally within 25 cm (10 inches) of the ground surface for two to twelve months of the year. MATERIALS AND MerHops—Surface water runoff sampling was conducted at the middle drainage sub-basin discharge into the detention pond and at the system discharge structure in the final pond during the period (May, 1986 to December, 1987). Boynton Beach Mall water quality analyses and laboratory methodologies are presented in Table 1. Episodic storm event sampling was conducted, based upon the following minimum hydro- logic criteria: (1) Minimum precipitation rate of 0.64 cm/hr (0.25 in/hr), sustained over a two-hour period. (2) An antecedent dry period of at least 4 days, where antecedent dry day is defined as a 24- hour period with less than 0.64 cm (0.25 in) of total rainfall. (3) Sufficient laboratory analysis capacity. Study instrumentation included one ISCO Model 2700 discrete sampler at the detention pond inflow sampling site (BOYIN), which was digitally linked to a Marsh-McBirney Model 265 flow meter. In addition to measuring surface runoff inflow entering the system, the Model 265 flow meter possessed the capability to activate the inflow sampler based upon a predetermined runoff flow increment of 37,850 L/sample. The flow meter installed at the inflow station (BOYIN) was originally intended to measure runoff flows for that sub-basin (29% of the total developed area). These flows were to be extrapo- No. 1, 1989] HOLLER—EFFICIENCY OF STORMWATER MANAGEMENT 53 TaBLeE 1. Boynton Beach Mall Study Laboratory Methods. Z Parameter Reference Code Units Total Kjeldahl nitrogen EPA Meth. 351.2 TKN mg/L-N Ammonia Std. Meth. 417G NH, mg/L-N Nitrite + nitrate Std. Meth. 418F NO, mg/L-N Nitrate Calculated NO, mg/L-N Nitrite Mod. Std. Meth. 418F NO, mg/L-N Total phosphate Std. Meth. 424CIII TPO, meg/L-P Orthophosphate Std. Meth. 424G o-PO, mg/L-P Total suspended solids Std. Meth. 209C TSS mg/L lated to other sub-basins within the watershed drainage area. However, this approach later proved to be neither accurate nor representative of actual recorded stage and precipitation mea- surements. Instead, a mass balance approach was used to compute surface stormwater runoff inflow quantities entering the detention ponds, as described in a subsequent hydrology/hydrau- lics section of this paper. Detention pond discharge was sampled by a Sigamotor 6200 series automatic wastewater monitor. The monitor was activated by an electrical contact float device, positioned on an adjust- able steel tape at ambient water level elevation. Any subsequent rise in pond level due to precipi- tation resulted in outflow sampler activation. This sampler was programmed to collect discrete samples on a 30-minute time step interval for 12 hours after activation (time proportional). A weather station was equipped with two rain gauges; a Belfort Model PL-15605 weighing bucket type drum chart recorder and a tipping bucket rain gauge. The tipping bucket gauge was linked to a Stevens Digital Model 7000 series punched tape recorder (U.S. Patent 3,588,888) and served as the primary rainfall record. This particular recorder was equipped with a fast cycling punch motor to minimize the possible problem of the bucket tipping while the previous tip was being recorded. Rainfall was measured in 0.03 cm (0.01 in) increments and recorded every five minutes. Surface water elevation was measured by a stilling-well float and data were recorded with two additional Stevens Digital Model 7000 series recorders on 16-track paper tape at pre-selected five minute time intervals. One recorder was located in the final discharge pond and the other in the C-16 canal. All data were processed using IBM Personal Computers, Models AT and XT. Rainfall and stage data, along with dates and times, were stored in a random access master data file. Several computer programs were written in BASIC to check the raw data and insert data into the master file. An additional program was written to accomplish the hydrology/hydraulics computations and to produce a hard copy print-out of computed results. Mass loadings—To estimate surface water treatment efficiencies accurately, it was first neces- sary to compute the mass of constituents entering and leaving the detention lake system. Since inflow water quality samples were often collected at uneven time increments (flow proportional), raw concentrations data were averaged over five-minute intervals (when necessary) in order to relate discrete sample concentrations to individual flow records. Mass loads were calculated at each five-minute interval during an event period, (Eqn. 1). Whe OOF 10 (1) where M is the mass of the constituent; Q is flow; C is the concentration of pollutant; and T is the time factor. Pollutant constituent mass loads were totaled for each event period (usually 12 hours) and treatment efficiency was calculated as percent (Eqn. 2). ([Massin—Massout]/Massin) * 100 (2) where Massin is the constituent load inflow, and Massout is the constituent load outflow. Hydrology/hydraulics—The Boynton Beach Mall detention pond discharge structure was de- scribed previously in detail. During the entire study period, the majority of surface water dis- charged from the site occurred through the bleeder mechanism (V-notch), which was designed to return the lake system to its normal elevation after storm events. As previously mentioned, this mechanism consisted of an inverted triangle, 0.36 m (1.17 ft) in height, with a notch angle of 60 degrees and a notch elevation of 2.5 m (8.2 ft NGVD). 54 FLORIDA SCIENTIST [Vol. 52 If the lake system elevation was not greater than the top of the bleeder slot, discharges were computed according to the V-notch weir equation (Eqn. 3). Q = 2.5 tan (e/2) 2 (3) where Q is the discharge flow; ¢ is the notch angle; and H is the head on the notch. If the lake system elevation was greater than the top of the bleeder slot, discharges were computed according to the orifice equation (Eqn. 4). Q = Ca (2gh)°° (4) where Q is the discharge flow; C is the discharge coefficient; a is the area of the orifice; g is the acceleration due to gravity; and h is the orifice head. When the weir elevation was exceeded, the orifice discharge computed as described by equa- tion 4 was added to the weir discharge, as computed using the weir equation (defined by Eqn. 5). © = CL" (5) where Q is the discharge flow; C is the weir coefficient; L is the weir length; and H is the head over weir. For both bleeder slot flow and weir flow, reduced discharges due to submergences was taken into consideration, when applicable. Submergence occurs when the tailwater elevation (C-16) exceeds the bleeder notch invert or the weir elevation. The submergence weir equation was used (Eqn. 6). Q = QI (1 (H2-H1)!9)95 (6) where Q is the discharge flow; Q1 is the unsubmerged discharge; H1 is the headwater head; and H2 is the tailwater head. Under high discharge conditions, it is theoretically possible for a weir structure to pass more flow than the attached culvert can accommodate. This situation is known as “culvert control’, and when it occurs, the discharge from the site is the lesser of the two values, which would be the culvert discharge (Eqn. 7). 2g¢H 1+ KE + 29.2 rh R43) where Q is the discharge flow; A is the culvert x-sectional area; g is the acceleration due to gravity; H is the head on culvert; KE is the entrance loss coefficient; n is the Manning’s n; L is the culvert length, and R is the hydraulic radius. Surface inflow to the detention lake system was computed, based on a mass balance approach shown in Eqn. 8: Ores — (8) where I is the surface inflow; Q is the surface outflow; S is the change in storage; and R is the direct rainfall on the lake. Change in storage and rainfall were converted as averages for five-minute time increments. Evapotranspiration (ET) has not been taken into consideration, due to the short-term episodic duration of storm events analyzed. ET is an important component in long-term water budgets; however, when short periods of storm event generated runoff data are examined, it becomes insignificant. RESULTS AND Discussion— Eight distinct rainfall events were sampled for water quality trends. Critical rainfall information associated with these events is provided (Table 2). Each individual storm event rainfall volume is provided to represent the magnitude of precipitation. Maximum five-minute rainfall intensity is char- acteristic of the erosion and wash-off potential of pollution constituents present on the watershed prior to rainfall. Storm duration is included to provide some indication of the storm intensity and magnitude of runoff po- tential. Inter-event time is the time subsequent to an event until immediately No. 1, 1989] HOLLER—EFFICIENCY OF STORMWATER MANAGEMENT 55 TABLE 2. Boynton Mall storm event rainfall data. Event Number Storm Attribute 1 2 3 4 5) 6 7 8 Date 6SE 150C 270C 6DE 7MAR~ 15 AP aw 22 JU Volume 0.88 1, 78) 1S W530) 2.88 0.76 0.58 M65 (inches) Intensity Omri, 0223 0.26 Opza 0.35 ONS) OF 0.49 (5-minute maximum) Duration 4 ie 1 6 11 0.75 1 2 (hours) Inter-event ] 3 3 5 8 13 1 y time (days) Long term 3.89 5.65 5.65 Sal 6.9] 1.01 5.65 5.65 monthly total rain (inches) TABLE 3. Boynton Beach Mall treatment efficiencies; all values percent. Storm Event Number Parameter 1 2 3 4 5 6 i 8 Mean NO,-N 96 17 78 — 62 90 87 35 66 NO,-N 99 56 90 133 88 99 99 87 87 NH,-N 81 36 68 42 6 98 84 21 55 TKN 84 18 4] Sill 67 9] 86 46 58 o-PO,-P 90 42 68 4] i 9] 93 56 69 T-PO,-P 95 56 59 69 90 94 94 50 76 TSS 98 94 49 96 98 99 99 94 91 preceding another significant raintall. Long term monthly rainfall total rain values represent a 47 consecutive year period and are from the weather sta- tion at the Palm Beach International Airport (PBIA), the nearest National Weather Service Station to the study site. Comparisons between the event volume and long term total rainfall values provide a good means of charac- terizing storm events monitored in relation to typical rainfall patterns charac- teristic of this geographic region. In other words, these attributes are shown to characterize the normality of each individual storm event monitored dur- ing the study period. Surface water treatment efficiency—The Boynton Beach Mall surface water management system provided excellent removal efficiencies of nutri- ents. Phosphorus was removed from raw stormwater runoff an average of 69 percent and 76 percent for o-PO,-P and T-PO,-P, respectively. Nitrogen spe- 56 FLORIDA SCIENTIST [Vol. 52 TABLE 4. Reported detention pond treatment efficiencies for select nutrient parameters; all values percent. Study Site TSS T-PO,-P o-PO,-P NO,-N TKN NURP! 45 22 25 33 14 Orlando? 12 67 43 — 20 Orange Co.3 85 61 93 92 91 Timbercreek* 68 55 93 93 -31 Springhill® 0 64 98 98 MT Boynton Beach al 76 69 87 58 1USEPA, 1983 27Martin & Smoot, 1986 3ECFRPC, 1983 4Cullum, 1984 5Holler and Gregg, 1987 cies were removed an average of 87 percent, 58 percent, and 55 percent NO,- N, TKN, and NH.-N, respectively. Relatively lower efficiencies were realized for nitrogen than for phosphorus. This observation has been shown previ- ously in SFWMD studies, as weli as in other regional and national studies. Boynton Beach Mall surface water constituent treatment efficiencies are pre- sented in Table 3. For purposes of comparison, Boynton Beach Mall data were compared with treatment efficiencies reported in the literature from several regional and national studies. These comparisons appear in Table 4. SUMMARY AND ConcLusions— Water quality and quantity investigations were conducted at the Boynton Beach Mall surface water management sys- tem, in south Palm Beach County, Florida. Treatment efficiencies were supe- rior to results indicated in the literature from similar land use studies. The average treatment efficiency during the eight events sampled between Sep- tember, 1986 and July, 1987 at the Mall for select nutrient constituents are as follows: NO,-N = 87 percent; NH,-N=55 percent; TKN =58 percent; o-PO,- P=69 percent; T-PO,-P = 76 percent; TSS =91 percent. Diversion of stormwater runoff from a commercial parking lot area into a detention basin was found to be an effective means of reducing stormwater nutrient mass loads entering receiving waters. Since this research did not examine constituent mass losses and gains to and from groundwater seepage, the observed results can be attributed to sedimentation and settling mecha- nisms involving only the water column. Nutrient surface water treatment efficiancies presented herein may be interpreted as yielding a substantial ben- efit to receiving waters which would not have occurred without stormwater detention. ACKNOWLEDGMENTS— This study was supported by the South Florida Water Management District. Jeffrey Dee Holler was a Research Environmentalist with the South Florida Water Management District throughout the period of time relevant to data presented in this paper. No. 1, 1989] HOLLER—EFFICIENCY OF STORMWATER MANAGEMENT Si LITERATURE CITED AMERICAN Pusiic HEALTH AssociATION. 1980. Standard Methods for the Examination of Water and Wastewater, 15th ed., Washington, D.C. Cote, G. E. 1979. Textbook of Limnology. C. V. Mosby Co., St. Louis. CuLiuM, M. G. 1984. Evaluation of the Water Management System at a Single-Family Residen- tial Site: Water Quality Analysis for Selected Events at Timbercreek Subdivision in Boca Raton, Florida, So. FL. Water Mgt. Dist. Tech. Pub. 84-11, Vol. II, West Palm Beach. East CENTRAL FLORIDA REGIONAL PLANNING CounciL (ECFRPC). 1983. Best Management Prac- tices, Stormwater Management Report, 208 Area-Wide Water Quality Management Planning Program, Winter Park. ENVIRONMENTAL PROTECTION AGENCY. 1974. Methods for chemical analysis of water and wastes. Environmental Protection Agency, Water Quality Office, Analytical Quality Control Laboratory, Cincinnati. Ho ter, J. D., AND J. R. Grecc. 1987. Water Quality and Hydrological Characteristics of a Wet Detention Stormwater Management System in South Florida. Presented at the 14th An- nual ASCE Water Resources Planning and Management Div. Conference, Kansas City. Kunisu1, H. M., A. W. Taytor, W. R. HEALp, W. J. GrueK, AND R. N. Weaver. 1972. Phos- phorus movement from an agricultural watershed during two rainfall periods. J. Agricul. and Food Chem. 20:900-905. Martin, E. H., anp J. L. SMoor. 1986. Constituent-Load Changes in Urban Stormwater Runoff Routed Through a Detention Pond-Wetlands System in Central Florida. U.S. Geolog. Survey Water-Resources Report 85-4310, Tallahassee. Mattraw, H. C., Jr., J. HARDEE, AND R. A. MiLuer. 1978. Urban Stormwater Runoff Data for a Residential Area, Broward County, Florida. U.S. Geolog. Survey Water-Resources Report 78-324, Tallahassee. Novotny, V., H. Sunc, R. BANNERMAN, AND K. Baum. 1985. Estimating nonpoint pollution from small urban watersheds. J. Water Poll. Cont. Fed. 57(4): 339-348. Ritter, W. F., anp J. R. Ross. 1984. Nonpoint source nitrogen loads to Delaware lakes and streams. J. Agricul. Wastes, 9: 35-50. SCHREIBER, J. D., K. RauscH, AND A. OLNEss. 1980. Phosphorus concentrations and yields in agricultural runoff as influenced by a small flood detention reservoir. Pp. 20-46. In: STE- FAN, H. G. (ed), Proc. Symp. on Surface Water Impoundments, Minneapolis. SouTH FLoripA WATER MANAGEMENT District (SFWMD). 1987. Management and Storage of Surface Waters, Vol. IV, Permit Information Manual. West Palm Beach. UNITED STATES DEPARTMENT OF AGRICULTURE - SOIL CONSERVATION SERVICE. 1978. Soil Survey of Palm Beach County Area, Florida. Washington, D.C. Unitep STATES ENVIRONMENTAL PROTECTION AGENCY (USEPA). 1983. Final Report of the Na- tional Urban Runoff Program (NURP), Vol. I, USEPA, Washington, D.C. Youser, Y. A., M. P. WANIELISTA, AND H. H. Harper. 1985. Fate of Pollutants in Retention/ Detention Ponds. Presented at the meeting of the Florida Dept. of Environ. Reg. Annual Stormwater Mgt. Workshop, Orlando. Wanie.isTA, M. P., Y. A. Youser, AND J. S. Taytor. 1982. Stormwater Management to Improve Lake Water Quality. USEPA Report No. EPA-600/S2-82-048, Washington, D.C. Florida Sci. 52(1): 48-57. 1989. Accepted: June 17, 1988. Biological Sciences ALLOCATION OF ENERGY RESOURCES IN THE FRESHWATER ANGIOSPERMS VALLISNERIA AMERICANA MICHX. AND POTAMOGETON PECTINATUS L. IN FLORIDA. CLINTON J. DAWES AND JOHN M. LAWRENCE Department of Biology, University of South Florida, Tampa, FL 33620. USA AsstrRACT: Vallisneria americana Michx. and Potamogeton pectinatus L. were collected over a four-year period in the winter and summer from a spring-fed river in central Florida and analyzed for proximate constituents (dry weight, ash, protein, soluable carbohydrate, lipid, fiber and lignin) from which energy content was determined. The stolon of the freshwater species did not store carbohydrate seasonally, and kiloJoules per g dry weight were substantially lower in both winter and summer samplings in blades and stolons than reported for seagrasses of the same taxonomic families. Although leaf biomass increased significantly in the summer, a similar in- crease was not noted for the stolon again in contrast to their marine counterparts. Blade fiber levels in V. americana were higher (to 34%) than that reported for seagrasses or even some terrestrial grasses, suggesting that the broad, linear blades have adapted to high water movement by allocation of a large part of their organic content to structural carbohydrate. MonocotyLepons of the families Hydrocharitaceae and Potamogetona- ceae occur as submerged angiosperms in both freshwater and marine systems (Cronquist 1981). Thus similarities and differences might be expected in spe- cies found in the two aquatic environments. Seagrasses show distinct alloca- tion of energy at different times of the year to various plant organs (Dawes and Lawrence 1980; 1983). Information on energy allocation in freshwater angiosperms is more limited as shown in the reviews by Boyd and Scarsbrook (1975) and Little (1979). Boyd (1970) reported the levels of kilocalories per g dry weight for three species of Potamogeton but did not distinguish among plant components or seasonal variation. Donnermeyer and Smart (1985) av- eraged the energy levels for the different plant components for the entire year for Vallisneria americana Michx. from the northern U. S. and presented sea- sonal proximate composition and relative sizes of the plant components. The seasonal study presented here involved the rocted aquatic angio- sperms Vallisneria americana and Potamogeton pectinatus L., which are common submerged plants in northern and central Florida streams (Godfrey and Wooten 1979). We compare the seasonal composition and energy levels of the two species collected at the same site with Floridia seagrasses from the same taxonomic families, and also V. americana in Florida with populations from the northern U. S. MATERIALS AND METHODS—Summer and winter collections were made from January, 1982 to August, 1985. At each collection, entire plants of Vallisneria americana and Potamogeton pec- tinatus were collected from about 1 m depth of water in the Juniper Run approximately 10 km downstream from the spring source of the Ocala National Forest (29° N Lat. 82° W Long.), No. 1, 1989] DAWES AND LAWRENCE— ENERGY RESOURCES 59 Florida U. S. A. Eight plants from one population of each species were taken to the laboratory and cleaned of epiphytes and sand. Vallisneria americana was divided as described by Donner- meyer and Smart (1985) into the broad (1-2 cm) blades, a 2 cm section of stolon on each side of the shoot, and the fleshy root stock and roots if distinct. Potamogeton pectinaius was divided into the erect photosynthetic portion consisting of a green stem with the narrow (1-3 mm) linear leaves, a 2 cm section of stolon on each side of the stem, and the fleshy base or root stock. The roots of all plants were pooled. The plant samples were weighed, dried under vacuum over concentrated sulphuric acid at room temperature and reweighed. After grinding the dried tissue to a powder (Wiley Micromill, #40 mesh), it was analyzed for ash, protein, soluble carbo- hydrate, and lipid, and energy levels calculated as described by Dawes and Lawrence (1980; 1983) and Brody (1964). Insoluble carbohydrate was estimated by difference. Fiber and lignin were determined according to Goering and van Soest (1970) using the acid detergent method to remove hemicelluloses. All constituents are expressed as percent dry weight. Standard one and two way analyses (ANOVA’s) were used to ascertain statistical significance (P <0.05). TABLE |. Proximate constituents as percent of dry weight and energy levels (kiloJoules g dry wt!) from winter and summer collections over a 4-yr period of Vallisneria americana growing in Juniper River, Florida. N=8+1S.D. Blade Stolon Roots Root Stocks Winter Summer Winter Summer Winter Summer Winter Summer Dry Wt (%) Ik Oi 2 aaa Ue | oF 20 fool. 23.4 3 One One #302 eA eel Oy Reo. 2 Ash (% ) TA 5 Dl 78} 23 28 25 18 +9.4 +10.4 +9.7 +9.5 +10.1 +92 +15.2 +11.3 Sol Carbo. (%) D6e 4 SO oo 28 = 18 2S +11.6 +5.4 +17.1 +9.7 +13.0 +].1 +10.8 Insol Carbo. (%) 41 40 37,30 31 40 64 Protein (%) Sard Sr el (le ay 10° 1225 +3.8 +2.4 +3.7 + 3.5 SA +4.5 +8.8 Lipid (%) 1eG 3 D2 ele) 5.8 0 Levis 29 +0.4 +0.6 +().7 +0.6 +6.9 +0.7 Energy (kJ) ete 326 1S2oe es e7 13 See 107, Nee Sas} ResuLtts—The average water temperature at the Juniper Run site was 10.0 C° (winter) and 23.5 C° (summer). The percent dry weight of blades, stolons, and root stocks and roots of the two species were lower in the winter, but the differences were not significant (Tables 1, 2). Ash levels in the blades ranged from 21 to 29% (Potamogeton pectinatus, winter and summer, Table 2). Stolon ash levels ranged from 16% (P. pectina- tus, summer, Table 2) to 24% (Vallisneria americana, winter, Table 1). Ash levels in the root stock ranged from 18% (V. americana, summer, Table 1) to 27 % (P. pectinatus, winter, Table 2). Protein levels were not significantly different between winter and sum- mer collections or between blades and stolons for any species. Protein levels ranged from 8% (Vallisneria americana, winter, Table 1) to 10% (V. ameri- cana, summer; Potamogeton pectinatus, winter, Table 1 and 2 for blades and 60 FLORIDA SCIENTIST [Vol. 52 TABLE 2. Proximate constituents as percent of dry weight and energy levels (kiloJoules g dry wt!) from winter and summer collections over a 4-yr period of Potamogeton pectinatus growing in Juniper Run, Florida. N=8+1S.D. Blade Stolon Root Stock Winter Summer Winter Summer Winter Summer Dry Wt (%) He 12 Las, ah LO, 2 #776) 2 3:3 +320 (pan #2], = =23280 Ash (% ) 21 = =—29 PA ANG) oT 22 +6.6 +9.7 +11.8 +11.0 +10.5 +11.9 Sol Carbo. (%) 21, 23 29 40 23-23 +81 +702 +14.1 +9.7 +8.4 +1.4 Insol Carbo. (%) 48 35 40 38 Bono Protein (% ) tail) 7: 9 Tes +46 +1.9 +4.8 +3.0 +2.1 +5.0 Lipid (%) 125: 2256 Meche eta} 8,823 +0.8 +0.6 +1.] +0.6 +1.6 + ed Energy (kJ) 13.9 12.9 13295 W582 146° “Slifel stolons. Root stock protein levels ranged from 7 to 13% (P. pectinatus, winter and summer, Table 2). The roots and root stocks of V. americana had inter- mediate protein values. Soluble carbohydrate levels were highest in the stolon (30 to 40%) and root stocks (23 to 28 % ) for both species. The stolon had significantly higher levels of soluble carbohydrate when compared with the blades and root system of Pota- mogeton pectinatus. Summer collections of stolons of the two species had sig- nificantly higher levels than their blade counterparts. However, there were no significant seasonal differences in the levels of soluble carbohydrate between the winter and summer stolon samples for each species. Lipid levels were low in the blades, ranging from 1.5 to 2.6% (Potamoge- ton pectinatus, winter, Table 2) and 1.0% (Vallisneria americana, summer, Table 1) to 2.3% (P. pectinatus, summer) in the stolon. Lipid levels were highest in the root stock of P. pectinatus in the winter (9% , Table 2). Fiber in Vallisneria americana accounted for 29 and 30% in the blades and 30 and 40% in the stolon (winter and summer respectively). In Potamo- geton pectinatus, fiber levels were 31 and 23% in the blades and 39 and 22% in the stolon (winter and summer respectively). Lignin levels were highest in the winter collections of the blades of both species (V. americana: 6 and 2%; P. pectinatus: 4 and 3%, winter and summer) with lower levels in the stolons (V. americana: 0.2 and 2%; P. pectinatus: 3 and 2% , winter and summer). The blades of summer plants had significantly higher dry weights in Val- lisneria americana (27 and 7 g) and Potamogeton pectinatus (0.7 and 0.4 g) when compared to the winter levels, or the stolons of the same plants (V. No. 1, 1989} DAWES AND LAWRENCE—ENERGY RESOURCES 61 americana: 0.1 and 0.04 g; P. pectinatus: 0.1 and 0.02 g, summer and win- ter). Total energy levels in the stolon were low (V. americana: 0.6 and 0.2 kilojoules; P. pectinatus: 0.1 kilojoules, summer and winter) when compared with the blades of both species (V. americana: 206 and 39 kilojoules; P. pec- tinatus: 6 and 2 kilojoules, summer and winter). Energy values expressed per g dry weight did not differ much seasonally for the blades of both species (Tables 1, 2), but did show a summer high for stolons of Potamogeton pectinatus and for the root stocks of both species. The roots of Vallisneria americana had lower kilojoules in the summer. DiscussioNn— Differences in allocation of energy are evident when com- parisons are made between Vallisneria americana and the seagrass Thalassia testudinum Banks ex Konig, both in the family Hydrochayitacae, and be- tween Potamogeton pectinatus and the seagrass Halodule wrightii (Ascher- son) Ascherson, both in the family Potamogetonaceae (Cronquist 1981). Un- like the marine macrophytes found at the same latitude, there is no seasonal shift in constituents in the freshwater macrophytes. Due to the influence of the Ocala Springs, the Juniper Run has a very limited temperature range, and the seasonal changes in irradiances are assumed to be similar when com- pared with the marine environment on the west coast of Florida although water transparency can vary in seagrass beds. Blade biomass did increase in the summer in both species in a manner similar to that reported for seagrasses (Dawes and Lawrence 1980), and for Vallisneria americana from the Mississippi River (Donnermeyer and Smart 1985). However, there was never complete die back to a “winter bud” in the Florida V. americana as in the Mississippi plants. In fact, the plants showed new leaf growth in winter collections, unlike their marine counterparts. Sto- lon biomass did not show any major increase in dry weight and the level of soluble carbohydrate did not increase significantly in the summer. The stolon of these freshwater macrophytes apparently does not function as a storage organ for growth as shown in the marine macrophytes (Dawes and Lawrence 1980) although soluble carbohydrate is higher than in the blades. The blades, in fact, contain the majority of the plant’s organic constituents in both sum- mer and winter, unlike the seagrasses. Protein levels of blades and stolons in the freshwater macrophytes were similar or less than (Dawes and Lawrence 1980; Dawes 1986) levels found in the two seagrasses. The root stocks, when a fleshy stem base was separable, had the highest protein levels. The generally lower levels of protein and ash in the freshwater macrophytes over their marine counterparts suggest a lower energy allocation in growth. Blade protein levels reported here are less than half when compared with the data presented by Donnermeyer and Smart (1985). Perhaps it is because these authors calculated protein based on total nitrogen (Kjeldahl method) while we report soluble protein based on the Folin Ciocalteau reaction. 62 FLORIDA SCIENTIST [Vol. 52 Except for the winter blades of Vallisneria americana, the energy values of blades were 90 to 92% of their marine counterparts (Dawes and Lawrence 1980). The stolons of the freshwater plants also had lower energy levels than their marine counterparts; 78% for winter stolons of Potamogeton pectinatus when compared with Halodule wrightii and 95% for summer stolons of Val- lisneria americana when compared to Thalassia testudinum. Furthermore, if fiber is considered as a non-nutritive component, then the blades and stolons of the freshwater macrophytes have almost a third less of the nutritively available energy constituents than their seagrass counterparts (Dawes 1986). The levels of insoluble carbohydrate determined by difference ranged from 24 to 47% for blades and 33 to 40% for stolons while fiber levels ranged from 23 to 34% and 14 to 39% respectively. Thus, fiber does account for up to 90 % of the calculated insoluble carbohydrate. Lignin and fiber levels of winter blades of Vallisneria americana and Po- tamogeton pectinatus were higher than those reported for the marine coun- terparts while the summer levels were similar. Fiber levels in both freshwater macrophytes were also higher than true terrestrial grasses in Florida (in Dawes 1986) such as Festuca arundinacea (tall fescue) and Cynodon dactylon (Bermuda grass) but lower than the range reported for a population of V. americana in Mississippi by Donnermeyer and Smart (1985). However, these authors used a neutral detergent analysis which will include hemicelluloses while this study employed the acid detergent which removes the hemicellu- loses. The higher fiber levels of the broad, linear blades of V. americana, when compared to the morphologically similar blades of Thalassia testu- dinum as well as terrestrial grasses may be indicative of adaptation to current drag in a stream. The narrow blades of P. pectinatus would not experience the same amount of drag. Conc.usions—The stolons of Vallisneria americana and Potamogeton pectinatus do not function as storage organs in contrast to the seagrasses. This difference may be due in part to the stable year-around production of new blade material. The higher levels of fiber found in V. americana blades is probably due to the current of the stream habitat. LITERATURE CITED Boyp, C. E. 1970. Amino acid, protein, and caloric content of vascular aquatic macrophytes. Ecology. 51:902-906. , AND E. ScarsBrook. 1975. Chemical composition of aquatic weeds. Pp. 144-150. In: Bresonic, P. L., anp J. L. Fox, (eds.). Proc. Symp. on Water Quality Management through Biological Control. Environmental Protection Agency, Corvallis Oregon. ENV- 07-75-1. Bropy, S. 1964. Bioenergetics and Growth. Hafner, New York. 1023 pp. Croneuist, A. 1981. An integrated System of Classification of Flowering Plants. Columbia Uni- versity Press. New York. 1262 pp. Dawes, C. J. 1986. Seasonal proximate constituents and caloric values in seagrasses and algae on the west coast of Florida. J. Coastal Res. 2:25-32. , AND J. M. Lawrence. 1980. Seasonal changes in the proximate constituents of the seagrass Thalassia testudinum, Halodule wrightii, and Syringodium filforme. Aq. Bot. 8:371-380. No. 1, 1989] DAWES AND LAWRENCE—ENERGY RESOURCES 63 , AND J. M. Lawrence. 1983. Proximate constituents and caloric content of seagrasses. Mar. Technol. Soc. J. 17:53-58. DoNNERMEYER, G. N., AND M. M. Smart. 1985. The biomas and nutritive potential of Vallisneria americana Michx in Navigation Pool 9 of the upper Mississippi River. Aq. Bot. 22:33-44. Goprrey, R. K., anp J. W. Wooren. 1979. Aquatic and Wetland Plants of the Southeastern United States. Univ. Georgia Press, Athens. 933 pp. Goerine, H. K., anp P. J. VAN Soest. 1970. Forage Fiber Analyses. U. S. Department of Agricul- ture. Handbook No. 319. 20 pp. Littte, E. C. S. 1979. Handbook of Utilization of Aquatic Plants. A Review of World Literature. Food and Agriculture Organization of the United Nations, Rome. FAO Fish. Tech. Pap. No. 187. 176 pp. Florida Sci. 52(1):58-63. 1989. Accepted: July 8, 1988. 64 FLORIDA SCIENTIST [Vol. 52 ACKNOWLEDGMENT OF REVIEWERS It is a pleasure to acknowledge the service, dedication, and cooperation of the following persons who gave generously of their time and expertise in reviewing manuscripts for Volume 51 of the Florida Scientist. Some reviewed more than one manuscript. W. L. Adair, Jr. Jesse S. Binford, Jr. John V. Betz Joan Chernela Bruce Cowell Jeff C. Davis, Jr. Clinton J. Dawes Max Dertke Patricia M. Dooris Kent Fanning Jack W. Frankel Roger T. Grange Michael J. Hansinger Gertrude Hinsch Harold Humm Yasar Iscan Joseph C. Joyce F. Wayne King John M. Lawrence James N. Layne Leslie Sue Lieberman William Loftus Jeffrey M. Mitchem Paul Moler Thomas Morosko Henry Mushinsky John S. Osborne George M. Padilla Anthony Paredes M. J. Perez-Cruet Paul L. Shafland Joseph L. Simon Gregory J. Stewart I. J. Stout Walter K. Taylor Diane TeStrake S. G. Tolley Curtis Wienker Richard P. 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Florida's Estuaries —Management or Mismanagement? — Academy Symposium FLorRIDA SCIENTIST 37(4) — $5.00 Land Spreading of Secondary Effluent —Academy Symposium FLORIDA SCIENTIST 38(4) — $5.00 Solar Energy — Academy Symposium FLORIDA SCIENTIST 39(3) — $5.00 (includes do-it-yourself instructions) Anthropology — Academy Symposium FLORIDA SCIENTIST 43(3) — $7.50 Shark Biology — Academy Symposium FLoRIDA SCIENTIST 45(1) — $8.00 Future of the Indian River System — Academy Symposium FLoriDA SCIENTIST 46(3/4) — $15.00 Individual orders should be sent with payment. A statement will be sent in re- sponse to a bona fide purchase order over $10.00 from a recognized institution. Ad- dress all orders to: The Florida Academy of Sciences, Inc. c/o The Orlando Science Center 810 East Rollins Street Orlando, Florida 32803 Phone: (305) 896-7151 ISSN: 0098-4590 ‘Florida Scientist Volume 52 Spring 1989 Number 2 CONTENTS The Sand Pine Scrub Community: An Annotated Bibltography .......5...... PIMPS SETAE A a) WEIN UU Anica eagle Gh ied Donald R. Richardson 65 Calculated X-Ray Data Aid in Collecting High Quality X-Ray Powder Data— Oxytetracycline Dihydrate, C,,H,,N,O,e2H,O, 20 EDD Zymeeayol (5 A) aR see eee a a Frank N. Blanchard 94 Bostrichobranchus digonas: Confirmation of its Presence in MME MORO NICTACOS 21. 1 Sit 5 le Oeics slo ats | diem abi auaiaterele va ale eo Gerald E. Walsh 100 Community Waste to Energy System Technologies BE NG re a as el yk leer WW A) eal goats anal Alex E. S. Green 104 John B. Iverson and Cory R. Etchberger JUN) FE Sz Mm vl g - 193 ‘ cw 9 } QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES e. A FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1989 Editor: Dr. DEAN F. Martin _Co-Editor: Mrs. BARBARA B. 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EsTEVEZ Department of Physical Sciences Mote Marine Laboratory Seminole Community College 1600 City Island Park Sanford, Florida 32771 Sarasota, Florida 33577 Program Chairs: Dr. GEorcE M. Dooris Secretary: Dr. PATRICK J. GLEASON Dr. Patricia M. Doors 1131 North Parkway P.O. Box 2378 Lake Worth, Florida 33460 St. Leo, Florida 33574 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BaRrBarRa B. Martin, Co-Editor Volume 52 Spring 1989 Number 2 Biological Sciences THE SAND PINE SCRUB COMMUNITY: .. AN ANNOTATED BIBLIOGRAPHY CO ~ 3 id Se DONALD R. RICHARDSON 4 SD) Itin ¥ on f j, At f) Department of Biology, University of South Florida, Tampa, FL/33620 | i te PEL, 5 p pee. Vj ep, FESO Ee CO oy me, HE vA om Pe ee v Asstract: This annotated bibliography with 316 references represents an initial effort to compile the available material dealing, either primarily or peripherally, with the sand pine scrub community in Florida. The literature survey utilized a computer search of four data bases, a hand search of local libraries, references contained in pertinent articles, unpublished studies and research projects, and resource information from private, state, and federal agencies. References concerning taxonomic botany and faunal associates are for the most part not included. For the past decade, there has been a growing concern for the rapid disappearance of the sand pine scrub community in Florida. Many southern Florida counties (Palm Beach, Broward, and Dade) have lost as much as 90- 95% of this upland plant community to urban development. Even in the less populous counties of central Florida (Polk and Highlands), as much as 65 % of the land has been claimed recently for expansion of the citrus industry and real estate development, a result of several hard freezes and growing compe- tition from foreign markets. In Florida, about 300 vascular plant species (excluding weeds) have been collected from areas described as scrub vegetation. Estimates indicate that at least 10 % —may be as high as 40% (depending on your interpretation of the habitat) are endemic to the scrub. Many of these narrow endemics are re- stricted to the Lake Wales Ridge District in Polk and Highlands counties. Presently, the rapid demise of scrub habitat has prompted the Federal listing of many scrub endemics. In the past 3 years, 10 scrub species have been listed by the U.S. Fish and Wildlife Service Endangered Species Pro- gram and many more are under review. Recent interest has sparked a rapid increase in land acquisition for nature preserves around the state. Except for a few large holdings such as the Ocala National Forest (210,000 acres), Archbold Biological Station (4,300 acres), Jonathan Dickinson State Park (1,895 acres) and numerous smaller parcels scattered in many of the Florida State Parks, most scrub sites have been developed or are vulnerable to development. Of those that remain, several thousand acres in the panhandle have been purchased by the CARL program 66 FLORIDA SCIENTIST [Vol. 52 (Conservation and Recreation Land, Department of Natural Resources). The Nature Conservancy is actively pursuing the purchase of scrub tracts and many local governments are now requiring developers to preserve at least 25 % of each habitat type, including sand pine scrub. For animals, about 65-70 species of vertebrates regularly occur in the Florida scrub. Of these, 9 are considered obligate scrub endemics and occur nowhere else in the world. More recently, 56 species of ants have been ob- served in scrub vegetation with only 6 endemic to scrub. With the exception of a few examples in southern Alabama, sand pine scrub occurs primarily on Pleistocene shorelines of the eastern Atlantic Coastal Ridge, central Lake Wales Ridge, Mount Dora Ridge, Ocala Trail Ridge, western Brooksville Ridge, Gulf Coastal Lowlands and the northern Coastal Beach Dunes of the panhandle (Alt and Brooks 1965, White 1970, Davis 1943). However, numerous small islands of scrub occur throughout the peninsula. Scrub vegetation is often described as a xerophytic evergreen plant com- munity characterized by sand pine (Pinus clausa) as the dominant tree with rosemary (Ceratiola ericoides) or scrub oaks (Quercus geminata, Q. chapma- nii, QO. myrtifolia) comprising a major component of the understory. The herbaceous layer is often sparse with numerous open, sandy areas devoid of cover or blanketed by lichens. The scrub is a fire maintained association adapted for long infrequent (40-60 year intervals), intense crown fires. The presence of serotinous cones results in the regeneration of even-aged stands throughout most of central and southern Florida although in the panhandle stands with open cones and mixed age distributions tend to dominate. Fire history, edaphic conditions, and allelopathy are all thought to be significant factors in determining patterns of distribution of scrub, succession within scrub and the physiognomy of the community. References to scrub vegetation date back to the late 1600s. Many early explorers noted the plant communities they encountered on their journeys, but few detailed descriptions remain. The first detailed account of scrub vegetation was given by Dickinson (1699) where he described scrub as “sand hills with shrubby palmetto.” First attempts to order Florida’s upland pine communities were published by DeBrahm (1773) and Romans (1775). Al- though considerable data on southern Florida had been gathered by the close of the Indian Wars (1859), the publication by J. C. Ives (1856) was by far the best report of the time. This was one of the first works to recognize scrub as a major plant community. For the next 100 years, many descriptive studies and various names were used to depict the various types of sand pine scrub in Florida, e.g., sand hill (Dickinson 1699), low pine (DeBrahm 1773), pine barrens (Romans 1775), scrub (Ives 1856; Williams 1870; Austin 1976), sand pine (Harshberger 1914), sand pine scrub (Harper 1927; Davis 1943; Laessle 1958; Myers 1985), and sand scrub (Craighead 1971). Due to the recent demise of this upland community throughout Florida, considerable research is necessary to determine the management methods for No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY 67 preservation of the remaining fragmented scrubs. One purpose of this anno- tated bibliography is to provide convenient access to a summary of the litera- ture pertaining to scrub ecology. Hopefully, the summary will facilitate re- search that leads to productive management and preservation of this forest resource. A portion of the information provided in this paper (1978-1988) was gath- ered through reviewing a literature search of Biological Abstracts, Forestry Abstracts, Biosis, and Agriculture Index. The brief synopsis of each paper was excerpted from the abstract of the paper or written by the author after read- ing the original publication. Emphasis was placed on papers dealing with ecological, geological, and floristic components of the scrub and to a lesser degree on faunal associates. Taxonomic or systematic studies have been largely ignored for this publication due to the large number of plant species. Scrub vegetation is often described as a xerophytic evergreen plant commu- nity characterized by sand pine (Pinus clausa) as the dominant tree with rosemary (Ceratiola ericoides) or scrub oaks (Quercus geminata, Q. chapma- nii, Q. myrtifolia) comprising a major component of the understory. The herbaceous layer is often sparse with numerous open, sandy areas devoid of cover or blanketed by lichens. The scrub is a fire maintained association adapted for long infrequent (40-60 year intervals), intense crown fires. The presence of serotinous cones results in the regeneration of even-aged stands throughout most of central and southern Florida although in the panhandle stands with open cones and mixed age distributions tend to dominate. Fire history, edaphic conditions, and allelopathy are all thought to be significant factors in determining patterns of distribution of scrub, succession within scrub and the physiognomy of the community. References to scrub vegetation date back to the late 1600’s. Many early explorers noted the plant communities they encountered on their journeys, but few detailed descriptions remain. The first detailed account of scrub vegetation was given by Dickinson (1699) where he described scrub as “sand hills with shrubby palmetto”. First attempts to order Florida’s upland pine communities were published by DeBrahm (1773) and Romans (1775). Al- though considerable data on southern Florida had been gathered by the close of the Indian Wars (1859), the publication by J. C. Ives (1856) was by far the best report of the time. This was one of the first works to recognize scrub as a major plant community: For the next 100 years, many descriptive studies and various names were used to depict the various types of sand pine scrub in Florida, e.g., sand hill (Dickinson 1699), low pine (DeBrahm 1773), pine barrens (Romans 1775), scrub (Ives 1856; Williams 1870; Austin 1976), sand pine (Harshberger 1914), sand pine scrub (Harper 1927; Davis 1943; Laessle 1958; Myers 1985), and sand scrub (Craighead 1971). Due to the recent demise of this upland community throughout Florida, considerable research is necessary to determine the management methods for preservation of the remaining fragmented scrubs. One purpose of this anno- tated bibliography is to provide convenient access to a summary of the litera- 68 FLORIDA SCIENTIST [Vol. 52 ture pertaining to scrub ecology. Hopefully, the summary will facilitate re- search that leads to productive management and preservation of this forest resource. A portion of the information provided in this paper (1978-1988) was gath- ered through reviewing a literature search of Biological Abstracts, Forestry Abstracts, Biosis, and Agriculture Index. The brief synopsis of each paper was excerpted from the abstract of the paper or written by the same author after reading the original publication. Emphasis was placed on papers dealing with ecological, geological, and floristic components of the scrub and to a lesser degree on faunal associates. Taxonomic or systematic studies have been largely ignored for this publication due to the large number of plant species. ABRAHAMSON, W. G. 1984. Species responses to fire on Florida Lake Wales Ridge. Amer. J. Bot. 71(1):35-43. [The responses of native plant species to prescribed and natural burns were recorded for scrub, sandhill, scrubby flatwoods, flatwoods, swales, and seasonal pond communities. These frequently burned associations recover via sprouting in contrast to species in the sand pine scrub that are killed and recover via seedling. ] . 1984a. Fire: Smokey Bear is wrong. Bioscience. 34(3):179-180. [A brief condensation of the 1984 paper emphasizing that fire is a normal environmental feature in many Flor- ida ecosystems. | . 1984b. Post-fire recovery of Florida Lake Wales Ridge vegetation. Amer. J. Bot. 71(1):9-21. [Community response to fire of five vegetation types (sandhill, sand pine scrub, scrubby flatwoods, flatwoods, swales) showed that recovery occurred in less than 2 years for poorly drained sites and between 1-4 years for more xeric sites. Species composi- tion following fire did not change from preburn conditions. ] , A. F. Johnson, J. N. Layne, and B. Peroni. 1984. Vegetation of the Archbold Biologi- cal Station, Florida: an example of the southern Lake Wales Ridge. Florida Scient. 47(4):209-250. [A floristic analysis of the ABS is described. Species diversity, successional changes and the effects of disturbance in relation to each community is provided, as well as a vegetation map and a brief discussion of endemic ridge species. | ALT, D. ano M. K. Brooks. 1965. Age of Florida marine terraces. J. Geol. 73:406-411. [Topo- graphic features produced by prolonged stands of sea level are recognized at elevations of 215-250 and 90-100 feet. Stratigraphic evidence indicates that the 215-foot shore line was occupied during the Upper Miocene and the 90-foot shore line during the Pliocene. Briefer stands have been distinguishable at elevations of 45-55 and 70-80 feet. These shore lines can be interpreted as being either Pliocene or earlier Pleistocene interglacial. | ANDERSON, L. C. AND L. L. ALEXANDER. 1985. The vegetation of Dog Island, Florida. Florida Scient. 48(4):232-251. [Nine plant communities (including sand pine scrub) are presented in this study, in addition to an annotated checklist of the vascular flora. ] ANonyMovus. 1942. Birds in Florida. Florida Department of Agriculture. Pp. 130-131. [Briefly describes the habitat, size, food, and nest building attributes of the Florida scrub jay. | ANonymMovs. 1963. Sand pines do best on sandhills, USDA reports. Florida For. Reporter 6(2):4. [The results of a 10-year study in four separate blocks on the Chipola Experimental Forest showed that the growth of sand pine was virtually unaffected by site preparation, and exceeded that of both longleaf and slash pine. | Anonymous. 1964. A pine for hot sandhills? For. Farmer 23(8):12. [Research finding indicate that sand pine may be the best choice for use on sandhill soils. In tests against other southern pines, sand pine was much superior as far as cost, survival, response to fertiliza- tion, disease susceptibility, and reported insect enemies. | ANonyMovus. 1981. Southeast sandhills can be productive. Management alternatives for pines (slash, longleaf, and sand). USDA Forest Serv. 5 pp. [On cleared sandhill land in north Florida, only longleaf pine and Choctawhatchee sand pine have survived in sufficient numbers and grown at an acceptable rate. Choctawhatchee sand pine is the most promis- . ing as it will produce twice the volume of wood in 25 years as will longleaf pine on sandhill sites. Yields of at least 1 cord per acre per year from CSP plantations are pre- dicted. ] No. 2, 1989] RICH ARDSON— SAND PINE SCRUB COMMUNITY 69 AUFFENBERG, W. 1981. Florida environments and their herpetofaunas. Part I. Environmental characteristics. Florida Herp. 2:1-36. [Part I describes the most important plant commu- nities (including scrub) which harbor amphibians and reptiles of the state. | . 1982. Florida environments and their herpetofaunas. Part II. Changes in the environ- ment. Florida Herp. 3:1-25. [Part II discusses the fate of the Florida herpetofauna and how sea level changes have molded each community. | AND J. B. Iverson. 1979. Demography of Terrestrial turtles. Pp. 541-569. In: Har.ess, M. anv H. Mor tock (eds.), Turtles. John Wiley and Sons, New York. [Contains gopher tortoise densities for several plant communities (sand pine scrub, longleaf pine-turkey oak, and xeric hammock) in Florida. | Austin, D. F. 1976. Florida Scrub. Florida Nat. 49(4):2. [A descriptive account of the Florida sand pine scrub community with reference to important plant and animal species. Succes- sion and fire cycles are briefly examined. | . 1977. Vegetation of southeastern Florida, III: Yamato scrub. Florida Scient. 40:338- 345. (The history, soils, and vegetation of the Yamato region is discussed with emphasis as to recent changes of the original associations. ] , F. R. Postn, anv J. N. Burcu. 1986. Scrub species patterns on the Atlantic Coastal Ridge, Florida. J. Coastal Res. (3)4:491-498. [Seven major scrub ridges were studied and mapped for differences in species composition. Scrub community indicators decrease in frequency and richness to the east and west of the highest dune ridge system. | BaBer, D. W. Anp J. G. Morris. 1980. Florida scrub jays (Aphelocoma coerulescens) foraging from feral hogs. Auk 97(1):202. [Florida scrub jays on Merritt Island were observed forag- ing for ectoparasites on the backs of hogs in a scrub flatwood community. | Baker, J. B. AND R. M. BRENDEMUEHL. 1972. Soil phosphorus level adequate for growth of Ocala sand pine seedlings: A greenhouse evaluation. Soil Soc. Amer. Proc. 36(4):666-667. [The deficiency level of available soil phosphorous for sand pine seedlings grown in Lakeland sandhill soils is about 1 ppm extractable P and that levels less than 1 ppm showed a marked reduction in dry matter production after 10 months. | Baumer, W. E. 1973. A close look at sand pine. USDA Forest Serv., State and Priv. For., South. Area, Forest Management Bull. 6 pp. [This article highlights the information presented at the Sand Pine Symposium held in Panama City, Florida on December 5-7, 1972. | Baumer, W. A., K. A. Utz, AND R. M. Burns. 1977. Planting Guide and economic analysis for sand pine. USDA Forest Serv., State Priv. For., South. Area, Misc. Publ. 8 pp. [Planting sand pines in sandhill sites throughout northern Florida was found to be economically feasible with limited site preparation techniques. | BARNARD, E. L. 1979. Resin-soaked root disease of sand pine. U.S. Forest Serv. Pest Alert, Forest Bull. SA-FB. p. 12. [A root disease is killing sand pine throughout much of Florida for both the Ocala and Choctawhatchee races. The roots and root collar area of affected trees are soaked with resin resulting in crown thinning and discolored foliage. ] . 1980. Phytophthora root rot of sand pine. Florida Department of Agriculture Con- sumer Services, Division Plant Industry. Plant Pathol. Circ. 214. 3 pp. [The origin, dis- semination, detection and control of P. cinnamomi on sand pine roots is discussed. | , R. L. ANperson, J. T. ENGLISH, AND G. M. BLAKESLEE. 1982. Sand pine root disease survey: Florida-1980. U.S. Forest Serv., Forest Pest Management, Asheville, N.C. Field Office Report 82-1-30. 21 pp. [Sixty thousand sand pines in 200 natural and planted stands distributed in north and central Florida were examined for symptoms of root dis- ease. Overall root disease amounted to 4.5% of the trees examined. Incidence varied with stand type, stand age and host variety reaching a high of 42% for one Ocala sand pine plantation. Six known or suspected root disease fungi were isolated from roots of diseased trees. | , G. M. BuakesLeE, J. T. ENGLISH, AND S. W. Oak. 1985. Pathogenic fungi associated with sand pine root disease in Florida. Plant Dis. 69(3):196-199. [Eight known or sus- pected pathogenic fungi were isolated from roots of diseased sand pine from planted and natural stands in Florida. | , J. T. EnNcuisu, R. L. ANDERSON, AND G. M. BLAKESLEE. 1982. Fungi associated with root disease of sand pine (Pinus clausa) in Florida. Phytopathology. 72(7):979. [An evalua- tion of more than 200 sand pine stands showed that root disease from at least 6 known fungi causes substantial damage to both Ocala and Choctawhatchee sand pines in Flor- ida. | 70 FLORIDA SCIENTIST [Vol. 52 BarneETT, J. P. 1970. Germination inhibitors unimportant in dormancy of southern pine seeds. BARRY, U.S. Forest Res. Note SO-112, South. Forest Exp. Stn., New Orleans. 4 pp. [Bioassays of seed extracts and leachates revealed no evidence of a germination inhibitor in seeds of slash, shortleaf, spruce, or Ocala sand pine. | . 1970. Storage of sand pine seeds. USDA Forest Serv., Tree Plant. Notes. 21(4):11-12. [Temperature and moisture levels were tested to determine the storage capacity of Ocala and Choctawhatchee sand pine seeds. Viability of Ocala seeds stored for 5 years was unchanged when compared to initial collections. Sand pine stored well over a wide range of conditions, but moisture contents of 10% or less and subfreezing temperatures were recommended for long-term storage. Choctawhatchee seeds were more sensitive to stor- age conditions with reduced viability occurring after 1-3 years. ] AND B. F. McLemore. 1965. Cone and seed characteristics of sand pine. U.S. Forest Serv. Res. Pap. SO-19, South. Forest Exp. Stn., New Orleans. 13 pp. [Information was gathered on cones and seeds of both races of sand pine, with a view to developing recom- mendations for handling and storage. Seed extraction, specific gravity, yields, preger- rmination treatment, seed viability, and storage are discussed in this paper. | . 1966. Sand pine cones and seed. USDA Forest Serv., Tree Plant. Notes 76:15-16. [Cone and seed characteristics (cones per bushel, seeds per pound, requirements for seed extraction, degree of dormancy) of both races of sand pine was studied in the winter of 1962 in Florida. ] AND J. M. McGitvray. 1976 Storing sand pine seeds and cones. USDA Forest Serv., Tree Plant. Notes 27(2):10 and 13. [Viability of seeds from new and 1-year old Ocala sand pine cones remained unchanged after 10 years of storage at all combinations of moisture and at two different temperatures. Average germination rates for new Ocala sand pine was 97 and 72 percent, respectively. Viability of seed dropped severely after 6 years for both pines. ] J., J. Wonc, P. Kenney, L. BarBer, M. FLAKE, AND R. ExBLap. 1981. Deposition of pesticide drops on pine foliage from aerial application. J. Appl. Entomol. 92(3):224-232. [A few basic application methods of pesticides are summarized. Spray coverage is deter- mined for both slash pine and sand pine in the Withlacoochee State Seed Orchard. The results indicate a greater amount of spray drops in the upper crown than in the lower crown for both species, suggesting that tree size and dosage may be important consider- ations. | Barry, J. W., L. R. BARBER, P. A. KENNEY, AND N. OvERGAARD. 1984. Feasibility of aerial spraying of southern pine seed orchards. South. J. Appl. For. 8(3):127-131. [This test demonstrated the utility of using aircraft to apply water-base sprays over slash and sand pine orchards. The helicopter achieved higher quantities of spray to the crown where the majority of the cones are than the biplane. The higher spray drop density on slash pine suggests that more spray volume and possibly more active ingredients would be required to achieve compara- ble insect control on sand pine. ] BEAL, J. F. 1951. A study of the wilting percentages of sandy soils of the Ocala National Forest and their effect on the regeneration of sand pine. Masters thesis, Univ. Florida. Gaines- ville. 77 pp. [The purpose of this study was to determine the various soil types in sand pine forests, determine the permanent wilting percentages of these soils by direct methods, determine whether sand pine seedlings of different ages wilt simultaneously with other plants growing in the same soil, and to ascertain the maximum age at which sand pine seedlings show visible signs of wilting. ] BELCHER, E.. W. AND L. Miter. 1975. Influence of substrate moisture level on the germination of sweetgum and sand pine seed. Proc. Assoc. Offic. Seed Analysts 65:88-89. [Sand pine germination was significantly reduced at all moisture levels of 36 percent and greater. Optimum substrate moisture levels on cellulose paper for sand pine was about 10 percent and for sweetgum 45-75 percent. | Bengtson, G. W. 1976. Comparative response of 4 southern pine species to fertilization effects of P, NP, and NPKMgS applied at planting. Forest Sci. 22(4):487-494. [Seedlings of sand, slash, longleaf and loblolly pine planted on prepared sandhills in west-central Florida were treated immediately after planting with varied rates of fertilizer. Three years after fertilization, survival of sand pine was slightly lower (86%) than slash (88%) or loblolly (91%) pine. Species height was greatest for sand pine, followed by slash, loblolly, and longleaf pines. The data suggests that K deficiency may limit the growth potential of these four species and their capacity to respond to NP fertilizer. ] No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY GA BuamTiA, V. K. anp J. Kacan. 1970. A photochemical synthesis of 2’, 6’-dihydroxy-4’-methoxy and 2’, 4’-dihydroxy-6’-methoxy-chaclones. Chem. Sci. 37:1203-1204. [The isolation of this compound from Lindera was shown to be identical with those previously isolated from the fern Pityogramma and from Pinus clausa. | Bian, M. V. anp G. Davis. 1979. Sand pine: Why use as a Christmas tree? Amer. Christmas Tree J. 23(2):43-44. [When compared to shortleaf and loblolly pines as a possible Christ- mas tree species, sand pine has several desirable characteristics, such as the ability to maintain color, flexibility, and moisture within its needles. ] BLAKESLEE, G. M., S. W. Oak, AND S. M. Kratxa. 1980. Shoot dieback of planted sand pine caused by Fusarium moniliforme var. subglutinans. Plant Dis. 64(7):703-704. [Reports on the isolation of pitch canker disease from sand pine, describes the symptomatology of the naturally occurring infections, and establishes proof of pathogenicity. ] Bracu, V. 1976. Notes on the life history of Onthophagus aciculatulus (Scarabeidae: Coprini). Florida Entomol. 59(1):106. [Fourteen beetles were collected from beneath scat in the white sand scrub at Archbold Biological Station. ] Bray, M. W. ann J. S. Martin. 1942. Pulping Florida sand pine. South. Pulp Pap. J. 5(1):7-14. [Sand pine was found to be adaptable to the sulfate process for the production of strong kraft and bleachable pulps. | BRENDEMUEHL, R. M. 1968. Research progress in the use of fertilizers to increase pine growth on the Florida sandhills. Pp. 191-196. In: Forest fertilization, Theory and Practice, Sympo- sium on Forest Fertilization. Tennessee Valley Authority, National Fertilizer Development Center, Muscle Shoals, Alabama. [Slash, sand and longleaf pines were grown in green- house and under field conditions with various levels of fertilizer in order to maximize growth. Greenhouse data suggest that sand pine growth was increased appreciably when N was applied alone. A further response resulted when N was applied in combination with P, but all other elements showed no increase in growth. Placement of fertilizer, such as soil disking or banding, proved better than normal surface application methods. | . 1971. The phosphorus placement problem in forest fertilization. Pp. 43-50. In: YOUNGBERG, C. T., AND C. B. Davey (eds.), Tree Growth and Forest Soils. Proc. 3rd North Amer. Forest Soils Conference; held at North Carolina State University at Raleigh in August 1968. Oregon Stat Univ. Press, Corvallis, Oregon. [Ocala sand pine showed the greatest response (height at age 3) to application of superphosphate when applied by broadcasting in strips and disking or by banding. | . 1972. Fertilization review—sand pine (Pinus clausa Chapm., Vasey). South. Cooper- ative Serv. Bull. 158:48-52. [This paper gives a basic overview of the work in progress dealing with fertilization of sand pines in natural or pine plantations. | . 1974. Choctawhatchee rootstock recommended for sand pine seed orchards. USDA Forest Serv., Tree Plant. Notes 25(4):25-27. [Choctawhatchee sand pine rootstocks are preferable for sand pine seed orchards, but grafting has proved difficult. Growth stages of scion and rootstock when grafted appear to be the key to success. A success rate of 60-75 percent has been achieved for these grafting techniques. | . 1981. Options for management of sandhill forest land. South. J. Appl. For. 5(4):216- 222. [Thirty eight species of pine have been included in trial plantings on sandhill lands in northwest Florida for the past 50 years. Of these, longleaf pine (Pinus palustris) and Choctawhatchee sand pine (Pinus clausa var. immuginata) are recommended for sandhill reforestation. Choctawhatchee sand pine produced twice the volume of wood in 25 years as did longleaf pine when growing on similar sites and in stands of comparable density. ] AND L. Mize. 1978. Nursery practices for Choctawhatchee sand pine. USDA For. Serv., Tree Plant. Notes 29(1):8-11. [The paper provides the base-line information needed to establish a sand pine nursery in Florida. Important factors affecting overall success include: nursery site selection, soil management and fertilization methods, and handling and lifting of Choctawhatchee sand pines prior to planting. ] Britt, R. W. 1973. Management of natural stands of Choctawhatchee sand pine. Pp. 135-143. In: USDA Forest Service, Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE- 2. [This paper describes the important factors that have influenced the formation of stands of Choctawhatchee sand pine, discusses the characteristics of the species, and ex- plores some management dynamics and techniques used for natural stands. | Brooks, M. K. 1972. The geology of the Ocala National Forest. Pp. 81-92. In: SNEDAKER, S. C. AND E. A. Luco, (eds.), Ecology of the Ocala National Forest. USDA Forest Serv., South- ern Region, Atlanta, Ga. [Discusses the geologic formations that play an important role in ( FLORIDA SCIENTIST [Vol. 52 shaping the various plant communities of the forest, especially the sand pine scrub com- munity. | Bryan, O. C. ann R. Srouramire. 1940. Soils of Florida and their utilization. Department of Agriculture, State of Florida, No. 42. 36 pp. [Briefly describes the dominant vegetation type (rosemary and sand pine) associated with St. Lucie and Lakewood sands in Florida. | Burns, R. M. 1966. Tip moths in the Sandhills. For. Farmer 26(1):12-14. [The conclusions from a South-wide study comparing the growth of unprotected pines (Choctawhatchee sand pine, slash pine, loblolly pine, shortleaf pine) with that of pines chemically treated to control tip moth damage showed that the increased growth of some of the pines did not warrant the expense of repeated applications of chemical. Loblolly pine showed the great- est damage, while tip moth infestation had minimal effect on shortleaf and sand pine. Untreated Choctawhatchee sand pine was recommended for conversion of scrub oak- wiregrass sites in northwest Florida. ] . 1968. Sand pine: Florida’s much maligned pioneer. South. Lumberman 217(2704):170-181. [Reforestation of west Florida sandhills with both varieties of sand pine far exceeded comparative plantings of slash, longleaf, loblolly, Virginia, and shortleaf pines. Its rapid growth and survival make it a valuable species for pulpwood production in Florida. ] . 1968. Sand Pine: a tree for west Florida’s sandhills. J. For. 66:561-562. |The perform- ance of sand pine plantings or seeding on sandhill soils is discussed. Data showed that the Ocala race grew larger, but sand pines of the Choctawhatchee race had better survival and appeared best adapted to sandhill soils when tested over a 12 year period. Four other pines (shortleaf, Virginia, loblolly and longleaf) competed poorly. | . 1971. Enzyme activity as an index of growth superiority of Pinus clausa var. clausa on two soils. Ph.D. dissert., Univ. Florida, Gainesville. 129 pp. [Suggests a method for se- lecting superior Ocala sand pines by comparing levels of enzymes that catalyze growth processes. | . 1972. Choctawhatchee sand pine, good prospect for Georgia-Carolina sandhills. J. For. 70(12):741-742. [Choctawhatchee sand pine appears to be a good prospect for con- verting sandhill sites dominated by hardwoods to pine. In five years, it outgrew loblolly, slash, longleaf, and Ocala sand pines in the presence or absence of regrowth by hardwood sprouts. | . 1973. Sand Pine: distinguishing characteristics and distribution. Pp. 13-27. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [The morphological characteristics that differentiate sand pine from other southern pines that have been introduced into the sandhills of Florida is discussed. | . 1973. Comparative growth of planted pines in the sandhills of Florida, Georgia and South Carolina. Pp. 124-134. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [The performance of longleaf, slash, loblolly, shortleaf, and Ocala and Choctawhatchee sand pines were compared on several soils in the sandhills. Choctawhatchee sand pine appears best suited for planting on sandhill soils in Florida and Georgia. | . 1974. Survival of sand pine seedlings affected by time of lifting and bale-storage period. USDA Forest Serv., Tree Plant. Notes 26(1):8-11. [Ocala sand pine require a longer period of preconditioning than those of Choctawhatchee sand pine to attain a comparable state of dormancy. Planting should be scheduled so that seedlings are lifted in January or early February in quantities that can be planted in one week. Seedlings should not be stored more than a few days, however, bales may be stored up to 8 days if lifted in late January. Site preparation does increase survival of planted seedlings but should be done well in advance of planting to allow soil compaction. | . 1975. Tip moth control pays off. For. Farmer 34:13. [Choctawhatchee sand pine was far superior to loblolly pine and shortleaf pine when planted on sandhill soils. This was due to its superior growth rate and relative indifference to tip moth attack. Following 10 successive years of chemical treatment, unprotected sand pines averaged only 1.5 feet less in height when compared to protected trees of the same d.b.h. Further, growth of CSP was substantially unaffected by methods of site preparation. | . 1975. Sand pine: fifth-year survival and height on prepared and unprepared sandhill sites. USDA Forest Serv. Res. Note SE-217. 5 pp. [Survival of Chochtawhatchee sand pine was higher and trees were taller 5 years after planting on sandhill sites prepared by double-chopping than on adjacent wooded sites. Planting on unprepared sites also low- No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY 73 ered survival and reduced height growth of Ocala sand pine, but time of lifting and length of seedling storage affected their survival as well. ] . 1978. Evaluation of a Choctawhatchee sand pine plantation at age 35. USDA Forest Serv. Res. Pap. SE-183. 13 pp. [Detailed measurements (trees per acre, height, dbh, specific gravity, reproduction, pulpwood volume, extractives, and soil organic matter) were made for a 35 year old sand pine plantation on Elgin Air Force Base, Okaloosa County. Seed dispersal and establishment of sand pine into the adjacent scrub hardwoods was also reported. | AND R. M. BRENDEMUEHL. 1969. Yield of a Choctawhatchee sand pine plantation at age 28. USDA Forest Serv. Res. Note SE-103. 4 pp. [Choctawhatchee sand pine is well- suited to the infertile, droughty soils common to the sandhills of Florida. Since published yield data for plantation grown Choctawhatchee sand pine is not available, a search for an existing plantation resulted in the location of one 28 year old stand. Wood production was estimated at 32 cords per acre for this plantation. | AND R. M. BRENDEMUEHL. 1978. Sand pine: early responses to row thinning. USDA Forest Serv. Res. Note SE-262. 8 pp. [Row thinning of Choctawhatchee and Ocala sand pine stands provided a means of overcoming growth stagnation in crowded plantations in northwest Florida. Thinning stimulated diameter growth and reduced the time needed for sand pine to reach marketable size. | AND E. A. Hess. 1972. Site preparation and reforestation of droughty, acid stands. USDA Agric. Handb. 426. 61 pp. [A thorough study of the sandhills of the southeast with reference to conversion of scrub oaks and wiregrass areas to pine forests. Methods of site preparation (mechanical, chemical and fire) and their effects on survival of planted pines (Ocala and Choctawhatchee sand pine, longleaf, slash and loblolly) is discussed. | AND R. D. McReyno ps. 1975. Planting dates for longleaf, slash, and sand pine seeds in the southeastern sandhills: effect on temperature and rainfall. USDA Forest Serv. Res. Pap. SE-143. 17 pp. [This paper reports the seeding success of various pines in the sand- hills of northwest Florida. Seedling establishment was best when daily high temperatures averaged no more than 22°C and daily lows no less than 3°C during the 10 weeks after seeding. Seeding guidelines are provided for several panhandle counties. | CAMPBELL, M. W. AND S. P. CuristMAN. 1982. The herpetological components of Florida sand- hill and sand pine scrub associations. Pp. 163-171. In: Scott, N. J., (ed.), Herpetological Communities: A Symposium of the Society for the Study of Amphibians and Reptiles and the Herpetologist’s League, August 1977, U. S. Fish and Wildlife Service, Wildlife Re- search Report 13. [Investigations of the amphibian and reptile species in the sandhills and scrub revealed a diverse complex composed of a minority of xeric-adapted species com- bined with an array of wide-ranging and aquatic species that can be found in many Florida habitats. Sand pine scrub contained more species than any other vegetation type in Florida. | CampBELL, R. B. 1940. Outline of the geological history of peninsular Florida. Proc. Florida Acad. Sci. 4:87-105. [Presents data on the distribution of subsurface geological features in a north-south direction the length of the Florida peninsula. | CHELLMAN, C. W. 1969. New record of Acantholyda circumcincta (Hymenoptera: Pamphi- liidae) in Florida. Florida Entomol. 52(1):51. [The web-spinning sawfly was collected from a natural stand of sand pine in Okaloosa County. The infestations varied from light to heavy, and were scattered over a 90 thousand acre area. | CurisTENSEN, N. L. 1979. The xeric sandhills and savanna ecosystems of the southeastern Atlan- tic Coastal Plain. Veroff. Geobot. Inst. ETH Stiftung Rubel Zurich 68:246-262. [The sand pine scrub ecosystem is discussed with reference to succession, soils, and fire. Sandhill, savannas and flatwoods are also described. | . 1981. Fire regimes in southeastern ecosystems. Pp. 112-136. In: Mooney, M. A., T. M. BonnicusEn, N. L. CuHrIsTENSEN, J. E. Loran, AND W. A. REINERS. Fire Regimes and Ecosystems Properties. USDA Forest Serv. Gen. Tech. Rep. WO-26. [Briefly describes the sand pine scrub community and discusses fire frequency and flammability. Suggests that the nutrient poor soils of the scrub favor shrub growth over herbs which would diminish fire probability. | CurisTMAN, S. P. 1988. Endemism and Florida’s interior sand pine scrub. Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program, Project # GFC-84-101. (in press). [This report will locate and discuss the biology of the last remaining scrubs along the Lake Wales Ridge in Highlands and Polk counties, Florida. | 74 FLORIDA SCIENTIST [Vol. 52 , H. L. Kocuman, H. W. Campse i, C. R. Smiru anp W. S. Lippincott, Jr. 1979. Unpublished report. Successional changes in community structures: Amphibians and rep- tiles in Florida sand pine scrub. [Herpetofaunal community structure changes through time as Florida sand pine scrub matures. Total amphibian and reptile abundance de- creases with time since last major environmental perturbation. The herpetofauna of Flor- ida sand pine scrub is a function of the physical characteristics of the habitat rather than the specific plant association, and a mosaic of successional stages is necessary to support the entire scrub community. | CxiarK, A. AND M. A. Taras. 1969. Wood density surveys of the minor species of yellow pine in the eastern United States. Part II. Sand pine (Pinus clausa (Chapm.) Vasey). U.S. Forest Serv. Res. Pap. SE-51. 14 pp. [Wood specific gravity was determined for Ocala and Choctawhatchee sand pines throughout central and northwestern Florida. | CLEWELL, A. F. 1981. Natural setting and vegetation of the Florida panhandle, an account of the environments and plant communities on northern Florida west of the Suwannee River. Report prepared under Contract No. DACWO1-27-0-0104, U. S. Army Corps of Engi- neers, Mobile, Al. [A thorough overview of the sand pine scrub community with special reference to the panhandle of Florida. Includes a discussion on the soils, species richness, physiognomy, fire, mineral nutrients, maintenance, origin, and response to disturbance. ] AND J. E. PoppLeron. 1983. Sand pine restoration at a claimed phosphate mine, Florida. Proceedings of the Florida Institute of Phosphate Research Conference, “Recla- mation and the Phosphate Industry,’ Jan. 27, 1983, Clearwater, Florida. [Sand pine scrub communities can be restored on reclaimed phosphate lands using the mulching tech- nique. | Coun, E. anp D. T. Kapuan. 1983. Parasitic habits of Trophotylenchulus floridensis (Tylenchuli- dae) and its taxonomic relationship to Tylenchulus semipenetrans and allied species. J. Nematol. 15(4):514-523. [Parasitism by T. floridensis, a nematode, was found on the roots of sand pine from central Florida. It was previously an unrecorded host for sand pine in Florida. Other species such as slash pine, loblolly pine, red oak and post oak were also found to be infected at the locations surveyed. | , D. T. Kapian, AND R. P. Esser. 1984. Observations on the mode of parasitism and histopathology of Meloidodera floridensis and Verutus volvingentis (Heteroderidae). J. Nematol. 16(3):256-264. [M. floridensis, a sedentary nematode, induced formation of single giant cells in the stelar parenchyma tissue of sand pine roots. | Connery, C. B. 1984. Factors that influence plant species richness on habitat islands of sand pine scrub. Masters thesis, Univ. Central Florida, Orlando. 92 pp. [Evidence is presented which suggests that there is no relationship between sand pine species and area, isolation of scrub communities and species, soil types and species or fire and species. The data indicates that the equilibrium theory of island biography does not apply to the sand pine scrub community in Florida because most plant species reproduce vegetatively and rarely undergo extinction even following catastrophic fires. | Cooke, C. W. 1939. Scenery of Florida interpreted by a geologist. Florida Geol. Surv. Bull. 17:1- 118. [Outlines the geological history of Florida with reference to the various marine ter- races. Briefly discusses the coastal dune community but no general description of the sand pine scrub is provided. | . 1945. Geology of Florida. Florida Geol. Surv. Bull. 29:1-339. [This treatment of the geology of Florida includes the composition and structure of the Floridian Plateau, topog- raphy, stratigraphy, and a description of the various deposits of the paleocene, eocene, oligocene, miocene, pliocene, and pleistocene series. ] Cooper, R. W. 1951. Release of sand pine seed after a fire. J. For. 49(5):331-332. [Seedfall was studied in a 40 year old stand of sand pine that burned in February near the Lake Doerr area of the Ocala National Forest. Seeds began to fall less than 9 hours after the wildfire and at the end of 3 weeks about 1,092,000 seeds per acre had fallen. Seed dispersal was completed by the end of the third week following the fire. Seedlings germinated immedi- ately, but were killed by high surface temperatures and drought. | . 1951. Foresters finding how to reproduce seed-hoarding sand pine. For. Farmer 10(9):14. [A practical way of getting seed dispersal of sand pines without fire has been devised. Branches are cut from the crowns of harvested trees and scattered over the area so as to place the seeds near the soil layer. As summer soil temperatures approach 160°F, the seeds are released and natural regeneration becomes possible. Additional treatment may be necessary to prevent losses from predators. | No. 2, 1989] RICHARDSON—SAND PINE SCRUB COMMUNITY Wo . 1953. Prescribed burning to regenerate sand pine. USDA Forest Serv., Southeast. For. Exp. Stn., Res. Notes 22. 1 p. [ Various types of fires (backfire, headfire) were used on test plots in the Ocala National Forest to determine if fire can be controlled in the sand pine scrub and used as a management tool without loss of wood. ] . 1957. Silvical characteristics of sand pine (Pinus clausa Chapm. Vasey). USDA Forest Serv., Southeast. For. Exp. Stn., Pap. No. 82. 8 pp. [The life history, distribution, and associated vegetation of sand pine for Florida is discussed. ] , AND C. S. SCHOPMEYER, AND W. H. D. MacGrecor. 1959. Sand pine regeneration on the Ocala National Forest. USDA Forest Serv., Prod. Res. Rep. No. 30. 37 pp. [The serotinous nature of the cones of sand pine, limited soil moisture, high surface soil temper- atures and predators have restricted adequate regeneration of this species in Florida. Data showed that germination and survival is directly correlated with increased levels of shade. Mechanical alteration of scrub areas was also suggested for regeneration of natural stands. | Cox, J. A. 1981. Status and distribution of the Florida scrub jay. A Report to the Florida Game and Freshwater Fish Commission. 92 pp. [Scrub habitats occur in a variety of types throughout Florida. This paper discusses the various habitats utilized by scrub jays in 43 Florida counties. ] . 1984. Distribution, habitat, and social organization of the Florida scrub jay, with a discussion of the evolution of cooperative breeding in new world jays. Ph.D. dissert. Univ. of Florida. Gainesville. 271 pp. [Excellent survey of the distribution and habitat require- ments of the Florida scrub jay in Florida. Distribution of scrub is documented for 39 counties, with a brief description of the type of scrub. | CRAIGHEAD, F. C. 1971. The Trees of South Florida. Vol. 1, Univ. of Miami Press, Coral Gables. [Briefly describes the sand scrub of south Florida. | Davis, J. M. 1943. The natural features of sothern Florida. Accompanied by vegetation map of southern Florida. Florida State Geol. Surv. Bull. 25:1-311. [The sand pine scrub forests of south Florida are characterized for both inland and coastal scrubs. The transition from coastal dune vegetation into scrub oak and tropical hammocks is also presented. | DeBrauM, J. W. 1773. Report of the general survey in the southern district of North America and map of the general survey of east Florida. Reprint 1971. Univ. South Carolina Press, Columbia. [Three types of pinelands are described: low pine-wiregrass, middling pine- bunch grass, high pine-bunch grass. From his description of each pineland, it seems possi- ble that the low pine community is equivalent to scrub pine since he mentioned that the trees are not good for lumber and that high pines are what is referred to as slash pine. | DE LA PENA, A. C. 1985. Terpenoids from Conradina canescens (Labiatae) with possible allelo- pathic activity. Masters thesis, Louisiana State Univ. Baton Rouge. 64 pp. [This thesis briefly describes the ecological interaction of the Florida sand pine scrub and sandhill communities. Bioassay evidence is provided which indicates that secondary metabolites (monoterpenes, sesquiterpenes, and triterpenes) in the leaves of Conradina causes inhibi- tion of lettuce and native sandhill grasses. De VatL, W. B. 1944. A bark character for the identification of certain Florida pines. Proc. Florida Acad. Sci. 7(3):101-103. [Bark characters are presented in the form of a key to distinguish eight species of Florida pines. The key includes longleaf pine, swamp pine, shortleaf pine, loblolly pine, slash pine, pond pine, spruce pine and sand pine. ] Deyrup, M. 1986. Observations on Mantoida maya (Orthoptera: Mantidae). Florida Entomol. 69(2):434-435. [The little Yucatan Mantid was collected in dense sand pine scrub at Arch- bold Biological Station using Townes traps. Capture records (198 individuals over 3 years) indicate that the flight period is concentrated in July and August. | AND D. Man _ey. 1986. Sex-biased size variation in velvet ants (Hymenoptera: Mutilli- dae). Florida Entomol. 69(2):329-335. [The relative sizes of male and female Mutillidae were studied at the Archbold Biological Station in an area of sand pine scrub and adjacent firelanes. Females are larger than males. Sex-biased size variation may be associated with courtship behavior or host-seeking behavior. ] AND J. Tracer. 1986. Ants of the Archbold Biological Station, Highlands County, Florida (Hymenoptera: Formicidae). Florida Entomol. 69(1): 206-228. [The 102 species of ants known to occur on the ABS are listed with annotations on their habitat and microhabitat preferences and timing of nuptial flights. Several species are endemic to the xeric scrub of central Florida, which was an insular refugium during Pleistocene or Plio- cene flooding. About 20 species are exotic, of which 6 have invaded scrub habitats. ] 76 FLORIDA SCIENTIST [Vol. 52 , J. Tracer, AND N. Caruin. 1985. The Genus Odontomachus in the southeastern United States (Hymenoptera: Formicidae). Entomol. News. 96(5):188-195. [The ants Odontomachus brunneus, O. clarus, and O. ruginodis are reported from Florida. At the Archbold Biological Station, O. clarus was usually found in well drained sites of sandhill and sand pine scrub. | Dickinson, J. 1699. God’s Protecting Providence: Being The Narrative of a Journey from Port Royal in Jamaica to Philadelphia. Reprint 1945. ANpREws, E. W. anp C. M. ANDREWS (eds.). Yale University Press. New Haven. [Briefly describes the coastal scrub along the southeastern coast of Florida as sand hills covered with shrubby palmetto. ] Dixon, W. N. anv D. L. Ensmincer. 1982. Foliar damage of sand pine by Systena marginalis, a cypress leaf beetle. Florida Entomol. 65(2):289-290. [Feeding and mating by the cypress leaf beetle on sand pine needles was observed near Tennille, Florida. Symptoms were partial to entire reddening of the crowns. Surveys showed that all infected sand pine stands were adjacent to cypress ponds where similar damage to cypress had occurred. | Doren, R. F., D. R. RICHARDSON AND R. E. Roserts. 1987. Prescribed burning of the sand pine scrub community: Yamato scrub, a test case. Florida Scient. 50(3):182-192. [Sand pine scrub tends to burn only under extreme fire-weather conditions, thus exhibiting extreme (uncontrollable and unpredictable) fire behavior. A prescription for burning this fuel type was written from a fire spread model to predict fire behavior and then tested on two fires in the Yamato scrub. The results suggest that an effective and safe means exists to burn sand pine scrub. ] Dorman, K. W. 1976. The genetics and breeding of southern pines. USDA Forest Serv., Agric. Handb. No. 471. [The geographical distribution of the two races of sand pine is provided. Cones and seeds, root rot susceptibility, needles, oleoresin composition, growth rates, and wood specific gravity is also discussed for sand pine in Florida. | Duever, L. C. 1981. The Florida scrub: misunderstood habitat. ENFO 81:1-3. [Overview article on the ecology of the Florida scrub community. | EcoImpact, Inc. 1981. The feasibility of restoring xeric forest ecosystems on mined lands. Un- published report for International Minerals and Chemical Corporation, Bartow, Florida. 98 pp. [Three experimental study plots were established on varying mined soil types (sand tailings, overburden/tailings mix, and overburden) at the Kingsford Mine in Polk County to study the establishment of scrub vegetation on previously mined sites. The study indi- cates that managed establishment of xeric ecosystems appears feasible, and warrants fur- ther investigation. | EISENBACK, J. D., B. YANG, AND K. M. Hartman. 1985. Description of Meloidogyne pini n. sp., a root-knot nematode parasitic on sand pine (Pinus clausa), with additional notes on the morphology of Meloidogyne megatyla. J. Nematol. 17(2):206-219. [Root-knot nematode infestations of sand pine stands were reported on several occasions in southern Georgia. Symptoms were chlorotic, dwarfed, and tufted foliage and stunting of trees. This is the first report of a root-knot nematode attacking sand pine. | Eveuterius, L. N. 1979. A phytosociological study of Horn and Petit Bois Islands, Mississippi. Final report to coastal Field Research Laboratory, Southeastern Regional Office, National Park Service. 192 pp. [Ceratiola ericoides is a frequent component of the relict dunes on the central and north side of the islands. Bioassays to test the allelopathic effects of rose- mary were conducted. Results showed that toxins leached from the leaves did not inhibit the growth of lettuce seeds. | Eyre, F. M. 1980. Forest cover types of the United States and Canada. Soc. Amer. Foresters, Washington, D. C. [The geographic distribution and ecological relationships of sand pine in Florida are discussed. | FERNALD, R. T. 1988. Preservation and management of coastal xeric scrub communities of the Treasure Coast Region, Florida. Florida Game and Fresh Water Fish Commission, Non- game Wildlife Program. (in press). [This report is primarily a synthesis of published and unpublished research on scrub biology, an inventory of existing scrub tracts within the Treasure Coast Region and recommendations for the preservation and management of remaining scrubs. ] Fiscuer, N. H., N. TANRISEVER, A. DE LA PENA, AND G. B. WILLIAMSON. 1987. The chemistry and allelopathic mechanisms in the Florida scrub community. Proc. 1987 Plant Growth Regu- lator Society of America 14:192-208. [Discusses the phytotoxic properties of chemicals from Ceratiola and several mint species found in the sand pine scrub. ] , N. TANRISEVER, AND G. B. WiLLiAMson. 1988. Allelopathy in the Florida scrub com- No. 2, 1989] RICHARDSON—SAND PINE SCRUB COMMUNITY 77 munity as a model for natural herbicide actions. In: Curuer, H. (ed.), Natural Products: Potential in Agricuture. Amer. Chem. Soc. Symp. Series XXX (In press). [A review of the allelopathic properties of several shrub species from the scrub and how the delivery and activation mechanisms of phytotoxins may be useful for herbicide applications. ] FoweE ts, H. A. 1965. Silvics of Forest Trees of the United States. USDA Agric. Handb. No. 271, Washington, D. C. [Life history of sand pine with reference to soils and topography. | Frampton, L. J., K. R. Roeper, D. L. Rockwoop, anp C. A Ho.uis. 1982. Genetic variation in Choctawhatchee sand pine pollen shed and viability. South. J. Appl. For. 6(4):225-228. [Initiation and duration of viable pollen production differed significantly among clones of Choctawhatchee sand pine. Clonal differences also occurred for pollen germinability, pollen conductivity, and catkin moisture content, but these traits were influenced by the stage of pollen production. Viable pollen could be collected during any period of pollen shed. Information will aid tree breeders in establishing seed orchards. | AND D. L. Rockwoop. 1983. Genetic variation in traits important for energy utiliza- tion of sand pines and slash pines. Silv. Genetica 32(1-2):18-23. [There is sufficient genetic variation in sand and slash pines to warrant selection for increased productivity in planta- tions. Heritability of tree stem, branch and total biomass were moderate to high. | FRONCZEK, F. R., N. TANRISEVER, AND N. K. FiscHer. 1987. Structure of 2’, 4’-Dihydroxychalone. Acta Crystallogr. C43:158-160. [The flavonoid (2’, 4’-dihydroxychalone) was isolated from Florida rosemary (Ceratiola ericoides), a dominant shrub in many Florida scrubs. | Garren, K. M. 1943. Effects of fire on vegetation of the southeastern United States. Bot. Rev. 9:617-654. [The role of fire in the origin and perpetuation of scrub and its effect on species composition is summarized. Genys, J. B. 1971. Hybrids of Virginia pine x sand pine, Pinus virginiana x clausa, in Maryland. Chesapeake Sci. 12(3):188-191. [A cross of Virginia pine with sand pine was produced in 1953. Hybrid trees were significantly larger, more uniform in height, produced larger pollen and a high seed set. | Givens, K. T. 1980. Succession in the vegetation of the Lake Wales Ridge: An analysis of observed changes in five plant communities at the Archbold Biological Station over a ten-year period. Honors Thesis, Bucknell University, Lewisburg. [A successional study of five sand ridge plant communities (sand pine scrub, scrubby flatwoods, low flatwoods, bayheads, and slash pine/turkey oak) which had been kept free of fire for almost 50 years was conducted in natural examples of these habitats along the Lake Wales Ridge. ] , J. N. Layne, W. G. ABRAHAMSON, AND S. C. WHITE-SCHULER. 1984. Structural changes and successional relationships of five Florida Lake Wales Ridge Plant Communi- ties. Bull. Torrey Bot. Club. 111(1):8-18. [The most striking result of this study is the relative lack of change in species composition for the five communities. A change in canopy and species richness was greatest for the southern ridge sandhill association, fol- lowed by the sand pine scrub and scrubby flatwoods. | Gopparb, R. E. AND R. K. StRicKLAND. 1972. Genetics of sand pine and the program for superior tree selection. Pp. 226-235. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [Choctawhatchee sand pine has better tree form, a higher survival rate, less susceptibility to mushroom root-rot, and several other features that are desirable for outplanting in sandhills. Numerous selections of both varieties are now avail- able for tree improvement programs. | AND D. L. Rockwoop. 1981. Cooperative forest genetics research program. Twenty- third progress report. Research report, Univ. Florida, School of Forest Resources and Conservation. 17 pp. [Briefly discusses the formation of a long-term program for the improvement of Choctawhatchee sand pine via control-pollination. The use of sand pine progenies which may be usefull in the Christmas tree market are mentioned. ] , D. L. Rockwoop snp H. Kox. 1982. Cooperative forest genetics research program. Twenty-fourth progress report. Research Report No. 33, Univ. Florida, School of Forest Resources and Conservation. 23 pp. [Pollen shed and viability were determined for Choc- tawhatchee sand pine in north Florida. Conductivity as a measure of pollen viability was also tested. | Grecor, H. J. 1982. Pinus aurimontana—A new species of Pinus from the Neogene of the Goldberg (Ries). Stuttgarter Beitr. Naturk., Ser. B (Geol. Palaeontol.) 83:1-19. [Relates this new fossil pine with sand pine which seems rather tenuous. ] Har.ow, R. F., B. A. Sanpers, J. B. WHELAN AND L. C. Cuapre.. 1980. Deer habitat on the Ocala National Forest: Improvement through forest management. South. J. Appl. For. 78 FLORIDA SCIENTIST [Vol. 52 4(2):98-102. [The longleaf pine-turkey oak and sand pine scrub communities are de- scribed for the Ocala National Forest. Acorn production and available forage was assessed for each community type. Deer food was most abundant in young stands and least abun- dant in 25-40 year old stands. Availability of deer food increased with canopy thinning. | Harms, W. R. 1969. Sand pine in the Georgia Carolina sandhills: Third year performance. U.S. Forest Serv. Res. Note SE-123. 3 pp. [Survival and height of Choctawhatchee and Ocala sand pine, longleaf pine, slash pine, and loblolly pine were compared. The Choctawhat- chee race survived better than the Ocala race on two different soils. Both races of sand pine grew to greater heights than the other pine species. | Harper, R. M. 1914. Geography and vegetation of northern Florida. Ann. Rep. Florida State Geol. Surv. No. 6:163-391. [This report discusses the more striking natural features of several vegetation types (including sand pine scrub) of 20 different regions of Florida. | . 1915. Vegetation types: Natural resources in an area in central Florida. Ann. Rep. Florida State Geol. Surv. No. 7:135-188. [The sand pine scrub community is discussed in reference to soils and fire. A brief checklist of the scrub of Marion and Citrus counties is given. | . 1921. Geography of central Florida. Ann. Rep. Florida State Geol. Surv. No. 13:71- 301. [The characteristic species of the sand pine scrub community and the relationship of fire to their survival is discussed for central Florida. | . 1927. Natural resources of southern Florida. Ann. Rep. Florida State Geol. Surv. No. 18:27-206. [This paper describes the scrub and the transition between scrub and high pine lands in south Florida. A brief checklist is also provided. ] . 1940. Fire and forests. Amer. Bot. 46:5-7. [Indicates that fires burn only once in a lifetime for the sand pine scrub of Florida. ] . 1955. Historical notes on the relation of fires to forests. Proc. Tall Timbers Fire Ecol. Conf. 9:11-29. [The sand pine scrub is depicted as a fire sensitive community that proba- bly burns only once in a lifetime. He describes the Florida scrub as an area of bare sand with no grass to carry a fire. ] HARSHBERGER, J. W. 1914. The vegetation of southern Florida south of 27 30’ N, exclusive of the Florida keys. Trans. Wagner Inst. Sci. Philadelphia 3:51-189. [The sand pine scrub com- munity is discussed for south Florida, with emphasis on the distribution of rosemary. | Hartnett, D. and D. R. RicHarpson. 1988. Life history and population biology of Bonamia grandiflora in central Florida. Am. J. Bot. In press. [The role of fire and community maturity on several populations of Bonamia was investigated for central Florida. ] Hess, E. A. 1972. Resistance to ice damage—a consideration in reforestation. USDA Forest Serv., Tree Plant. Notes 22(2):24-25. [The effect of ice damage was monitored for several trees (Ocala sand pine, Choctawhatchee sand pine, slash pine, longleaf pine, loblolly pine) within the Sandhills State Forest, South Carolina. Ocala sand pine showed the most damage (37 % ) and loblolly the least (13%), immediately following the storm. Because of its prominence in sandhill plantings, any damage to slash pine is of considerable signifi- cance. The resistance of loblolly to ice damage suggests it as a substitute, although it grows only in better soils. Although no more resistant than slash, Choctawhatchee sand pine might also be considered because it is well suited to poor soils. | . 1981. Choctawhatchee sand pine growth on a chemically prepared site—10 year results. South. J. Appl. For. 5(4): 208-211. [Site preparation using pelleted herbicides (fenuron) caused a three fold increase in sand pine volume per acre on treated vs un- treated plots. Tree height was also greater on treated plots. ] . 1982. Sand pine performs well in the Georgia-Carolina sandhills. South. J. Appl. For. 6(3):144-147. [On the Georgia and South Carolina sandhill, planted Choctawhatchee sand pine grew better than loblolly, longleaf and slash. Heights at age 15 averaged 35.4 feet for the Choctawhatchee variety, 32.8 for the Ocala variety and 22.6, 23.8 and 24.0 feet for loblolly, longleaf and slash pine, respectively. Ocala sand pine had good height but poor initial survival. Comparison with Florida stands of similar age shows the Choc- tawhatchee variety growing as well in Georgia and South Carolina. ] . 1982. Cold storage of Choctawhatchee sand pine seedlings: Effect of lifting date and length of storage. South. J. Appl. For. 6(4):229-231. [Cold storage was found to keep Choctawhatchee sand pine seedlings in good condition for planting for up to 12 weeks. Survival was lower for seedlings stored longer. Lifting seedlings before dormancy reduced overall survival. Differences in survival could not be correlated with seedling size, mor- phology, or nutrient content. ] No. 2, 1989] RICH ARDSON—SAND PINE SCRUB COMMUNITY 79 AND R. M. Burns. 1973. Methods and goals in preparing sand pine sites. Pp. 82-92 In: USDA Forest Serv., Sand Pine Symp. Proc. USDA For. Serv. Gen. Tech. Rep. SE-2. {Mechanical site preparation of sandhills using chopping and strip site methods are dis- cussed. Choctawhatchee sand pine appears best suited for planting on partially prepared sandhill sites. | HENDERSON, J. R. 1939. The soils of Florida. Univ. of Florida, Agric. Exp. Stn. Bull. 334, Gaines- ville, Florida. 67 pp. [The dry sand soils of the Lakewood, St. Lucie, and Dade series that support scrub vegetation are discussed. | Henprickson, W. M. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the Environment Symp. Proc., May 1-5, 1972, Denver, Colorado, USDA Forest Serv. [Spe- cies of pines with serotinous cones and a possible explanation for its occurrence is men- tioned. | Hernpon, A. 1985. Fire in natural communities. Palmetto 5(3):4-5. [Briefly explains that fire is necessary for the continued existence of scrub and that endemics require openings created by fire. ] . 1986. Native plants for fire protection. Palmetto 5(4):6-7. [Suggests that homes should not be built in the sand pine scrub community because fires are usually uncontrol- lable. Fires are described as too hot to attack, can jump great distances in the canopy, and the rate of spread is rapid. | Hopces, J. D. AND R. L. ScHEER. 1962. Soil cover aids germination of pine seed on sandy sites. USDA Forest Serv., Tree Plant. Notes 54:1-3. [In the sandhills of west Florida, pine seeds covered with a thin layer of soil germinate better than those lying on the surface. Direct- seeding tests indicate that sowing depths up to three-fourths of an inch are best for slash and sand pine. For longleaf pine, depths should not be greater than one-half inch. ] Houeu, W. A. 1973. Fuel and weather influence wildfires in sand pine forests. U.S. Forest Serv. Res. Pap. SE-106. 11 pp. [Seasonal trends in fuel characteristics indicate that crown fires in sand pine have the highest probability of occurring in late February or early March when needle moisture content is low and ether extractive content is highest. Extended drought conditions, frontal movements, and low-level airmass instability were suggested as indicators of potential development of large fires and erratic fire behavior. | Hv, S. C. ann P. Y. Burns. 1979. Testing sand pine for Christmas tree production in Louisiana. LSU For. Notes. 2 pp. [The results indicate that sand pines can be succcessfully grown and marketed as Christmas trees within five years in southeastern Louisiana. Survival of hand-planted seedlings in plantations lacking site preparation was poor, but improved with mowing and shearing. | Husse., T. M. 1985. Unfinished business and beckoning problems. Florida Entomol. 68(1):1- 10. [A brief description of the Florida scrub in the 1920s suggests that the boundaries between the scrub and other communities is often sharp with much exposed sand. An account of several endemic insects that occupy central and north Florida scrubs is pro- vided. | Ives, J. C. 1856. Memoirs to Accompany a Military Map of Florida. Wynkoop Co., New York. [Several Florida plant communities are discussed: swamps, prairies, marshes, pinelands, hammocks, and scrub. This is probably the first work to recognize scrub as a major plant community. | Jackson, J. F. 1973. Distribution and population phenetics of the Florida scrub lizard, Scelo- porus woodi. Copeia. 1973:746-761. [The relationship between the distribution of sand pine scrub and the scrub lizard was found only on the Lake Wales Ridge and northern Bartow Ridges, in the Ocala National Forest region, along the central and southern Atlan- tic coast, and on the southwestern Gulf coast. Habitat restriction was related to poor dispersal by the scrub lizard. | AND S. R. TeLForp, Jr. 1974. Reproductive ecology of the Florida scrub lizard, Scelo- porus woodi. Copeia. 1974:689-694. [The data collected from populations in the Ocala National Forest indicate that the Florida scrub lizard is an early-maturing, multiple- brooded, small clutch species that may occur at densities of 10.0 lizards/hectare or greater in optimal habitat. | James, C. W. 1961. Endemism in Florida. Brittonia. 13:225-244. [Distribution of a few scrub endemics and their survival in Florida is discussed. | JOHANSEN, R. W. anv R. W. Cooper. 1965. Aerial attacks on sand pine crown fires. South Lumberman 211(2632):105-106. [A water drop of 250 gallons on the head of an advanc- ing sand pine crown fire was found to be a successful method for knocking the blaze to the 80 FLORIDA SCIENTIST [Vol. 52 ground where conventional fire fighting methods can be used. Ammonium phosphate formulations were also effective in stopping advancing fires even when sprayed 24 hours prior to contact. | Jounson, A. F. 1982. Some demographic characteristics of the Florida rosemary Ceratiola eri- coides Michx. Amer. Midl. Naturalist. 108:170-174. [Seed productivity and survivorship patterns were determined for rosemary in different aged stands. Seed germination oc- curred 10 years after a fire with virtually no seed store in the soil under nonreproductive shrubs. Self-thinning occurred 10-20 years after a fire, when densities decreased from 5 to 0.5 shrubs per meter. ] . 1986. Recipe for growing Florida rosemary, main ingredient: patience! Palmetto. 6:5. [Briefly describes preferred habitat and methodology for growing rosemary from seed in a greenhouse. | AND W. G. ABRAHAMSON. 1982. Quercus inopina: A species to be recognized from south-central Florida. Bull. Torrey Bot. Club. 109(3):392-395. [Morphological traits sug- gest that Q. inopina, a distinct species endemic to the central ridge scrub, is different from QO. myrtifolia. | , W. G. ABRAHAMSON, AND K. D. McCrea. 1986. Comparison of biomass recovery after fire of a seeder (Ceratiola ericoides) and a sprouter (Quercus inopina) species from south-central Florida. Amer. Midl. Naturalist. 116(2):423-428. [The results of this study suggest that the seeder (rosemary) and sprouter (oak) are indeed occupying the same niche in their respective habitats. After ca. 30 years postfire the biomass of the rosemary stands (1400 g/m) nearly equalled the average biomass of the oak stands (1500 gm). | Joye, N. M. Jr., A. T. PRoveaux, AND R. V. Lawrence. 1972. Composition of pine needle oil. J. Chromatogr. Sci. 10(9):590-592. [Several volatile compounds were identified from pine needles of sand pine and four other southern pines. | Kars, A. G. AND G. A. SNow. 1972. Host response of pines to various isolates of Cronartium quercuum and Cronartium fusiforme. U.S. Dep. Agric. Misc. Publ. 1221:495-503. [Sand pine was found to be moderately susceptible to the North Carolina isolates (C. quercuum) but extremely resistant to the Mississippi and Wisconsin isolates, as well as inoculations of C. fusiforme. | Kauisz, P. J. 1982. The longleaf pine islands of the Ocala National Forest, Florida: A soil study. Ph.D. dissert., Univ. Florida, Gainesville. 143 pp. [Concluded that the separation of the sand pine scrub and the sandhill communities was not correlated with any differences in soil physical or chemical properties within the upper 5m depth. Evidence is presented to support the hypothesis that the longleaf pine islands in the Ocala National Forest may be the result of burning by early man and may therefore be mere human artifacts. | AND E. L. Stone. 1984. Soil mixing by scarab beetles Peltotrupes youngi and pocket gophers Geomys pinetus in north central Florida. Soil Sci. Soc. Amer. J. 48(1):169-172. [Scarab beetles are an important mixing agent that burrows to 360 cm or deeper and can transport as much as 8 metric tons of subsoil to the surface per hectare per year. Its association with other burrowing animals such as the pocket gopher, influences the course of soil genesis and hence profile morphology. Populations of both species are much re- duced or absent from sand pine scrub compared to longleaf pine areas. | AND E. L. Stone. 1984. The longleaf pine islands of the Ocala National Forest, Flor- ida: a soil study. Ecology 65(6):1743-1754. [Longleaf pine-wiregrass-turkey oak occurs as isolated islands in a matrix of sand pine scrub on deep sands of the Ocala National Forest. Striking contrasts in physiognomy and species composition, and sharp, stable boundaries suggested that soil differences determined vegetation boundaries. Examination of soils at 131 locations revealed no consistent differences in profile morphology, particle size distri- bution, or extractable nutrients. Thus the hypothesis that soil differences were responsible for the distribution of the two communities was not sustained. ] Kem, M. H. anv I. J. Stour. 1987. Longevity record for the Florida Mouse, Peromyscus flori- danus. Florida Scient. 50(1):41. [A captive Florida mouse lived for a period of 7 years, 4 months and 2 days; the longest on record for this species. The mouse was captured in the sand pine scrub from the University of Central Florida campus at an estimated age of 3 weeks. | Kine, J. A. 1968. Biology of Podoyms (Rodentia). Special Publication No. 2, Amer. Soc. Mam- mologists. [This book is a comprehensive treatise on the biology of the genus Peromyscus in North America with various references to the Florida mouse, a scrub endemic. | Komarek, E. V. 1974. Effects of fire on temperate forests and related ecosystems: Southeastern No. 2, 1989] RICHARDSON — SAND PINE SCRUB COMMUNITY 81 United States. Pp. 251-277. In: AHLGREN, C. E. anv T. Koz.owski (eds.), Fires and Ecosystems. Academic Press, New York. [Briefly describes the distribution of sand pine and other common associates in Florida. Explains that sand pines have serotinous cones and that severe fires are required for regeneration. However, if man-caused or lightning fires occur in the wrong season or if reburned too quickly, the forest would ultimately come back very slowly. | Kossutu, S. V. AND R. H. Biccs. 1981. Role of apophysis and outer scale tissue in pine cone opening. Forest Sci. 27(4):828-836. [Removing the apophysis and outer part of the cone scale of sand pine reduced the time for the cones to begin opening, to complete opening, and shed seed. | AND E. L. Barnarb. 1983. Monoterpene content of healthy sand pine and sand pine with root disease. Forest Sci. 29(4):791-797. [The amount and composition of monoter- penes in branches, scions, rootstocks and roots were made from healthy and diseased Choctawhatchee and Ocala sand pine in natural stands. The quantity of monoterpenes was higher in roots and rootstocks of diseased grafts than of healthy grafts. Composition in healthy and diseased grafts was similar for sand/sand grafts and sand/slash grafts. In the stem xylem samples from natural stands, no monoterpene differences were detected be- tween healthy and diseased Ocala sand pine. | Kurz, H. 1942. Florida dunes and scrub, vegetation and geology. Florida Geol. Surv. Geol. Bull. 23:1-154. [Discusses the distribution of scrub and its relationship to the high pine land, water and chemical relations, geological aspects and succession. | LagssLe, A. M. 1942. The plant communities of the Welaka Area. Univ. Florida Publ., Biol. Sci. Series 4(1):95-109. [Description of the sand pine scrub community at Welaka, with refer- ence to soils, fire, and succession. | . 1958. A report on successional studies of selected plant communities on the University of Florida Conservation Reserve, Welaka, Florida. Quart. J. Florida Acad. Sci. 21:102- 112. [Documents the invasion of sand pine into longleaf pine/turkey oak sandhills. ] . 1958. The origin and successional relationship of sandhill vegetation and sand pine scrub. Ecol. Monogr. 28:361-387. [Good discussion of the presence of scrub on old shore- lines, characteristics of the soils, successional relationship of sandhill and scrub, and a brief description of several scrubs throughout Florida. ] . 1965. Spacing and competition in natural stands of sand pine. Ecology. 46(1 and 2):65-72. [Stands under 23 years old were found to be aggregated or randomly distrib- uted. Most stands 23 or more years old showed regular spacing. In spite of the large number of variables that effect spacing in competing sand pine stands, the comparison of live-and-dead with live-only spacing shows that such competition results in a more even overall pattern of surviving trees. | . 1967. Relationship of sand pine scrub to former shorelines. Quart. J. Florida Acad. Sci. 30(4):269-286. [Lack of soil differences between scrub and sandhill does not account for their sharp boundaries. The role of fire and the similarity of scrubs throughout the state and their occurrence on old shorelines is discussed. ] LatuaM, P. J. 1985. Structural comparisons of sand pine scrubs of east-central Florida. Masters thesis, Univ. of Central Florida, Orlando. [This study provides a quantitative analysis of selected sand pine scrubs of east-central Florida with emphasis on structural comparisons within tree, shrub, and ground level plant layers. Stands of sand pine scrub characterized by high similarity in plant species occurred on St. Lucie fine sand, Pomello sand, and Paola sand. Tree layers were dominated by sand pine, shrub layers of myrtle oak and ground cover by a mixture of 16 woody andd 8 herbaceous species. ] Layne, J. N. 1963. A study of the parasites of the Florida mouse Peromyscus floridanus, in relation to host and environmental factors. Tulane Stud. Zool. 11:1-27. [Thirty-two spe- cies of parasites were recorded from populations of the Florida mouse inhabiting various habitats: sand pine scrub, longleaf pine/turkey oak, slash pine/turkey oak, upland ham- mock, and pine flatwoods. A brief description of each vegetation type is provided. It appears that parasitism has little direct role in the regulation of Florida mouse popula- tions in north Florida. ] . 1966. Postnatal development and growth of Peromyscus floridanus. Growth. 30:23- 45. [Data are given on morphological and behavioral development of the young, maternal behavior, and related aspects of the reproductive biology of a population of the Florida mouse occurring in xeric pine-oak woodlands and scrub habitats in north-central Flor- ida. | 82 FLORIDA SCIENTIST [Vol. 52 . 1970. Climbing behavior of Peromyscus floridanus and Peromyscus gossypinus. J. Mammol. 51(3):580-591. [Field observations show that the cotton mouse is a much better climber than the Florida mouse. Layne suggests that psychological differences in the mice are probably more important than structural features in accounting for their different climbing tendencies. ] . 1978. Peromyscus floridanus. Pp. 21-22. In: Layne, J. N. (ed.), Rare and Endan- gered Biota of Florida, Vol 1. University Presses of Florida, Gainesville. Pp. 21-22. [A brief description, range, habitat, life history, and a basis for status classification in Florida is discussed for this species. | . 1979. Natural features of the Lake Annie tract, Highlands County, Florida. Archbold Biological Station, Lake Placid, Florida. [Lake Annie is the southernmost lake along the southern axis of the peninsula with an interesting geological and ecological history. The major vegetation type associated with the lake is the sand pine scrub that is particularly rich in both endemic plants and animals. | Lee, D. S. 1969. Flying squirrels feeding on the cones of the sand pine. Florida Natural. 42(1):41. [Flying squirrels in Lake county were supplementing their diet by eating sand pine seeds from the nearby sand pine-rosemary scrub. | Litt.e, E. L., Jk. AND K. W. Dorman. 1952. Geographic differences in cone-opening in sand pine. J. For. 50:204-205. [Cone opening is reviewed for the Ocala and the Choctawhat- chee races of sand pine. It has been suggested that the open cone race of sand pine in western Florida may have developed in response to surface fires (increased fuel loads) while the closed cone race has developed to conditions favoring less frequent but more destructive fires. | Lourer, F. E. 1980. Eastern coachwhip predation on nestling blue jays. Florida Field Naturalist. 8(1):28-29. [Briefly records the predation of nestling blue jays in sand pine scrub by an eastern coachwhip snake. | . 1985. Ontogeny of thermoregulation in the eastern screech owl. J. Field Ornithol. 56(1):65-66. [The results of a field study of thermoregulation in nestling Eastern Screech- owls (Otus asio) in sand pine scrub at Archbold Biological Station is presented. | Lone, R. W. 1974. The vegetation of southern Florida. Florida Scient. 37(1):33-45. [Indicator species of the scrub community in southern Florida is presented. | Luco, A. E. anv C. P. Zucca. 1983. Comparison of litter fall and turnover in 2 Florida ecosys- tems. Florida Scient. 46(2):101-110. [Litter-fall rates in the sand pine and sandhill were low (330 and 278 g/m2/yr, respectively), litter standing crop was high (averaging 1367 and 842 g/m, respectively) and litter turnover rate was low (0.21 and 0.33 times/yr, respec- tively). Patterns of litter fall were different for the two forests, probably reflecting differ- ent responses to fire frequency and nutrient availability. | Luzzi, N. A. AND A. C. Tarjan. 1982. Vector and transmission studies on the pinewood nematode Bursaphelenchus xylophilus in Florida. J. Nematol. 14(4):454. [Eighteen month old seed- lings of sand pine were inoculated with the pinewood nematode. All seedlings died within 40 days. | MacNEIL, F. S. 1949. Pleistocene shore lines in Florida and Georgia. U. S. Geol. Surv. Prof. Pap. 221-F:95-107. [This paper determines what terraces and shore lines existed in Florida and the coastal plain of Georgia in order to discover any possible relationship between the phosphate beds and pleistocene terraces. | Martin, J. S. 1962. Kraft pulping of west Florida sand pine and longleaf pine. USDA Forest Serv. For. Prod. Rep. 2248, Madison, Wis., U.S. Forest Products Laboratory. 12 pp. [Kraft pulps from the open-cone form of sand pine grown in western Florida were higher in overall strength and brightness than those made from longleaf pine of equal growth rate grown in the same area. | Marx, D. H., W. C. Bryan ann C. E. CorpnE.u. 1976. Growth and ectomycorrhizal development of pine seedlings in nursery soils infested with the fungal symbiont Pisolithus tinctorius. Forest Sci. 22(1):91-100. [In the Florida nursery, Pisolithus did not dominate sand pine seedling roots or stimulate seedling growth as it did in the North Carolina nursery. Lateral movement of the inoculum in the soil was 120 cm from its original introduction into nursery beds in Florida and North Carolina. | , W. C. Bryan AND C. E. Corpe.u. 1977. Survival and growth of pine seedlings with Pisolithus ectomycorrhizae after 2 years on reforestation sites in North Carolina and Flor- ida. Forest Sci. 23(3):363-373. [Two-year field data indicated that ectomycorrhizae formed by Pisolithus increased survival and growth of the Choctawhatchee sand pine in addition to four other southern pine species. ] No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY 83 , C. E. Cornet, D. S. KENNeEy, J. G. MEXAL, J. D. ARTMAN, J. W. RIFFLE, AND R. J. Mo ina. 1984. Commercial vegetative inoculum of Pisolithus tinctorius for development of ectomycorrhizae on bare-root tree seedlings. Forest Sci. Monogr. 25:1-101. [Choc- tawhatchee sand pine seeds treated with Pisolithus inoculum were planted at the Andrews Nursery. Results indicate that the Pisolithus inoculum significantly increased seedling growth compared to the controls. Results were consistent with other pines from other locations. | McConneLL, W. V. 1984. Options in energy wood farming. South. J. Appl. For. 8(3):149-152. [A U.S. Air Force study identified 5 potential management options for woody biomass pro- duction in Florida. The prefered option was short rotation of Choctawhatchee sand pine on lakeland soils. It was suggested that this low energy option may have application on much of the 8 million acres of sandhill in the south. | McGee, C. E. 1964. Species test in the sandhills. For. Farmer 24(2):10, 15. [Eleven species of pine and red cedar were planted in South Carolina sandhills to provide landowners with a better basis for choosing a pine for converting cutover areas to timber production. Sand pine did not fair very good in this study. Five species of pine and red cedar did better on the Americus and Lakeland soils than sand pine. | McGovern, J. N. anp E. L. Keer. 1943. Sulfite pulping experiments on sand pine. USDA Forest Serv., Forest Products Lab Reports R-1429. 6 pp. [Samples from the heartwood and sapwood of sand pine were completely pulped by the sulfite process to determine its paper-making qualities. The pulps were decidely inferior compared to other pines and the results suggested that this species may not be a promising source of sulfite pulp. | McNas, W. H. Aanp R. H. BRENDEMUEHL. 1983. Choctawhatchee sand pine survival, height, and biomass in South Carolina: third-year results. USDA Forest Serv. Gen. Techn. Rep. Se- 24. Pp. 96-100. [Choctawhatchee sand pine seedlings from Florida nurseries were planted on scrub oak sites in South Carolina to determine the effects of source, soil amendments, planting method, survival and height. Nursery source and soil treatments had no effect on overall survival or height. Machine planting had significantly greater survival than those planted by dibble-bar. Morphological grade had more of an affect on height than on survival. | AND A. R. Carter. 1981. Sand pine on South Carolina sandhills. South. J. Appl. For. 5(2):84-88. [Fifteen plantations of Choctawhatchee and Ocala sand pine in South Caro- lina sandhills produced an annual increment of wood approximately twice that of other commonly planted pines. ] , K. W. OuTcaLt AND R. H. BRENDEMUEHL. 1985. Weight and volume of plantation- grown Choctawhatchee sand pine. USDA Forest Serv. Res. Pap. SE-252. 44 pp. [The above ground green weight of the total tree and its major components, the main stem and crown, were determined in eight stands of planted Choctawhatchee sand pine ranging in age from 7 to 27 years. | McReyno.ps, R. D. anp R. M. Burns. 1975. Three machines for planting pine seeds in the sandhills. USDA Forest Serv. Res. Note-SE-218. 6 pp. [In sandhills, longleaf and Choc- tawhatchee sand pines can be successfully established from seed if sites are intensively prepared and if the seeds are covered with a thin layer of soil. Cultipackers satisfactorily covered seeds that had been broadcast on prepared sites cleared of debris, and well- stocked stands developed. The row-seeding machine produced comparable results in one operation using less than half as many seeds. | MILLER, G. J. ANDS. B. Jones, Jr. 1967. The vascular flora of Ship Island, Mississippi. Castanea. 32:84-99. [The eastern end of the island is covered by a small forest of slash pine and live oak with scattered shrubs of rosemary (Ceratiola ericoides). | Mitter, R. 1950. Ecological comparisons of plant communities of the xeric pine type on sand ridges in central Florida. Masters thesis, Univ. Florida, Gainesville. [The geologic, edaphic, and successional relationships of the sand pine scrub and sandhill communities in central Florida is presented. | Minno, M. 1987. The ecology and pollination biology of Curtiss’ milkweed (Asclepias curtissi). Florida Game and Fresh Water Fish Commission, Division of Wildlife, Nongame Wild- life Section, Project # GFC-86-027, Project Director, J. Putz. 77 pp. [The purpose of this project is to determine the nature and causes of rarity for the Curtiss’ Milkweed in Flor- ida. The Curtiss’ milkweed is endemic to Florida and restricted to scrub communities. No conclusive evidence for its restriction to scrub is provided, however, its high survival rate 84 FLORIDA SCIENTIST [Vol. 52 in spite of a low reproductive effort may have allowed this species to persist in the harsh scrub environment. Management recommendations for its preservation are also dis- cussed. | AND MInno. 1984. The proposed Fort Lauderdale executive airport gopher tortoise preserve. Pp. 38-48. In: Jackson, D. R. anv R. J. Bryant (eds.), The gopher tortoise and its community. Proc. 5th annual meeting, Gopher Tortoise Council. Univ. Florida, Gainesville. [Recommendations for the preservation of a rosemary dominated scrub at the Executive Airport in Broward county is discussed. This scrub represents one of the best developed scrubs left in Broward county. The vegetation and animal life is included in the paper, in addition to a simplified outline for its preservation. ] AND R. Myers. 1986. Archbold Biological Station: Its history and its biology. Palmetto. 6(4):3-7. [Briefly discusses the vegetation and wildlife of the station with some emphasis on the scrub community. The early history and current research goals of the station are also discussed. | Minter, D. W. ann I. A. S. Gipson. 1978. Ploioderma lethale. Commonwealth Mycological Institute descriptions of pathogenic fungi and bacteria No. 570:561-570. [Sand pine is indicated a host to this organism which causes pine needle blight. | Mirov, N. T. 1967. The Genus Pinus. The Ronald Press Company, New York, N.Y. [Range- information is presented for sand pine and several other pine species. ] MOoHLENBROCK, R. H. 1976. Woody plants of the Ocala National Forest, Florida. Castanea. 41:309-319. [A checklist of the sand pine scrub community known as the Big Scrub in the Ocala National Forest is provided. | Monk, C. D. 1967. Successional and environmental relationships of the forest vegetation of north central Florida. Amer. Midl. Natural. 79(2):441-457. [The successional, environmental, and geological relationships of the sand pine scrub community is discussed in this paper. Six other community types of north Florida are also discussed. | . 1967. Tree species diversity in the eastern deciduous forest with particular reference to north central Florida. Amer. Mid]. Natural. 101(918):173-187. [Tree species diversity in the sand pine scrub community was intermediate when compared to pine flatwoods, southern mixed hardwoods, mixed hardwood swamps, sandhills, cypress swamps, and bayheads. | Moore, D. M., J. B. HARBORNE AND C. A. WituiAMs. 1970. Chemotaxonomy, variation and geographical distribution of the Empetraceae. J. Linn. Soc., Bot. 63:277-293. [Flavonoids and other related phenolics are discussed for rosemary (Ceratiola ericoides). | Morais, J. E. 1967. Racial variation in sand pine seedlings. Proc. Southern Conference on Forest Tree Improvement. 9:132-136. [The objective of the study was to investigate possible variation of certain sand pine seedling characteristics, especially growth rate and response to fertilization. Seed color, seed size, cotyledon number, deficiency symptoms, and shoot weight for both races are discussed. | . 1967. Physiological and morphological traits of sand pine (Pinus clausa (Engelm.) Sarg.). Masters thesis, Univ. Florida, Gainesville. [This study was to investigate possible racial variation in certain characteristics of sand pine, primarily seedling growth rate and response to fertilizer. ] Mutvanlia, M. 1932. Ecological survey of a Florida scrub. Ecology. 12:528-540. [One of the first numerical surveys of the Florida scrub near Windermere, Orange county. Provides a brief description of the soils, water content, water holding capacity, and the often distinct separation between scrub and sandhill. Concludes that water relations are the major factors in determining scrub. | Muna, M. H. 1968. Phytoseiidae of sand-pine litter. The Florida Entomol. 51(1):37-44. [Eight- een species of mites were collected from pine litter of various scrubs throughout Florida. |] MussELMAN, L. J. AND W. F. Mann, Jr. 1979. Notes on seed germination and parasitism of seedlings of Buckleya distichophylla Santalaceae. Castanea. 44(2):108-113. [Sand pine, although never found in nature with Buckleya, supported vigorous haustoria. ] Myers, R. L. 1985. Fire and the dynamic relationship between Florida sandhill and sand pine scrub vegetation. Bull. Torrey Bot. Club 112(3):241-252. [Evidence suggests that the boundary between these two communities is dynamic, and the areal extent of each may have changed through time in response to fluctuations in their respective fire regimes. | AND N. D. Deyrup. 1983. The dynamic relationship between Florida sandhill and sand pine scrub vegetation. Bull. Ecol. Soc. Amer. 64(2):62. Abstract. [Sandhill and sand pine scrub form mutually exclusive, fire-dependent ecosystems on Florida’s sandridges. No. 2, 1989] RICHARDSON—SAND PINE SCRUB COMMUNITY 85 NasH, NEILL, Historic and prehistoric shifts in fire regimes may account for the distribution of the two types. | a AND N. D. Deyrup. 1984. Fire history and age structure relationships of three pine species on Florida’s sand ridges. Bull. Ecol. Soc. Amer. 65(2):258. Abstract. [Sand ridges in south-central Florida support a mosaic of xeric pine forests dominated by either sand pine, south Florida slash pine and longleaf pine, or a combination of the three. Sertinous- coned sand pine forms even-aged stands following intense fires. Both slash and longleaf pine occur as uneven-aged stands maintained by short interval, low intensity fires. Juxta- position of these pines in the landscape is a function of each species’ longevity and fire tolerance and of recent fire history. | AND P. A. Peroni. 1983. Commentary: Approaches to determining aboriginal fire use and its impact on vegetation. Bull Ecol. Soc. Amer. 64(3):217-218. [This paper suggests that longleaf pine sites have been replaced by south Florida slash pine or sand pine scrub vegetation. The vegetation changes suggest shifts in the fire regimes, possibly due to a decrease in Indian burning. | G. V. 1895. Notes on some Florida plants. Bull. Torrey Bot. Club. 22:141-161. [The relationship between the high pine and scrub communities is discussed. ] W. T. 1957. Historical biogeography of present-day Florida. Bull. Florida State Mus. 2(7):175-200. [This paper is concerned with patterns of animal and plant distribution in Florida and the ways in which these patterns may have come about. A discussion of rosemary scrub and how it is replaced by high pine and other associations is also pre- sented. | NEWMAN, C. AND E. GrirFin. 1950. Deer and turkey habitats of Florida. Florida Game and Fresh Water Fish Commission, Tech. Bull. No. 1. [General vegetation map of Florida showing scrub and other communities. | Oak, S. W., R. L. ANDERSON, G. W. Ryan, R. J. UHLER, AND J. W. Lewis. 1981. Sand pine root disease on the Ocala National Forest, Florida. Incidence and Losses, 1980. USDA Forest Serv. Area State and Priv. For. Rep. No. 81-1-35. 11 pp. [Root diseases associated with sand pine collected from 30 stands on the Ocala National Forest showed that at least 3 pathogens were responsible for tree death; however, there was no significant difference in the percent of suppressed and non-suppressed trees attacked. The percent of trees affected varied from stand to stand, ranging from 0 to 28 % . Losses due to sand pine root disease in the forest were estimated at $500,000 in 1980. Losses tend to increase with age and appear to accelerate in stands older than 25 years. | Ovutca.t, K. W. 1983. A comparison of sand pine varieties in central Florida. South. J. Appl. For. 7(1):58-59. [In a comparison planting of the Ocala and Choctawhatchee varieties of sand pine in central Florida, survival was equal 10 years after establishment. On the basis of height and volume of wood after 10 years, the Ocala variety is far superior to the Choctawhatchee in central Florida. | . 1985. Direct seeding versus planting for establishment of pines on west Florida sand- hills. Pp. 122-124. In: Proc. Third Biennial Southern Silvicultural Research Conference. USDA South. Forest Exp. Stn., Gen. Tech. Rep. SO-54. [Survival and growth of direct seeded longleaf, slash, and Choctawhatchee sand pines were compared with that of nur- sery grown 1-0 seedlings on a typical sandhill site. Choctawhatchee sand pine had the best growth in both direct-seeded and planted sites. In addition, sand pine was the only tested species where adequately stocked stands were established by the direct-seeding method of regeneration. | AND W. E. Barer. 1983. Sand pine. Silvicultural systems for the major forest types of the United States, USDA Forest Serv. Agric. Handb. No. 445. Pp. 170-171. [Summarizes the distribution of the two races of sand pine in Florida, their occurrence in various forest cover types, morphological traits, and the differences that fire, insects, and disease play in their management. ] AND R.H. BRENDEMUEHL. 1984. Sand pine survival and growth on prepared and un- prepared sites. South. J. Appl. For. 8(2):96-99. [Sand pine on unprepared areas had ac- ceptable stocking at age 10. Thus, both varieties can be established by underplanting among scrub hardwoods which exist on many sandhill sites. Choctawhatchee sand pine had better survival and seems to suffer less from mushroom root rot. Double chopping significantly increased height, diameter and volume growth. | AND R. H. BRENDEMUEHL. 1985. Growth of choctawhatchee sand pine plantations in Georgia. South J. Appl. For. 9(1):62-64. [Surveys of plantations of Choctawhatchee sand 86 FLORIDA SCIENTIST [Vol. 52 pine on sandhills in 10 Georgia counties showed that the growth is nearly comparable to that observed on sandhills in Florida. Damage from insects, disease, and cold weather was not evident. | PaRHAM, G. 1982. Improved reforestation with seedbed scarification device. USDA Forest Serv., Tree Plant. Notes 23(2):26-28. [Reforestation problems in the Ocala National Forest are discussed for the Ocala sand pine. A seedbed scarifier developed by a private seeding contractor resulted in much improved seed germination and establishment. Seeding fail- ures have been reduced substantially, even at reduced seeding rates. | Peacock, C. H. anp R. S. Wess. 1984. Screening potential causes of pine tree decline on Florida golf courses. Soil Crop Sci. Soc. Florida. 43:34-35. [Symptoms of yellowing and needle loss from slash pine, sand pine and longleaf pine on golf courses in central Florida may be the result of changes in soil pH and liming that may be interfering with the tree’s ability to uptake phosphorus. | Peron, P. A. 1983. Vegetation history of the southern Lake Wales Ridge, Highlands County, Florida. Masters thesis, Bucknell Univ., Lewisburg, PA. [A vegetation history was con- structed for the southern Lake Wales Ridge of Archbold Biological Station whose major vegetation associations include southern ridge sandhills, rosemary and sand pine scrubs, scrubby flatwoods, flatwoods, bayheads, swales, and seasonal ponds. Aside from habitat destruction, reduction of fire frequencies and alterations of fire seasonality represent the most marked effects of post-1920 development on the southern Lake Wales Ridge. | AND W. G. ABRAHAMSON. 1985. Vegetation loss on the southern Lake Wales Ridge. Palmetto. 5(3):6-7. [The results indicate that of the 36,121 ha of scrub and sandhills, 64% has been lost due to citrus, real estate, and improved pasture. | AND W. G. ABRAHAMSON. 1985. A rapid method for determining losses of native vegetation. Natural Area J. 5(1):20-24. [A method involving the use of microcomputer graphics to measure habitat area, soil maps to determine the presettlement extent of native vegetation, and current aerial photographs to provide data on habitat losses is discussed for the central ridge of Florida. Data shows that 64.2% of the scrub, scrubby flatwoods and southern ridge sandhills have been lost to pasture, development and culti- vation. | AND W. G. ABRAHAMSON. 1986. Succession in Florida sandridge vegetation: A retro- spective study. Florida Scient. 49(3):176-191. [Successional changes on Lake Wales Ridge vegetation at eight study sites is provided. Three findings were linked to reduced fire frequencies: 1) expansion of mesic broadleaf evergreen bayhead vegetation into adjacent flatwoods and some seasonal ponds, 2)substantial slash pine regeneration in flatwoods, and 3) increased growth of scrub oaks and hickory in xeric sandhills accompanied by low slash and longleaf pine regeneration. Scrubby flatwoods, sand pine and rosemary scrubs remained relatively stable except where repeated burns were responsible for the loss of sand pine. | Persons, J. L. 1897. A chemical study of some typical soils of the Florida peninsula. Florida State Exp. Stn. Bull. 43:601-714. [The chemical characteristics of several sand pine scrub soils is discussed. ] Puanris, R. P. anp F. W. Woops. 1960. Effects of temperature upon photosynthesis and respiration of Choctawhatchee sand pine. Ecology. 41(4):797-799. [Rates of photosynthesis and respi- ration were measured on seedlings at temperatures between 18°-48°C, while light re- mained constant. The maximum rate of photosynthesis was near 23°C and was negligible between 43°-48°C. Respiration reached a maximum at 48°C. The results indicate that sand pine seedlings may be more efficient than loblolly pine, but more work is needed. | Pire., J. A. AND D. J. Durzan. 1978. Chromosomal proteins of conifers, Part 1, Comparison of histones and nonhistone chromosomal proteins from dry seeds of conifers. Canad. J. Bot. 56(16):1915-1927. [Twenty bands of nonhistone chromosomal proteins were isolated from sand pine seeds. | Poponog, J. 1981. Vascular plants of Jonathan Dickinson State Park. Fairchild Tropical Garden, Miami. [Checklist includes plants that occur in the sand pine scrub community. ] Popp.eTon, J. E., A. G. SHury, AnD H. C. Sweet. 1977. Vegetation of central Florida’s east coast; A checklist of the vascular plants. Florida Scient. 40(4):362-389. [A checklist of the plants found on the Merritt Island peninsula and offshore barrier island complex encom- passing Cape Canaveral is presented. The paper does not mention the various plant asso- ciations, however, many of the species common to sand pine scrub are included in the checklist. | No. 2, 1989] RICHARDSON—SAND PINE SCRUB COMMUNITY 87 Price, M. B. 1973. Management of natural stands of Ocala sand pine. Pp. 144-151. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [This paper presents information on establishing, growing, and managing the Ocala variety of sand pine under the standard practices of timber management. Direct seeding on ade- quately prepared soils was the best method of reconverting scrub vegetation to Ocala sand pine. | PrircHett, W. L. anp J. W. Gooninc. 1975. Fertilizer recommendations for pines in the south- eastern Coastal Plain of the United States. Agric. Exp. Stn., IFAS, Bull. 774, Univ. Flor- ida, Gainesville. [Fertilizers are not generally recommended for pines in the sandhills due to site moisture limitations. However, 150-200 lb DAP/acre (168-225 kg/ha) may be used on young sand pine, while urea is sometimes used for older stands. ] AND J. L. Gray. 1974. Is forest fertilization feasible? For. Farmer. 23(8):6-7, 14, 16. [ The role of fertilizer and its use on sterile sandhill soils was discussed. An application rate was recommended for sand pine plantations in the southeastern coastal plain. | RicHarpson, D. R. 1977. Vegetation of the Atlantic Coastal Ridge of Palm Beach County, Flor- ida. Florida Scient. 40:281-330. [The major plant communities of PBC, including sand pine scrub, were correlated with geological history in order to show how the physical and biological parameters affect vegetation. Predrainage maps of the vegetation patterns were also produced to study successional changes with regard to species composition. | . 1985. Allelopathic effect of species in the sand pine scrub of Florida. Ph.D. dissert., Univ. of South Florida, Tampa. 121 pp. [Evidence is provided that allelochemicals re- leased from scrub plants would deter the invasion of sandhill herbs into the scrub and periodic surface fires in the sandhill would preempt the success of invaders from the scrub. | AND G. B. Wituiamson. 1979. Allelopathy between Florida’s scrub and sandhill com- munities. Florida Scient. 44(Suppl. 1):17. Abstract. [Allelochemicals may be an impor- tant factor in the maintenance of the sand pine scrub community. | AND G. B. Wituiamson. 1987. Chemical inhibition of sandhill grasses by sand pine scrub. Amer. J. Bot. 74:656-657. Abstract. [Allelochemicals released from several scrub species inhibit the growth of sandhill herbs in bioassays, greenhouse and field studies. | AND G. B. Wituiamson. 1988. Allelopathic effects of shrubs of the sand pine scrub on pines and grasses of the sandhills. Forest Sci. (in press). [Laboratory bioassays of sand pine scrub species were tested for inhibitory effects against native grasses and lettuce. Green- house studies showed that leachates from rosemary and scrub mint were inhibitory to a native broomsedge grass. Field transplant studies also indicated that pines and wiregrass grew more slowly in scrub soils than in sandhill. Allelochemicals may reduce fuel loads in the sand pine scrub community. | Rockxwoop, D. L., L. F. ConpE, ANp R. H. BRENDEMUEHL. 1980. Biomass production of closely spaced Choctawhatchee sand pines. USDA Forest Serv. Res. Note SE-293. 6 pp. [High planting densities (spacing less than 3 feet) of Choctawhatchee sand pine produced ade- quate production levels (volume, green weight, dry weight) for at least a 10-year period. Only minimal levels of site preparation are required to establish sand pine plantations on sandhill soils. | AND R. E. Gopparp. 1980. Genetic variation in Ocala sand pine and its implications. Silv. Genetica. 29(1):18-22. [Genetic potential for improving wood volume, form and fecundity appears good for Ocala sand pine. However, in spite of these generally positive indications, the overall potential of the Ocala race for large scale reforestation is suspect. The race’s poor survival and relatively low wood density, with little likelihood of genetic improvement, make it compare unfavorably to Choctawhatchee sand pine. | AND H. R. Kox. 1978. Which sand pine to plant in Florida? Fifth-year test results. South. J. Appl. For. 2(2):49-50. [After five years, Choctawhatchee sand pine had good survival but the lowest height. Ocala sand pine had the poorest survival and the greatest height. Choctawhatchee sand pine seems best suited for large scale reforestation. | Romans, B. 1775. A concise history of east and west Florida. Accompanied by maps of part of the province of east Florida. (R. W. Patrick, ed. Florida Facsimile Reprint, 1962.) [Described in detail the species composition of certain habitats, including the white sands of the pine barrens or scrub. | Ross, E. W. 1968. Sand pine, a new host of Fomes annosus. Plant Dis. Reporter 52(8):635. [First known report of sand pine as a host for Fomes in Florida. Infection of the roots apparently took place by contact with infected longleaf pineroots, following planting. ] 88 FLORIDA SCIENTIST [Vol. 52 . 1969. Clitocybe tabescens root rot of Pinus clausa. Phytopathology. 59(8):1047. [Choctawhatchee sand pine seemed resistant to mushroom root rot in Florida. Infection of sand pine in plantations was related to hardwood debris left in the soil following site preparation. | . 1970. Sand pine root rot—pathogen: Clitocybe tabescens. J. For. 68:156-158. [Mush- room root rot has caused up to 25% mortality in some Ocala sand pine plantations in Georgia and Florida. Advanced symptoms include chlorotic foliage and reduced shoot growth. | AND D. H. Marx. 1972. Susceptibility of sand pine to Phytophthora cinnamomi. Phytopathology 62(10):1197-1200. [P. cinnamomi is reported as a virulent pathogen of sand pine seedlings; both mycorrhizal and nonmycorrhizal seedlings of the Choctawhat- chee and Ocala races were attacked readily. | RvuEHLE, J. L. 1972. Response of sand pine to parasitism by lance nematode. Plant Dis. Reporter 56(8):691-692. [Seedlings of the Ocala and Choctawhatchee races of sand pine were grown in the greenhouse and inoculated with lance nematodes. After 4 months, all seed- lings had less weight, less foliar-stem weight and less root weight than controls, and there was no difference between the two races in numbers of nematodes recovered. | . 1971. Reaction of two races of sand pine to parasitism by the lance nematode, Haplo- laimus galeatus. J. Nematol. 3(4):328-329. [A greenhouse study showed there were no differences in numbers of nematodes recovered from the roots of Ocala and Choctawhat- chee sand pine seedlings. | AND R. H. BRENDEMUEHL. 1981. Performance of Choctawhatchee sand pine seedlings inoculated with ectomycorrhizal fungi and outplanted in the sandhills of north Florida. USDA Forest Serv. Res. Note SE-301. 7 pp. [This study indicates that inoculation with Pisolithus tinctorius can improve nursery grade of bare-root stock and suggests that more research into inoculation, increased colonization, and handling of Choctawhatchee sand pine seedlings with Pisolithus ectomycorrhizae is needed. | Sampson, O. R. 1973. Nursery practices used for sand pine. Pp. 67-73. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [Nursery practices such as site selection, soil management and fumigation, seed treatment and sowing, seedbed mulching and weeding, insect control, and growth and lifting are summarized for sand pine. | SANDIFER, P. A., J. V. MIGLARESE, D. R. CAupEr, J. J. MANzI, AND L. A. Barcxay. 1980. Ecologi- cal characterization of the Sea Island Coastal region of South Carolina and Georgia. Vol. III, Biological Features of the Characterization Area, Coastal Ecosystems Project, Office of Biological Sci., Fish and Wildlife Service, Washington, D. C. [Briefly compares the scrub forest communities in Georgia, Florida and North Carolina. | SayLor, L. C. AND R. L. Koenic. 1967. The Slash x Sand Pine Hybrid. Silv. Genetica. 16:134- 138. [Slash pine and sand pine were successfully crossed at the Union Camp Paper Corpo- ration seed orchard in Savannah, Georgia. the hybrids were intermediate between their parent controls in seven of nine morphological characteristics. ] ScHEER, R. L. 1957. Sand pine—scrub or timber tree? South. Lumberman. 195(2441):191-193. [The use of sand pine in converting sandhill lands was discussed in this paper. Several good traits were pointed out for this species, such as high survival rate, hardiness, competitive ability and its use as a source of pulp and knotty-pine paneling. | . 1959. Comparison of pine species on Florida sandhills. J. For. 57(6):416-419. [This paper deals with slash pine, loblolly pine, shortleaf pine, Monterey pine, and longleaf pine to determine suitability for planting on prepared sandhill sites. Sand pine was not used in this experiment because planting stock was unavailable. However, reference is made to the fact that sand pine has consistently produced better early height growth than the other pines, but its survival has been erratic. ] AND J. D. Honces. 1960. Planted sand pine grows well on unprepared Florida sand- hills. USDA Forest Serv., South. For. Res. 1(1):7-8. [On the deep sands of western Flor- ida, sand pine planted on unprepared sites grew well with little or no release from com- peting vegetation. Other pines (longleaf, slash, shortleaf) did poorly without release. | SCHMALZER, P. A. AND C. R. HInk te. 1983. Preliminary observations on scrub vegetation at Kennedy Space Center, Merritt Island, Florida. Unpublished report. [Stands with differ- ent fire histories but on similar sites were selected for comparison at the Kennedy Space Center. The scrub was defined as mixed oak scrub in association with saw palmetto and staggerbush. Dominance and structure varied with the time since last burning. Mean No. 2, 1989] RICHARDSON—SAND PINE SCRUB COMMUNITY 89 vegetation height declined from oldest to youngest, while above ground live biomass in- creased with stand age. Management implications suggested that 1) frequent burning (<5 years) will reduce height of vegetation below optimum for scrub jays, 2) frequent burning increased dominance of saw palmetto, 3) litter removal is significant, 4) the ratio of dead fuel to live fuel may be shifted with increased burning and 5) frequent burning will adversely affect rosemary. | AND C. R. HINKLE. 1987. Effects of fire on composition, biomass, and nutrients in oak scrub vegetation on John F. Kennedy Space Center, Florida. NASA Technical Memoran- dum 100305. 134 pp. [Four stands of scrub vegetation (2, 4, 8, and 25 years since last fire) were sampled for species composition, above ground biomass, and nutrient pools. | SELLARDS, E. H. 1912. The soils and other surface residual materials of Florida. Ann. Rep. Florida State Geol. Surv. No. 4:7-79. [The soils and character of the scrub is briefly discussed. | SMALLEY, G. W. AND R. L. SCHEER. 1963. Black root rot in Florida sandhills. Plant Dis. Reporter. 47(7):669-671. [Black root rot caused severe mortality in young slash pines of west Florida and showed root symptoms on planted longleaf pine, sand pine, and loblolly pine. |] SNEDAKER, S. C. 1963. Some aspects of the ecology of the Florida sandhill. Masters thesis, Univ. Florida, Gainesville. [Soil characteristics and their effect on species composition were determined for a variety of sandhills in six counties of north central Florida. Succession in the sandhills and its relationship to sand pine scrub are also discussed. | AND E. A. Luco. 1972. Ecology of the Ocala National Forest. USDA Forest Serv.., South. Region, Atlanta, Georgia. 209 pp. [The major plant communities of the Ocala National Forest are described based on the characteristics of the typical vegetation and certain aspects of the environment (soils, fire, succession). The sand pine scrub ecosystem is thoroughly discussed. ] STALLCuP, J. A. AND G. E. WooLFENDEN. 1978. Family status and contributions to breeding by Florida scrub jays. Animal Behav. 26(4):1144-1156. [Nestling care and breeding behavior of Florida scrub jays inhabiting low open scrub oak vegetation at Archbold Biological Station is discussed. | STEINBERG, B. 1980. Vegetation of the Atlantic Coastal Ridge of Broward County, Florida. Flor- ida Scient. 43:7-12. [Ten major plant communities of Broward county are mapped and described, including sand pine scrub. Characteristic plant species are given for each com- munity. | Stout, I. J. 1975. Structural attributes of scrub communities in relation to spatial distribution and competition among small mammals. Florida Scient. 38(Suppl. 1):7-8. [This study provided data on the question of how scrub community structure may relate to resource division, habitat selection, and species diversity of small mammals on the University of Central Florida campus. | . 1976. Response of small mammals in a scrub community to supplementary food. Florida Scient. 39(Suppl. 1):8. [This study presents data on the response of Peromyscus floridanus, P. gossypinus, Ochrotomys nuttalli, and Sigmodon hispidus to supplementary food. | . 1979. Efforts to inventory wildlife habitat, progress toward a terrestrial ecosystem monitoring program for the U.S. Space Shuttle Program. Trans. 44th North Amer. Wild- life and Natural Resources Conf. Pp. 457-465. [This paper outlines the objectives of the baseline inventory of the plant and animal populations on Merritt Island, inclusive of sand pine scrub community. | . 1981. Conservation biology of Peromyscus floridanus. Florida Scient. 44(Suppl.1):37. [Problems in the conservation biology of Peromyscus floridanus are high- lighted. The Florida mouse is a Florida endemic largely confined to sand pine scrub habitats. Destruction, fragmentation, and isolation of these habitats pose serious threats to the continued existence of this species over its former range. Long-term studies in central Florida provide evidence that isolation of habitat islands may lead to local extinc- tion. An incidence function which depicts the probability of occurrence in habitat islands should be developed for this species. ] . 1982. Small mammal community in sand pine scrub. Bull. Ecol. Soc. Amer. 63(2):68- 69. [Small mammal community structure was investigated over 9 years in sand pine scrub habitats in central Florida. ] , C. L. Connery, AND P. L. Kuincer. 1984. Preliminary findings on endangered plants in sand pine scrubs of east central Florida. Florida Scient. 47(Suppl. 1):45. [Ecological 90 FLORIDA SCIENTIST [Vol. 52 assessments of more than 30 relatively undisturbed sand pine scrubs of east central Florida were made. Data were also gathered on plant species currently included on state or fed- eral lists as having some potential level of endangerment, with emphasis on the problems of preservation. | AND R. J. DEMMer. 1982. Cotton rat Sigmodon hispidus invasion of sand pine scrub habitat. J. Mammol. 63(2):236-242. [Cotton rats invaded sand pine scrub in central Flor- ida in 1974. The sand pine scrub appeared to have acted as a dispersal link to the more optimal surrounding pine flatwoods. | AND M. H. Kem. 1981. Variation in number of plantar tubercles in Peromyscus flori- danus. Florida Scient. 44:126-128. [Plantar tubercles ranged from 3-6 per hind foot, typically 5. A tendency was shown for an additional pad to be present on mice of sand pine scrub habitats. The extra pad may be adaptive where climbing is advantageous. | , B. C. Mapsen, AND L. A. CHyNowetu. 1976. Field exposure of two upland cover types to solid rocket motor fuel emissions. Pp. 419-426. In: Southeastern Asso. Game and Fish Comm. Thirtieth Annual Conference. [Exposure of two upland plant communities (pine flatwoods and scrubby flatwoods) to solid rocket motor fuel emissions did not reveal any impact on the plant species. Soil pH, chloride concentrations, root biomass, and litter fractions remained unchanged. ] SULLIVAN, E. T. ANpD P. W. Frazer. 1969. Needle drop in sand pine Christmas trees. Sunshine State Agric. Res. Rep. 7(4):18-19. [The ability of sand pine to hold its needles when cut shortly before use and its value in plantation production makes it a good choice for the Florida Christmas tree producers. ] TANRISEVER, N., N. H. FiscHer, AND G. B. WiLuLrAMson. 1988. Calaminthone and other mentho- furans from Calamintha ashei; their germination and growth regulatory effects on Schi- zachyrium scoparium and Lactuca sativa. Phytochemistry. (In press). [Results of investi- gations of Calamintha phytotoxins reveal a novel monoterpene and several known menthofurans. Their biological activities are assessed in petri dish assays with a native grass and lettuce. | , F. R. Fronczek, N. H. FiscHer anp G. B. WILLIAMSON. 1986. Ceratiolin and other flavonoids from Ceratiola ericoides. Phytochemistry. 26:175-176. [Rosemary, a sand pine scrub associate in Florida, was chemically studied for its possible allelopathic activity. Chemical analysis of ground aerial parts of rosemary yielded the two known dihydro- chalcones angoletin and 2’,6’-dihydroxy-4-methoxy-3’,5’-dimethyldihydrochalcone, as well as 2’,4’-dihydroxychalcone. From water washes of freshly harvested leaves a novel dihydochalcone, ceratiolin, was isolated. ] Taras, M. A. 1973. Properties, uses and potential market of sand pine. Pp. 28-54. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [The morphological features, chemical, mechanical, and physical properties, and wood quality characteristics of both varieties of sand pine are discussed. Studies indicate that sand pine can be utilized as pulp under the same conditions as slash and longleaf pines without affecting paper properties. | . 1980. Aboveground biomass of Choctawhatchee sand pine in northwest Florida. USDA Forest Serv. Res. Pap. SE-210. 24 pp. [Choctawhatchee sand pine trees 4-14 inches DBH were selected from an uneven aged stand to determine weight and volume of above- ground biomass. | THompson, C. R. 1984. Association of Paratrechina arenivaga (Hymenoptera: Formicidae), with nymphs of Oecleus borealis (Homoptera: Cixiidae). J. New York Entomol. Soc. 92(1):35-41. [The nymphs of the planthopper (O. borealis) are thought to feed on the roots of sand pine, saw palmetto or turkey oak. | USDA Forest Service. 1973. Sand Pine Symposium Proceeding. USDA Forest Serv. Gen. Tech. Rep. SE-2. 253 pp. [Compilation of papers presented at the sand pine symposium. Topics include: sand pine characteristics, establishment and early growth, management and as- sociated problems, and tree improvement. | VAN Pett, A. F. 1958. The ecology of the ants of the Welaka Reserve, Florida (Hymenoptera: Formicidae). Part II. Annotated list. Amer. Midl. Nat. 59:1-57. [Exhaustive survey of the ants of the Welaka scrub habitats. ] VANDER Kioer, S. 1979. Florula archboldensis: Being an annotated list of the vascular plants of the Archbold Biological Station, Lake Placid, Florida. 72 pp. [Complete list of the flora of the different vegetation types (scrub, sandhill, bayheads, swale, wet prairies and ponds, pine flatwoods and scrubby flatwoods) at the Archbold Station. ] No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY 9] Veno, P. A. 1976. Successional relationships of five plant communities. Ecology. 57:498-508. [Five plant communities (mesic hammock, xeric hammock, bluejack oak sandhill, turkey oak sandhill, and scrub) in the Welaka Conservation Reserve in Putnam County were inventoried to determine successional changes. No species change was observed for the scrub community, except for size and number. | VIGNOLES, C. 1823. Observations upon the Floridas. Bliss, New York, N.Y. 197 pp. [The scrub lands vary but very little in their general appearance wherever found. Rules out the use of the scrub by the farmer, but indicates that it is excellent ground for hog farming. | Vozzo, J. A. 1984. Evaluating seed leachate measurements relative to imbibition time and solute volume. J. Seed Tech. 9(1):54-59. [ Leachate conductivity (total solute concentration) may be a useful measurement of seed vigor comparable to conventional seed germination tests. Measurements were taken from six southern pine species (sand, shortleaf, slash, scotch, loblolly and Virginia pines). Sand pine conductance followed the expected pattern of greater conductance over a longer imbibition time. ] Wane, D., J. Ewet anp R. Horstetter. 1980. Fire in south Florida ecosystems. USDA Forest Serv. Gen. Tech. Rep. SE-17, Southeast. For. Exp. Stn., Asheville, N.C. [Makes reference to an isolated pocket of sand pine in Collier county, Florida, which may be its one time southern limit. | Warp, B. W. anp C. J. CHAPMAN. 1973. Military herbicide driftage and Florida vegetation. Florida Scient. 36(2):110-122. [Spray equipment designed to apply herbicides and other biologically active materials was tested in a sandhill community in Eglin Air Force Base. Descriptions of the test area indicate that sand pine scrub had invaded the once longleaf pine forest, now dominated by turkey oaks. | Warp, D. B. 1964. Contributions to the flora of Florida 2, Pinus (Pinaceae). Castanea. 28:1-10. [Briefly discusses the habitat and distribution of sand pine in Florida with reference to the differences between the two varieties. | Watts, W. A. 1971. Postglacial and interglacial vegetation history of southern Georgia and central Florida. Ecology. 52(4):676-690. [Evidence is presented that in the period from about 8,500 to 5,000 B.P., the coastal plain of Georgia and north-central Florida had a vegetation that was a mosaic of sclerophyllous oak woodland and small patches of prairie. The presence of sand pine scrub surrounding Mud Lake in Florida may suggest that the early flora was a mixture of sand pine or longleaf pine among several scrubby oak species. ] . 1975. A late Quaternary record of vegetation from Lake Annie, south central Florida. Geology. 3:344-346. [Pollen from Lake Annie indicated that from 37,000 to 13,010 B.P., Ceratiola and Polygonella spp. were abundant, implying a very dry environment. From 13,010 to 4,715 B.P., oaks and ragweed dominated and from 4,715 to present, the modern flora prevailed. | Wess, J. W. AND J. D. Dopp. 1981. Tree and shrub establishment and survival on sandy dredged, material. Southwest Naturalist 26(4):345-352. [Sand pine was one of eight tree species used to revegetate dredged spoil material from the intracoastal waterway on Bolivar Pe- ninsula, Texas. | Wesser, H. J. 1935. The Florida scrub, a fire-fighting association. Amer. J. Bot. 22:344-361. [The general character of several interior scrubs of Florida are discussed. Soil characteris- tics, fire regimes, successional trends and disturbance are presented in an effort to parti- ally explain the abrupt separation between the scrub and the high pine land or sandhill. ] Westcott, P. W. 1970. Ecology and behavior of the Florida scrub jay. Ph.D. dissert., Univ. Florida, Gainesville. [Nesting scrub jays are invariably located in types of sand pine scrub vegetation. Preferred habitat is discussed for several populations throughout Florida. ] Wuire, D. L. anv W. S. Jupp. 1985. A flora of Gold Head Branch Ravine and adjacent uplands, Clay County, Florida. Castanea. 50(4):250-261. [Xeric oak scrub occurs as an interface between the mixed hardwood community of the ravine and the sandhill at the higher elevations. A checklist of the oak scrub and sandhill is provided. ] White, W. A. 1970. The geomorphology of the Florida peninsula. Bu. Geol., Div. Interior Resour., Florida Dept. Nat. Resour., Geol. Bull. No. 51. 164 pp. [The major relict shore- lines supporting sand pine scrub today are discussed, in addition to several other forma- tions. | Wuirney, M. 1898. The soils of Florida. USDA Div. Soils Bull. 13:14-27. [Chemical analysis of scrub soils and a possible explanation for the sharp ecotone between the sand pine scrub and the high pine land is discussed. ] Winnie, L. P. ann R. P. Scuuttz. 1969. Summer planting of sand and longleaf pines fail. USDA 92 FLORIDA SCIENTIST [Vol. 52 Forest Serv., Tree Plant. Notes. 20(1):7. [Sand pine and longleaf pine seedlings were planted in the summer months in 3 Florida counties. Each site received ample rainfall during July, August and September, however, survival was less than 9% for the planted stock. On the basis of these results, sand pine is not suited for summer planting. | Wiikens, K. T. 1982. Systematics and zoogeography of fossil and recent pocket gophers in Flor- ida. Ph.D. dissert., Univ. Florida, Gainesville. [The distribution of Geomys in Florida has been strongly influenced by repeated Pleistocene and recent sea level changes. Four primary habitats are used by this species in Florida: sandhill, xeric hammock, longleaf pine flatwoods and sand pine scrub. The role of fire in the maintenance of Geomys popu- lations is also discussed. | Witkinson, R. C. 1971. Neodiprion excitans (Hymenoptera: Diprionidae) on sand pine in Flor- ida. Florida Entomol. 54(4):343-344. [Larvae of this sawfly caused severe defoliation of a natural stand of sand pine in west Florida. The outbreak resulted in 10% mortality of mature trees. | AND C. W. CHELLMAN. 1978. A new sawfly on sand pine in west Florida. Florida Entomol. 61(1):26. [The sand pine sawfly has caused defoliation of both Ocala and Choc- tawhatchee sand pines 10 years or older. Defoliation occurred along stand edges and within those stands with stockings of 750 or less trees per acre. | WiuuiaMson, G. B. 1988. Allelopathy: Fiction, crucifixion, and the neck riddle. In: Grace, J. B. AND G. D. TitMAN (eds.), Perspectives in Plant Competition (in press). [Provides a brief history of the growth of allelopathic studies over the last thirty years, and then summa- rizes generalizations from phytotoxic investigations of the scrub. ] , N. H. Fiscuer, N. TANRISEVER, AND A. DE LA PENA. 1987. Ecological chemistry of inhibition of grasses and pines by shrubs of the sand pine scrub community. Amer. J. Bot. 74:660. Abstract. [Brief review of the degradation of ceratiolin from Ceratiola and the toxicity of the degradation product, hydrocinnamic acid. | AND D. R. RicHaArpson. 1988. Bioassays for allelopathy: Measuring treatment re- sponses with independent controls. J. Chem. Ecol. 14:181-187. [A response index was derived to compare bioassay treatment effects against those of controls. To illustrate the use of this index, source leachates from ten scrub species collected over a winter and summer season were compared against distilled water controls. Three target grasses and lettuce were placed in the leachates and distilled water to determine germination rates and radicle growth. | AND D. R. RICHARDSON, AND N. H. Fiscuer. 1988. Allelopathic mechanisms in fire- prone communities. In: Rizvi, S. J. H. (ed.), Frontiers of Allelochemical Research. Mar- tinus Nijoff Publishers (in press). [The role of allelopathic as a mechanism to reduce fuel around fire-sensitive shrubs is reviewed in Florida scrub and compared to the California chaparral. | Woops, F. W. 1959. Converting scrub oak sandhills to pine forest in Florida. J. For. 57:117-119. [Survival and growth of pine seedlings planted on west Florida sandhills are best on completely cleared sites, where available soil moisture is most abundant during critical dry periods. Slash pine was recommended for planting the first year following clearing. The author indicates that sand pine shows promise as a potential species for revegetating sandhills in Florida. ] . 1983. Operation sand-hills. South. Lumberman. 187(2345):150-151. [Describes a method for clearing scrub oaks and wiregrass in Florida sandhills. ] WooLFENDEN, G. E. 1969. Breeding-bird censuses of five habitats at Archbold Biological Station. Audubon Field Notes. 23(6):732-738. [A bird census was conducted in the sand pine scrub, scrubby flatwoods, slash pine/turkey oak, recently burned scrubby flatwoods, and low flatwoods communities at Archbold Biological Station. ] . 1973. Meeting and survival in a population of Florida scrub jays. Living Bird. 12:25- 49. [Nesting behavior, reproductive success, and habitat preference were studied for the Florida scrub jay at the Archbold Biological Station, Lake Placid, Florida. } . 1978. Growth and survival of young Florida scrub jays. Wilson Bull. 90(1):1-18. [Growth of young scrub jays was measured in a marked population inhabiting sparse oak scrub at Archbold Biological Station, Florida. The overall growth index was similar to that of pinon jays and the young grow only slightly slower than expected for their body size. | AND J. W. Fitzpatrick. 1977. Dominance in the Florida scrub jay. Condor. 79:1-12. No. 2, 1989] RICHARDSON— SAND PINE SCRUB COMMUNITY 93 [Detailed observations of dominance among 21 families of Florida scrub jays inhabiting oak scrub at Archbold Biological Station were obtained. | AND J. W. Firzpatrick. 1978. The inheritance of territory in group breeding birds. Bioscience. 28(2):104-108. [Dispersal strategies, territory size and growth of scrub jays occupying oak scrub in central Florida is discussed. | AND J. W. Firzpatrick. 1984. The Florida scrub jay: Demography of a cooperative- breeding bird. Monogr. Pop. Biol. 20. Princeton University Press, Princeton, N.J. [Florida scrub jays are an excellent example of a cooperative breeding species in which adult birds often help raise offspring not their own. After several years of study, habitat restraints rather than kin selection are the main source of the behavior of Florida scrub jays. The study tract consisted of primarily scrub oak and scrubby flatwoods at the Archbold Bio- logical Station. A thorough description of the various habitats utilized by scrub jays at the station is provided. | Younc, B. L. 1983. Food supplementation of small rodents in sand pine scrub. Masters thesis, Univ. of Central Florida, Orlando. 76 pp. [Peromyscus gossypinus responded to added food by slight increases in density and immigration. In contrast, Ochrotomys nuttalli appeared to have improved survival, but densities were not changed. P. floridanus did not respond to the added food. | AND I. J. Stour. 1983. Food supplementation of small rodents in sand pine scrub. Florida Scient. 46(Suppl. 1):26. [A two-year study was conducted to evaluate population fluctuations of sand pine scrub small rodents in response to supplementary food. | AND I. J. Stout. 1984. Diets of small rodents in the Florida sand pine scrub: Are rodents food-limited? Bull. Ecol. Soc. Amer. 65(2):97. [Diets of cotton mice, golden mice, and flying squirrels were determined by fecal analysis during a 1-year supplementary food study in sand pine scrub habitats of central Florida. Rodent abundance was not altered by the added food. | ANDI. J. Stour. 1986. Effects of extra food on small rodents in a south temperate zone habitat: Demographic responses. Canad. J. Zool. 64:1211-1217. [Demography of Pero- myscus gossypinus, Ochrotomys nuttalli, P. floridanus, and Glaucomys volans was studied for two years on replicate grids in sand pine scrub habitat in central Florida. | ZELAWSKI, W. AND R. K. STRICKLAND. 1973. Temperature effects on growth, assimilation, and bud development of sand pine. Pp. 73-82. In: USDA Forest Serv., Sand Pine Symp. Proc. USDA Forest Serv. Gen. Tech. Rep. SE-2. [The Choctawhatchee and Ocala varieties of sand pine differ in their response to temperature. Choctawhatchee sand pine was more tolerant of low temperatures but showed slightly lower dry matter accumulation at higher temperatures than did the Ocala variety. Higher temperatures were also needed to break dormancy in the Choctawhatchee variety. The net assimilation rate of sand pine was comparable to other conifers. | ACKNOWLEDGMENTS—I especially honor the late Dr. Albert Lassele for his fruitful discussions and support of this document. I would also like to thank the many persons who have contributed reprints and articles over the many years. This material was based upon work supported in part by the U.S. Department of Agriculture, Competitive Research Grants Program for Forest and Rangeland Renewable Resources under Agreement No. 85— FSTY-9-0139. Florida Sci. 52(2):65-93. 1989. Accepted: July 16, 1988. Chemical Sciences CALCULATED X-RAY DATA AID IN COLLECTING HIGH QUALITY X-RAY POWDER DATA— OXYTETRACYCLINE DIHYDRATE, C,,H,,N,O,.2H,O, AS AN EXAMPLE FRANK N. BLANCHARD Department of Geology, University of Florida, Gainesville, Florida 32611 Asstract: Observed X-ray powder diffraction data were interpreted with the aid of a calcu- lated pattern based upon the known crystal structure. The results consist of a high quality stand- ard powder pattern with the usual supporting data for an important compound which is not presently included in the X-ray Powder Diffraction File. TETRACYCLINES comprise an important group of compounds of which some are useful as antibiotics. X-ray powder diffraction serves as a rapid and accu- rate method of identification of the various tetracyclines, either in pure form or in mixtures. In addition, deterioration of the therapeutic usefulness of the drug with shelf aging can be detected easily by X-ray powder diffraction. Unfortunately, few of the tetracyclines are characterized by published ray powder data. In this report, I present indexed X-ray powder diffraction data and related data for oxytetracycline dihydrate C,,H,,N,O,.2H,O. It is emphasized that crystal structure information made it possible to calcu- late a powder diffraction pattern, which aided greatly in processes involved in treating the observed raw data. MATERIALS AND METHODs—Experimental Conditions—The powder diffraction data were collected on a Philips APD 3600 vertical diffractometer (173 mm), with a Cu-target tube oper- ated at 45 KV and 40 MA. Instrumental conditions include a © -compensating slit (13 mm irradiated length), Soller slits on the incident and diffracted beams, and a 6° take-off angle. Because the a,-a, doublet was not resolved in the diffraction lines for the tetracycline, a wave length of 1.54178 (unresolved CuKa) was used to obtain d-values from 2 angles. A receiving slit of 0.2 mm was used in front of a curved graphite monochromator and a scintillation detector No. 2, 1989] BLANCHARD— X-RAY DATA AID 95 with pulse height discrimination was used. No temperature control was used and the estimated temperature range during data collection was 25+ 2° C. All scans were from 2° to 60° 26, with a step size of 0.01° and a count time of 2 seconds per step. The instrument profile breadth was determined using -20 um silicon at 56.12° 26. Three repeated measurements each yeilded a FWHM (full-width half-maximum) value of 0.13°. Sample, methods, and data reduction—The sample consisted of a yellow powder (finely divided enough so that additional grinding was not necessary), produced by the Sigma Chemical Company; the powder was vertically packed into a cavity-type specimen holder. One scan was made with fluorophlogopite (NBS SRM-675) blended with the sample in a “Spex” mixer-mill as an internal standard, and measurements from this mixture were used to obtain corrected 26 angles for all of the strong peaks of the tetracycline. Eleven selected (and corrected) peaks of the tetracycline, chosen at approximately uniform intervals through the 26 range scanned, were used to produce a calibration curve to correct the peaks in a sample of the pure tetracycline. Three scans were made with pure samples and the data from these were used to obtain average intensities. The reference intensity ratio, an instrument-dependent constant, is useful for quantitative X- ray powder diffraction. If the reference standard is corundum, the ratio is abbreviated I/Ic (Hubbard et al., 1976), which means the intensity of the strongest line in the powder diffraction pattern of a substance divided by the strongest line for corundum (d= 2.085) in a 50-50 mixture (by weight) of the two phases. With known I/Ic values for crystalline phases the “matrix-flush- ing’ method of quantitative analysis may be used (Chung, 1974). Even though I/Ic varies some- what from one instrument to another, experimental values are now included in the X-ray Powder Diffraction File (PDF). The reference intensity ratio (I/Ic) was obtained from a sub-sample consisting of oxytetracycline dihydrate mixed in 50-50 weight proportion with 1 ym size corun- dum powder (Linde C). Eight scans, covering the angular ranges required for the most intense peaks of the tetracycline (d=7.93) and corundum (d=2.085), were made on separately loaded sample holders in order to obtain an average I/Ic value (Ig_7.93(tetracyctine)/Ta=9.085 (corundum))» In order to be comparable with other data in the PDF, I/Ic was corrected to a the value which would have been obtained with a 1° divergence slit from the © -compensating slit actually used (based on a 13 mm sample length). ; The Philips analytical software was used for peak searching. Where inspection indicated that a visual estimate of peak position seemed better than that found by the computer, the peak was relocated visually. Indexing and refinement of cell parameters was accomplished using the pro- gram of Appleman and Evans (1973) with the starting parameters and space group of Stezowski (1976). The averages of three observed intensities were converted to those that would have been obtained with a 1° fixed divergence slit (as described above). UsE oF CALCULATED PATTERNS— From the known crystal structure of oxy- tetracycline dihydrate (Stezowski, 1976) a calculated pattern was obtained (using the program of Smith et al., 1983), and this permitted pre-indexing of most of the reflections for a final least-squares refinement of unit-cell param- eters. This greatly reduced the number of reflections that were rejected in the least-squares refinement, leading to a better set of unit-cell parameters and to confirmation of the indexing. A segment of the experimental pattern (5° to 30° 2e) with a simulated diffractogram (calculated pattern) shows the close correspondence between the two (Fig. 1). Included in the figure are possible hk1’s for each reflection. Below about 15° 2¢ all reflections are unambiguously indexed, however, at higher 2.6 angles in some instances there is more than one set of lattice planes which may account for the reflection. This problem becomes increasingly severe as 2© increases. These reflections which cannot be unambiguously indexed were rejected (R in Fig. 1) in the refinement of unit-cell parameters. The pattern that is calculated from the known crystal structure includes cal- culated intensities for all possible reflections and this information makes it possible to pre-index most of the observed reflections. With pre-indexing the 13 “rejections” within this angular range no longer remain as “rejections”. 96 FLORIDA SCIENTIST [Vol. 52 a (oa) jo) i-4 oO N N 033 121 R 002 032 R 310 222 R311 320 R 141 123 R 240 232 R 042 319 R 203 213 R 241 051 R 150 133 R 400 231 R 041 140 Fic. 1. Simulated (top) and observed (bottom) diffractograms from 5° to 30° 26. This repre- sentative part of the pattern shows the close correspondence between the two. Possible hk1’s for each reflection are indicated and the “R’s” denote reflections “rejected” in the unit-cell refine- ment, unless pre-indexing is done based on the calculated powder data. No. 2, 1989] BLANCHARD— X-RAY DATA AID 97 For the entire pattern (to 60° 2 ) pre-indexing reduces the number of “rejec- tions” from 54 to 2 (out of a total of 84 observed reflections), resulting in 82 of the 84 reflections contributing to the least-squares refinement of unit-cell parameters; no pre-indexing results in only 30 of the 84 contributing to the refinement. In addition to the better lattice parameters from the greater number of used reflections, indexing is more reliable. ResuLts—Oxytetracycline dihydrate is orthorhombic with space group P2,2,2 (Stezowski, 1976), and a=12.081(1) A, b=15.886(3) A, and c=11.529(1) A (refined from the powder data presented in this report), Z=4, D (calculated) = 1.490, and I/Ic (observed) = 0.38. The figure of merit, which gives a measure of the quality of the data (Smith and Snyder, 1979), is F,,=50.7(0.011,52) computed from the output of the program used to refine lattice parameters. All differences between observed and calculated 26 an- gles are less than 0.05°. The quality evaluations just described indicate a high quality data set that should be useful to those involved with the production, use, and identification of tetracyclines. A summary of d, I, and hk1 values is given in Table 1. Multiple hkl values are assigned where two sets of lattice planes contribute significantly to a reflection; in some instances more than two sets of planes contribute and in these cases the “+” sign is used to indicate these additional planes, in keeping with the convention used on the PDF cards. Where possible, it is reeommended that investigators publishing new powder data make use of calculated patterns as an aid in indexing. ACKNOWLEDGMENTS— This work was supported in part by a grant-in-aid from the JCPDS - International Centre for Diffraction. Dr. Prof. Gus Palenik, Department of Chemistry provided the sample of oxytetracycline dihydrate. LITERATURE CITED APPLEMAN, D. E., AND H. T. Evans, Jr. 1973. Indexing and least-squares refinement of powder diffraction data. Geol. Surv. Contr., No. 20. Cuunc, F. H. 1974. Quantitative interpretation of X-ray diffraction patterns of mixtures. I. Matrix-flushing method for quantitative multicomponent analysis. J. AppL. Cryst. 7:519- 531. Hupparp, C. R., E. H. Evans, anp D. K. Smirn. 1976. The reference intensity ratio, I/Ic, for computer-simulated powder patterns. J. Appl. Cryst. 9:169-174. SmitH, D. K., M. C. NicHots, anp M. E. Zoenskr. 1983. POWD 10 a FORTRAN IV program for calculating X-ray powder diffraction patterns - version 10. The Penn. State Univ., University Park, Pennsylvania. SmiTH, G. S., AND R. L. Snyper. 1979. F(N): a criterion for rating powder diffraction patterns and evaluating the reliability of powder pattern indexing. J. Appl. Cryst. 12:60-65. StEzowskI, J. J. 1976. Chemical-structural properties of tetracycline derivatives. 1. Molecular structure and conformation of the free base derivatives. J. Amer. Chem. Soc. 98:6012- 6018. Florida Sci. 52(2): 94-99. 1989. Accepted: July 15. 1988. 98 FLORIDA SCIENTIST [Vol. 52 TABLE 1. List of d-values, relative intensities, hkl’s, and 26 angles for all observed reflec- tions. Also included are the three measured intensities for each reflection (© -compensating slit used) along with the average (AV) and relative standard deviation (RSTD). dA 11.49 961 9.32 8.35 7.93 7.38 6.54 6.044 5.754 5.642 5.350 5.202 5.073 4,939 4.849 4.802 4.664 4.436 4.352 4.172 4.028 3.970 3.899 3.763 3.693 3.588 3.429 3.323 3.275 3.230 3.179 3.155 3.108 3.087 3.061 3.048 3.009 2.967 2.920 Pecos fh 2.834 2.760 2.738 2.709 2.643 2.600 2.532 2.477 2.438 2.389 2.374 2.353 Fixed Slit I/Io — hkl 001 110 Ot 101 020 a ed 02) 200 002121 21.0 201 102 27 I Lie 30 220031 O22 OD 129 202 212 040 032310 23 1 NAVAS NEL 141320 SA 123 7320429 S162 213 142 033 Sol Odi S22 USES 15a] 401 ED) 014 114043 Sle3 024 13254 204 0344224 134 Souk 510 261 441413 7.692 9.494 10.596 PEG 11.992 13.536 14.655 15.400 15.705 16.569 17.044 17.480 17.959 18.294 18.474 19.026 20.014 20.406 Ane El 22.067 22.395 22.806 23.643 24.099 24.813 25.985 26.828 27.233 27.615 28.064 28.287 28.721 28.925 29.175 29.305 29.685 30.114 30.612 31.081 31.567 32.433 32.710 33.067 33.911 34.496 35.456 36.260 36.862 37.652 37.899 38.250 Intensities 1(2) 100 22 32 Z 95 49 5 8 43 7 23 1(3) I(AV) 93 23 33 8 98 51 6 9 44 8 25 I(RSTD) 11.2% 6.1% 4.3% 10.2% 2.4% 3.3% 8.3% 10.9% 3.2% 6.1% 6.5% 0.0% 5.1% 6.1% 7.4% 0.0% 0.0% 7.8% 5.6% 8.3% 4.3% 7.0% 10.1% 4.1% 6.0% 6.8 % 8.0% 6.7% 4.9% 7.9% 3.3% 5.8% 4.4% 1.9% 4.2% D2 8.2% 30.0 % 0.0% 3.7% 6.4% 4.7% 7.9% 10.9% 5.8% 5.7% 4.4% 6.8% 0.0% 4.9% 5.7% T.% No. 2, 1989] TABLE 1. Continued. Fixed Slit I/Io mb DR rR RE ENR RR RE EY NWN DD WH UAN®D UUORBK ®D CO BLANCHARD— X-RAY DATA AID 99 Intensities hkl 20 La) 2) 136)" TAV)) 1 RSTD) YS al 38.579 6 Y) 0.0% 144352 39.339 23 20 21 DD) 5.8% DOS 39.731 22 17 19 19 10.6% ILS 40.220 14 IY IY tS 7.4% 442 40.656 10 9 9g 9g 5.1% 244 41.482 20 18 19 19 4.3% 054215 42.334 11 10 10 10 4.6% 072035 42.808 12 10 9g 10 WONG 154 43.015 2 11 10 lig 7.4% 9h, NS) 43.502 18 16 Wi ING 4.8% 362540+ 43.837 11 11 10 11 4.4% LOS HG” 44.132 10 10 9 10 4.9% 424 44.979 13 13 13 13 0.0% DB) 45.445 15 14 13 14 5.8% Grleleie4s5 46.124 7 7 5 6 14.9% 5 ALY NG OT 8 i 6 o 11.7% 3638 47.388 7 i 5 6 14.9% 354 48.285 4 4 y 3 28.3% 335 48.631 4 2 y 50.0 % 182264 49.126 5 5 3 4 21.8% 216415 50.127 4 3 3 3 14.1% 5 al 50.586 3 4 3 3 14.1% 425255+ lll i qf 6 i Col % STSESO4 52.064 4 4 5 4 10.9% 435306 52.842 2 3 2 Y 20.2% 146 53.014 3 3 2 3 17.7% 290 04.107 1 il 1 1 0.0% 7 L210 316 50.896 4 4 4 4 0.0% 643156+ 56.540 4 4 4 4 0.0% 5) 57.928 5 4 5) 3) 10.1% 1100703+ 58.628 8 i ef q 6.4% Be 59.215 4 4 4 4 0.0% Biological Sciences BOSTRICHOBRANCHUS DIGONAS: CONFIRMATION OF ITS PRESENCE IN THE GULF OF MEXICO CERALD E.. WALSH U. S. Environmental Protection Agency, Environmental Research Laboratory, Gulf Breeze, Florida 32561 Asstract: Although Bostrichobranchus pilularis has been reported from the Gulf of Mexico, descriptions are similar to those of B. digonas. Developmental stages of B. digonas are described and shown to be distinct from those of B. pilularis. Published literature and information pre- sented here indicate that B. digonas is present in shallow water between central and northwest- ern Florida. THERE is confusion about distribution of the ascidian genus, Bostricho- branchus, in the Gulf of Mexico. Van Name (1945) stated that Bostrichobran- chus (Eugyra) pilularis Verrill is distributed from the St. Lawrence estuary and the banks of Newfoundland to the Gulf of Mexico off the central coast of Florida. Plough (1978) stated that B. pilularis is a cold-water species found in the Apalachee Bay region of northern Florida, but did not make the species diagnosis himself. Plough (1978) also mentioned, without description or ref- erence, a related form, Eugyra arenosa padrensis, from Padre Island, Texas. Bostrichobranchus and Eugyra belong to the Eugyrinae, a subfamily of the Molgulidae, which contains some species that have a single gonad on the left side and direct larval development, rather than the usual ascidian urodele tadpole metamorphosis. Van Name’s (1945) distribution of B. pilularis in the Gulf of Mexico was based on a single individual that had gonads on both sides of the body. He did not comment on the possible significance of two gonads and incorporated his specimen into the species pilularis, even though the species was described by Verrill (1871) as being monogonadal. Following Van Name (1945), Plough (1978) reported B. pilularis from the northern coast of Florida, and Cooley (1978) described Bostrichobranchus from the vicinity of Pensacola (north- western Florida), accepting the species name pilularis, even though two go- nads were present (personal communication). Abbott (1951), however, described individuals with gonads on both sides of the body from a beach on the western coast of Florida along the Peace River estuary, Charlotte Harbor. He compared them to van Name’s (1945) description of B. pilularis and stated that the digonadal form represented a distinct species based upon the number of gonads, mantle musculature, and structure of the dorsal tubercle. He named the species Bostrichobranchus digonas. The report describes development of B. digonas from the northern Gulf of Mexico and examines its relationship to the monogonadal species, B. pilularis. No. 2, 1989] WALSH—B. DIGONAS IN THE GULF OF MEXICO 101 y Yyff iff Wi) mm Fic. 1. Development stages of B. digonas. Early zygote (la), zygote with small ampullae (1b), later stage in development of ampullae (lc), further developmental stages showing resorption of ampullae and development of siphons and tunic (ld,e,f), juvenile ascidian with endostyle (en) and early branchial basket; the circulatory system is functional at this stage (lg), and adult (lh). Scale bar =0.1 mm. amp = ampullae; os=oral siphon; as atrial siphon. 102 FLORIDA SCIENTIST [Vol. 52 MetrHops— Adult ascidians were collected by hand from sand in settling tanks at the Environ- mental Research Laboratory, Gulf Breeze (87° 09° WL, 30° 20’ NL), between November 1985 and February 1986. Adult taxonomic characteristics were compared with those of B. pilularis (van Name 1945) and B. digonas (Abbott 1951). Young were extruded from the brood sac through the atrial siphon into filtered (0.45 um) seawater of 20 parts per thousand (ppt) salinity by gentle pressure on both sides of the animal. They were examined microscopically and photomicrographs were taken of developmental stages from early zygote to early larva. Early larvae were maintained in filtered seawater in Petri dishes through the juvenile stage (completely metamorphosed) and were fed marine unicellular algae, Skeletonema costatum, Thalassiosira pseudonana, Isochrysis galbana, and Chlorella sp. once a day. Developmental stages were compared with those of B. (Eugyra) pilularis and E. arenosa given by Berrill (1931). Temperature and salinity were monitored continuously by automatic sensors during the col- lection period (Table 1), and grain size analysis of sand was performed with a graded series of geological sieves. TABLE 1. Salinities and temperatures of water in the ascidian settling tanks. Salinity (ppt) Temperature (°C) Month Average Range Average Range November 1985 21.3 16.9-27.5 18.6 14.2-22.2 December 1985 22.4 10.8-32.9 1 7.9-22.6 January 1986 25.2 16.0-31.9 12.8 10.0-15.4 February 1986 24.2 172-3135 14.8 9.8-17.7 amp. Fic. 2. Early developmental stages of B. pilularis given by Berrill (1931). amp = ampullae. No. 2, 1989] WALSH—B. DIGONAS IN THE GULF OF MEXICO 103 RESULTS AND DiscussioNn—Ascidians were buried in the sand, and tips of the siphons protruded from the surface. Ninety-eight per cent of the sub- stratum was between 250 and 710 um in size (medium to coarse sand) and 50.7% was retained on a sieve of 550 um pore size (medium sand). The species conformed to that of B. digonas (Abbott 1951), and could be collected only during the coldest months, November through February. Its developmental stages from early zygote to juvenile and adult are shown in Figure 1. Early developmental stages of B. digonas differ from those of B. pilularis (Fig. 2): body form of B. digonas is more angular than that of B. pilularis, and ampullae of B. digonas are club-shaped, whereas those of B. pilularis tend to be of the same diameter throughout their length. Develop- mental stages of B. digonas resemble those of E. arenosa given by Berrill (1931) except the tunic is more highly developed and ampullae are present. Also, E. arenosa has a single gonad. ConcLusIon—The data presented here support Abbott’s (1951) separa- tion of B. digonas from B. pilularis by anatomical differences among adults. Differences occur in developmental stages as well. It is possible that B. pilularis does not occur in the northern Gulf of Mex- ico. Only two publications (van Name 1945, Cooley 1978) other than Abbott (1951) reported collections of Bostrichobranchus from the Gulf of Mexico, and they were of a digonadal species. Although lack of evidence does not prove that B. pilularis is absent from the Gulf of Mexico, information in the published literature indicates the presence of B. digonas from central to northwestern Florida. Bostrichobranchus digonas has not been reported from southern Florida, perhaps because it requires the cooler temperature of higher latitudes in winter. Contribution No. 643 from the Gulf Breeze Laboratory. LITERATURE CITED AssorTrT, D. P. 1951, Bostrichobranchus digonas, a new molgulid ascidian from Florida. J. Wash. Acad. Sci. 41:302-307. Berriti, N. J. 1931. Studies in tunicate development. Part II. Abbreviation of development on the Molgulidae. Phil. Trans. Roy. Soc. London, Ser. B. 219:281-346. Coo ey, N. R. 1978. An Inventory of the Estuarine Fauna in the Vicinity of Pensacola, Florida. Florida Dept. Nat. Resources, St. Petersburg, Florida. Publ. No. 31. 199 p. PLoucu, H. H. 1978. Sea Squirts of the Atlantic Continental Shelf From Maine to Texas. The Johns Hopkins University Press, Baltimore, Maryland. 118 p. vAN Name, W. G. 1945. The North and South American ascidians. Bull. Am. Mus. Nat. Hist. 84:1-476. VeRRILL, A. E. 1871. Description of some imperfectly known and new ascidians from New En- gland. Amer. J. Sci., Ser. 3, 1:54-446. Florida Sci. 52(1):100-103. 1989. Accepted: June 22, 1988. Engineering Sciences COMMUNITY WASTE TO ENERGY SYSTEM TECHNOLOGIES (CWEST) ALEX E.. S. GREEN Clean Combustion Technology Laboratory, Department of Mechanical Engineering, University of Florida, Gainesville, FL 32611 Asstract: The problem of solid waste disposal by small and medium size communities in low population density regions is addressed in the context of major national problems. In particular, the merits of coburning community waste with locally grown or gathered biomass together with sewage generated biogas and natural gas and coal as needed are examined. What emerges is a strategic approach in which local public problems are solved in a fashion which helps alleviate national problems rather than in isolation. In this approach, factory-fabricated modular waste and biomass to energy systems are installed in local communities and operated by well-trained personnel. This distributed solution has many advantages with respect to a central landfill or mass burn facility including: (1) increased incentives for recycling and separation at the source of toxic and hazardous waste, (2) reduced capital costs, (3) reduced transportation cost, (4) reduced emissions, (5) lower ambient pollutant concentrations, (6) reduced ash toxicity, and (7) the avail- ability of inexpensive energy in the form of steam, hot water, or hot gas. Several possible uses for this energy, which can stimulate the local economy, are discussed. Our work points to the near- term availability of community waste to energy system technologies (C WEST) and also the need to further develop this approach to energy security and environmental protection. THE behavior of the stock market commencing on October 19, 1987 (Black Monday) is without doubt an expression of our economic insecurity because of national problems which have developed over decades. At this time, when faced with a specific public policy problem, it behooves us to take a strategic approach and seek solutions which also help mitigate our major national problems. Regional waste disposal is an example of a specialized problem which can be addressed by elected officials from such a strategic standpoint rather than the usual tactical approach which considers the local problem in isolation. This report describes CWEST, a strategic approach to waste disposal in low population density regions. In this approach small com- munities or groups of communities coburn their waste with locally grown or gathered biomass, along with domestic natural gas and coal as needed, in factory built modular community waste to energy systems (CWES) in the 10- 100 ton per day (tpd) range. This approach is a viable alternative to waste disposal in local garbage dumps—or regional landfills. Table 1 lists some of our major national economic, technological, and environmental problems, not necessarily in order of seriousness. These are manifesting themselves in social problems such as family breakdown, drugs, homelessness, etc. This accumulation of major problems may be due to our current weakness in strategic solutions and our concentration on tactical solu- tions to public problems and certainly to private problems. This tactical reli- ance is fostered by fragmentation of administrative missions, increasing reli- ance on internal adversarial systems, overspecialization, and acceptance of No. 2, 1989] GREEN —CWEST 105 TABLE 1. Major economic, technological, and environmental problems. Problems Energy insecurity Decline in farming Trade deficit Waste disposal Budget deficit Urban ozone Shortage of liquid fuels Water degradation Decline in competitiveness Acid rain Decline in manufacturing Greenhouse effect things as they are rather than as they could be. If, for example, we take a strategic approach to a local or regional waste disposal problem we would consider it in the context of these national problems. For example, we would then take advantage of the fuel value and recoverable assets of waste by utilizing the latest developments in clean combustion and resource recovery technologies. To do this we must maintain a balanced assessment of the bene- fits along with the potential risks of waste combustion and energy and re- source recovery. It would be an insuperable obstacle to the utilization of waste energy to insist upon zero risks, or risks which are infinitesmally small compared to risks we accept in our daily lives, e.g. automobile use, sports, ultraviolet sunshine, radon, alcohol, red meat, sweets, medicine, etc. Today, programs and regulations fostered by our Department of Energy to achieve energy security are often negated by programs and regulations fostered by our Environmental Protection Agency to achieve neglible environmental risk. TABLE 2. Proven fossil fuel reserves in Q = quads = 1015 BTU Oil Gas Coal Oil Gas Coal United States 200 200 7,300 #£Middle East 2,310 900 = Mexico 280 80 50 ~=—s Africa 330 190 1,650 Canada 50 ~=100 170 = India 20 20) Wao South America 200 110 310 ~=Indonesia 80 8130 — Australia 10 20 840 China 120 30 =©2,790 Europe 140- 210 2,590 USSR 370 1,510 8,190 World 4,110 3,500 25,000 Our lack of energy security is perhaps our most serious long range prob- lem since energy is to a nation what blood is to a living being. Table 2 summa- rizes the world reserves of fossil fuels (Leibson, 1985). As shown in Table 2, the United States is well endowed with coal; unfortunately, it is a major consumer of oil, largely by a transportation infrastructure that goes back to the beginning of the Twentieth Century. Table 3 gives the U.S. consumption of energy in 1987 (EIA, 1988). Our total energy use was about 79 quads, of which 41% was from oil, 23% from coal, 22% from natural gas, 6% from nuclear, 4% from hydroelectric and 4% from biomass and other renewable energy sources. Our large reliance on oil energy is a basic cause of our energy insecurity. Crude oil prices since 1970 106 FLORIDA SCIENTIST [Vol. 52 TaBLE 3. US consumption of energy in 1987 (quadrillion BTUs) (all 79 quads, all fossil 67.8 quads) Natural Coal Gas Oil Nuclear Hydro Biomass* Residential 0.1 4.4 1.4 — — 0.7 Commercial 0.1 2.5 12 — — 0.3 Industrial Mandl 6.9 8.1 -- 0.3 1.3 Electric utilities 15.1 ene] 1.3 4.9 3.0 0.7 Transportation = 0.5 20.6 — — Total 18.0 ie2 32.6 4.9 3.3 3.0 Percentage 22.8 21.8 41.3 6.2 4.] 3.8 *Estimated (personal communication R. L. San Martin, 1988) 70 Outbreak of lran/Iraq War U.S. Oil Price Decontrol OPEC Decision 1985 Iranian Revolution To Regain Dollars 40 per Barrel* Market Share UN ee Embargo ios ll a 1970 1972 1974 1976 1978 1980 1982 1984 1986 * Average quarterly cost of crude oil imported by U.S. refiners. Fic. 1. Oil Prices and International Events (EIA 1988) in 1985 dollars per 42 gallon barrel (EIA, 1988) shows three significant fea- tures (Fig. 1). The steplike jump (1) reflects the first OPEC initiated oil shock of 1973. The larger increase (2) beginning in 1979 has been called the second oil shock. The precipitous decline in 1986 (3) caused by the OPEC decision to regain market share has been called the third oil shock. The slow decline beginning in 1981 was due in part to the excess oil capacity which was stimu- lated by the high oil prices and also due to U.S. oil price decontrol. Our production and consumption of the major fossil fuels are compared (Fig. 2). It is noteworthy that our domestic production of gas has been in No. 2, 1989] GREEN — CWEST 107 Consumption Quadrillion Btus Production £970 1974 1978 1982 1986 Fic. 2. Production and Consumption of Fossil Fuels in the United States. Downward arrows indicate extent of petroleum deficit. Upward arrows, coal export. balance with our consumption. Our production of coal has exceeded our consumption and we are exporting a small fraction of our production. Our big problem has been the large gap between our production of oil and our consumption (see arrows). Some progress was being made in developing envi- ronmentally sound domestic alternatives to oil in the utility and industrial sectors (FEA 1974, Rider 1981, Green 1981, DOE 1983, Green and Pamidi- mukkala 1984, Green et al. 1986). However, the third oil shock at the begin- ning of 1986 took the momentum out of this trend. Abelson (1987) has re- cently noted that “The United States is failing to ensure against a major economic disaster that might be brought on by a severe shortage of liquid fuels.” He also notes that “the free market is not functioning very well to enhance the energy security of this country.’ Community waste to energy system technologies (CWEST) can help alleviate this liquid fuel crisis, first by providing alternative energy sources for perhaps 10 oil quads used in the industrial, commercial and residential sectors (see Table 3). Perhaps later CWEST can serve as a source of liquid fuels to reduce oil consumption in the transportation sector. 108 FLORIDA SCIENTIST [Vol. 52 DiscussioN—Co-combustion in Energy Recovery Systems—Prior to the Clean Air Act of 1970 and the first oil shock of 1973, it was customary to design furnaces and boilers to use the least expensive and most available fuel. With the high cost of capital equipment and with programs extending boiler life from 30 years to 50 years or so, there is now a great need to develop the technologies of retrofitting and fuel blending for maximum energy efficiency and minimum harmful emissions. An incinerator with energy recovery is intrinsically a versatile fuel utilization system which can accommodate a large number of co-combustion combinations. The technology and economics of co-combustion as a strategy for reduc- ing our reliance on imported oil, improving combustion and reducing harm- ful emissions are still in early stages of development. To a large extent the need will be dictated by the available combustion system and the characteris- tics and complementarities of available fuels or wastes. A number of qualities of fuels including various waste categories that should influence co-combus- tion choices are illustrated (Fig. 3A). From a technological standpoint, heat- ing value is a primary variable and it is thermochemically useful to blend lower heating value waste with higher heating value fuels. Heating values of standard trash components along with those of representative biomass and fossil fuels are listed elsewhere (Niessen 1978, Singer 1981). The hydrogen/carbon (H/C) atomic ratio (Fig. 3B) is a thermochemical variable which influences the speed of the flame chemistry. The increased hydrogen fosters the formation of free radicals such as H and OH, which are important intermediate links in reaction chains. Also, the larger the H/C ratio, the larger the water vapor/carbon dioxide fraction in the exhaust. The H/C ratio is closely correlated with physical phase (e.g. solid, liquid or gas) which governs the macroscopic characteristics of the combustion process. The moisture content of the fuel is a major factor in burning. Heating values of standard trash components (Fig. 3C) are influenced by their mois- ture content. For the sake of comparison it takes 1000 Btu to evaporate one pound of water. Today, the emissions of fuels and waste and the toxicity of the ash residues are important variables in the co-combustion equation. Figure 3D gives a pictorial presentation of environmental factors. Biomass occurs between No. 2 oil and No. 6 residual oil. Community and institutional waste are placed to the left of coal by virtue of the heavy metals, halogenated plastics and other hazardous or toxic materials now in these waste streams. This position could be shifted to the right by recycling, separation of toxics at the source and by hazardous waste reduction programs (OTA, 1987). Co-combustion with bio- mass, No. 2 oil or natural gas also shifts the position to the right. Market price is the dominant decision variable today and hence the costs of the fuels useful in co-combustion with institutional or urban wastes must be considered. For most waste the disposer pays a tipping fee, so the incinera- tor operator can list waste itself as a negative fuel cost or as an income. For example, a $20/ton tipping fee with waste averaging 5000 Btu/lb corresponds No. 2, 1989] GREEN —CWEST 109 H G_ Re Ru, T ets OILS Wes Wd S AB B €@ 5421 P m h alee ca) i 0 5 10 15 20 25 60 62 SOLIDS LOU sS—— — a GASES ----- es @ A B: 28 6 W 21 Ge by fp e m h : 1 es ; (B) H/C 0 1 2. 3 4 @ ee ee ee a Wa Coal 6 Wd 2 M,E m G | | | | | | (D) Emission HIGH ah Fig. 3. Fuel Properties. (3A) Heating value of various fuels in 1000 BTU/lb. The symbols denote the following: H - Human and animal remains, G - Garbage, Re - Refuse, Ww - Wet wood, Ru - Rubbish, L - Lignite, T - Trash, Wd - Dry wood, S - Subbituminous, A - Anthracite, B - High volatile Bituminous, B - Low volatile Bituminous, 6 - No. 6 oil, 5 - No. 5 oil, 4 - No. 4 oil, 2 - No. 2 oil, 1 - No. 1 oil, p - Propane, m - Methane, h - Hydrogen. (3B) Representative Hydrogen/Carbon atomic ratios for various hydrocarbons. The additional symbols denote the following: C - Coke, W - Wood, g - gasoline, b - Butane, e - Ethane. (3C) Moisture contents of various wastes in percentage. S] denotes Sludge. (3D)Qualitative environmental emission factors for various categories of fuels. Wa represents waste as a broad category, M - Methanol, E - Ethanol. to a negative fuel cost or income of $2/MMBtu. Figure 4 shows negative costs as downward arrows along with typical cost or prices of biomass energy, fossil fuel energy and electrical energy for mid 1985, 1986, 1987 and a projection to 1992 (AGA, 1987). The high variability of energy prices (illustrated in Fig. 4) is a major source of confusion and uncertainty in waste to energy decisions and indeed all alternative energy programs in this country. For example, as of mid-1985, coal had a substantial energy price advantage, although natural gas was more advantageous from the standpoint of reduced emissions. In 1987, after the drastic reduction of crude oil prices by the OPEC cartel, natural gas became the most favored co-combustion fuel, from both stand- points. Thus, from the market price standpoint, the choice of the optimum co-combustion fuel in energy recovery incineration may change rapidly with time. Accordingly, there is a strong need to incorporate co-fuel flexibility in new or retrofitted waste to energy systems, and indeed other steam genera- tion systems, to accommodate various fuel combinations, fuel qualities, fuel prices and tipping fees. Thermodynamic losses associated with the conversion of heat to mechani- cal energy and then to electrical energy account for the high cost of electrical energy (see E in Fig. 4). If the heat is used directly (e.g. in sludge drying) or converted to process steam which is used directly (e.g. in district heating) much greater benefits are obtained from the waste-biomass energy. 110 FLORIDA SCIENTIST [Vol. 52 Coal NG 6 Z E Coal 9G NG 2 1986 $3 Coal ~=sCWW N 6 : 1987 33 Cgal “? N 6 E 1 2 3 4"wa F992 5 S10 15 20 Dollars per Million BTU Fig. 4. Fuel Prices. Economic comparison of fuel costs to industrial market ($/MMBTU) for mid-year 1985/6/7 and anticipated 1992. Tipping fees for waste, (5M BTU/lb) assumed to go from $20/ton to $40/ton in 1992, are presented below the line to represent a gain. E denotes electrical energy (note factor of 10 scale change). Cultivated Biomass for Energy and Combustion Support—Much of the co-combustion fuel for supporting the burning of the waste from small com- munities or for supplementing waste energy can be furnished by locally har- vested high-yielding energy crops. Table 4 provides a list of short rotation intensive culture (SRIC) crops which show the greatest promise for annual dry biomass yields (Rockwood, 1986). These can roughly be translated into barrels of oil equivalent by multiplying by 3. Prine and Mislevy (1983) have pointed to perennial annual regenerating plants as a source of biomass for energy use in colder subtropics and/or warmer temperature climates of Flor- ida. Two types of these plants should be particularly useful. The first is tall C, grasses with full season growth potential such as napiergrass and sugarcane. The second is perennial tropical shrubs such as leucaena (Othman and Prine, 1986). Both the grass and the leucaena are normally killed by freezes and the plants are regenerated in the spring from underground plant parts. A disad- vantage of leucaena is that harvesting can take place only in late fall and winter at the end of the growing season and some method of storing the biomass until it is needed is necessary. In the case of the grasses, harvesting can be carried out while growing. If harvested green, then some method of removing part of the water is necessary for most energy uses. A drier using heat from a waste to energy system arranged to coburn biomass and institu- tional waste should serve well in this regard (White and Plaskett, 1981). Waste Biomass for Energy and Combustion Support—In addition to cul- tivated biomass, waste biomass which is abundantly available in agricultural regions can greatly broaden institutional waste and biomass co-combustion options and provide a more continuous fuel supply. The forms of waste bio- mass include: (1) sawdust, a by-product of log and lumber cutting operations No. 2, 1989] GREEN—CWEST Ii TaBLeE 4. Approximate yields from high biomass production species Common name tpa-yr? Common name tpa-yr Leucaena 13 Sugarcane ap) Eucalyptus 10 Napier grass 20 Melaleuca 5 Kenaf 20 Casuarina 5 Sudan grass (sorghum) 19 Slash pine 5 Corn (zea mays) 11 Sand pine 4 Alfalfa 10 tpa-yr is dry tons per acre per year varying in size from a fine dust to pieces nearly as large as a small finger nail; (2) wood chips consisting of larger chunks of wood from tree limbs and main stems available from line clearing operations; (3) scrap lumber from fabrica- tors of roof trusses and from local home builders; (4) logging debris, the residue left over from logging of tree farms, a massive source in agricultural regions such as north Florida; (5) urban tree trimmings, which represent a steady source of waste biomass; (6) Christmas trees which become abun- dantly available after New Years, when heat demands are high; (7) weeds, undesirable shrubs, lawn and garden trimmings; (8) crop residues including corn and grain stalks and spoiled hay and (9) paper, when energy demands are high and recycling value is low. It is estimated that such forms of biomass are available at costs in the range from $0.3 to $3/MMBTU. Currently fossil fuels cost between $1.5 to $4/MMBTU (see Fig. 4). Sewage from community waste water treatment plants or septic tanks can be a large source of biomass which can be used in a variety of ways. It can be a major feedstock for the anaerobic production of biogas (Chynoweth et al., 1983). This in turn can be used to support the burning of community waste in waste to energy systems. The residual sludge, after drying with the help of settling ponds or tanks and belt presses and subjected to high temperatures for evaporation of water and sterilization can be pelletized as a fertilizer (Public Works, 1981). If the original sewage is free of hazardous industrial waste it can be used for food crops. If not, it can still be used for energy crops and horticulture applications. When the pelletized sludge exceeds market de- mands it can be used as an inexpensive coburning fuel to support the combus- tion of ordinary community waste and for supplementary energy. Aquatic plants, primarily water hyacinth, are being used in several areas of the southern United States for nutrient removal from secondarily treated sewage effluent. The potential yields of water hyacinth exceed the biomass yields of many subtropical terrestrial, salt water and freshwater plants (Bagnall, 1986). However because of its high water content (>95% ) the use of water hyacinths to produce methane by anaeorobic digestion (Chynoweth et al., 1983) or to produce compost is more advantageous than its direct use as a fuel. Indeed using sewage and aquatic weeds in anaerobic digesters to gen- erate biogas which is then used for coburning with community waste and 112 FLORIDA SCIENTIST [Vol. 52 TO BOILER fi FOSSIL FUEL BURNER SECONDARY CHAMBER = aNd, DSO ee PRIMARY CHAMBER TX KID Bean, 106:a6 A ass SEs Mos tes a vas ee: ieaseseaeaa ie ete’) a sei ‘ orscive FEED RAM Ze ee &a ae Peet oer alg Bae St 5S iy jie, jee ASH TRANSFER RAMS lor arent ASH DISCHARGE RAM — aa ASH CHUTE ASH QUENCH Fig. 5. Typical Modular Incinerator (adapted from DER 1983) wood in waste to energy systems could be an important part of integrated waste disposal approach. Finally we should mention plastics which is now developing into a large component of the every day waste stream. Hydrocarbon plastics free of halo- gens have high heating values and H/C ratios and can serve as excellent and environmentally benign support fuels for garbage incineration. Halogenated plastics, unfortunately, produce high outputs of hydrochloric acid and when improperly burned produce other hazardous emissions and ash. Both sets of problems could be avoided or minimized if halogenated plastics were kept out of the everyday waste stream or subjected to strong recycling incentives. The recycling of plastics is now developing rapidly (Weis et al., 1988). Uses for Energy from Waste-Biomass to Energy Systems— Heavily popu- lated regions of the country, because of shortage of landfill space, are already resorting to large mass burn or refuse derived fuel (RDF) incineration for waste volume reduction. The present trend is to equip these large units to produce steam for electricity generation. Sparsely populated, forested and agricultural regions of the country can also help in reducing oil imports and reducing the use of non-renewable energy sources. Modular incinerators in the 10-100 ton per day range, which usually have a starved air first chamber and an afterburner second chamber (DER, 1983) can be prefabricated at reasonable cost. Figure 5 illustrates the main features of a typical modular incinerator. No. 2, 1989] GREEN —CWEST 113 In modular thermal systems, biomass can be co-combusted with commu- nity waste to provide a useful fuel multiplier to bring the output levels of the energy system to a critical economic level. Uses must be found for the heat, low grade steam or hot water. Industrial processes, crop drying, distillation of methanol and ethanol, desalinization, fish and alligator ponds, dairy farms, laundries, and district heating or cooling (with the aid of absorption chillers) could be large outlets. Biomass production, sludge drying, heat treatment, and conversion to organic fertilizer at municipal waste water treatment plants are also ecologically sound uses for surplus energy. For medium sized communities, a useful strategy might be to locate modular CWES at several locations e.g. industrial parks and water treatment plants where the low grade steam can be used directly. For larger communities, tire collection and restaurant grease collection centers could be located at one or two sites with more stringent emission controls to convert these wastes to energy. The pyro- lytic first stage and afterburner second stage of a typical modular incinerator make possible a level of emission control which is difficult to achieve in a mass burn type incinerator (Makansi 1987, Oppelt 1987, Howes et al. 1987). Electrical energy can be produced with reasonable efficiency if the energy recovery boiler is equipped with a superheater to produce high pressure-high temperature steam. However, unless this energy is used locally it would be difficult to recover capital costs at current utility avoided costs. A comprehensive small community waste to energy and recycling pro- gram is illustrated (Fig. 6). Toxic and hazardous materials are first separated from the waste output for disposal at a hazardous waste depository. Sewage is channeled to an anaerobic digester to produce biogas and residual sludge. Both of these products are sent to the waste to energy system (WES). Materi- als which can be recycled after cleaning or repair are disposed of by garage sale or Goodwill type outlets. Other materials can be blended with virgin feedstocks with the help of recycling centers. The waste to energy channel displays a number of illustrative uses for the energy recovered from the resid- ual community waste and energy obtained from locally cultivated or waste biomass. The waste disposal and multifuel capability of an incinerator makes it economically competitive with specialized wood chip or gas boilers. Advantages of CWEST—A recent cost estimate of a 1200 tpd two unit mass burn facility to serve 13 counties in North Central Florida was $112,800,000 (CDM, 1987). This is roughly consistent with the $100,000/tpd used to estimate capital cost of current mass burn units which generate elec- tricity and have extensive emission controls. Modular incinerators range in capital costs from $10,000/tpd to about $80,000/tpd depending upon the au- tomation of feeding and ash removal, the energy recovery system, the use of a superheater to obtain high pressure-high temperature steam, and the exten- siveness of the post combustion emission controls. If qualified for hazardous waste destruction, a 100 tpd unit could cost as much as $50,000,000 corres- ponding to $500,000/tpd. 114 FLORIDA SCIENTIST [Vol. 52 WASTE OUTPUT SEWAGE BIOMASS NATURAL GAs| |_Digester COAL Biogas Sludge bole Werte to Energy Clothes Newspaper Cardboard Toys Fumiture Aluminum Building Steel Industrial Steam Heating & Cooling Local Electricity Biogas Production Sludge Treatment Biomass Drying Desalinization Distillation Mineral Drying Special Crops Landfill Fig. 6. Proposed comprehensive community waste to energy and recycling program. Materials | | Copper Appliances | | Glass Electronics | | Plastics Rather than a mass burn unit for a large region in low population density agricultural area, the obvious alternative is to use several appropriately sized community waste to energy systems at various locations. The advantages of this distributed solution are manifold. In the first place package unit prices for 25 to 50 tpd modules with low grade steam output and simple emission controls usually range below $50,000/tpd, about half that of a large field constructed unit built for electricity generation. The lower transportation cost for CWES is an obvious additional benefit. The emissions from a refrac- tory walled modular CWES are also less harmful than the emissions of a water walled mass burn unit (Oppelt 1987, Howes et al. 1987). Because of the low emission output for each large geographic region the ambient concen- tration of pollutants will certainly be much lower than the ambient concen- No. 2, 1989] GREEN —CWEST ls trations in the vicinity of a centralized mass burn facility or possibly a cen- tralized landfill. Furthermore, having each small community burn its own waste in its own backyard provides a tremendous incentive for recycling and separation at the source of hazardous and toxic substances. These incentives can be reinforced in a community by charging districts which deliver a good waste fuel, free of hazardous material, a low garbage collection fee. Such measures should hold the emissions of pollutants and the ash toxicity to low levels and greatly reduce the cost of emission control and ash disposal. In the absence of such environmental concern, local residents would find the haz- ardous waste they send to their CWES would return in the air they breathe or the water they drink. In addition to direct earnings from recyclables the extension of landfill lifetime is an additional incentive to conduct a vigorous recycling program. The availability of low cost energy can also revitalize many small commu- nities in agricultural regions. It will take planning, ingenuity and some risk taking to cultivate such outlets. The replacement of food or tobacco crop agriculture with energy crop agriculture can alleviate some of our national problems. In this connection the production of alcohol fuels, methanol (wood alcohol) or ethanol (grain alcohol) would be very appropriate outlets for bio- mass and waste energy generated in small CWES. Congress is in the process of legislating incentives to encourage use of methanol, ethanol or natural gas in vehicles to mitigate our liquid fuel problem and to reduce urban ozone and acid rain. Farm vehicles converted to methanol might lead this country in successful oil backout and environmental protection movements. A more immediate impact of the development of CWES is the possibility of displacing oil in the industrial, commercial, and residential sectors (see Table 3). In 1987 these sectors consumed 14% of our total energy use and about 33% of our petroleum. Replacing this oil with as much renewable energy as possible would bring our consumption of oil much closer to our domestic production (see Fig. 2). This approach together with the develop- ment of alcohol or natural gas fueled vehicles is probably our shortest path to energy security. It is estimated that renewable energy could provide 10% to 20% of our total energy needs by the year 2000 AD. Southeastern states and particularly Florida with longer growing seasons should even do better. Conc.usions—In high population density regions landfills are too costly and waste to energy systems are almost an inevitable accommodation to real- ity. This possibly explains why Japan and Europe are now leaders in waste to energy technologies (Makansi, 1987). In low population density regions the landfill option may calculate to be the cheapest solid waste disposal method from a tactical or local standpoint. However, from a strategic or overall na- tional standpoint this mode of waste disposal has few redeeming features and makes little contribution to the solution of other national problems (Table 1). Furthermore, this solution seriously limits the future growth of energy farms 116 FLORIDA SCIENTIST [Vol. 52 and waste wood utilization which are natural economic outlets for such re- gions. The decline in income potential and the consequent sacrifice of quality of life might then outweigh the immediate financial savings to this region. Modular CWES which yield the least stack emissions of refuse to energy systems (Howes et al., 1987) are also favored by obvious considerations of pollution dispersion. When used in the biomass and waste co-combustion mode and operated by well trained personnel, this sensible solid waste man- agement option can contribute to the solution of many national problems (Table 1). If source separation is not fully adequate, future systems could incorporate new emission control and ash stabilization technologies devel- oped in our national programs on clean coal technologies (Blackmore and Leibson, 1986), hazardous waste (NUS, 1987) and resource recovery (Clarke, 1987). In final summary, this work proposes the use of community waste to en- ergy system technologies (CWEST) for low density population regions. Our CWEST has several major features (see Figure 6): (1) toxic and hazardous substances, usually a small component of the everyday waste stream, are first removed from the waste output for disposition at a depository, (2) recyclable materials, a much larger component, are also removed for direct or material reuse, (3) sewage is anaerobically digested to produce biogas which is burned in the thermal system for combustion support and energy recovery. The sludge is dried, sterilized, and used as a fertilizer or burned for energy, (4) the residual non-hazardous-waste (NHW) is coburned together with biomass and biogas along with the domestic fuels natural gas and coal (as needed) to support thermal waste destruction and to meet energy needs. The facilities consist of factory-fabricated modular thermal systems in the 10-100 ton per day range (or larger as the technology advances) which are located close to where the energy can be utilized. Components of CWES are already avail- able. However, there is a great need to further develop this multidimensional approach to energy security and environmental protection. Since CWES maximizes the use of renewable energy and minimizes the use of fossil fuels it mitigates several national problems, including the greenhouse effect. CWES, a strategic solution of waste disposal problems for small and mid- dle sized communities could, with some modifications, also be applied in large communities. Thus institutions, condominiums, apartment complexes, hotels, office buildings and shopping centers etc. could incorporate an inte- grated waste to energy heating and cooling system with provisions for recy- cling and hazardous waste separation. In Florida, since most populous re- gions are along the coast and inland regions are agricultural, nearby sources of biomass for supplementary fuel are available. Furthermore, natural gas and a coal stockpile would provide energy during periods of low renewable fuel availability. Heavier waste disposal requirements, say in 400-600 tpd range could be handled by multiple combinations of smaller units— an ar- rangement which provides useful redundancy and opportunities for preven- tative maintenance. No. 2, 1989] GREEN — CWEST W177 Recent state legislation (Kirkpatrick et al., 1988) provides a strong vehicle for Florida to assume national leadership in waste disposal. If officials at the city, county and state levels make a dedicated commitment to strategic solu- tions of public problems their example might be followed by private citizens in approaching other problems. The combined impact of such strategic solu- tions could mitigate many national problems, reverse our national decline and help us reassume our role of world leadership. ACKNOWLEDGMENTS— This proposed solution to solid waste disposal by small communities draws extensively upon pilot studies (Green et al. 1986a, b, 1987, 1988) at an institutional level. The author would like to thank his collaborators and the sponsors (the Florida Governor’s Energy Office, the Tennessee Valley Authority's Southeastern Regional Biomass Energy Pro- gram, the University of Florida, and the Sunland Training Center) for the knowledge gained while executing these studies. He would also like to thank Robert San Martin, Dean F. 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Power. 11-30. Niessen, W. 1978. Combustion and Incineration Processes. Applications in Environmental Engi- neering. Marcel Dekker, Inc., New York, NY. Nus Corp. 1987. Second International Conference on New Frontiers for Hazardous Waste Man- agement. Sept. 1987. EPA/600/9-87/018F. OppeELT, E. T. 1987. Incineration of hazardous waste, a critical review. JAPCA. 37:558-586. OTA. 1987. From Pollution to Prevention: a progress report on waste reduction. OTA-ITE-347, Congress of the United States, Office of Technology Assessment. OTHMAN, A. B. ANDG. M. Prine. 1986. Biomass Production and Nutrient Removal by Leucaena in Colder Subtropics. ibid. Biomass Energy Development. pp. 95-102. Prine, G. M. AND P. Misevy. 1983. Grass and Herbaceous Plants for Biomass. Proc. Soil and Crop Sci. Soc. Fla. 42:8-12. Pusuic Works. Oct. 1983. New Use of Incinerator Cuts Cost of Sludge Drying. Riper, D. 1981. Energy: Hydrocarbon Fuels and Chemical Resources. Wiley, New York, NY. Rockxwoonp, D. L. 1986. Development of woody biomass cultural systems for Florida. Pp. 85-94. In: Smitu, W. H., (ed.) Biomass Energy Development. Plenum Press, New York, NY. SAN Martin, R. L. 1988. Assistant Secretary for Biomass, Department of Energy, Washington, D. C. Personal Communication. SinceER, J. 1981. Combustion-Fossil Power Systems. Combustion Engineering, Inc., Windsor, CT. Weis, R. S., W. E. Pearson, AND D. R. Morrow. 1988. Plastic Recycling. A Strategic Vision. Plastic Recycling Foundation. Washington, DC. Wuite, L. P. AND L. G. PLasketr. 1981. Biomass as Fuel. Academic Press. New York, NY. p. 211. Florida Sci. 52(2):104-118. 1989. Accepted: August 23, 1988. Biological Sciences THE DISTRIBUTIONS OF THE TURTLES OF FLORIDA JOHN B. IVERSON AND Cory R. ETCHBERGER ()Department of Biology, Earlham College, Richmond Indiana 47374, and 2) Department of Biology, Indiana University, Bloomington, Indiana 47405 Asstract: The distributions of all 25 turtle species recorded from Florida (including 17 fresh- water, five marine, one estuarine, and two terrestrial species) are mapped and reviewed. Fresh- water turtle species density in Florida is among the highest in the world, with 13 species known to co-occur in the Escambia River, and possibly also in the Apalachicola, Wakulla, and Suwannee river basins. Although at least eight exotic species have been introduced into Florida, the only confirmed breeding populations of non-native forms are of Trachemys scripta elegans. ALTHOUGH the highest species density of turtles in the western hemisphere occurs in the southeastern United States (Iverson, 1989), the distributions of turtles in Florida have not been mapped in detail since Archie F. Carr’s dissertation fifty years ago (Carr, 1937). This paper examines the distribu- tions of Florida’s 25 species of turtles, and points out the major distibutional and taxonomic problems. Fossil sites beyond the present range of a species are also reviewed. The pattern of turtle species density in the State’s major drain- age basins is also examined. MeEtHops— Distribution maps of each turtle species found in Florida were prepared from a combination of museum and literature records, the field notes of colleagues, and our field work. Although not all records were verified by examination of every voucher specimen, specimens from the margins of the species range (especially outliers) were usually examined. Localities were plotted on a base map with both hydrographic and political features (Fig. 1), because turtle distributions tend to correspond to hydrography and topography even though they are often presented with respect only to political boundaries (e.g. Collins, 1982; Ashton and Ashton, 1985). Taxonomy and common names follow Iverson (1986) and Meylan (1987; i.e., Apalone for Tri- onyx), and subspecies are not discussed here. Museum designations follow Leviton and co-work- ers (1985), except for the Florida State University collection (FSU), George Mason University (GMU), and the Tall Timbers Research Station collection (TT), which has recently been donated to the Florida Museum of Natural History (UF) collection. Each species name is followed by a habitat code: FW, freshwter (/R, primarily lotic;/L, primarily lentic); M, marine; E, estuarine; or T, terrestrial. REsSuLTS—Species Accounts. Family Chelydridae Chelydra serpentina (FW/L). The common snapping turtle (Fig. 2) is distributed virtually throughout Florida, excluding the Keys. The lack of records from the upper Suwanee river basin and the drainages between the Apalachicola and the Yellow river basins may reflect inadequate collecting efforts in those areas. Macroclemys temminickii (FW/R). The alligator snapping turtle (Fig. 3) is distributed across the Panhandle of Florida but ranges no farther on to the peninsula than the Suwannee River basin (Allen and Neill, 1950; Pritchard, 120 FLORIDA SCIENTIST [Vol. 52 Fic. 1. Drainage map of Florida (counties borders indicated by dashed lines) with major rivers numbered near their mouths: 1) Perdido, 2) Escambia, 3) Blackwater, 4) Yellow, 5) Choc- tawhatchee, 6) Econfina, 7) Apalachicola, 8) Chipola, 9) New, 10) Ochlockonee, 11) Wakulla, 12) Aucilla, 13) Steinhatchee, 14) Suwannee, 15) Santa Fe, 16) Gainesville interior drainage basins, 17) Waccasassa, 18)Withlacoochee, 19) Hillsborough, 20) Alafia, 21) Peace, 22) Kissim- mee, 23) St. Johns, and 24) St. Marys. 1982). The hiatus in the range between the Escambia and Apalachicola ba- sins in the Panhandle may or may not reflect inadequate field work in that area. The Pleistocene fossil record extends the former range of the species in Florida to the region of Tampa Bay (Dobie, 1968; P.A. Meylan, 1987). A record (KU 61844) from the Oklawaha River (Marion Co., 6.4 km E Silver Springs) is considered an introduction. Family Kinosternidae Kinosternon baurii (FW/L). The striped mud turtle (Fig. 4) occurs throughout the Florida peninsula and Keys (Ernst, 1974; Iverson, 1978). The western limit of its range in the Panhandle is uncertain, in part because it is so similar morphologically to the non-peninsular subspecies of Kinosternon sub- rubrum occurring there. A multivariate statistical analysis of morphometric data from both species (e.g. see Lamb, 1983) from the Suwannee River basin westward is needed to clarify their distributions in the state. Kinosternon subrubrum (FW/L). The common mud turtle (Fig. 5) occurs throughout Florida excluding the Keys (Iverson, 1977a). Previous references to its presence in the Keys (e.g., Conant, 1975) were based on misidentified, darkly pigmented specimens of K. baurii (Iverson, 1978). No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA DA | Fic. 2. Distribution of Chelydra serpentina in Florida. Sternotherus odoratus (FW/L). The common musk turtle (Fig. 6) occurs virtually statewide, excluding the Keys (Reynolds and Seidel, 1982). The scarcity of records from west of the Apalachicola basin may reflect lack of collecting there. Sternotherus minor (FW/R). The loggerhead musk turtle (Fig. 7) occurs across the entire Panhandle and southward on the peninsula to the Withla- coochee and St. Johns drainages (Iverson, 1977b, 1977c). Within its general range it appears to be absent from the Waccasassa River basin (Iverson, EO he): Family Emydidae Clemmys guttata (FW/L). The range of the spotted turtle (Fig. 8) in Florida has received a lot of recent attention (e.g. Ernst, 1972; Berry and Gidden, 1973; Banicki, 1981); however, its secretive habits suggest that much remains to be learned about its Florida distribution. Isolated records are known from the St. Mark’s National Wildlife Refuge in Wakulla Co. to at least the lower St. John’s basin. The validity of the two early records from central Florida (Marion Co., Lake Weir; USNM 52394; Carr, 1940; and Polk Co., near Winter Haven; Neill, 1954; no voucher specimen known) remains 122 FLORIDA SCIENTIST [Vol. 52 Ne coeee | Fic. 3. Distribution of Macroclemys temminckii in Florida. Solid square indicates introduc- tion discussed in text. os Ay HY v \ ; op ete, Ip a Fic. 4. Distribution of Kinosternon baurii in Florida. No. 2, 1989] IVERSON AND ETCHBERGER—TURTLES IN FLORIDA 123 Fic. 6. Distribution of Sternotherus odoratus in F lorida. 124 FLORIDA SCIENTIST [Vol. 52 Fic. 8. Distribution of Clemmys guttata in Florida. Squares indicate problematic localities discussed in text. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA jae) questionable, although a recent record from Polk County (2.3 km NW I-4 on SR 557; UF 66605) suggests that the species may occur naturally in central Florida. Deirochelys reticularia (FW/L). The chicken turtle (Fig. 9) is known vir- tually statewide, excluding the Keys (Schwartz, 1956; Zug and Schwartz, 1971). Field work in the Peace River basin may reveal that the species is more common there than current records indicate. Graptemys barbouri (FW/R). Barbour’s map turtle (Fig. 10), the last species of turtle in Florida to be described (Carr and Marchand, 1942), is known in Florida only from the Apalachicola River and its major tributary, the Chipola River (Sanderson and Lovich, 1988). Its dependence on mollusk populations seems to restrict its distribution to the main channels of these rivers. Pleistocene fossils from the Santa Fe River are similar to if not identi- cal with this species (Jackson, 1975), and suggest that its distribution was once more extensive in Florida. Graptemys pulchra (FW/R). The Alabama map turtle (Fig. 10) is known in Florida only from the Escambia and Yellow River basins (Shealey, 1976; Lovich, 1985). These basins need to be surveyed thoroughly to determine the extent of the range of this species (and also Apalone mutica) and potential threats to its existence in the state. Its closely allopatric distribution with respect to Barbour’s map turtle reflects their closé ancestry (Cagle, 1952). Malaclemys terrapin (EF). The diamondback terrapin (Fig. 11) is an estua- rine species preferring mangrove swamps and salt marshes along almost the entire coast of Florida (Schwartz, 1955; Ernst and Bury, 1982). It is probably absent along the coast only where such habitats are lacking (e.g. the coast of southeast Florida), but the apparent range gaps along the east and panhandle coasts need to be investigated. Pseudemys concinna (FW/R). The river cooter (Fig. 12) ranges across the Gulf Coast drainages of Florida from the Perdido to the Alafia basin (Lithia Springs) near Tampa Bay (Ward, 1984; Iverson 1986). It is also known from the interior drainage basins near Gainesville (Alachua County) and at Silver Springs (Marion County), but its presence there is probably due to introduc- tion. The presence of this turtle in Lithia Springs (Crenshaw, 1955) is prob- lematical because it is unknown from the Hillsborough River drainage, which lies between the Alafia basin and the remainder of the species range. The possibility exists that this turtle was introduced into Lithia Springs; in fact, its present existence there should be confirmed. Surprisingly, no speci- mens document the ocurrence of the river cooter in the Suwannee River above the mouth of the Santa Fe. Pseudemys floridana (FW/L). The common cooter (Fig. 13) occurs virtu- ally statewide, except in the Keys (Duellman and Schwartz, 1958; Ward, 1984). Carr (1935) reported the species from Key Largo, but did not repeat the record in 1937; it is presumably unfounded. Blaney (1971) reported this species on St. Vincent Island, Franklin County, but his record is apparently based on P. nelsoni (Dobie, 1985). A specimen from Stock Island, Monroe 126 FLORIDA SCIENTIST [Vol. 52 Fic. 9. Distribution of Deirochelys reticularia in Florida. Fic. 10. Distributions of Graptemys barbouri (solid circles) and G. pulchra (stars) in Florida. IVERSON AND ETCHBERGER— TURTLES IN FLORIDA i Fic. 12. Distribution of Pseudemys concinna in Florida. Solid squares mark problematic localities discussed in text. 128 FLORIDA SCIENTIST [Vol. 52 soveera: Bt ae oe , oo Fic. 13. Distribution of Pseudemys floridana in Florida. Solid square marks introduced speci- men discussed in text. Ee Pa wS- he Fic. 14. Distribution of Pseudemys nelsoni in Florida. Solid squares mark problematic locali- ties discussed in text. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 129 County (MCZ 86984) apparently represents an escaped captive (J.T. Collins, 1987). Turtles of the genus Pseudemys have also been introduced into Blue Hole Pond on Big Pine Key, Monroe County (Iverson, pers. observ.), but the species identification is uncertain; they are probably this species or P. nelsoni. Pseudemys nelsoni (FW/L). The Florida red-bellied turtle (Fig. 14) prob- ably ranges continuously over the Florida peninsula (but not the Keys) from the St. Marys, St. Johns and Waccasassa basins southward (Jackson, 1978); however, confirmed records are still lacking from the Peace River system. Isolated problematic records exist from the Apalachicola, New, Wakulla, Ocklocknee, Steinhatchee, and Suwannee basins. Despite Ward’s (1984) opinion that this species does not occur in the Apa- lachicola basin, specimens from Calhoun County (between Scotts Ferry and Blountstown on the Chipola River; Carr and Crenshaw, 1957; UF 440-443; identification verified by Dobie, 1985 and D.R. Jackson), and Franklin County (near Apalachicola; Carr and Crenshaw, 1957; USNM 101090; and St. Vincent’s Island; UF 56347, AUM 32541, AUMP 2679; Dobie, 1985), Gulf County (base of Wewahitchka dam on Chipola River, UF 67757; and Dead Lakes near Dam, UF 67750), and the field observations of Means (1977) seem to confirm its presence there. The species may also occur in the New River basin immediately to the east of the Apalachicola basin in Franklin County, based on a specimen from “New River” (FSU 379) reported by Dobie (1985). Another specimen from the vicinity of this basin (Carabelle Beach, on beach 0.8 km SE US Hwy 98, TT 691, now UF 67791) consists only of a preserved head, and despite its apparent identification as P. nelsoni by Dobie (1985), its striping pattern is more like that of P. alabamensis. Because the specimen was found dead on the beach, and because the prevailing currents along the Panhandle are from the west, it is possible that the specimen represents an P. alabamensis that died, drifted eastward and finally washed ashore in Franklin County. Three supposed records exist for this species in the upper Ochlockonee River basin in Leon County (just outside west city limits of Tallahassee by Pritchard, 1980a; and between Tallahassee and St. Mark’s Wildlife Refuge by Pritchard, 1980a, but based on a personal communication from R. Seigel (1980); and 8 km N Tallahassee, Lake Iamonia, USNM 95765). The only record for the Wakulla River (Wakulla Springs; Carr and Cren- shaw, 1957; apparently based on UF 13545; a skeleton identified as P. alaba- mensis in the UF catalog; Auth, 1987) is a misidentified Pseudemys concinna according to Dobie (1985). The next more easterly record is from the Steinhatchee basin in Lafayette County (Hwy 51, 9.6 km N Jct. Hwy 19; UF collection fide D. R. Jackson; presently unlocatable: D. Auth, 1987). The Recent natural presence of the Florida red-bellied turtle in the Su- wannee River basin is problematic (Pritchard, 1980a), despite the existence of four Recent records and abundant Pleistocene material from at least 11 sites in the Santa Fe River basin (Jackson, 1978). Two of the records are from springs in Levy County that were well known tourist attractions early in the 130 FLORIDA SCIENTIST [Vol. 52 century (Ferguson, et al., 1947): Fannin Springs (Jackson, 1964, UF 14166 and 21474, verified by Iverson) and near Manatee Springs (Carr, 1937, 1940, UF collection; presently unlocatable; Auth, pers. comm.). The third record is a shell only (without scutes) from near the upper Santa Fe River in Alachua County, 3.2 km S Brooker (UF 7212); although its identification appears to be correct, it may have been transported there by humans for food. The fourth record is from the Okefenokee Swamp in Ware Co., Georgia in an area near the headwaters of both the Suwannee and the St. Marys rivers (Vitt and Dunham, 1980); the species is definitely known from the latter basin (Powers and Smith, 1977; Shoop and Ruckdeschel, 1986; D. R. Jackson, 1987). Pleistocene fossils of this species from Colleton County, South Carolina were reported by Dobie and Jackson (1978), who suggested that the range of P. nelsoni may once have been continuous with that of P. rubriventris. If this scenario is correct, a natural, relict distribution might be expected in north- west Florida. Thus the accurate identification of the problematic specimens mentioned above is critical. The relationship between P. nelsoni and P. alabamensis remains a prob- lem. External color characters are insufficient to separate all specimens of the two taxa, suggesting (as it did to Carr and Crenshaw, 1957) that the two forms may be related subspecifically. Field surveys between the Suwannee and Mobile River basins are badly needed, followed by detailed systematic analysis (e.g. morphometric study of scute shape variation and/or studies of biochemical variation) of the entire red-bellied turtle complex (i.e., the P. rubriventris group). Terrapene carolina (T). The common box turtle (Fig. 15) ranges virtually throughout Florida, including both the upper and lower Florida Keys (Mil- stead, 1969), and thus has the most extensive range in the state of any turtle. Trachemys scripta (FW/L). The common slider (Fig. 16) ranges across the Florida Panhandle southeastward to the base of the peninsula in the Suwan- nee River basin and the interior basins near Gainesville (Iverson, 1986). It is unknown farther south on the peninsula, except where introduced (see be- low). No records exist for the St. Marys River basin on the Georgia border in northeast Florida (despite the map in Conant, 1975), and the single record for the St. Johns River basin (UF number uncertain; Duval Co., creek near St. Johns River, near Bishop Kenny Park fide D. R. Jackson; specimen pres- ently unlocatable: Auth, 1987) is probably an introduction. The present dis- tribution of this species is surprising considering the abundance of Pleistocene fossil material for the species throughout the peninsula (Jackson, 1988). Its near parapatry with the distantly-related Florida red-bellied turtle suggests that the two species may be competitors. Family Testudinidae Gopherus polyphemus (T). The gopher tortoise (Fig. 17) is found throughout the Panhandle and most of peninsular Florida (Auffenberg and Franz, 1978, 1982), wherever well-drained soils permit it to dig its burrows. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 131 Fic. 15. Distribution of Terrapene carolina in Florida. aie Fic. 16. Distribution of Trachemys scripta in Florida. Solid squares mark introductions of Trachemys scripta elegans discussed in text. 132 FLORIDA SCIENTIST [Vol. 52 Its range is shrinking rapidly as populations are destroyed by human activi- ties, principally habitat destruction and collection by humans (Auffenberg and Franz, 1982). Family Trionychidae Apalone ferox (FW/L). The Florida softshell turtle (Fig. 18) probably occurs virtually statewide, except for the Keys (Webb, 1973a), although its distribution in west Florida deserves special attention. It has been recently introduced into Blue Hole Pond on Big Pine Key (Iverson, pers. observ. Dec. 1986). Apalone mutica (FW/R). The smooth softshell turtle (Fig. 19) is known in Florida only from the Escambia River, at the southeast margin of its range (Webb, 1973b). Apalone spinifera (FW/R). The spiny softshell turtle (Fig. 19) reaches the southeastern limit of its extensive range in northern Florida (Webb, 1973c). It is known from the St. Marys basin (UF 45478, Nassau Co., Campbell and Christman, 1980; UF 51115, Baker Co.; both verified by Iverson) on the Georgia border, and westward from the Ocklocknee basin in the Panhandle. Its absence from the Suwannee River basin in Florida suggests that Knepton’s (1956) records (as Trionyx ferox agassizi = A. spinifera fide Webb, 1962) from that basin in Berrien and Irwin counties in Georgia are in error. Possible competitive relationships among the three Florida softshell turtles in the Pan- handle deserve study. Family Cheloniidae Caretta caretta (M). The loggerhead turtle (Fig. 20) is the marine turtle most likely to be seen along any part of the Florida coast (Dodd, 1988). It still has an extensive nesting range in Florida, primarily along the Atlantic coast and secondarily along the Gulf coast. The most important nesting beach is in south Brevard County (Caldwell et al., 1959; Gallagher et al., 1972; Conley and Hoffman, 1986), but extensive nesting occurs between Cape Canaveral and Jupiter Inlet (Brevard to Martin Counties). Gordon (1983) summarizes all U.S. nesting records. The absence of nesting records along the Gulf Ham- mock coast presumably reflects the absence of suitable nesting beaches. Chelonia mydas (M). The green turtle (Fig. 21) ranges throughout Flori- da’s coastal waters (Hirth, 1980). Its primary nesting beaches in Florida are between Cape Canaveral National Seashore, southern Volusia County and Key Biscayne, Dade County (Dodd, 1982; Conley and Hoffman, 1986). A single nesting occurrence in the Panhandle (on Eglin Air Force Base property) has recently been reported, but remains unconfirmed (D.R. Jackson, 1987). Eretmochelys imbricata (M). The hawksbill turtle (Fig. 22) can be ex- pected anywhere off Florida’s coasts, but most commonly in more southerly areas associated with coral reefs (Witzell, 1983). This tropical turtle rarely nests in Florida, but isolated records are available from Juno Beach, Palm No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 133 Fic. 17. Distribution of Gopherus polyphemus in Florida. frate ADT III CC PIQENT I VOI 77 Qn ZNO FRO OOOO Ly 3 LE ee os i ben 2 IW Wace Y; ‘ ~ Yor Ns ea S CHT IYN : Sek a aie Fic. 18. Distribution of Apalone ferox in Florida. Solid square indicates introduced locality discussed in text. 134 FLORIDA SCIENTIST [Vol. 52 . als ) Fic. 19. Distributions of Apalone mutica (star) and A. spinifera (star and solid circles) in Florida. Fic. 20. Distribution of Caretta caretta in Florida. Solid circles mark nesting records; solid squares mark non-nesting records. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 135 (aS ——p hee | i ed C ob ie <\ ue Fic. 21. Distribution of Chelonia mydas in Florida. Solid circles mark nesting records; solid squares mark non-nesting records. Ai Fic. 22. Distribution of Eretmochelys imbricata in Florida. Solid circles mark nesting re- cords; solid squares mark non-nesting records. 136 FLORIDA SCIENTIST [Vol. 52 Beach County (Carr et al., 1966), Soldier Key, Dade County (Dalrymple et al., 1985); Elliott Key, Dade County and Jupiter Island, Martin County (Carr et al., 1982; Pritchard and Trebbau, 1984; Lund, 1985); and J. U. Lloyd State Recreation Area, Brevard County, Lantana Beach, Palm Beach County, and Longboat Key, Manatee County (W. J. Conley, 1987). Lepidochelys kempi (M). Although rarely seen, Kemp’s ridley turtle (Fig. 23) probably ranges throughout Florida’s marine waters (Groombridge, 1982). It nests almost exclusively on a single beach on the Gulf of Mexico in Tamaulipas, in northeastern Mexico, with small numbers apparently also nesting in Veracruz and Tabasco, Mexico and on Padre Island, Texas (Prit- chard and Marquez, 1973; Marquez et al., 1976; Smith and Smith, 1980; Groombridge, 1982). It does not nest in Florida. I have not been able to verify the Dade County record reported by Ashton and Ashton (1985). Family Dermochelyidae Dermochelys coriacea (M). The leatherback turtle (Fig. 24) probably ranges throughout Florida’s coastal waters; however, it nests only occasion- ally in Florida today (Pritchard, 1980b; Conley and Hoffman, 1986), in Walton and Okaloosa Counties along the Gulf coast (Yerger, 1965), and from St. Johns County to Dade County along the Atlantic coast (Allen and Neill, 1957; Carr, 1952; Caldwell et al., 1957; Caldwell, 1959). Other possible species—The painted turtle Chrysemys picta (FW/L) ranges within 35 km of the west Florida border in the Mobile River basin of Alabama (Mount, 1975; Ernst, 1971) and may eventually be found within the state. It is the only unrecorded turtle species that may possibly be found in Florida naturally. Obviously introduced specimens of C. p. bellii have been collected in Jackson County in the Chipola River near Marianna (in 1938; UF 1898) and in Orange County in Lake Conway (Bancroft et al., 1983). The black-knobbed map turtle Graptemys nigrinoda (FW/R) is also found in the Mobile basin in Alabama within 30 km of Florida, but despite Cagle’s (1952) erroneous record of the species in the Escambia River in Flor- ida (corrected by Dobie, 1972), it is almost certainly found only in the Mobile drainage (Lahanas, 1986). Likewise, the Alabama red-bellied turtle Pseude- mys alabamensis also occurs only in the Mobile River basin within 30 km of Florida (McCoy and Vogt, 1979, 1985; Dobie, 1985). Introduced species—Several species of freshwater turtles have been intro- duced into Florida, but only one (Trachemys scripta elegans) is known to have established breeding populations (Miami, Dade Co., Wilson and Porras, 1983; and Lake Conway, Orange Co., Bancroft et al., 1983). Iverson has also collected a single specimen in Rainbow Springs Run, Marion County (Fig. 16). The following exotic species have also been introduced in south Florida near Miami (King and Krakauer, 1966; Smith and Kohler, 1977): Chelus fimbriatus, Kinosternon scorpioides, Chrysemys picta, Graptemys kohni, several Central and South American subspecies of Trachemys scripta, No. 2, 1989] IVERSON AND ETCHBERGER—TURTLES IN FLORIDA 137 Fic. 24. Distribution of Dermochelys coriacea in Florida. Solid circles mark nesting records; solid squares mark non-nesting records. 138 FLORIDA SCIENTIST [Vol. 52 Fic. 26. Freshwater turtle species density in Florida. This map assumes that the distribution of Pseudemys nelsoni does not include the Suwannee through Apalachicola River basins. Only a single species is found in the Keys. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 139 Trachemys malonei (=T. stejnegeri), Podocnemis lewyana, P. sextubercu- lata, and P. unifilis. DISTRIBUTIONAL PATTERNS— General—Florida turtles include 17 fresh- water, five marine, one estuarine, and two terrestrial species. Six species of freshwater turtles range virtually statewide, except for the Keys; this pattern reflects the scarcity of permanent fresh water on the Keys and its abundance on the mainland. No Keys’ freshwater species is also found statewide; how- ever, the terrestrial box turtle (Terrapene carolina) ranges across the entire mainland and Keys. Florida’s freshwater and terrestrial turtles can be classified zoogeographi- cally (Table 1) as 1)wide-ranging eastern United States forms (five species), 2) coastal plain forms (ten species), with five distributed primarily along the Gulf coast and one primarily along the Atlantic Coast, and 3) primarily pe- ninsular species (four). However, despite its peninsular topography and pa- leogeographic island history (Iverson, 1977c; Neill, 1957; among others), the state of Florida has no endemic species of turtles, although Pseudemys nelsoni is nearly so, ranging only 20 km into Georgia. A composite map of all collecting localities of all freshwater turtle species in Florida (Fig. 25) reveals the areas of the state needing systematic surveys. The most significant gaps are in the upper Suwannee, the area just east of the Apalachicola basin, the Panhandle west of the Apalachicola, and the Peace River basin. Density patterns—The highest turtle species density (here synonymous with species diversity) in the world occurs in the lower Mobile basin area in Alabama with 15 freshwater species (Mount 1975; Iverson, 1986, 1989) and the lower Ganges-Brahmaputra river basin in India and Bangladesh with 17 freshwater species (12 batagurines, including the estuarine Batagur, and five TABLE 1. Distributional affinities of the freshwater and terrestrial turtle species of Florida. Coastal plain Wide-ranging Eastern US Gulf coast Atlantic coast Both coasts Peninsula Chelydra Macroclemys Clemmys Deirochelys Kinosternon serpentina temminckii guttata reticularia baurii Kinosternon Sternotherus Pseudemys Pseudemys subrubrum minor concinna floridana Sternotherus Graptemys Gopherus Pseudemys odoratus barbouri polyphemus nelsoni Terrapene Graptemys Apalone Apalone carolina pulchra spinifera ferox Trachemys Apalone scripta mutica 140 FLORIDA SCIENTIST [Vol. 52 trionychids; Iverson, 1986). Within the United States, only Texas includes more total turtle species (27) than Florida (25), but the former covers nearly five times the area of the latter. Thus, despite the fact that it is not located in the Tropics, northern Florida boasts one of the richest turtle faunas in the world. The highest diversity of freshwater turtles in Florida (i.e., excluding Ma- laclemys, Terrapene, Gopherus, and sea turtles) in the state (Fig. 26)occurs in the Escambia basin (13 species). Only one fewer species (12) is known from the Yellow, Apalachicola, Wakulla to upper Suwannee, and interior basins near Gainesville. However, if the natural range of Pseudemys nelsoni extends west across the Suwannee river to the Apalachicola basin, then the Suwan- nee, Apalachicola, and Wakulla basins would each equal the Escambia basin with 13 species present. This high diversity of turtles in Florida can be attributed in part to the warm temperate, maritime climate with abundant rainfall. In a worldwide analysis of correlates of emydid turtle species density (examining mean Janu- ary and July maximum and minimum temperatures, basin area and dis- charge, latitude, and annual and seasonal rainfall), I found that rainfall was the only significant correlate (Iverson, 1989). However, Florida’s location at the crossroads of the Gulf Coast, the Atlantic seaboard, and the peninsula itself probably also contributes to the high species diversity. Species from each of these regions have penetrated various distances onto or beyond the penin- sula, respectively, but the pattern is a general decline in diversity as one moves south along the peninsula (Fig. 26). This “peninsula effect” has been noted for all components of Florida’s herpetofauna (Means and Simberloff, 1987). ACKNOWLEDGMENTS— We acknowledge the inspiration provided by the late Dr. Archie F. Carr, the father of Florida turtle biology. For the contribution of specimens and/or locality lists we thank R. Ashton, R.L. Bezy (LACM; all acronyms follow Leviton et al., 1985), J.T. Collins (KU), W.J. Conley, J. Crenshaw, J. Diemer, J.R. Dixon, W.E. Duellman (KU), H.A. Dundee (TUL), L. Ehrhart, C. Ernst (GMU), R. Franz (UF), the Florida Natural Areas Inventory, D. Hoffmeister (UIMNH), D.R. Jackson, A.G. Kluge (UMMZ), A.E. Leviton (CAS), H. Marx (FMNH), C.J. McCoy (CM), D.B. Means (TT), P.A. Meylan (UF and AMNH), R.H. Mount (AUM), C.W. Myers (AMNH), M.A. Nickerson (MPM), L. Ogren (National Marine Fisheries Service), G.K. Pregill (SDNHM), P.C.H. Pritchard (PCHP), J. Rosado (MCZ), D.A. Rossman (LSU), B. Schroeder and the National Marine Fisheries Service, E.E. Williams (MCZ), J.W. Wright (LACM), G.R. Zug (USNM), R.G. Zweifel (AMNH), and especially D. Auth (UF) and his assistants. R. Ashton, W. Auffenberg, D. Auth, J.T. Collins, R. Conant, C.K. Dodd, C.H. Ernst, M.A. Ewert, D.R. Jackson, A.B. Meylan, P.A. Meylan, P. Moler, P.C.H. Pritchard, and H.M. Smith provided comments on early drafts of the manuscript. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 141 LITURATURE CITED ALLEN, E. R., AnD W. T. NEILL. 1950. The alligator snapping turtle Macrochelys temminicki in Florida. Spec. Publ. Ross Allen’s Rept. Inst. 4:1-15. . 1957. 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GALLAGHER, R. M., M. L. Houuincer, R. M. INGLE, AND C. F. Hutcu. 1972. Marine turtle nesting on Hutchinson Island, Florida, in 1971. Florida Dept. Nat. Res., Marine Res. Lab. Contrib. 207:1-11. Gorpon, W. G. 1983. National report for the United States of America to the Western Atlantic Turtle Symposium. San Jose, Costa Rica. 17-22 July 1983. Unpaginated. [cited by Prit- chard and Trebbau, 1984]. GrooMsriDcE, B. 1982. The IUCN Amphibia-Reptilia Red Data Book. Part 1. Testudines, Croco- dylia, Rhynchocephalia. Internatl. Union Conserv. Nat. 426 pp. Hirtu, H. F. 1980. Chelonia mydas. Cat. Amer. Amphib. Rept. 249:1-4. IversON, J. B. 1977a. Kinosternon subrubrum. Cat. Amer. Amphib. Rept. 193:1-4. . 1977b. Sternotherus minor. Cat. Amer. Amphib. Rept. 195:1-2. . 1977c. Geographic variation in the musk turtle, Sternotherus minor. Copeia. 1977(3):502-517. . 1978. Variation in striped mud turtles Kinosternon baurii. J. Herpetol. 12(2):135- 142. . 1986. A checklist with distribution maps of the turtles of the world. Iverson, Rich- mond, Indiana. . 1989. Global correlates of species density in freshwater and terrestrial turtles. J. Biogeography. Submitted Jackson, C. G. 1964. A biometrical study of form and growth in Pseudemys concinna suwan- niensis Carr (Order Testudinata). Unpubl. Ph.D. Dissert., Univ. Florida, Gainesville. 76 pp. Jackson, D. R. 1975. A Pleistocene Graptemys(Reptilia: Testudines) from the Santa Fe River of Florida. Herpetologica. 31:213-219. . 1978. Chrysemys nelsoni. Cat. Amer. Amphib. Rept. 210:1-2. . 1987. Florida Natural Areas Inventory, Tallahassee, Pers. Comm. . 1988. A reexamination of fossil turtles of the genus Trachemys (Testudines: Emydi- dae). Herpetologica. 44(3):317-325. Kinc, W., AND T. Krakauer. 1966. The exotic herpetofauna of southeast Florida. Quart. J. Florida Acad. Sci. 29(2):144-154. No. 2, 1989] IVERSON AND ETCHBERGER— TURTLES IN FLORIDA 143 KnepTon, J. C. 1956. County records of Testudinata collected in Georgia. J. Tennessee Acad. Sci. 31(4):322-324. Lananas, P.N. 1986. Graptemys nigrinoda. Cat. Amer. Amphib. Rept. 396:1-2. Lams, T. 1983. The striped mud turtle (Kinosternon bauri) in South Carolina, a confirmation through multivariate character analysis. Herpetologica. 39:383-390. Leviton, A. E., R. H. Grsss, Jr., E. HEAL, AND C. E. Dawson. 1985. Standard in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia. 1985:802-832. Lovicu, J. E. 1985. Graptemys pulchra. Cat. Amer. Amphib. Rept. 360:1-2. Lunp, P. F. 1985. Hawksbill turtle (Eretmochelys imbricata) nesting on the east coast of Florida. J. Herpetol. 19:164-166. Marquez M., R., A. VILLANUEVA O., AND C. PENAFLORES S. 1976. Sinopsis de datos biologicos sobre la tortuga golfina, Lepidochelys kempi (Eschscholtz, 1829) en Mexico. INP Sinop. Pesca 2:1-61. McCoy, C. J., AnD R. C. Voct. 1979. 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John Johnson, North Bennington, Vermont. 144 FLORIDA SCIENTIST [Vol. 52 Vitt, L. J., AND A. E. DunHam. 1980. Geographic distribution: Chrysemys nelsoni. SSAR Herp Review 11(3):80. Warp, J. P. 1984. Relationships of the chrysemyd turtles of North America (Testudines: Emydi- dae). Spec. Publ. Mus. Texas Tech. Univ. 21:1-50. Wess, R. G. 1962. North American Recent soft-shelled turtles (Family Trionychidae). Univ. Kansas Publ. Mus. Natur. Hist. 13:429-611. . 1973a. Trionyx ferox. Cat. Amer. Amphib. Rept. 138:1-3. . 1973b. Trionyx muticus. Cat. Amer. Amphib. Rept. 139:1-2. . 1973c. Trionyx spiniferus. Cat. Amer. Amphib. Rept. 140:1-4. Witson, L. D., AND L. Porras. 1983. The ecological impact of man on the south Florida herpe- tofauna. Univ. Kansas Spec. Publ. 9:1-89. WirzE..L, W. N. 1983. Synopsis of biological data on the hawksbill turtle, Eretmochelys imbri- cata (Linnaeus 1766). FAO United Nations Fish Synopsis 137:1-78. YERGER, R. W. 1965. The leatherback turtle on the Gulf Coast of Florida. Copeia 1985:365-366. Zuc, G. R., AnD A. ScHwartz. 1971. Deirochelys, D. reticularia. Cat. Amer. Amphib. Rept. 107:1-3. Florida Sci. 52(2):119-144. 1989. Accepted: August 24, 1988. INSTRUCTIONS TO AUTHORS Individuals who publish in the Florida Scientist must be active members in the Florida Academy of Sciences. Submit a typewritten original and two copies of the text, illustrations, and tables. All type- written material—including the abstract, literature citations, footnotes, tables, and figure legends — shall be double-spaced. Use one side of 8% x 11 inch (21% cm X 28 cm) good quality bond paper for the original; the copy may be xeroxed. Margins should be at least 3.cm all around. Number the pages through the Literature Cited section. Avoid footnotes and do not use mimeo, slick, erasable, or ruled paper. Use metric units for all measurements. 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Also, contributions will be considered which present new applications of scientific knowledge to practical problems within fields of interest to the Academy. Articles must not duplicate in any substantial way material that is published elsewhere. Contributions are accepted only from members of the Academy and so papers submitted by non-members will be accepted only after the authors join the Academy. Instructions for preparation of manuscripts are inside the back cover. Officers for 1989-90 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. ERNEstT D. ESTEVEZ Treasurery: Dr. ANTHONY F.. WALSH Mote Marine Laboratory 5636 Satel Drive 1600 City Island Park Orlando, Florida 32810 Sarasota, Florida 33577 Executive Secretary: Dr. ALEXANDER DICKISON President-Elect: Dr. FRED BUONI Department of Physical Sciences Operations Research Program Seminole Community College Florida Institute of Technology Sanford, Florida 32771 Melbourne, FL 32901 Program Chair: Dr. PAULA THOMPSON Division of Natural Science Secretary: Dr. PATRICK J. GLEASON Florida Community College 1131 North Parkway 11901 Beach Blvd. Lake Worth, Florida 33460 Jacksonville, FL 32216 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BARBARA B. MartTIN, Co-Editor Volume 52 Summer 1989 Number 3 Social Sciences BOOSTING SCIENCE CAREERS FOR THE PHYSICALLY HANDICAPPED STUDENT LuURETHA F. Lucky Division of Curriculum and Instruction, Florida International University, Miami, FL 33199 Asstract: Equal access to education, employment opportunities, career options and indepen- dent functioning has long been the aspiration of individuals with physical handicaps. Histori- cally, physically handicapped persons have been denied access to public education and employ- ment opportunities commensurate with their abilities. This study supported these contentions and further indicated that barriers still exist for individuals with physical handicaps. The purpose of this study was to identify physically handicapped scientists in the State of Florida and to determine the extent to which businesses, agencies and organizations were employing scientists with handicaps. From the 608 businesses, agencies, and organizations surveyed, only 16 physi- cally handicapped scientists were identified. A questionnaire was then sent to these 16 scientists to obtain specific information related to their educational training and professional career develop- ment. Analysis of the results revealed that, overall, few scientists with handicaps reside within the State of Florida; businesses, agencies and organizations are not employing the physically handicapped in scientific areas; and there is still gross underrepresentation of physically handi- capped scientists in scientific fields. THE physically handicapped comprise heterogeneous groups with vary- ing abilities (Kirk and Gallagher, 1986). They may sometimes be referred to as the physically disabled and/or physically impaired (Sirvis, 1978). This par- ticular population includes individuals with functional limitations related to their physical disability (e.g., hand control), and medical conditions (e.g., strength and stamina). Having a physical disability does not, however, limit an individual’s ability to be a productive member of society in career areas such as science, medicine, or industry (Sirvis 1978; Stearner, 1981). The occupational outlook for the physically handicapped, however, is complex. Persons with mild or transitory handicapping conditions may not be affected in their occupational choice. Others with severe physical handi- caps who are able to use their intelligence, social skills, and residual physical abilities to the fullest, have been denied full access to public education (Hal- lahan and Kauffman, 1978; Stearner, 1981). Reference to the physical handi- capped in this study will include individuals with orthopedic and neurologi- cal impairments (cerebral palsy, muscular dystrophy, spinal bifida, 146 FLORIDA SCIENTIST [Vol. 52 osteogenesis imperfecta, limb deficiency, spinal cord injury), and other health impairments (epilepsy, juvenile diabetes mellitus, cancer, cardiac or heart conditions, sickle cell anemia, hemophilia, cystic fibrosis) (Sirvis, 1978). BackcrouND— Until recently, one of America’s largest minorities, the physically handicapped, have faced many obstacles in trying to remain inde- pendent or become independent. There are approximately 36 million dis- abled individuals living in America today (Bruck, 1978). People with disabili- ties who try to function outside their homes have faced many difficulties ranging from transportation, education, and recreation to employment. In spite of their physical disabilities, some of the physically impaired have con- tinued to function effectively as citizens, consumers, taxpayers, jobholders, and students. Many of the physically handicapped attending public schools today are not aware of the many career options available to them (Anony- mous, 1978). History shows that children with handicaps have been denied a free public education for more than 200 years, and traditionally, the judicial system has been called upon to remedy the situation (Ricker, 1978). One area in which the physically handicapped are still denied complete access is in science education. The physically handicapped have not had full access to public school science programs, higher education in science, or pro- fessional science training programs (Anonymous, 1981; Ricker, 1978; Stearner, 1981). In a conference held by the American Association for the Advancement of Science to identify barriers to handicapped persons entering careers in science, (Redden, Davis, and Brown, 1979), it was reeommended that science education should involve more than removing obstacles (widen- ing doors and redesigning physical spaces) to make science more accessible, but should also involve making a commitment to the idea that handicapped students, like all other students, have a basic right to learn science content and skills provided they have the intelligence and interest in the field (Anony- mous, 1978, 1981). In addition, the handicapped should receive similar en- couragement and opportunities as the non-handicapped to pursue studies that will lead to science-related careers (Ricker, 1978). Ricker (1978) further indicated that within the past few years, scientists and science educators have recognized the challenges of making science education and science related careers accessible to handicapped persons. Efforts by such organizations as Science for the Handicapped Association (SHA), the Foundation for Science and the Handicapped, the American Chemical Society, the American Association for the Advancement of Science (AAAS) and other special projects for the visually and hearing impaired have brought attention to the needs of the physically handicapped in science edu- cation (Ricker, 1978). However, Ricker (1978) relates that these efforts have only been slightly effective in providing physically handicapped persons full access to science education. Results of these efforts have been in the form of one or two courses or a special science section. No. 3, 1989] LUCKY—SCIENCE CAREERS FOR HANDICAPPED 147 According to Vermeij (1978), education conveys knowledge and skills that enable one to (1) deal with life, pay taxes and hold jobs, and (2) live and function in the society in which one grows up. The scarce data in the area of science education for the physically handicapped have begun to show that the physically handicapped can be taught chemistry (Anonymous, 1980), medi- cal terminology (Braverman, 1979); paramedical skills (Beller, 1978); biology (Francoeur and Eilar, 1975); physics (Baughman and Zollman, 1977; Weems, 1977) and physical science (Eichenberger, 1974; Hadary, 1975; Walsh, 1977). The above programs have demonstrated that, with adaptation in equipment and modification in laboratory space and teaching techniques, the physically handicapped can be successfully integrated into science programs. Many of the physically handicapped either are not aware of the many career options available to them (Bruck, 1978; Anonymous, 1978; Rawls, 1978) or they have been discouraged from entering the field of science (Walsh, 1977). Short-term and long-term information in the area of teaching science to the physically handicapped is definitely needed for these individ- uals to advance in this field (Ricker, 1978). Another need of the physically handicapped is role models. According to Stern and Redden (1978), role models are needed to break down the stereo- typed idea that handicapped people need to be told what career fields are best for them. This study was designed to identify physically handicapped scientists in the State of Florida and to determine the extent to which busi- nesses were employing scientists with handicaps. MetuHops— To begin this project, directories and resource guides for various agencies, busi- nesses, hospitals, industries, and institutions throughout the State of Florida, with a high proba- bility of utilizing the services of a scientist, were obtained. Six hundred and eight businesses, agencies and organizations throughout the State of Florida were randomly selected for the study. A “Letter of Inquiry” was sent to each of these agencies, businesses and organizations explain- ing the purpose of the project. Contact was made through the president of the agency, business or company and/or the personnel director. Space was provided at the end of each letter for listing the names of physically handicapped scientists employed in the business, agency or organization, providing consultant work to the organization, or working as a volunteer or trainee in the organi- zation. An alternative suggestion was to pass the letter along to the physically handicapped employee to complete voluntarily, thereby protecting the privacy of the employee. Some 608 “Letters of Inquiry” were distributed. Of these, 460 (76%) were returned. From this distribution, only 16 physically handicapped scientists were identified. Some respondents listed names of other individuals who may have known scientists with handicaps or have them as employees in their businesses, agencies or organizations. A follow up contact was made to these individuals. A number of the forms returned also contained the following comments: “The nat- ure of our business does not lend itself to the employment of a scientist;” “We do not have a physically handicapped scientist currently employed nor have we employed one in the past;” “I am sorry we do not know any;” and, finally, “A physically handicapped person cannot function in our type of business.” A questionnaire was constructed (Gay, 1976) to obtain specific information related to the educational training and professional career development of these 16 scientists. Validation of the instrument was by students in graduate classes in science education at Florida International University. The questionnaire was then mailed to each scientist. For the purpose of this project, a scientist was broadly defined as a biologist, chemist, com- puter programmer, computer technician, engineer, environmentalist, laboratory technician, mathematician, medical doctor, nurse, paramedic, pharmacist/pharmacologist, physicist, sci- ence teacher, social scientist, and any of the related fields in behavioral, natural, physical, and 148 FLORIDA SCIENTIST [Vol. 52 social sciences. A scientist was considered to be physically handicapped if she or he had functional limitations related to physical disabilities (e.g. hand or leg control) and/or medical conditions (e.g. strength and stamina). TaBLE 1. Educational training and professional career development of physically handi- capped scientists. Question Scientist % Type of Education received in elementary school Regular 13 81.2 Special — — Combination 2 12.5 Other 1 6.2 Type of education received in high school Regular 12 75.0 Special — — Combination 3 18.8 Other 1 G22 Individual most influential in helping to make career choice Another handicapped individual 2 12.5 Teacher 3 18.8 Counselor 3 18.8 Parent 2 12.5 Peer 3 18.8 Other 3 18.8 Information on career choices provided to handicapped on the same basis as to the non-handicapped. Yes 6 37.5 No 5 31.2 Other a 12.5 Doesn’t apply 3 18.8 Experienced physical barriers during professional training Yes 6 37.5 No 6 37.5 Other 1 6.2 Doesn't apply 3 18.8 Experienced academic barriers during professional preparation Yes 4 25.0 No 8 50.0 Other — — Doesn’t apply 4 25.0 Experienced difficulty finding a job after completing training Yes 4 25.0 No y) 56.2 Doesn’t apply 3 18.8 Type of degree currently held High school diploma = = Bachelor of Art i 6.2 Bachelor of Science 4 25.0 Master of Science 1 6.2 Master of Social Work il 6.2 Doctor of Education — -_ Doctor of Medicine 2 12.3 Doctor of Philosophy 1 6.2 Other 1 6.2 No. 3, 1989] LUCKY—SCIENCE CAREERS FOR HANDICAPPED 149 Table 1. Continued. Question Scientist Jo Adaptation needed in physical environment to function on job Yes 8 50.0 No 8 50.0 Active with professional groups Yes a 43.8 No 9 56.2 Active with group that works directly with children Yes 6 37.5 No 10 62.5 Active with group that works directly with handicapped children Yes 4 25.0 No 12 75.0 Willing to serve as a resource person to classes serving children with physical handicaps Yes iL 68.7 No 4 25.0 Other 1 6.2 Permit name to be included in a resource directory Yes 13 81.2 No 2 12.5 Other 1 6.2 Age when disability developed 0-5 9 56.2 6-11 1 G22 12-17 1 6.2 over 18 5 31.2 Age group 20-29 4 25.0 30-39 5 Se? 40-49 3 18.8 over 50 4 25.0 Sex Female 5 31.2 Male 11 68.8 Ethnic group American Indian —_ == Asian/Pacific Islander Black 1 6.2 Hispanic aes rea White 15 93.8 Other = as N=16 SaAMPLE— This study emphasized identification of scientists with physical handicaps. Sixteen scientists with handicaps were identified from the 608 businesses, agencies and organizations surveyed throughout the State of Florida. Each of these scientists was sent a questionnaire with a cover letter explaining the purpose of the study and directions for completing the questionnaire. All 16 of these scientists returned the questionnaire. 150 FLORIDA SCIENTIST [Vol. 52 RESULTS AND Discussion—Table 1 presents a summary of the questions responded to by handicapped scientists. Analysis of the questionnaire indi- cated that the majority of the physically handicapped scientists in this study received regular education training during their elementary (88% ) and high (81%) school days (see Table 1). Those who received a combination (special and regular) type of education indicated that they were mainstreamed into regular classes. One scientist related that he transferred from a public school into a private school and hired a tutor in order to continue in a regular education program. In terms of the most influential person on the scientists in making a career choice, the breakdown was more even distributed between such individuals as another handicapped person, teacher, counselor, parent, peer and others. One respondent indicated that he alone was instrumental in his pursuit of a career. Approximately 38% of the physically handicapped scientists indicated that information on career choices was provided on the same basis for them as for their non-handicapped peers. Even though this 38 % indicated that physically handicapped scientists did receive career coun- seling, the fact that 31% felt that they did not receive adequate career coun- seling reveals that counselors, whether at the high school, college or voca- tional rehabilitation level, are still very unaware of the capabilities of individuals with handicaps, their contributions to the field of science, and their motivation to pursue their interest in the field of science. As indicated in Table 1, 38% of the scientists surveyed indicated that physical barriers presented problems for them during their professional train- ing. Another 38% indicated that they experienced no physical barriers. The 19% who checked the “doesn’t apply” category related that they experienced no physical barriers because their disability occurred after completing gradu- ate school or in their last year of professional training. The types of physical barriers experienced included such problems as: the elevators breaking down; bathrooms too small for wheelchairs; parking behind buildings in “service areas’ for access to ramps to get to classes; difficulty in reading printouts and no provisions for interpreters; difficulty reaching classrooms and science lab- oratories on the second floor of older buildings; and, for those who could walk without aides, being forced to use the stairs when elevators would not work. Some 50% of the physically handicapped scientists experienced no aca- demic barrier during their professional training (see Table 1). The 25% who did, however, listed such negative problems as discouragement from entering the field of science, negative attitudes of some of the professors, counselors and advisors, and encouragement by counselors to enter such fields as clerical work for the females and drafting for the males. One scientist related how he was discouraged from entering medical school by advisors and the medical school committee where he had already been accepted into the program. Note and test taking presented problems for some of the scientists which resulted from attitude of the professors. As summed up by one scientist, the biggest problem facing the handicapped student in graduate school is the No. 3, 1989] LUCKY — SCIENCE CAREERS FOR HANDICAPPED 151 prevailing attitude by college advisors and professors of “Why are you here anyway? , “With your handicap, what have you to offer anyone?” Concerning finding a job after completing professional training, Table 1 shows that 56% of the physically handicapped scientists experienced no diffi- culty finding a job. The reason given by one scientist for not experiencing difficulty finding a job was because he started his own business, thereby, negating the search for a job. Another scientist entered medical school imme- diately after completing graduate school. One scientist related that she went to work for the same agency where she had done her internship. She received her first promotion nine years later. One scientist related that his disability occurred during his senior year in college. After the graduate school into which he was accepted found out that his disability was progressive, it was no longer interested in him. Approximately 31% of the handicapped scientists hold master and above degrees. Half (50 % ) of the scientists indicated that no adaptation in the phys- ical environment was needed in order for them to function and half (50%) indicated some adaptation was needed. However, those who did require ad- aptation in the physical environment listed such areas as: installation of ramps to the building; providing larger examination rooms for seeing and examining patients; installation of handrails on stairways and steps; designat- ing parking space close to the building; purchasing dictating machines; secur- ing secretarial services; installation of a light in the classroom to remain on when using audio-visual equipment, and installation of rugs in the office. The job responsibilities of the scientists in this study span such areas as: office gynecology; normal obstetrics; obesity research; teaching science and serving as the director of a transitional living facility for the physically dis- abled; directing a comprehensive rehabilitation center; adjudicating SSI and SSI (Title II and XVI) social security disability claims according to the Fed- eral regulations; serving as computer engineer involved with program design, coding, testing, debugging and other applications of computer science and mathematical concepts to computer engineering; involvement with research at a blood bank directed toward extending the viability of stored platelets from three hours to three days; working at a university as professor of physics and physical science; and serving as the chairman of the Chemistry/Earth Science Department in a major university. As indicated by Table I, approximately 44% of the physically handi- capped scientists were active with a professional group. Not only were the scientists active in professional organizations, but many of them were also active with groups that worked directly with children. The percentage of scientists (38 %) with handicaps working with organizations that serve chil- dren with handicaps was much lower than with organizations serving non- handicapped children. According to one scientist, “these children need so much attention and encouragement to pursue goals of their own choice.” However, most of the scientists indicated a willingness to work with organiza- tions serving children with handicaps. 152 FLORIDA SCIENTIST [Vol. 52 Disability among most of these scientists developed at a young age: 56% of the scientists developed their disability between the age of 0-5, and 31% were over 18 years of age when the disability developed. The types of disabili- ties reported by the scientists included: spina bifida; neuromuscular disorder; weaknesses in peripheral muscles; mild to moderate cerebral palsy with speech impediment; spinal cord injury resulting in paralysis; severe learning impairment; paraplegia; arrhythmia hypertension; profound deafness; blindness; rheumatoid arthritis; and progressive muscle weakness (a variation of muscular dystrophy). Of the scientists with handicaps, 31% responding to the questionnaire fell within the 30 to 39 year old age range, and the majority (69%) were males. About 94% were white and 6% were black, no other ethnic group was iden- tified. Discrimination was also listed as a barrier to entering the scientific field. This discrimination was exhibited in the form of denial of opportunity to participate in laboratory activities, lack of interpreters and readers, attitudes of professors toward the physically handicapped, lack of promotion on the job, and counseling to enter other fields such as clerical or drafting rather than science. As stated earlier, making the public more aware of the abilities of the physically handicapped can help alleviate these problems. Data gath- ered from the 16 scientists with handicaps identified in this study were com- piled and profiled into a resource guide. SUMMARY AND CoNCLuSION—As revealed in this study, few scientists with handicaps were located within the State of Florida. Based on the small num- ber of physically handicapped scientists identified as a result of this study, three conclusions can be drawn: (1) employers and colleagues did not choose to identify the physically handicapped scientist employed in the business or give the physically handicapped scientists the opportunity to identify them- selves; (2) industries and businesses are not employing the physically handi- capped in scientific areas, and (3) there is gross underrepresentation of physi- cally handicapped scientists in the scientific fields. Even though only a small number of physically handicapped scientists was identified, there is a high probability that the letter of inquiry to the various agencies, businesses and industries made them aware of the need to provide employment opportunities for scientists with handicaps. It is antici- pated that efforts such as this and efforts by other groups working with the handicapped will help make the public more aware of the abilities of physi- cally handicapped individuals. Through cooperation from the handicapped, classroom teachers, counselors, college advisors, professors, and employers, attitudinal, academic, communication, and physical barriers can be re- moved, resulting in more accessibility to the field of science for individuals with handicaps. Considering the demands and challenges that are placed upon an individ- ual entering the field of science, respect must be given to the individual with a handicap who makes the decision to meet these challenges and to pursue a No. 3, 1989] LUCKY—SCIENCE CAREERS FOR HANDICAPPED 153 career in the field of science. It is recognized that not all physically handi- capped individuals should enter the field of science, but those with the ability and determination to be successful should be encouraged to actively pursue their goals. Counselors at all levels, peers, parents, college advisors and pro- fessors should not view the physical limitation of the individual as a main obstacle toward pursuing a career in the field of science, especially since there are numerous career options to select from within the field. As indicated by one scientist, support and encouragement should begin at the elementary level for handicapped students and continue throughout their academic prep- aration. LITERATURE CITED .ANoNyMous. 1981. Guide to teaching chemistry to the handicapped. Chem. Eng. News. 39 (26)223. . 1980. Ideas explored to teach handicapped chemistry. Chem. Eng. News. 58(19):24. . 1978. Project boosts science careers for handicapped. Chem. Eng. News. 56(17):20. ____«. 1978. Opening doors for handicapped youth. Sci. Teacher. 45(7):24. BAUGHMAN, J., JR., AND D. ZoLLMAN. 1977. Physics labs for the blind. Physics Teacher. 15:339- 342. BELLER, J. 1978. Handicapped students learn a helping way of life. Sci. Teacher. 45:25-26. BRAVERMAN, B., J. EGELSTON-Dopps, AND R. EGELSTON. 1979. Cue utilization by deaf students in learning medical terminology. J. Resea. Sci. Teach. 16:91-103. Bruck, L. 1978. Access, the Guide to a Better Life for Disabled Americans. Random House. New York. EICHENBERGER, R. 1974. Teaching science to the blind student. Sci. Teacher. 41:53-54. FRANCOEUR, P., AND B. Eriar. 1975. Teaching the mammalian heart to the visually handicapped. Sci. Teacher. 42:8-11. Gay, L. R. EpuCATIONAL RESEARCH. 1976. Charles E. Merrill. Columbus, Ohio. Hapary, D. 1975. Picking up good vibrations from science for the handicapped. Sci. Teacher. 42:12-13. HALLAHAN, D. P., AnD J. M. KaurrMan. 1978. Exceptional Children, Introduction to Special Education. Prentice-Hall. Englewood Cliffs, New Jersey. Kirk, S. A., AND J. GALLAGHER. 1983. Educating Exceptional Children. Houghton Mifflin. Bos- ton. Raw .s, R. Easing the way for the handicapped in science. Chem. Eng. News, 56:(4)25-27. REDDEN, M., C. Davis, AND J. Brown. 1979. Science for Handicapped Students in Higher Educa- tion. American Association for the Advancement of Science, Washington, D. C. Ricker, K. S. 1978. Science and the physically handicapped. Viewpoints Teach. Learning. 55:67- 76. Sirvis, B. 1978. The physically disabled. In: Meyen, E. (Ed.), Exceptional Children and Youth, An Introduction. Love Publishing Co., Denver, Colorado. STEARNER, S. P. 1981. Overview of the National Science Foundation (NSF) projects on the handi- capped in science. J. College Sci. Teach. 10:352-354. STEARN, V., AND M. ReEppEN. 1978. Role models for the handicapped. National Elementary Prin- cipal. 58:43-45. VeRME]J, G. J. 1978. On teaching the blind student. Today’s Educ. 67:77-78. Wausu, E. 1977. The handicapped and science moving into the mainstream. Science. 196:1424- 1426. Weems, B. 1977. A physical science course for the visually impaired. Physics Teacher. 15:333-338. Florida Sci. 52(3):145-153. 1989. Accepted: August 23, 1988. Atmospheric and Oceanographic Sciences FATE OF SATELLITE-TRACKED BUOYS AND DRIFT CARDS OFF THE SOUTHEASTERN ATLANTIC COAST OF THE UNITED STATES GeorcE A. MAUL AND NIcoLas J. BRAvo Atlantic Oceanographic and Meteorological Laboratory, National Oceanic and Atmospheric Administration, 4301 Rickenbacker Causeway, Miami, Florida 33149 Apstract: Twelve satellite-tracked drifting buoys were released, one each month between April 1984 and March 1985, at the 75 m isobath, 70 km southeast of Cape Canaveral, FL. Starting in May 1984, 500 drift cards were also released at each buoy deployment from the same site (28.0°N, 80.0°W). In addition, satellite-tracked buoys from May 1979, September 1983, and February 1985, which drifted into or through the area, were studied for information about surface currents and particle trajectories. All 12 buoys from the 28°N/80°W launch site drifted north past Cape Canaveral (28.5°N); only the May 1984 buoy drifted into the coastal zone off Mayport, FL (30.4°N), although several others showed a tendency to do so in that area. Every buoy deployed within 5 km of the western edge of the Gulf Stream was entrained into the current, and some as far as 25 km were also entrained. The September 1983 bouy, which was deployed slightly west of 28°N/80°W, and the February 1985 buoy, which was deployed in the Gulf of Mexico, both came ashore near St. Lucie Inlet, FL (27.2°N), 140 km south of Cape Canaveral. Drift cards from May, September, October, and November 1984, and February 1985 were recovered west and north of 28°N/80°W. Forty percent of the cards recovered were from south of Cape Canaveral; Cocoa Beach, FL (28.3°N), and Pontevedra Beach, FL (30.2°N), reported most of the returns. For those months when drift cards were returned, buoy trajectories showed little correlation with drift card vectors. Drift cards established the possibility of materi- als coming ashore along the Florida Atlantic coast, both north and south of Cape Canaveral, particularly in the autumn. THE Florida Atlantic Coast Transport Study (FACTS) was a program to study the circulation north of the Straits of Florida, with emphasis on the potential impact of offshore minerals mining on Florida waters. The region to the north of Cape Canaveral, the South Atlantic Bight (SAB), has been studied extensively (Lee, 1983), but the SAB studies did not include surface drifters. Drift cards and satellite-tracked drifting buoys are one aspect of FACTS which is described in more detail by Maul (1985), and in Rinkel e¢ al. (1986). For an overview map see Fig. 1, where the very different tracks of two buoys that were released in the Gulf of Mexico, independent of FACTS, are also shown. Although FACTS was centered on the coastal zone from approximately Jupiter, FL to the Georgia border (31°N) most buoys drifted through the SAB, and a discussion of their trajectories is included. In addition to drifters in FACTS, current meters were moored along the 75 m isobath off Florida at 27°N, 27.5°N, 28°N, 29°N and 30°N; along 29°N six additional current meter moorings were deployed from the 15 m to the 800 m isobath (Smith, 1987). To supplement the 29°N current meters, periodic cruises were conducted to make direct measurements of Gulf Stream No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 155 CARE aro a ON —— MAY 1979 Ps FEB 1985 7 oe 34 ; LP OY CAPE FEAR oy 2 CAPE ROMAIN ig cuarceston, sc ' 32 ig} SAVANNAH, GA ee — SEPT. 1983 FEB. 1985 --- MAY. 1979 é 155 ON 28 -#) MAYPORT, FL 30 ST. LUCIE INLET ‘, GRAND BAHAMA ISLAND 75m 82 80 8 Fic. 1: Gulf Stream region of the Florida Atlantic Coast Transport Study. Solid line shows the track of buoy 1776 and dotted line shows buoy 5501, both released in the eastern Gulf of Mexico. Dashed line locates the Jupiter-Grand Bahama Island submarine cable used to measure trans- ports shown in Fig. 4a. Star at 28°N/80° W locates FACTS buoy and drift card launch site. Inset: Detail of buoys 2194 (September 1983), 5501 (February 1985) and 1776 (May 1979). Asterisks mark the launch sites of buoy 2194. STRAITS OF FLORIDA 156 FLORIDA SCIENTIST [Vol. 52 velocity, and inverted echo sounders with pressure gauges were moored on the bottom. Although conclusions herein take into account prior studies (Leming, 1979) and other FACTS results (cf. Zantopp et al., 1987), this ar- ticle emphasizes drift cards and satellite tracked buoys. Several drift card studies (e.g. Murphy et al., 1975; Maul and: Molinari, 1975; Williams et al., 1977; Parker, et al., 1979) show drift cards which were released in the Gulf of Mexico or the Caribbean Sea that came ashore on the east coast of Florida. Most Gulf or Caribbean drift cards that come ashore in Florida do so between Key West and Palm Beach, although a few are ob- served to come ashore between Cape Canaveral and Mayport, FL. On the other hand, some satellite-tracked buoys deployed in the northeastern Gulf of Mexico came through the Straits of Florida averaging over 150 cm/s, and continued into the SAB. These studies suggest that in the FACTS area, the fate of surface drifters is highly variable. In order to study this variability and the potential for oil spills coming ashore, twelve buoys were deployed from above the current meter mooring at 28°N latitude, 80° W longitude, at the 75 m isobath, 70 km southeast of Cape Canaveral. One buoy was deployed each month from April 1984 through March 1985. For 11 months, starting in May 1984, 500 drift cards were also deployed from the 28°N/80°W site whenever a buoy was deployed. In addi- tion, data from three buoys from other sources were incorporated into this study because they provided information that was not possible to obtain from those launched at the 28°N/80° W site. DaTA SouRCES AND Processinc—The drift cards were standard 7.6 x 12.7 cm (3” x 5”) post- cards encased in flat plastic envelopes. Persons recovering the cards were asked to fill out the information and mail them (post paid) to the Atlantic Oceanographic and Meteorological Labo- ratory (AOML). The cards were very light and floated in the upper centimeter of the sea; they seemed to be more affected by the wind and waves than the satellite-tracked buoys. Satellite-tracked buoys were of three types. The May 1979 buoy (Vukovich and Maul, 1983) was a cylinder with a flotation collar and a cross-plane drogue; total length was 2 m. The September 1983 buoy was a 3 m long undrogued PVC pipe; it is described in detail by Bitterman and Hansen (1986). The Shell Offshore Co. buoy, for which data from 1 January 1985 through 22 March 1985 were provided, is a Polar Research Inc. design of about 3 m length; it was found undrogued when recovered approximately 140 km south-southeast of Cape Canaveral. All buoys deployed at the 28°N/80° W site were of the same type as the September 1983 buoy. All but the May 1979 buoy were tracked by Service ARGOS using the random access receivers aboard the TIROS series of National Oceanic and Atmospheric Administration (NOAA) satel- lites. The data processing algorithms are described by Herman and Hansen (1977). The May 1979 buoy was Research Triangle Institute (RTI) sponsored, and was located using the Tracking and Data Relay Experiment (T&DRE) on the NIMBUS-6 satellite; processing of the T&DRE buoy location data was identical to that of the ARGOS located buoys. REesuLts—The buoys not released from the 28°N/80°W site are listed in Table 1 along with their deployment dates and positions. Buoy 2194 is listed as two separate deployments because it was picked up by the U.S. Coast Guard and brought ashore from 18-21 September 1983, and then redeployed approximately where it was picked up. Buoy 5501 was deployed by Shell Offshore Inc. in early 1984 in the Gulf of Mexico; data supplied to FACTS was from | January 1985 onward. The inset to Fig. 1 shows details of the tracks of buoys 1776 (solid), 2194 (dotted), and 5501 (dashed). - No. 3, 1989] MAUL AND BRAVO—BUOYS AND DRIFT CARDS 157 TaBLE 1. Buoy deployment summary from sites other than 28°N/80°W during the Florida Atlantic Coast Transport Study. Buoy Date Lat(°N) Lon(° W) Tracking Sponsor 1776 79May21 26.70 83.20 T&DRE RTI 2194 83Sep09 2H 80.26 ARGOS NOAA/AOML 2194 83Sep21 27.50 80.20 ARGOS NOAA/AOML 5501 85Jan01 26.00 85.40 ARGOS Shell The first deployment of 2194 (9-18 September 1983) was about 27 km west-southwest of the 28° N/80° W site, and the track initially showed anticy- clonic curvature then a southerly flow of approximately 50 cm/s. Winds dur- ing this period were variable at about 2 m/s and there seemed to be good correlation between winds and the buoy trajectory. The second deployment (21-26 September 1983) showed a cyclonic flow in the same area less than a week later, and the northward and southward speeds were each approxi- mately 40 cm/s. Winds were southeast at about 1 m/s during 21-22 Septem- ber, and from the northeast at 6-7 m/s from 23-25 September. Winds once again seemed to dominate the buoy motion, driving buoy 2194 towards shore on 26 September. This buoy was recovered floating northward inside the Indian River Lagoon, having entered the estuary through the St. Lucie Inlet. Buoy 1776 passed through the Straits of Florida on 2 June 1979 at speeds in excess of 150 cm/s, and eventually left the Gulf Stream south of Cape Fear, NC, after having advected into a warm-tongue filament; the T&DRE loca- tion ceased after 16 June 1979. The other Gulf of Mexico launched drifter, buoy 5501, passed north of Jupiter in late January 1985. After it detrained from the Gulf Stream, it first turned anticyclonically, and then cyclonically, and it showed speeds averaging 25 cm/s, but with northward and southward bursts over 40 cm/s. Buoy 5501 ran aground north of St. Lucie Inlet on 12 February 1985. Both the May 1979 buoy and the February 1985 buoy passed through the Straits along almost identical trajectories between about 26.0°N and 26.5°N. Once north of 27°N, their trajectories diverged, but this was not uncommon in the Gulf Stream (e.g. Chew et al., 1985), and is indicative of the complex processes along the Stream’s cyclonic shear front. Satellite images (Lee and Atkinson, 1983) often indicate that there are warm-tongues of Gulf Stream water extending southward from onshore meander crests in the SAB, and this was indeed the case for February 1985 off Florida. Apparently, the detraining process was not dissimilar from that described by Vukovich and Maul (1983) off North Carolina. Whether or not detraining was linked to cold-domes, as Chew and co-workers (1985) argue, is not clear, as both cyclonic and anticy- clonic motion are observed in this region. Buoy 5501 passed about 2 km east of the 27°N current meter mooring on 31 January 1985. Buoy speed was approximately 50 cm/s northward, yet the mooring showed about 25 cm/s southward at 10 m water depth. Buoy 5501 158 FLORIDA SCIENTIST [Vol. 52 continued north almost to 28°N on 3 February, when it rather abruptly turned south. At the 30 m water depth current meter on the 28°N mooring (the 27.5°N mooring was not operational), currents during 3-5 February were weak and changing from northward to southward; buoy 5501 was heading south at speeds up to 40 cm/s. February 5-7 had buoy 5501 heading north while the 30 m depth current meter at 28°N measured a burst of south- ward current up to 100 cm/s. From 8-10 February, when 5501 was heading south as fast as 60 cm/s, both the 27°N and 28°N upper layer current meters showed southward flow. Comparison of the motion of buoy 5501 with 40-hour low-pass filtered local wind shows a rather different picture. On 31 January, when 5501 passed the 27°N mooring, winds were from the north at 4 m/s. From 1-7 February, winds were from the southeast with maximum speed over 8 m/s, and the 27°N and 28°N near-surface current meters measured northward speeds of 100 cm/s and 50 cm/s respectively; conversely the drift of 5501 was to the north from 1-3 February, to the south from 3-5 February, and to the north again from 5-8 February. Strong north and northeast winds from 9-11 Febru- ary accompanied the final southward drift of 5501 into the coast. It is difficult to discern much correlation in the track of buoy 5501 with either wind and/or current meter observations. Current meter moorings along the 75 m isobath were located at the shelf edge and currents there are often dominated by Gulf Stream meanders whereas the coastal waters seem to respond to a mixture of wind and Gulf Stream forcing, particularly that due to meanders and eddies along the western boundary. The trajectory of buoy 5501 illustrates the complexity of the near-shore circulation in the offing of Cape Canaveral, and the difficulty in nowcasting its motion. TABLE 2. Summary of drift card returns during the Florida Atlantic Coast Transport Study. Date Total Primary % Time Winds (m/s) Number Location Easterlies* North East pees May 20 Cocoa Beach 44 0.1 2.6 June 0 37 0.4 0.9 July 0 32 2 2.4 August 0 36 0.4 0.4 September 2 Fernandina Beach 55 1.1 3.5 & Flagler Beach October 210 Pontevedra Beach 53 0.3 5.0 November 75 Cocoa Beach 46 a3 2.4 December 0 36 3 3.1 1985 January 0 On -2.1 0.2 February 34 Flagler Beach 30 0.6 3.2 March 0 30 -0.3 8) *Measured in the offing of Cape Canaveral; from the U.S. Navy (1974) Atlas as the composite of winds with easterly (NE, E, and SE) components, using the “from” (meteorological) directional sense. No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 159 3! 123 FERNANDINA { BEACH 38 MAYPORT 3 ATLANTIC BEACH JACKSONVILLE BEACH * PONTEVEDRA BEACH PALM VALLEY 30 SOUTH PONTEVEDRA 7 TOTAL NUMBER OF CARDS RETURNED FROM ALL LAUNCHES. ST. AUGUSTINE BEACH 2 19 FLAGLER BEACH ORMOND by SEAQ™ 10 ORMOND BEACH DAYTONA BEACH PONCE INLET 29 NEW SMYRNA BEACH a! EDGEWATER ox ! 87 i 3 CAPE CANAVERAL COCOA BEACH SATELLITE BEACH INDIAN HARBOUR BEACH\E TNDIALANTIC\RC! 1 28 MECBOURNE BEACH IK Launch Site \ \\ \ \ SEPT 1983 BUOY JENSEN BEACH \ =\we SHELL OIL CO. BUOY 82 8| 80 79 78 Fic. 2a: Summary of all drift card returns in FACTS regardless of the month during which they were deployed. Number at end of bar indicates total returns from that geographic area. The circulation along the Florida Atlantic coast was also investigated with drift cards. The total number of cards found and returned from loca- tions along the Florida coast, without regard for their date of launching is given in Fig. 2a. Two cards not included in the figure were found: one near Rodanthe, NC (35.6°N) in April 1985, two months after being launched, and the other from “somewhere near” Titusville, FL (28.6°N). Only cards launched during the months of May, September, October, and November 1984, and February 1985 were returned; with the exception of February, it was during these months that the winds generally exhibited the most pro- nounced easterly components (Table 2). 160 FLORIDA SCIENTIST [Vol. 52 3l LOCATIONS AT WHICH CARDS LAUNCHED FERNANOINA DURING INDICATED MONTHS WERE FOUND BEACH ; NESS MAY 1984 EBSA : MAYPORT Zs Scopes SEPT 1984 Wass Soe OCT 1984 be NOV 1984 30 wee ST. AUGUSTINE BEACH & ~~ ga FEB 1985 FLAGLER BEACH ORMOND BEACH 29 NEW SMYRNA BEACH CAPE CANAVERAL p cocoa BEACH \ 28 MELBOURNE BEACH JENSEN wet SEPT. 1983 BUOY SHELL OIL CO. BUOY 27 82 8! 80 79 78 Fic. 2b: Stippling shows months for which the drift cards were returned and the predominant location(s). The final location of buoys 2194 (September 1983) and 5501 (February 1985; Shell Oil Co.) are shown at the bottom of the map. ' Drift card returns for February warrant further comment. Buoy 4480 was launched on 15 February 1985, at which time 500 drift cards were released, and buoy 5501 ran aground on 1] February. Winds for 16-20 February were from the northeast at about 4 m/s. Satellite infrared imagery showed a 70 km- long warm-tongue juxtaposed to the Gulf Stream that must have influenced the trajectory of buoy 5501 (q. v. Fig. 1 inset). The combination of northeast winds and the south-southwestward advection in the warm-tongue could ac- count for the drift card returns. However, there were other releases of cards that were accompanied by equally strong northeast winds, but no cards were returned. Apparently a sensitive combination of wind, current, and release site determine drift card recovery probability and location in this region. No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 161 The stretches of coastline along which cards were found are shown in Fig. 2b. It can be seen, for instance, that for the coast between and including Cape Canaveral and Cocoa Beach, cards launched in May 1984, November 1984, and February 1985 were received. On the other hand, only cards launched in September 1984 were received from Fernandina Beach. Cards launched in September 1984 and in February 1985 were returned from Flagler Beach. Because cards deployed in October 1984 were found both north and south of Flagler Beach, it is assumed that the October event repre- sented by these cards would have been observed at Flagler Beach also. As one would expect, all of the months that had drift card returns (q.v. Table 2) had strong easterly winds. Comparing the observed winds with the “percent time easterlies” from historical records suggests that the 1984-1985 returns are representative. There are notable exceptions, and the results sum- marized in Fig. 2 and Table 2, must be used with caution. However, it is clear that the area north of Cape Canaveral is subject to significant returns, a fact not documented in previous studies. These drift card return results must be contrasted with the twelve buoys released from 28°N/80° W, which are sum- marized in Table 3 and in Figure 3. The trajectories of buoys 2083, 4470, and 4471 deployed in April, May, and June 1984 respectively, are given in Fig. 3a. Off Mayport buoy 2083 took a cyclonic turn followed by an anticyclonic turn; this pattern was typical of the small meanders in this vicinity. On Julian Day (JD) 480 (referenced to 1 January 1983), the speeds exceeded 150 cm/s, and buoy 2083 left the area on a northeasterly heading. Buoy 4470 left the launch site at speeds of less than 100 cm/s, and at about latitude 30°N detrained from the Gulf Stream. After an initial cyclonic turn as buoy 4470 left the direct influence of the current, it showed a wandering pattern for about a month with speeds of about 15 cm/s. The speeds and directions after JD 532 (16 June) indicated that buoy 4470 was picked up by a boat (in fact, buoy 4470 was found in a seafood restaurant near Mayport, still transmitting). After being launched and initially drifting northward, buoy 4471 also left the direct influence of the Gulf Stream off Mayport. Contrary to 4470, after about 10 days, buoy 4471 was re-entrained in the current and accelerated to over 180 cm/s as it headed northeastward. As with the May buoy, there was a short speed burst of about 100 cm/s just before detraining, and the sense of rotation was cyclonic as it left the Stream. Buoys 4472, 4473, and 4474, launched in July, August, and September 1984 respectively, are shown in Fig. 3b. Buoy 4472 left the launch site in a north northwesterly direction, but with speeds in excess of 125 cm/s it tran- sited the FACTS and SAB area in less than a week. At JD 564, the effect of the “Charleston Bump” (a shallow bottom area on the Blake Plateau centered near 32°-33°N, 77°-79°W) was seen on its trajectory. For the August 1984 deployment, speeds of almost 200 cm/s were observed under the influence of the bottom topography off Charleston. This buoy passed through the FACTS area in one day, but the buoy stopped transmitting after JD 591 and had the shortest record: one week in duration. Buoy 4474, after leaving Florida wa- 162 FLORIDA SCIENTIST [Vol. 52 TABLE 3. Buoy deployment/lifetime summary from the 28°N/80°W site during the Florida Atlantic Coast Transport Study. Buoy Lifetime Deployment Deployment JD* GMT Lat(°N) Lon(°W) 2083 84Apr06-85Apr29 462 1658 28.00 Tooee! 4470 84May11-84Junl5 497 1524 28.00 80.00 447] 84Jun12-85Nov11 529 1600 28.00 80.00 4472 84Jull2-85Dec10 559 1402 28.00 80.00 4473 84Aug07-84Aug14 585 1402 28.21 79.88 4474 84Sep13-86May07 622 1411 28.00 79.99 4476 84Oct16-84Dec01 655 1357 28.00 80.00 4477 84Nov15-86Jan10 685 1557 28.00 80.00 4478 84Dec11-86Jun27 711 1457 28.00 80.03 4479 85Jan25-86Apr07 756 1510 28.00 80.03 4480 85Feb15-85Jul24 Ta 2300 28.00 80.03 448] 85Mar16-86Mar21 806 1300 28.00 80.03 *Continuous Julian Day (JD) with respect to 1 January 1983. —— APRIL. 1984 =—— JULY 1984 g2 80 78 76 Fic. 3(a) (left panel): Tracks of buoy 2083 (solid), 4470 (dotted) and 4471 (dashed). Crosses along the track are one per day, with every fourth Julian Day numbered from 1983. 3(b) (right panel): Same as Fig. 3a for buoy 4472 (solid), 4473 (dotted), and 4474 (dashed). No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 163 ters at over 150 cm/s, turned east off Charleston, made a full-circle cyclonic loop off Cape Romain, and continued northeastward past Cape Hatteras, NC. At about latitude 30°N, buoy 4474 made an anticyclonic turn with a radius of curvature of approximately 300 km, exhibiting the opposite behav- ior to that of the August buoy. The buoy tracks in October, November, and December 1984 (buoys 4476, 4477 and 4478 respectively), are shown in Fig. 3c. Buoy 4476 showed initial behavior at the launch site similar to that of the buoy deployed in September, that is, the trajectory was initially cyclonic with a large radius of curvature, and speeds for the first few days were only about 60 cm/s. Off Florida, buoy 4476 showed no tendency to detrain from the current, but off the Carolinas there were two events, one at JD 664 (where the speed decreased almost to 0 cm/s) and another between JDs 666-671, where a large cyclonic turn was completed full circle. The November 1984 buoy had the same initial east- ward trajectory as did the October buoy; speeds were somewhat higher through the FACTS area than in the previous month, averaging about 100 cm/s. Passage of buoy 4478 through FACTS waters was fairly rapid except for an almost complete stop at JD 716 (17 December); however in the offings of Cape Romain and Cape Fear there were two remarkable detrainment/en- trainment events. Both events, centered at JDs 725 and 732 respectively, were cyclonic eddying characterized by highly varying speeds of + 25 cm/s. Each time, buoy 4478 made two complete circles before being entrained again. Trajectories with cyclonic turns such as this one and that of October 1984, are consistent with interpreting the juxtaposed circulation as resulting from en- trainment into cold, cyclonic frontal eddies (Lee and Atkinson, 1983). The January, February, and March 1985 buoys (4479, 4480, and 4481 re- spectively) are shown in Fig. 3d. Buoy 4479 showed a trajectory off Cape Canaveral somewhat like those of October and November 1984, and it smoothly passed the Florida Atlantic coast at about 100 cm/s. At about JD 765 (3 February 1985), a small cyclonic circle was completed at 32°N, very close to the southern one observed in December. As with the December buoy, once east of Cape Hatteras buoy 4479 showed a sustained period of speed near 200 cm/s. Buoy 4480 showed one of the most interesting trajectories of the series: centered at JD 783 (21 February) it made five rather regular speed changes from 25 cm/s to 100 cm/s and back again in 9 days; off Mayport there was a tendency for detrainment, but it continued on into the SAB area. In the offing of Cape Romain, buoy 4480 detrained from the Current and exhibited a random-ap- pearing trajectory for about a month (JDs 787-819; 25 February-29 March 1985). Speeds during this time were about 15 cm/s, followed by a rapid accel- eration to about 180 cm/s as buoy 4480 was re-entrained by the Gulf Stream and left the area. the last FACTS buoy was launched in March 1985 and at JD 814 (24 March) it made a small anticyclonic turn followed by a small cyclonic turn and a full circle four days later. Afterwards it accelerated to about 150 cm/s and then in the offings of Capes Romain and Fear (JDs 823 and 827 respectively) there were two more events tending to detrain the buoy. Upon 164 FLORIDA SCIENTIST [Vol. 52 —— OCT 1984 82 80 78 76 g2 80 7eaee 76 Fic. 3c (left panel): Same as Fig. 3a for buoy 4476 (solid), 4477 (dotted), and 4478 (dashed). Fic. 3d (right panel): Same as Fig. 3a for buoy 4479 (solid), 4480 (dotted), and 4481 (dashed). passing Cape Hatteras, buoy 4481 went northeastward at 25 cm/s and finally stopped transmitting at JD 851 after about 45 days of service. Discussion—Every buoy, including those originating in the Gulf of Mex- ico, was either launched near or was entrained into the cyclonic shear side of the Gulf Stream. All of the buoy trajectories show one common result: none crossed to the eastern side of the Gulf Stream until they passed Cape Hat- teras. This was also true, as best as is known, of the drift cards, i.e., except for the one card from Rodanthe, NC, there are no drift card returns from beyond Cape Hatteras. Drift cards were returned from five of the eleven deployments, but only the May 1984 buoy and drift cards were both found in coastal waters. The other four buoy trajectories that had associated drift card returns showed little obvious correlation to card returns. The windage on the cards must be significantly more than on the buoys, but it was suprising that these buoys, which were designed to be near-surface drifters, showed such behavior. None of the buoys from the 28°N/80°W site showed any tendency to enter the coastal zone south of Cape Canaveral. This is curious since satellite infrared images showed that the 28°N/80° W site is where the Gulf Stream’s No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 165 cyclonic edge has a standard deviation (a) of + 13 km about its mean position (Gaby and Baig, 1983), and Ekman drift is often westward. Buoy deploy- ment records also suggest, based on water color and temperature, that many buoys were not placed in the Gulf Stream. Apparently the 28° N/80° W site is close enough to the edge of the current that all but the shallowest surface drifters were entrained. Many of the buoy tracks suggest that such an entrain- ing process operates all along the Gulf Stream’s cylconic shear side. Table 4 summarizes the relationship between buoy speeds shortly after deployment and the distance to the cyclonic edge of the current as measured from satellite infrared observations (q.v. Gaby and Baig, 1983). There is a negative linear correlation (r=-0.7) between speed and distance to the Stream’s edge east (+) or west (—) of 28°N/80°W. On average, the edge of the Gulf Stream is about 5 km (+6 km) east of the deployment site. Because all the buoys eventually were entrained into the current, the entrainment zone juxtaposed to the cyclonic edge is therefore at least 5 km in this region. TABLE 4. Summary of Gulf Stream measurements during the Florida Atlantic Coast Transport Study. Date Buoy speed at Distance to deployment cyclonic edge* (cm/s) (km) 84Apr06 50 +4 84Mayl1] 85 +6 84Junl2 20 +13 84Jull2 115 ~2 84Aug07 150 fe 84Sep13 85 +4 84Oct16 40 +13 84Nov15 5 Ie 84Dec11 90 —2 85Jan25 135 i? 85Feb15 yp) +6 85Marl16 40 0 *Measured on infrared images from the Geostationary Environmental Operational Satellite as in Gaby and Baig (1983). With respect to drift card returns, data in Table 4 tend to support the notion that those months that had drift card returns were months when the Gulf Stream’s cyclonic edge was farthest east. Buoy 2194, September 1983, was deployed 25 km west of 28°N/80°W, and little direct influence of the current was evident. On the other hand, buoy 1776, May-June 1979, passed about 10 km east of 28°N/80°W at a speed of 150 cm/s. Although drift cards were not deployed with buoys 2194 and 1776, the ensemble of data suggests that an object deployed in the 5 km-25 km wide zone juxtaposed to the Gulf Stream tends to be entrained. Variability of initial buoy speeds with respect to volume transport of the Gulf Stream is summarized in Fig. 4a. Volume transports were obtained from voltages induced into the submarine cable (Larsen and Sanford, 1985) shown 166 FLORIDA SCIENTIST [Vol. 52 in Fig. 1. The large dots plotted on Fig. 4a are the speeds of the buoys in the half day or so immediately following deployment (q.v. Table 4). Except for March 1985, the initial buoy speeds follow the transports rather well. Spectra of transport from Fig. 4a (not shown) display organized energy at fortnightly, monthly, and longer periods. Similarly, spectra of lateral motion in the Gulf Stream’s cyclonic edge off Onslow Bay, NC (Maul, et al., 1978) show orga- nized energy at fortnightly and monthly periods, as well as at 6 and 10 days (cf. Johns and Schott (1987) who found spectral energy in volume transport from mooring data across 27°N at 5 and 12 days). 42 200 40 175 28 e I50 w > = " 25 2 L 34 | | | : o | | | loo & oe | | Lf e = g ac ye A i S | | 753 A ss | Wy) PAM Ay : a { . \ | \ 50 a sa 28 " {| & = a -25S 22 a MmO—-NMYGoORD HNO— © - se SESS Te ere ers = attt+t¥¢¢e¢¢ t+ 18 =79 O 100 200 300 400 500 600 700 800 900 1983 1984 1985 Fic. 4a: Cable transports at 27°N from the method of Larsen and Sanford (1985) during FACTS. Solid central line is the least squares annual plus semiannual harmonic fit. Dots are the buoy speeds in the first 12 hours after deployment. Buoy identification numbers mark each deployment date using 1 January 1983 for the datum. Fig. 4b shows the surface speed across the current at 27°N between Flor- ida and Grand Bahama Island as a function of volume transport, based on direct observations during 1982-1984 (Leaman et al., 1987). As the volume transport increases, the northward surface speed increases, particularly in the western half of the Straits of Florida. There also is a shift of the maximum surface speed from near mid-stream at low transports, to the extreme western side during high transports. Although data shown in Fig. 4b were not tempo- rally coincident with the buoys, the high correlation between transport and cross-stream average surface current speed (r=0.85) suggests that the rela- tionship between transport and current structure is significant. No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 167 Both the buoy data and the surface speed data support the notion that higher transports are associated with a westward shift of the Gulf Stream axis on both the long term and at higher frequencies. In a sense, Figs. 3a-3d, respectively, are representative of spring, summer, autumn, and winter tra- jectory conditions. In spring, transports are intermediate in strength, and buoy trajectories are moderately consistent. Summer transports are highest and the trajectories are almost identical. Autumn trajectories are the least consistent, and transports are lowest. Winter transports again have interme- diate strength, and the trajectories are more uniform than in autumn. Lower autumn transports and an associated eastward shift in the Gulf Stream, coup- led with higher percentages of winds from the east, make autumn a time of higher percent frequency of occurrence for flotsam and jetsam coming ashore from 28°N/80° W. 0 20 40 60 80 Distance Offshore (km) Fic. 4b: Northward surface speed (cm/s) in the Gulf Stream as a function of volume transport in Sverdrups (1 Sv = 106 m/s) and distance offshore of Florida at 27°N. 168 FLORIDA SCIENTIST [Vol. 52 SO°N 40° Otis ON Mie ‘ 49h 50: 70° 60° 210) 40° 30% 20° 10° Fic. 5: Composite of 15 satellite-tracked buoys analyzed in FACTS, plus buoy 4482 (May 1985-June 1986) released northeast of Grand Bahama Island, The Bahamas. The entire data set (Fig. 5), is consistent with the satellite study of Gaby and Baig (1983) that shows variance in the location of the western edge of the Gulf Stream from a minimum of o= +7 km off St. Lucie Inlet to a maximum of o= + 20 km off St. Augustine, another minimum (o= +13 km) off Savan- nah, and a maximum of o= +32 km off Frying Pan Shoals (33.5°N); the offing of Cape Hatteras shows another minimum. Three areas where detrain- ment seems to be most prevalent are: southeast of Cape Canaveral, east of St. Augustine, and southeast of Onslow Bay, NC. Although only one drift card was received from north of 31°N, both drift card and buoy data suggest that these three coastal areas seem most likely to be where detrainment will occur. If a buoy or a card is sufficiently west of the Stream, it then will be subject to the vagueness of local winds and it seems likely that the prevailing easterlies and Stokes drift will bring it ashore. Fig. 5 also shows the ultimate fate of all the FACTS buoys in the North Atlantic Ocean, and one released east of Grand Bahama Island. The average tracked lifetime of the 12 FACTS buoys was 10 months; the longest was 18 months. Buoy 4479, launched in January 1985, functioned for 14 months and arrived off the Iberian Peninsula in April 1986. All buoys that passed Cape Hatteras, except 4479 and 4473 (which lasted one week), ended their trans- missions in the central North Atlantic, south of the latitude of the Iberian Peninsula, i.e., more than 80% of the buoys released at the western edge of the Gulf Stream off 28°N/80°W ended south of the Gulf Stream System in the Sargasso Sea. No. 3, 1989] MAUL AND BRAVO— BUOYS AND DRIFT CARDS 169 ACKNOWLEDGMENTS—ARGOS-tracked buoys used in this study were designed by Mr. David S. Bitterman, and were built and launched by Mr. Robert J. Roddy. Processing of the ARGOS data was by Mrs. Mayra C. Pazos using programs developed by Mr. Alan Herman. Drafting was by Mr. David A. Senn; photography was by Mr. Andrew J. Ramsay, Jr.; Shell Offshore Inc. buoy data was obtained through the efforts of Dr. George Z. Forristall. Satellite infrared images of the Gulf Stream’s cyclonic shear edge were analyzed by Dr. Stephen R. Baig of NOAA’s National Weather Service. Volume transports from cable voltages were kindly provided by Dr. Jimmy C. Larsen of NOAA's Pacific Marine Environmental Laboratory; Mr. Mark H. Bushnell prepared the plot. Finally, we wish to express our appreciation to the Florida Institute of Oceanography, and particularly to Mr. Maurice O. Rinkel and Dr. Sandra L. Vargo, for their management of FACTS. This research was funded in part by the U.S. Department of the Interior, Minerals Management Service, under MMS Agreement No. 14-12-0001-30082. LITERATURE CITED BITTERMAN, D. S., AND D. V. Hansen. 1986. The Design of a Low Cost Tropical Drifter Buoy. Marine Data Systems International Symposium Proceedings, Mar. Tech. Soc., 575-581. Cuew, F., J. M. BANE, JR., AND D. A. Brooks. 1985. On vertical motion, divergence, and the thermal wind balance in cold-dome meanders: A diagnostic study. J. Geophys. Res., 90(C2):3173-3183. Gasy, D., ann S. Baic. 1983. Gulf Stream variability and width. Mar. Weather Log, 27(3):133- 134. HERMAN, A., AND D. V. HANSEN. 1977. Objective analysis of Lagrangian trajectory data. Pp 177- 185. In: Proceedings of the American Congress on Surveying and Mapping, 37th Annual Meeting, Washington, D.C. Jouns, W. E., AND F. Scuott. 1987. Meandering and transport variations of the Florida Current. J. Phys. Oceanogr., 17(8), 1128-1147. LaRSEN, J. C., AND T. B. SANForpD. 1985. Florida Current Volume Transports from Voltage Mea- surements. Science. 227:302-304. LeaMaN, K. D., R. L. Mouinari, AND P. S. Vertes. 1987. Structure and variability of the Florida Current at 27°N: April 1982-July 1984. J. Phys. Oceanogr., 17(5):565-583. Lee, T. N. 1983. Preface to a special issue on circulation in the South Atlantic Bight. J. Geophys. Res. 88(C8):4539. , AND L. P. Atkinson. 1983. Low-frequency current and temperature variability from Gulf Stream frontal eddies and atmospheric forcing along the southeast U.S. outer conti- nental shelf. J. Geophys. Res. 88(C8):4541-4567. Leminc, T. D. 1979. Observations of temperature, current, and wind variations of the central eastern coast of Florida during 1970 and 1971. NOAA Technical Memorandum NMFS- SEFC-6, 172 pgs. MauL, G. A., anD R. L. Mo.inari. 1975. Pollutant Trajectories. Pp 69-77 In: Compilation and Summation of Historical and Existing Physical Oceanographic Data from the eastern Gulf of Mexico. Final Report, Contract No. 08550-CT4-16 for the U.S. Bureau of Land Management from the State University System (SUS) of Florida Institute of Oceanogra- phy, St. Petersburg. , P. W. peWitt, A. YANAWAy, AND S. R. Batic. 1978. Geostationary Satellite Observa- tions of Gulf Stream Meanders: Infrared Measurements and Time Series Analysis. J. Geophys. Res. 83(C12): 6123-6135. . 1985. FACTS: The Florida Atlantic Coast Transport Study. EOS. 66(19):434-435. Murpny, E. B., K. A. Strerprncer, B. S. Roperts, J. WILLIAMS, AND J. W. Joey, fr. 1975. An explanation for the Florida east coast Gymnodinum breve red tide of November 1972. Limnol. Oceanogr. 20(3):481-486. RINKEL, M., S. Varco, T. Lee, F. ScHott, R. ZaAntopp, K. Leaman, N. Smiru, G. Maut, J. PRoNI, AND A. D. Kirwan. 1986. Final Report, Physical Oceanographic Study of Flori- da’s Atlantic Coast Region—Florida Atlantic Coast Transport Study (FACTS). SUS Flor- ida Institute of Oceanography, St. Petersburg, FL, Vols. 1-3. Parker, R. D., J. M. Morrison, anD W. D. Now tin, Jr. 1979. Surface drifter data from the Caribbean Sea and Gulf of Mexico, 1975-1978. Technical Report 79-8-T, Department of Oceanography, Texas A&M University, 157 pgs. 170 FLORIDA SCIENTIST [Vol. 52 SmiTH, N. P. 1987. Near-bottom cross-shelf heat flux along central Florida’s Atlantic shelf break: Winter months. J. Geophys. Res., 92(C10): 10843-10852. U.S. Navy. 1974. U.S. Navy Marine Climatic Atlas of the World, Volume 1, revised North Atlan- tic Ocean, NAVAIR 50-1C-528, Naval Weather Service Detachment, Asheville, NC. WiuuiaMs, J., W. F. Grey, E. B. MurpHy, AND J. J. CRANE. 1977. Drift bottle analyses of eastern Gulf of Mexico surface circulation. In: Memoirs of the HOURGLASS Cruise, Vol. IV(III), Marine Research Laboratory, Florida Department of Natural Resources, St. Petersburg, FL, 134 pgs. VuxovicH, F. M., anp G. A. Maut. 1983. An observation of the surface circulation in a Gulf Stream frontal perturbation. Geophys. Res. Ltrs., 10(7), 591-594. ZANTOPP, R. J., K. D. LEAMAN, AND T. N. LEE, 1987. Florida Current meanders: A close look in June-July 1984. J. Phys Ocean. 17(5):584-595. Florida Sci. 52(3):154-170. 1989. Accepted: September 2, 1988. Biological Sciences THE ABUNDANCE OF Aeromonas hydrophila L. AT LAKE HARNEY ON THE ST. JOHNS RIVER WITH RESPECT TO RED SORE DISEASE IN STRIPED MULLET (Mugil cephalus L.) JOHN A. OsBoRNE”), GERALD E. FENSCH ®, AND JULIUS F. CHARBA") (Department of Biological Sciences, University of Central Florida, P.O. Box 25000, Orlando, Florida 32816; Joyce Environmental Consultants, Inc., 414 Live Oak Blvd., Casselberry, Florida 32707 Asstract: The density of Aeromonas hydrophila L. in the St. Johns river at Lake Harney was examined within the environment (water and sediment) and on striped mullet (skin and stom- ach) between February and August, 1982. High densities of this pathogenic bacteria occurred within the environment during midsummer when sedimentary chlorophyll, and water tempera- ture were highest. Diseased striped mullet (fish with skin lesions), containing high densities of the bacteria within their stomachs and on their skin, were more abundant during summer. This suggests that mullet, browsing on bacteria-laden sediment for algae, accumulate bacteria within their gut and on their skin which, in turn, enhances infections of red sore disease. Significant correlation coefficients were not found between Aeromonas hydrophila L. abundance (water and sediment) and dissolved oxygen, pH, total alkalinity, specific conductivity, and planktonic chlo- rophyll,. Aeromonas hydrophila L. is a gram-negative, asporogenous, motile (monotrichous polar flagellations) rod (Schubert, 1974; Ewing and Hugh, 1975). Infection of this bacteria causes furunculosis or red sore disease; a worldwide disease of cultured and wild fishes. Furunculosis was described in 1894 after being isolated from diseased trout taken from a German hatchery (Emmerich and Weibel, 1894). It was first reported in the United States in brook trout (Salvenlinus fontinalis L.) in 1902 (Marsh, 1902). The disease causes generalized septicemia, skin lesions, termed furuncles, and in acute cases, death (Davis, 1967). Outbreaks of the disease in striped mullet (Mugil cephalus L.) within the St. Johns River and throughout Florida have been frequently publicized (Fogt, 1980). Degraded environmental conditions within the St. Johns River, by providing a more favorable environment for the growth of A. hydrophila L., may be responsible for the frequency of this disease. High bacteria counts for A. hydrophila L. are characteristic of eu- trophic environments having high chlorophyll, concentrations (Hazen and Fliermans, 1979; Rippey and Cabelli, 1980). The objective of this study was to examine the relationship between A. hydrophila L. infections of striped mullet (Mugil cephalus L.), the abun- dance of the bacteria within the environment, and environment conditions at Lake Harney on the St. Johns River. 172 1000 FLORIDA SCIENTIST COWHOUSE SLOUGH UNDERHILL SLOUGH LAKE HARNEY g Re, Wz 2 pl METERS N 1000 2000 t [Vol. 52 Fic. 1. The location of the sampling areas (Underhill Slough and Cowhouse Slough) at Lake Harney on the St. Johns River, Florida. No. 3, 1989] OSBORNE, ETAL. ABUNDANCE OF AEROMONAS HYDROPHILA 173 METHODS AND MATERIALS— Two study areas, within the St. Johns river at Lake Harney in Seminole County, Florida, were utilized. Underhill Slough is located in the northeast corner of Lake Harney and Cowhouse Slough is located approximately 3.7 km downstream from Lake Harney on the St. Johns River (Fig. 1). The water depth approximated 1.0 m within these areas. The sediment at Underhill Slough was predominately sand; at Cowhouse Slough the sediment was a dark brown silt-laden mud. Alligator weed (Althernanthera philoxeroides Mart.), hydrilla (Hydrilla verticillata (L.f.) Royle), waterhyacinth (Eichhornia crassipes (Mart.) Solms.), and coontail (Ceratophyllum demersum L.) were abundant at Underhill Slough; Cowhouse Slough did not contain vegetation. Physicochemical parameters and A. hydrophila L. were measured monthly between Febru- ary and August, 1982 at four stations within each study area. Chlorophyll, concentration and bacteria density were determined from water and sediment samples. These samples were col- lected with sterile 4.7 cm dia. plastic cylinders inserted into the sediment to a depth of 15 cm. Water samples (10 ml) were removed aseptically from just above the sediment prior to removing two | gsediment samples from the upper 1 cm. Enumeration of A. hydrophila L. from water and sediment samples was performed with the membrane filter technique (pore size=0.45n) of Rip- pey and Cabelli (1979). Membrane filters were incubated on 5 ml Rippey-Cabelli differential agar per plate and incubated for 20 hr at 37°C. The filters wee examined for A. hydrophila L. colonies (circular, convex and yellow); if colonies were present the filters were transferred to mannitol agar and incubated for 2 hr at 37°C. Only the colonies that remained yellow (i.e. those that fermented mannitol) were counted. The filters were then transferred to a culture pad satu- rated with phosphate-buffered saline (Levin and Cabelli, 1972) for 60 sec to neutralize the or- ganic acid end products which override the color reaction of the oxidase test. The colonies were tested for cytochrome oxidase with 1% oxidase reagent (N,N,N’,N’-tetramethy]l-p-phenylene- diamine dihydrochloride) and positive oxidase colonies were included in the count. A reference culture of A. hydrophila (American Type Culture Collection (ATCC, E9071)) was maintained as a check on the accuracy of the Rippey and Cabelli (1979) method. Chlorophyll, concentrations within the sediment (1 g samples) and water were determined with the Richard with Thompson (1952) method. Water samples were collected at mid-depth with a Kemmerer water sampler for turbidity, dissolved oxygen and total alkalinity (APHA, 1981). Water temperature and specific conductivity were measured with a Montedoro Whitney SCT meter at mid-depth. Striped mullet were captured by electrofishing (10 amp; 150 volts; AC) at night. Skin mucous samples were taken using an alcohol-flamed plexiglass template with a 6 cm? window. Two skin samples for a total of 12 cm? were taken per fish. A sterile cotton swab was used to obtain each sample; the swabs were transported in separate 4 ml sterile phosphate buffered saline solutions (Levin and Cabelli, 1972). A 100 mg sample from each stomach was removed and ground in 9.0 ml sterile distilled water to assay for A. hydophila L. (Rippey and Cabelli, 1979). Chlorophyll, concentrations within stomachs were determined with the Richards with Thompson (1952) method. The Spearman’s rank correlation coefficient was used to make statistical comparisons between bacteria levels and environmental parameters (Choi, 1978). RESULTS AND Discussion—The water depth at Underhill and Cowhouse Sloughs were at their lowest level in February, 1982; low water levels per- sisted until April 8, 1982, the first major rain event of the season. Even so, fluctuations in water depth were less than 1 m throughout the study (Table 1) and were not considered a major factor governing bacterial densities. Monthly mean water temperatures at Underhill and Cowhouse Sloughs were 18.1°C and 21.5°C, respectively, in February, 1982, increasing to their maxi- mum values (greater than 30°C) by July. Dissolved oxygen concentrations at the sampling stations were highest at the beginning of the study, then de- creased in March to remain low (3-4 ppm) throughout the remainder of the study. The lowest monthly mean dissolved oxygen concentration occurred in April (2.1 ppm) at Underhill Slough and in August (1.6 ppm) at Cowhouse Slough. Relatively constant pH values were found at both sites (Table 1). The 174 FLORIDA SCIENTIST [Vol. 52 pH ranged from 6.2 to 7.4 at Underhill Slough and from 6.6 to 7.5 at Cow- house Slough. The highest turbidity (24-62 NTU) was recorded in February and March (Tablel). Turbidity tended to remain fairly low (2-4 NTU) at the study sites after May, 1982. Total alkalinity ranged from 23.5 mg/1 CaCO, in April to 60.5 mg/1 CaCO, in February at Underhill Slough; alkalinity values were only slightly lower at Cowhouse Slough (Table 1). Specific conductivity were similar between the two study areas. Minimum values for specific con- ductivity were recorded in August (Table 1). TABLE 1. Monthly mean values for physicochemical parameters taken at Underhill and Cow- house Sloughs on the St. Johns River from February through August, 1982. Standard errors of the overall means are given in parentheses. Water Water Dissolved Total Specific Temperature Depth ‘Turbidity Oxygen Alkalinity Conductivity Month CC) (m) (NTU) (ppm) pH (mg/L CaCO,) (umhos/em @ 20°C) Underhill Slough February 18.1 0.28 4] 7.8 7.4 60.5 2962 March 24.8 0.37 24 3.9 6.6 315 1434 April 2321 0.76 15 2.1 6.2 23.5 720 May 24.9 0.60 24 3.8 Gaul 36.5 1396 June 29.4 O52 2 oe 6.6 33.2 1143 July 31.6 O72 4 320 6.8 43.0 765 August 28.6 0.94 2 3.7 6.8 47.0 635 Overall Mean 25.8 0.60 16 oy) 6.7 39.3 1294 (1.6) (0.08) (5) (0.6) = (0.1) (4.2) (635) Cowhouse Slough February NS) 0.34 60 8.3 ee) 57.8 3148 March 25.6 0.33 62 5.8 6.9 56.8 3168 April 25.2 0.44 41 3.8 6.6 37.0 1712 May 26.9 0.56 6 7.0 (3 44.2 2105 June 28.6 0.50 2 2.8 6.9 36.8 1380 July 31.9 0.78 D 3.2 6.8 49.0 780 August 28.8 0.85 3 1.6 6.7 46.5 565 Overall Mean 26.9 0.54 25 4.6 6.9 46.9 1837 te) (0.07) (10) (029) > 2(OR1) (3.0) (365) Planktonic chlorophyll, concentrations generally exceeded 30 pwg/L (eu- trophic waters) in May and June; these high values are attributed to an algal bloom, enhanced by increased runoff and warm water conditions. These values were accompanied by high sedimentary chlorophyll, concentrations even though the correlation coefficient for planktonic and sedimentary chlo- rophyll, was not significant (r=0.262, P=0.05, n=48). Chlorophyll, concen- No. 3, 1989] OSBORNE, ET AL. ABUNDANCE OF AEROMONAS HYDROPHILA aD TaBLE 2. Mean chlorophyll, concentration in water, sediment and within stomach samples from striped mullet taken from the St. Johns River, February through August, 1982. Standard errors of the means are given in parentheses. Chlorophyll, Concentration February March April May June July August Water* Underhill Slough 43.8 23.9 1532 45.7 142.4 20.0 28.3 Cowhouse Slough Slee 29.5 Aira 58.4 29.9 Aaa ORT Mean 47.8 26.7 28.2 52a 86.2 el 19.5 (2.8) (2.0) (9.2) (4.5) (39.8) (2a) (6.2) Sediment? Underhill Slough 555 18.3 16.1 14.3 20.8 9.5 10.2 Cowhouse Slough 28.6 11.6 13.1 23.4 24.7 16.2 23.0 Mean DOR 15.0 14.6 18.9 22.8 12.9 16.6 (4.6) (2.4) (1.1) (3.2) (1.4) (2.4) (4.5) Stomach content® $5.0 59.1 20501 63:5 “ug/L of river water. °mg/g dry weight of sediment. “mg/g dry weight stomach content of striped mullet, n= 27. trations in water and sediment were not significantly correlated with the chlorophyll, content in stomach samples of striped mullet, but often high values for each occurred simultaneously. For example, in June, 1982 the max- imum chlorophyll, concentration within striped mullet stomachs was 205.1 mg/g dry weight; this coincided with the highest mean chlorophyll, concen- trations recorded for water (39.8 mg/ml) and sediment (22.8 mg/g) (Table 2). The mean A. hydrophila L. density within the sediment was lowest in June (0.72 x 10° organisms/g dry weight); this low density corresponded with the minimum density of bacteria found in water (0.26 x 10° organisms/100 ml) (Table 3). The density of A. hydrophila L. within the water and sediment was not significant correlated with the bacteria density in the water (r=0.175, P=0.05, n=48). A significant correlation was determined be- tween the density of A. hydrophila L. in water and turbidity (r=0.351, P< 0.05, n=48). No significant correlation was determined between the abun- dance of sedimentary A. hydrophila L. and water temperature (r=-0.217, P=0.05, n= 48) in our study. The densities of A. hydrophila L. on the skin and in the stomachs of striped mullet were lowest in February; they were highest in fish stomachs in April (54.9 x 10° organisms/g dry weight) and on the skin in June (48 orga- nisms/cm’). The density of A. hydrophila L. in water was not statistically correlated (P=0.05) to the density of the bacteria on fish skin or within fish stomachs. The density of bacteria in stomach samples was statistically correl- ated (r=0.8, P<0.05, n=48) with the stomach chlorophyll, content. On the other hand, no significant correlations were found between bacterial densi- ties (water, sediment, stomach or skin) and water depth, dissolved oxygen, pH, total alkalinity, specific conductivity or planktonic chlorophyll,. The 76 FLORIDA SCIENTIST [Vol. 52 lack of a significant correlation between bacterial density and planktonic chlorophyll, was probably due to a plateau that generally occurs for A. hy- drophila L. density when planktonic chlorophyll, exceeds 30ug/L (Rippey and Cabelli, 1980). In our study areas, skin lesions on striped mullet became apparent between April and July when the fish had high bacteria levels on their skin and within their stomachs, the water was warm, and high concen- trations of sedimentary bacteria and chlorophyll, were found. Browsing mul- let, by grazing sedimentary algae laden with A. hydrophila L., probably created their own infections of red sore disease. Dense populations of benthic algae in eutrophic waters, stimulated by warm summer temperatures, pro- motes high levels of A. hydrophila L.; in turn, striped mullet are exposed to these bacteria by grazing the algae, their primary food. Since high densities of A. hydrophila L. are the result of eutrophication, the degradation of the St. Johns River will probably result in more frequent outbreaks of red sore disease in striped mullet. TaBLE 3. Mean number of Aeromonas hydrophila L. in water, sediment and on striped mullet at Underhill and Cowhouse Sloughs on the St. Johns River, February through August, 1982. Standard errors of the means are given in parentheses. Number of Bacteria February March April May June July August Water* Underhill Slough 32 4.80 4.03 1.08 0.46 0.98 DFA5 Cowhouse Slough 0.69 6.47 0.31 1.25 0.06 0.58 0.50 Mean 1.01 5.64 mn Mealy 0.26 0.78 1.48 (0.22) (0.59) (1.32) (0.06) (0.14) (0.14) (0.69) Sediment? Underhill Slough 2.56 BAL 10.34 3.94 0.70 4.06 13.94 Cowhouse Slough 6.67 6.56 5.16 9.23 0.74 3.82 3.62 Mean 4.62 4.99 TS 6.59 0.72 3.94 8.78 (1.45) (iit) (1.83) (1.87) (0.01) (0.08) (3:65) Stomach content® 0.29 54.90 4.82 DEAD, Skin® 3.0 25.0 48.0 4.0 a ‘per 100 ml river water x 10-3. per g dry weight river sediment x 10-3. er g dry weight stomach content of striped mullet x 10-3, n= 27. per cm? skin area on striped mullet, n= 27. Cc No. 3, 1989] OSBORNE, ET AL. ABUNDANCE OF AEROMONAS HYDROPHILA G7, LITERATURE CITED AMERICAN PuBLic HEALTH ASSsOocIATION (APHA). 1981. Standard Methods for the Examination of Water and Wastewater. APHA-AWWA-WPCF, Washington, D.C. 1134 pp. Cuo1, S.C. 1978. Introductory Applied Statistics in Science. Prentice-Hall, Inc., Englewood Cliffs, N.J. 278 pp. Davis, H.S. 1967. Culture and Diseases of Game Fishes. Univ. California, Berkley and Los Angeles. EMMERICH, R., AND C. WEIBEL. 1894. Ueber eine durch Bacterien erzeugte Seuche unter den Forellen. Arch Hyg. Bacteriol. 21:1-21. Ewinc, W.H., anp R. Hucu. 1975. Manual of Clinical Microbiology. Am. Soc. Microbiol., Washington, D.C. Focr, J. 1980. Fungus or pollution?, Here’s an exclusive investigative report on an alarming fish disease that may be far more ominous than the state maintains. Florida Sportsman. 26-32. Hazen, T.C., AND C.B. FLIERMANS. 1979. Distribution of Aeromonas hydrophila in natural and manmade thermal effluents. Appl. Environ. Microbiol. 8:166-168. Levin, M.A., AND V.J. CABELLI. 1972. Membrane filter technique for enumeration of Pseudo- monas aeruginosa. Appl. Microbiol. 24:864-870. Marsu, M. 1902. Bacterium truttae, a new species of bacterium pathogenic to trout. Science N.S. 16:706-707. RicHarp, F.A. witH T.G. THompson. 1952. The estimation and characterization of plankton populations by pigment analysis. II. A spectrophotometric method for the estimation of plankton pigments. J. Mar. Res. 2:156-172. Rippey, S.R., AND V.J. CABELLI. 1979. Membrane filter procedure for enumeration of Aeromonas hydrophila in fresh waters. Appl. Environ. Microbiol. 38:108-113. AND V.J. CABELLI. 1980. Occurrence of Aeromonas hydrophila in limnetic environ- ments: Relationship of the organism to trophic state. Microbiol. Ecol. 6:45-54. ScHUBERT, R.H. 1974. Bergey’s Manual of Determinative Bacteriology. Williams and Wilkins, Baltimore. Florida Sci. 52(3):171-177. 1989. Accepted: September 17, 1988. THE OCCURRENCE OF THE CRAYFISH Fallicambarus (Creaserinus) Fodiens IN FLORIDA— Barry W. Mansell, 2826 Rosselle Street, Jacksonville, Florida 32205. AsstTrAcT: On November 16, 1985, a specimen of the crayfish Fallicambarus (Creaserinus) fodiens (Cottle, 1863) was collected from the Escambia River flood plain south of Florida High- way 4, 6.6 km west of Jay, Santa Rosa County, Florida. This is the first identifiable specimen of F. (Creaserinus) fodiens from Florida. Hosss (1942) recorded an undescribed crayfish, Cambarus sp. incertis, from the Escambia River flood plain in Escambia and Santa Rosa Counties, Florida. He had not seen a first form male, and identification to species could not be made. However, he assigned it to the genus Cambarus on the basis of a second form male. In his revision of the genus Cambarus, Hobbs (1969) re- ferred this still unidentified crayfish to the newly erected genus Fallicam- barus and later assigned it to the subgenus Creaserinus (Hobbs, 1973). 178 FLORIDA SCIENTIST [Vol. 52 On November 16, 1985, while searching for specimens of Fallicambarus in the Escambia River flood plain south of Highway 4, 6.6 km west of Jay, Santa Rosa County, Florida, I dug a second form male of this crayfish from a burrow at the edge of an overflow pool. It was maintained in an aquarium where it molted to first form in late January, 1986. It was subsequently identified as F. (Creaserinus) fodiens (Hobbs, pers. comm.). This specimen provides the first record of F. (Creaserinus) fodiens from Florida (Hobbs, pers. comm.) and extends the known range to include the Escambia River flood plain in Escambia and Santa Rosa Counties, Florida. The species has been collected in the Chattachoochee drainage in Early and Seminole Counties, Georgia (Hobbs, 1981) and may eventually be found in the Chattahoochee/ Apalachicola drainage in Florida as well. The collection of this crayfish and its subsequent identification as F. fodiens clears up a long-standing mystery about the specific status of Hobbs’ Cambarus sp. incertis. ACKNOWLEDGMENTS— The author wishes to thank Paul Moler and Richard Franz for reading and commenting on the manuscript. I would also like to thank Paul Moler for his help in search- ing for Fallicambarus on many occasions. I would also like to thank Dr. Horton H. Hobbs, Jr. for reviewing the manuscript and guidance and encouragement while I searched for specimens of Fallicambarus. LITERATURE CITED Hosss, H. H., Jr. 1942. The Crayfishes Of Florida. Univ. Florida Publ., Biol. Sci. Ser. 3(2):179 Pp. , 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pp 93-178. In Hott, P. C., R. L. HorrmMan, ann C. W. Hart, Jr., (eds.) The distributional history of the biota of the southern Appalachians. Part 1: Invertebrates. Va. Polytechnic Inst., Re- search Div. Monograph 1. , 1973. New Species and Relationships of the Members of the Genus Fallicambarus. Proc. Biol. Soc. Washington, 86(40):461-481. , 1981. The Crayfishes of Georgia. Smithsonian Contrib. Zool. No. 318. 549 pp. Florida Sci. 52(3)177-178. 1989. Accepted: September 29, 1988. No. 3, 1989] PREECE— IMPACT OF ENGINEERING 179 WHAT IS THE IMPACT OF ENGINEERING ON FLORIDAS ECON- OMY—TODAY AND TOMORROW? -—Betty Preece, Melbourne High School, Melbourne, FL 32901. Asstract: The Engineering Sciences Section of FAS held a panel discussion at the 1988 An- nual Meeting to discuss ways in which engineers impact Florida’s Economy. Representatives of engineering education, construction engineering, high-technology industry, health and safety engineering, agricultural engineering, municipal engineering and space engineering all outlined specific impacts and made suggestions for increasing these impacts in positive ways. The panel and the FAS Section feel that these need wider dissemination to Florida's scientists and engineers. Durinc the FAS 1988 annual meeting, the Engineering Sciences Section sponsored a symposium with recognized leaders in Florida of important branches of engineering. The Moderator, George R. Knecht, P.E., President, Florida Engineering Society, Jacksonville, opened the discussion by stating the purpose of the Symposium was to arrive at some basic ways in which the panel agreed engineering was important to and will continue to impact Flori- da’s economy. EDUCATION was the first area, discussed by Melvin W. Anderson, Ph.D., P.E., Chairman, Department of Civil Engineering and Mechanics, Univer- sity of South Florida, Tampa. He cited three past engineering achievements that have had significant impact on Florida’s growth: (1) the rail transportation system from Atlanta to Key West which brought the first real opening of Florida to development; (2) the bridges and roads which have tied Florida people together; (3) the intro- duction of economically-priced residential air conditioning which opened Florida to year-round living. In the high-tech industry, engineering performance decides whether an industry withers or is prosperous. In 1980, only 16% of the engineers hired in Florida were educated in Florida; therefore, since the statistics have not greatly changed, Florida engineering schools are not meeting the state’s needs. (Olson, 1980; Anon, 1980; 1988b) Florida is a net importer of engi- neers. Florida engineering graduates also are now competing in a world-wide market for which the engineering schools trying to prepare them need greatly improved facilities (EDC, 1987). THE CONSTRUCTION INDUSTRY was represented by Peter D. Brown, Execu- tive Vice President, Peter Brown Construction Company, Tampa. In Florida, 7.5% of all jobs are construction-related and 34% of these are construction-mining related. The construction industry is a growth indicator in Florida. While nation-wide growth has been around 1% per year, in Flor- ida there has been as much as a 43% increase in growth in a single year. Engineers in construction should not be problem identifiers but problem solvers in a design-construction process that solves problems in design before construction begins. Changes in design are much cheaper than changes in a project under construction. Negotiated contracts allow this to be done more easily. Florida is increasingly turning to such contracts. The impact of engi- neers in construction on Florida’s economy is being improved by the engineer and architect working together as a team to evaluate designs and reduce both costs and legal liabilities. 180 FLORIDA SCIENTIST [Vol. 52 HicH TECHNOLOGY INDUSTRY was represented by Carmen Palermo, Vice President and Chief Scientist, Government Sector, Harris Corporation, Mel- bourne. In a comparison between US industry and both Japan and the four big European industrial countries (United Kingdom, France, Germany, Italy), the US has: (1) lower number of engineering graduates per capita; (2) lower academic ratings in all academic areas; (3) since 1960 dropped the rate of domestic patents issued by a factor of two while foreign patents issued in the US have increased; (4) dropped its productivity rate (as measured by eco- nomic growth and standard of living) while that of Japan has doubled. From the time of World War II to the 1960s, the US led in every produc- tivity indicator, but now the only area in which the US leads is research and development. Research and Development (R & D) is still the leading edge in productivity, but the US lags in the application of basic knowledge to indus- try—the role of engineering. Such engineering is important to Florida. Flor- ida must generate high-quality engineering, not necessarily in a high quan- tity, by: (1) recognizing the major burden for this falls on industry; (2) improving K-12 education; and (3) reducing serious regulatory, financial and legal restraints on engineering industries (Anon., 1988a). HEALTH AND SAFETY AND WOMEN IN ENGINEERING were represented by Sigrid Messmer, CSP, President, Commercial Safety Consultants, Winter Park, and President, Florida Section, Society of Women Engineers. Accidents occur because someone did not follow rules or prescribed proce- dures. Companies do not budget accidents so their costs come out of profits. When there is an accident, the company has many direct and indirect costs such as assistance to the worker and his family, expensive loss of time, equip- ment destroyed, lawyer and doctor fees, and services of a trained worker. If accidents could be prevented, lawyers and doctors would not have to deal with personal injury cases and insurance costs would go far down. No Florida college offers a degree in safety engineering or even a two-year safety technician program. Only 70 colleges in the whole United States do so. Companies can increase their profits by preventing accidents through safety engineering. Safety engineering can have an even greater positive ef- fect on improving Florida’s economy. AGRICULTURAL ENGINEERING was represented by Fedro Zazueta, Associate Professor of Agricultural Engineering, University of Florida, Gainesville. The impact of agricultural engineering on Florida’s economy can be mea- sured by: (1) the size of the agricultural work force which in Florida is 25% of the state’s total work force; (2) cash farm receipts on the farm itself which have more than doubled in the years 1974 to 1983 by rising from $2 to $5 billion; (3) stability of the agricultural work force that during the 1970's repression was great enough to maintain Florida at a level of “no growth” instead of at a decline; (4) size of the gross agricultural products produced, now around $39.6 billion, and agriculture is surpassed as the state’s most important industry only by tourism. No. 3, 1989] PREECE— IMPACT OF ENGINEERING 181 Agriculture consumes 43 % of the total water resources in Florida, so agri- culture is the most important user. Industry uses 21% of the total water resources and human consumption takes only about 20% (Shoemyen, 1987). A major contribution of agricultural engineering to Florida’s economy is to improve the efficiency of water use. The major target includes both quan- tity and quality. The most important area of agricultural engineering in the future is the implementation of traditional engineering systems to biological systems. MUNICIPAL SERVICES were represented by Michael Salmon, Administrator of Water Resources and Public Works, Tampa. The government engineer is a facilitator in Florida’s economy through: (1) construction of the infrastruc- ture of water, waste treatment buildings, roads; (2) being guardian of the gate to protect both the resources and the public safety. Engineers going into government need training in administration to work well with local governments and with construction/design industry. Agencies utilizing these engineers should allow the use of better purchasing guidelines. The City of Tampa may be the only city in the world to do a design/build on a multi-story high-rise office building. This has been so successful that the Florida Department of Transportation may also use design/build for road construction soon. This has potential for a profound impact on Florida’s economy. As a result of the immense growth of Florida, the role of the government engineer is increasing, especially in areas of legislated responsibility. The pub- lic is demanding both protection and responsibility from professional engi- neers. SPACE ENGINEERING was represented by George Mosakowski, Chief, Pro- jects Management Office, Kennedy Space Center, who said, “Engineering is the bridge between science and practicality.” Areas in which NASA engineering influences the nation’s economy in- clude space flight, space operations, aeronautics and space technology. In Florida, the economy is affected by the spaceflight operations at the John F- Kennedy Space Center. After a recent solicitation for space engineering projects to university re- search centers, no proposals from Florida schools were selected. Through its Office of Commercial Programs, NASA is encouraging Florida industry to get involved in space-related endeavors, particularly by high-tech small business. Sixteen private-sector R&D centers for commercial use of space have been established by NASA, but not one center is in Florida. About 50% of all US engineers work in one of ten states and Florida ranks 7th in those states; however, in ranking engineers per capita, Florida is 49th; in patents issued, Florida is 28th; in university R&D spending, Florida is 25th and in federal R&D spending, it is 23rd. (Schoemyn, 1987). The Florida university system is one of the top ten in the US but its K-12 educational system is near the bottom. A special Florida Chamber of Com- 182 FLORIDA SCIENTIST [Vol. 52 merce task group is now addressing both of these problems. One activity that can stimulate the schools towards improvement and study of space is the Young Astronaut Program that is now beginning to enter Florida schools. Florida has generally overlooked the opportunities afforded by the space program to develop technology and transfer it to the market place. Entrepre- nuerial scientists and engineers should begin using this vast resource to con- tribute to the economic well-being of Florida and of the country. CoNcLusiIon— Because the ways in which panel members have indicated that Florida engineers can impact Florida’s economy are all relatively easy to carry out, the panel recommended that they be communicated to a much wider audience in the Academy as a whole through publication in the Florida Scientist. LITERATURE CITED ANON. 1988a. Engineering degree statistics and trends— 1987, Engineering Manpower Bull. No 88:1-6. American Association of Engineering Societies, Inc., Washington, D.C. ANON. 1988b. Engineering enrollment highlights; Fall, 1988. Engineering Manpower Bull. No. 87-1-6. American Association of Engineering Societies, Inc., Washington, D.C. ANon. (1980). Recognization of Professional Schools of Engineering in the state of Florida. Prof. Policy. No. 12A, Florida Engineering Society No. 12A, Florida Engineering Society. ENGINEERING EDUCATION Comm iItTEE. 1987. Engineering Education in Florida—The Competi- tive Edge. Special Report Florida Engineering Society. Oxson, R. E. 1980, Prediction of success of graduate students in civil engineering. Civil Engn. Educ.. Fall 7-12: SHOEMYN, A. H. (ed) 1987. Florida Statistical Abstract, 21st Edition. University Presses of Flor- ida, Gainesville. Florida Sci. 52(3):179-182. 1989. Accepted: October 25, 1988. Symposium: “Groundwater Contamination and Protection in Florida” In this issue and the next one are collected papers presented at the Sym- posium held at the Fifty-Second Annual Meeting of the Academy. The Sym- posium was organized by Patrick Gleason, Ph.D., James M. Montgomery, Consulting Engineers, Inc. Owing to the press of time and other circum- stances, not all those who contributed to the symposium were able to contrib- ute to the collected papers. The importance and timeliness of the topic de- manded that deadlines be set and followed. We are grateful to those who contributed papers. We are grateful for support received from the Florida Academy of Sciences and from James M. Montgomery, Consulting Engineers, Inc., and we appreciate the services of Dr. Gleason, who served as Sympo- sium organizer and ad hoc Associate Editor for the contributed papers. DFM BBM Social Sciences EMERGING LEGAL ISSUES INGROUNDWATER CONTAMINATION CASES THOMAS J. GUILDAY AND RALPH A. DEMEO Huey, Guilday, Kuersteiner & Tucker, P.A., Tallahassee, Florida 32301 Asstract: Legal issues relating to groundwater contamination in Florida are rapidly chang- ing. The regulators responsible for protecting groundwater are increasingly more aggressive as scientific technique reveals the heretofore unknown scope of contamination caused by hazardous waste, hazardous substances, petroleum products, and pesticides. The states including Florida and private individuals recently have asserted common law remedies such as negligence, nui- sance, trespass, and indemnification to remedy groundwater contamination. Florida law is un- dergoing rapid metamorphosis and existing statutory and common law theories are being reeval- uated. Congress and the Florida Legislature should reexamine the legal issues and prepare new more meaningful remedies to groundwater contamination in Florida. Tuis article will outline emerging legal issues in groundwater contamina- tion cases. Recent trends include an increase in damages actions by govern- ments and private parties against active polluters and product manufacturers for environmental damage caused by toxic substances. These actions involve new applications of traditional legal theories. Theories of negligence, strict product liability, trespass and nuisance, along with statutory remedies, have emerged in recent suits by state and local governments against chemical man- ufacturers for environmental contamination. Florida law is clearly undergo- ing a rapid metamorphosis and existing statutory and common law theories are being reevaluated. Furthermore, the federal government is taking a more active role in the problem of groundwater contamination. Advances in scien- tific technique in part have made possible new theories of liability for expo- sure to toxic substances in the groundwater. Also, state and federal authorities only recently have begun to assess the scope of the groundwater contamina- tion problem in Florida. The thesis of this article is that Congress and the Florida Legislature should examine the legal and scientific issues in light of these developments and prepare more meaningful legislative remedies to groundwater contamination in Florida. FEDERAL Law—Congress has created a number of statutory schemes to address the problem of groundwater contamination. Chief among these are the Safe Drinking Water Act, 42 U.S.C. Section 200f et seg. (“SDWA’); the Clean Water Act, 33 U.S.C. Section 1251 et seq. (“CWA”); the Comprehen- sive Environmental Response, Compensation, and Liability Act, 42 U.S.C. Section 9601 et seq. (“CERCLA”); and the Resource Conservation and Re- covery Act, 42 U.S.C. Section 6901 et seq. (“RCRA”). The following is a brief analysis of the key provisions of each act. 184 FLORIDA SCIENTIST [Vol. 52 Safe Drinking Water Act The SDWA is drafted to provide assurances that public water systems meet certain minimum standards for the protection of public health. Because the majority of Florida residents rely on groundwater for their drinking wa- ter, the SDWA is an important tool in the protection of the groundwater supply. Congress enacted the original version of the SDWA in response to the detection of synthetic contaminants in public drinking water supplies. The original SDWA, intended to ensure the quality of water provided to con- sumers of public water systems,'* concentrated on measures necessary to “protect health to the extent feasible.” 42 U.S.C. Section 300g-1(a)(z) (1974) (emphasis added). Further, the SDWA contained provisions designed to pro- tect underground sources of drinking water,’ to the extent that those sources “can reasonably be expected to supply any public water system.” 42 U.S.C. Section 300h(d) (2). The provisions of the original SDWA, however, vested the EPA with discretion in deciding whether or not to set standards for cer- tain contaminants, this discretion again being triggered by health-based con- cerns.° Following the expiration of authorization of appropriations for carrying out the SDWA, and recognizing certain structural weaknesses in the original statute, Congress spent several years drafting what ultimately evolved into the 1986 Amendments.‘ The amendments dramatically change both the focus of the statute and the role of the federal government in promulgating and enforcing drinking water regulations. First, the amendments remove the dis- cretion the Administrator previously enjoyed in deciding whether or not to promulgate standards for drinking water contaminants.° Second, the author- ity of the EPA to force compliance with the provisions of the SDWA has been enhanced by the addition of statutory language authorizing the Administra- tor to issue administrative orders to violators, in lieu of the previous require- ment of filing suit in federal court.® Finally, recognizing the uncertainty involved in setting standards based on unknown or speculative health risks, the 1986 Amendments, while health-concerned, are technology based.’ The 1986 Amendments set forth numerous substantive changes that sim- plify and organize the federal drinking water program. The 1986 Amend- ments impact upon all areas of the previous statute, from regulation of con- taminants to enforcement of standards. Further, the SDWA as amended clarifies the roles of both government (at both the state and federal level) and private parties. The following is a brief outline of the effect of the 1986 Amendments upon government and industry. The Federal Role—The terms of the 1986 Amendments require, in addi- tion to those 80 contaminants specifically referenced by Congress, that the Administrator engage in a triennial review of contaminant regulations and list new contaminants that need to be regulated.’ “Each maximum contami- *Please see citations at end of article. No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 185 nant level goal established under this subsection shall be set at the level at which no known or anticipated adverse effects on the health of persons occur and which allows an adequate margin of safety.” 42 U.S.C. Section 300g-1(b) (4) (1986). Under the provisions of the 1986 Amendments, granular activated carbon is designated as a “feasible” treatment technique for removing syn- thetic organic chemicals.° However, another form of technology may be ap- proved as long as it is demonstrated to be “at least as effective in controlling synthetic organic chemicals as granular activated carbon.” 42 U.S.C. Section 300g-1(b) (5) (1986). Further, the 1986 Amendments dictate tha a regulation “shall not require that any specified technology, treatment technique, or other means be used for purposes of meeting such maximum contaminant level.” 42 U.S.C. Section 300g-1(b) (b) (1986). Thus, a system operator may choose among those treatment techniques that have proven feasible in the field to meet the designated maximum contaminant level. The 1986 Amendments also require EPA to promulgate regulations in two very significant areas. First, the language of Section 300g-1(b) (7) (c) requires the Administrator to promulgate primary drinking water regulations for sys- tem operators who draw their water from surface supply sources. These regu- lations will require filtration as a treatment technique. Second, within three years of the date of enactment of the 1986 Amendments,” “the Administrator shall propose and promulgate national primary drinking water regulations requiring disinfection as a treatment technique for all public water systems.” 42 U.S.C. Section 300g-1(b) (8) (1986) (emphasis added). Thus, the 1986 Amendments place a heavy burden on the EPA to promulgate numerous reg- ulations designed to ensure the future supply of contaminant-free drinking water. The State Role—Enforcement of Standards—The terms of the 1986 Amendments continue the provisions of the earlier SDWA in delegating pri- mary enforcement responsibility to the states to ensure that system operators comply with standards."! A state has primary enforcement responsibility when the Administrator determines that the state “has adopted drinking wa- ter regulations which are no less stringent than the national primary drinking water regulations in effect.” 42 U.S.C. Section 300g-z(a) (1) (1986) (emphasis added). In an attempt to “acquire primacy for the State of Florida under the Federal Act,’” Florida has enacted the Florida Safe Drinking Water Act, the provisions of which are set forth in Section 403.850, et seq., Florida Statutes (1987) (““FSDWA”), and Chapter 17-22, Florida Administrative Code. “These regulations adopt the national primary and secondary drinking water regula- tions of the federal government where possible and otherwise create addi- tional regulations fulfilling state and federal requirements.’ The addition of the approximately 80 newly listed contaminants," as well as those contaminants added through the triennial review process," there- fore, will place an increasing burden upon the states to monitor and enforce the provisions of the SDWA. This burden will manifest itself in the form of an 186 FLORIDA SCIENTIST [Vol. 52 increased workload for the state, and increased costs of testing to assure com- pliance. Much of this financial burden will probably be passed on to system operators in the form of fees and assessments for analysis of water samples, and review of construction plans. Further, it is likely that states will assess against system operators the cost of monitoring those systems found to violate standards. Legal Remedies—The 1986 Amendments substantially stiffen the penal- ties that may be assessed against violators, and give EPA the authority to enforce regulations by issuing administrative orders rather than suing in fed- eral court. Should the Administrator determine that a system operator is in noncompliance with any promulgated maximum contaminant level, he “may bring a civil action in the appropriate United States district court to require compliance.” 42 U.S.C. Section 300g-3(b) (1986). The court is not limited to issuing orders of compliance, however, as provisions for civil penalties also exist. The Amendments raise the maximum penalty that may be imposed upon violators from $5,000 per day to $25,000 per day.'® Moreover, and more significantly for system operators, the requirement in the original SDWA that the violation be willful has been deleted.” The Administrative enforcement section of the Amendments" authorizes the Administrator to issue orders requiring compliance with regulations. Safeguards are provided for affected system operators by requirements that EPA give notice and opportunity for a public hearing.'® Should the system operator violate or refuse to comply with the terms of an administrative en- forcement order, the Amendments authorize the Administrator to seek civil penalties. The civil penalty provisions subject violators to possible liability in the amount of “not more than $25,000 per day of violation.” 42 U.S.C. Sec- tion 300g-3(g) (3) (A) (1986). The statute authorizes the Administrator to assess civil penalties, following notice and opportunity for public hearing, in amounts not to exceed $5,000.” Should the Administrator seek to enforce a penalty greater than $5,000, however, EPA will have to sue in federal district court: The 1986 Amendments leave unchanged the citizen’s suit provisions of the original SDWA.” The original statute allows “any person” to commence a civil action on his own behalf against any person alleged to be in violation of the requirements of the SDWA.” Further, the statute requires that before commencing a civil action to require compliance by a violator, the citizen must first notify EPA, the State government of the state in which the violation occurs, and the alleged violator.“ This language provides governmental au- thorities with the first opportunity to “diligently pursue a civil action in a court of the United States to require compliance.” 42 U.S.C. Section 300g- 8(b) (1) (B) (1974). Finally, the statute preserves the right of any person to seek common law or other relief due to the presence of contaminants in drink- ing water.» Whether this savings clause extends so far as to allow property owners to sue for personal injuries resulting from exposure to regulated con- taminants is unclear and not developed by caselaw. No. 3, 1989] GUILDAY AND DEMEO—LEGAL ISSUES 187 The role of the State of Florida under the SDWA is discussed more fully below. Clean Water Act (C WA) The CWA reflects a Congressional effort to amend the Federal Water Pollution Control Act of 1948. “The objective of this Act is to restore and maintain the chemical, physical, and biological integrity of the Nation’s wa- ters.” 33 U.S.C. Section 1251(a). In an effort to achieve this objective, the CWA authorizes the Administrator of the EPA to establish effluent limitations for all point sources (defined very broadly in Section 1362(14)) from which pollutants are discharged. In order to avoid liability for the discharge of a substance regulated pursuant to Section 311 of the CWA, a source must ob- tain a National Pollution Discharge Elimination System (“NPDES”) permit. An NPDES permit, granted in accordance with the provisions of 33 U.S.C. Section 1342, allows the Administrator to “issue a permit for the discharge of any pollutant, or combination of pollutants, notwithstanding Section 301 (a) [33 U.S.C. Section 1311(a)],” which generally renders such discharge unlaw- ful. Although the provisions of the CWA apply primarily to navigable waters, which is defined to mean “waters of the United States,’ 33 U.S.C. Section 1362(7), the CWA has been utilized to establish liability for groundwater contamination.” In general, however, the provisions of the CWA will usually apply only where a facility discharges hazardous substances into flowing wa- ters by violating the provisions of an existing NPDES permit or by deliber- ately failing to obtain such a permit. The liability provisions of the CWA enable the Administrator to seek both civil and criminal penalties against alleged violators, as well as injunctive relief to restrain the violation. 33 U.S.C. Section 1319 provides: (b) Civil actions. The Administrator is authorized to commence a civil action for appropriate relief, including a permanent or tempo- rary injunction... (c) Criminal penalties. Any person who wilfully or negligently vio- lates Sections 301, 302, 306, 307, or 308 of this Act [33 U.S.C. Sec- tions 1311, 1312, 1316, 1317, or 1318], or any permit condition ...Shall be punished by a fine of not less than $2,500 nor more than $25,000 per day of violation, or by imprisonment for not more than one year, or by both. (d) Civil penalties. Any person who violates Sections 301, 302, 306, Mie Sole O40 Uso. G-. sections 1311, 1312, 1316, 1317, 1318, 1328, or 1345], or any permit condition...and any person who violates any order issued by the Administrator under subsection (a) of this Section, shall be subject to a civil penalty not to exceed $10,000 per day of such violation. (emphasis added) 188 FLORIDA SCIENTIST [Vol. 52 It is noteworthy that the civil penalty provisions of Section 1319(d) do not require a wilful or negligent violation of the CWA, as does the criminal pen- alty provision of Section 1319(c). Finally, although the federal government currently utilizes the provisions of CERCLA and RCRA, discussed below, as the primary means by which discharges of hazardous substances are remediated, owners and operators of hazardous waste facilities must continue to be mindful of the requirements of the CWA. Section 505 of the Act, 33 U.S.C. Section 1365, grants to citizens the right to sue to enforce provisions of the CWA. Although such citizen suits can only be to obtain injunctive relief, and not to obtain private damages, the CWA further preserves the right of any person to seek common law relief. 33 U.S.C, Section 1365(e). Thus, a citizen suing to obtain injunctive relief under the CWA is entitled to seek damages to private property based on nuisance, negligence, trespass, or strict liability theories, discussed below. CERCLA CERCLA is a liability statute designed to facilitate cleanup of facilities where “hazardous substances” were released into the environment, causing the incurrence of necessary response costs which are “consistent with the National Contingency Plan” (“NCP”). 42 U.S.C. Section 9607(a). CERCLA, 42 U.S.C. Section 9607(a), provides: (a) Notwithstanding any other provision or rule of law, and subject only to the defenses set forth in subsection (b) of this section: 2. any person who at the time of disposal of any hazardous sub- stance owned or operated any facility at which said hazardous substances were disposed of; 3. any person who by contract, agreement or otherwise arranged for disposal or treatment, or arranged with a transporter for transport for disposal or treatment, of hazardous substances owned or possessed by such person, by any other party or entity, at any facility or incineration vessel owned or operated by an- other party or entity and containing such hazardous substan- ces... from which there is a release, or a threatened release which causes the incurrence of response costs, of a hazardous substance, shall be liable for: (B) any other necessary costs of response incurred by any other person consistent with the national contingency plan... (emphasis supplied). “Hazardous substances” under CERCLA are defined to include many hazardous chemicals. 42 U.S.C. Section 9601(14). “Release” and “response” are defined at 42 U.S.C. Sections 9601(22), (25). The “environment” includes groundwater. 42 U.S.C. Section 9601(8). No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 189 The State of Florida and the U.S. Environmental Protection Agency (“EPA”) are authorized under CERCLA to bring an action in federal court to recover response costs incurred as a result of the release or threatened release of a hazardous substance. 42 U.S.C. Section 9607. CERCLA has also been held to provide a private right of action in appropriate circumstances.” CER- CLA also provides the right of a private party to recover response costs from the Hazardous Substances Response Trust Fund (“Superfund”), created un- der CERCLA. 42 U.S.C. Section 9611, 9631. The Florida Department of Environmental Regulation (“FDER’’) also has the right to bring an action to recover response costs under the Florida equivalent to CERCLA, Section 403.727(4), Florida Statutes (1987). It is well settled that parties seeking to recover response costs under CER- CLA must bear the burden to affirmatively demonstrate that they have in- curred necessary costs of response and that they are “consistent with the NCP’ CERCLA provides for the establishment of the NCP. 42 U.S.C. Sec- tion 9605. The NCP, found at 40 C.F.R. Section 300 et seq., effectuates the purposes of CERCLA, and is designed to provide for efficient, coordinated, and effective response to releases of hazardous substances. 40 C.F.R. Section 300.1, 300.3(b). Methods and criteria for determining the appropriate extent of a response authorized by CERCLA when there is a release or a threat of release of a hazardous substance into the environment are found at 40 C.F.R. Section 300.61, et seq. (Subpart F). In the appropriate case, CERCLA can be an effective remedy for address- ing groundwater contamination. However, it is generally reserved for major pollution incidents which meet the requirements under the statute. The EPA has stepped up its CERCLA enforcement actions, and intends to more fully utilize the remedies provided by CERCLA to address environmental contam- ination by hazardous substances. RCRA RCRA is a regulatory statute providing for a “cradle to grave” approach to the management of “hazardous waste.’ Hazardous waste is defined in RCRA as follows: (5) The term “hazardous waste” means a solid waste, or combination of solid wastes, which because of its quantity, concentration, or phys- ical, chemical, or infectious characteristics may: (A) cause, or significantly contribute to an increase in mortality or increase in serious irreversible, or incapacitating reversible, ill- ness; or (B) pose a substantial present or potential hazard to human health or the environment when improperly treated, stored, transported, or disposed of, or otherwise managed. 190 FLORIDA SCIENTIST [Vol. 52 RCRA, 42 U.S.C. Section 6903(5). RCRA is administered in Florida by the FDER, under delegation by the EPA, 52 Fed. Reg. 45634 (December 1, 1987) (except for the Hazardous and Solid Waste Amendments of 1984, Pub. L. No. 98-616, 98 Stat. 3221 (1984)). EPA developed regulatory standards for the treatment, storage and disposal of hazardous waste, found at 40 C.F.R. Part 260, et seq. FDER adopted virtually all of these standards in Chapter 17-30, Florida Administrative Code. The regulations promulgated under RCRA provide that a solid waste is a hazardous waste, unless otherwise excluded, if it exhibits any of the “char- acteristics” of hazardous waste identified in Subpart C of Part 261, or if it is “listed” in Subpart D of Part 261, and has not been excluded from the list in Subpart D of Part 261. 40 C.E.R. Section 261.3(a). If RCRA or FDER regulations are violated, the EPA and FDER are au- thorized to bring an administrative or civil action to force the violator to conduct a hazardous waste determination and remediate the contamination, and to impose civil penalties of up to $25,000 per day for noncompliance. 42 U.S.C. Section 6928; Sections 403.708, 403,141, Florida Statutes (1987). RCRA does not create private rights of action. Thus, its scope is somewhat limited to treatment, storage and disposal of hazardous wastes as defined in the statute. Like CERCLA, in the appropriate case RCRA can be an impor- tant remedy for cleaning up groundwater contamination. FLoriwa StaTuTORY LAw—The Florida Legislature has created a number of statutory schemes to address the problem of groundwater contamination. In addition to the SDWA, Florida is authorized to remedy groundwater con- tamination in the Florida Safe Drinking Water Act, Section 403.850-403.864, Florida Statutes (1987) (“FSDWA”); the Florida Air and Water Pollution Control Act, Section 403.011 et seq. (“FAWPCA”); and the Pollution Spill Prevention and Control Act, Section 376.011 et seq. (“PSPCA”). The follow- ing is a brief analysis of the key provisions of each act. SDWA As discussed above, the FDER has “primary” enforcement authority un- der the SDWA for public water systems in Florida. However, if EPA deter- mines that FDER has failed to assure timely and appropriate enforcement of drinking water regulations, EPA may bring a civil action in federal district court to require compliance with such regulations. 42 U.S.C. Section 300g-3. The current trend is for joint EPA-FDER enforcement action against viola- tion of the SDWA. FSDWA The FDER, in cooperation with the Florida Department of Health and Rehabilitative Services (“HRS”), regulates Florida “public water systems” under the FSDWA. Under the FSDWA, a “public water system” is defined as a “community” or “non-community” system for providing piped water to the No. 3, 1989] GUILDAY AND DEMEO—LEGAL ISSUES 191 public for human consumption for systems that have at least 15 service con- nections or regularly service at least 25 individuals daily at least 60 days out of the year, and includes any collection, treatment, storage and distribution facility or facilities under control of the operator of such system and used primarily in connection with such system, and any collection or pretreatment storage facility or facilities not under control of the operator of such system but used primarily in connection with such system. Sections 403.852(2) (a), (b), Florida Statutes (1987). The FSDWA further defines a “community water system” as a public water system which serves at least 15 service connections used by year-round residents or regularly services at least 25 year-round residents. Section 403.852(3), Florida Statutes (1987). A “non-community water system” is a public water system for provision to the public of piped water for human consumption which serves at least 25 individuals daily at least 60 days out of the year which is not a community water system. Section 403.852(4), Florida Statutes (1987). A “special non-community system” means a non-community system that serves a school, or a seasonal recreational vehicle park or mobile home park that serves 50 or more persons at least four consecutive months during the year. Chapter 17-22.103(30), Florida Administrative Code. Drinking Water Standards—a) Primary Drinking Water Regulations Pursuant to the SDWA and the FSDWA, the FDER has established “Pri- mary Drinking Water Regulations” which apply to all public water systems, except one which consists of distribution and storage facilities only, without collection or transmission facilities; which obtains all water from but is not owned or operated by, a public water system to which these regulations ap- ply; which does not sell water to any person; and which is not a carrier which conveys passengers in interstate commerce. Section 403.853, Florida Statutes (1987). The “Primary Drinking Water Regulations” are rules that specify contam- inants which in the judgment of the FDER after consultation with the HRS may have an adverse effect on the health of the public; specify for each contaminant either a Maximum Contamination Level (“MCL’) if it is eco- nomically and technologically feasible to ascertain a level of such contami- nant in public water systems, or specify each treatment technique known to the FDER which leads to a reduction in the level of the contaminants suffic- ient to satisfy drinking water standards if it is not economically or technologi- cally feasible to ascertain levels of such contamination; and contain criteria and procedures to assure a supply of drinking water which dependably com- plies with such maximum contaminant levels, including quality control and testing procedures to assure compliance at such levels and to ensure proper operation and maintenance of the system and which contains requirements as to minimum quality of water which may be taken into the system and siting for new facilities for public water systems. Section 403.852(12), Florida Stat- utes (1987). 192 FLORIDA SCIENTIST [Vol. 52 Primary Drinking Water MCLs or treatment techniques for “public water systems” are contained in the following regulations in the Florida Adminis- trative Code: 1. Inorganics (Rules 17-22.210(1) and 17-22.310(1)); . Organics (Rules 17-22.210(2) and 17-22.310(2)); . Turbidity (Rules 17-22.210(3) and 17-22.310(4)); . Microbiological (Rules 17-22.210(4) and 17-22.310(5)); . Radionuclides (Rules 17-22.210(5) and 17-22.310(6)); . Volatile Organics (applicable only to community systems and special non-community systems) (Rules 17-22.210(6) and 17-22.310(7)); 7. Trihalomethanes (Rules 17-22.210(2) (e) and 17-22.310(3)). These analyses must be performed every three years on finished water except when results or conditions warrant more frequent monitoring as determined by the FDER. Special non-community systems are also required to comply with standards requiring sampling. MH Ue W NS Drinking Water Standards—Secondary Drinking Water Regulations The FDER also has established secondary drinking water regulations which apply to all community systems. The secondary standards are found at Rule 17-22.220, Florida Administrative Code, and relate primarily to aes- thetic aspects of water, rather than health-based concerns. FAWPCA, PSPCA Liability— Florida law provides express statutory mechanisms to remedy groundwater contamination. Chapter 403, Florida Statutes (1987), provides: (1) It shall be a violation of this Chapter, and it shall be prohibited: (a) To cause pollution, except as otherwise provided in this chap- ter, so as to harm or injure human health or welfare, animal, plant, or aquatic life or property. Section 403.161 (1) (a), Florida Statutes (1987). Futhermore, Chapter 376, Florida Statutes (1987), provides: “The dis- charge of pollutants into or upon any waters of the state or lands, which discharge violates any departmental ‘standard’ as defined in Section 403.803(13), is prohibited.” Section 376.302, Florida Statutes (1987). “Waters” include “underground waters.” Section 403.031(12), Florida Statutes (1987). Groundwater has been held by the courts to be property of the State of Florida.” The FDER is authorized to bring a civil action against any person which “causes pollution” or any facility which “discharges a pol- lutant” within the meaning of Chapters 403 and 376 for cleanup of the pollu- tion. Section 403.121, Florida Statutes (1987). Classification of Groundwaters—Chapter 17-3, Florida Administrative Code, classifies groundwater of the state. This Chapter sets forth four catego- ries of groundwater, based on dissolved solids within the groundwater and whether the groundwater in question is potable. Rule 17-3.403(1), Florida Administrative Code. Groundwater is Class G-I (potable) if it is groundwater No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 193 in single source aquifers with a total dissolved solids content of less than 3,000 milligrams per liter. Class G-II groundwater (potable) is water in an aquifer with a total dissolved solids content of less than 10,000 milligrams per liter. Classes G-III and G-IV are non-potable waters. The Rule provides that FDER policy shall afford the highest protection to G-I groundwater, with FDER providing less protection in order of classification (G-I receiving the highest, G-IV the lowest). Rules 17-3.403(2) and 17-3.403(4), Florida Ad- ministrative Code. Procedures are provided for affected parties to petition for the reclassification of groundwaters. Rules 17-3.403(5) - 17-3.403(6), Florida Administrative Code. Remedies—As mentioned, the FDER has the authority to institute a civil action in a court of competent jurisdiction to recover damages and to assess a civil penalty for “causing pollution” under Chapter 403 or “discharging pol- lutants” from a facility under Chapter 376. Section 403.121(1), Florida Stat- utes (1987). “Damages” include reasonable costs and expenses in tracing the source of the discharge and in controlling and abating the source and the pollutants, costs of restoration of portions of the environment which were damaged, and a judicially imposed civil penalty not to exceed $10,000 dollars per offense (each day during any portion of which such violation occurs con- stitutes a separate offense). Section 403.121, Florida Statutes (1987). Florida law also authorizes the FDER to bring a similar action for injunc- tive relief. Section 403.131, Florida Statutes (1987). Further, the FDER may issue a Notice of Violation to the alleged violator which includes Orders for Corrective Action. Section 403.860, Florida Statutes (1987). The alleged vio- lator may enter into a Consent Order or petition for an administrative hear- ing under Section 120.57, Florida Statutes (1987). Id. FDER is also autho- rized to bring criminal actions for willful violations of Chapter 403. Sections 403.161(3), (4), Florida Statutes (1987). Municipalities, other local government bodies, and citizens of Florida have the authority to enjoin the FDER to enforce the State’s environmental laws and regulations. Section 403.412, Florida Statutes (1987). Common Law—Notwithstanding the availability of statutory remedies for groundwater contamination, the State of Florida and FDER have re- cently asserted common law damages theories to remedy groundwater con- tamination. Four common law causes of action have emerged in environmen- tal litigation: negligence, strict product liability, nuisance, and trespass. In addition, two theories, enterprise liability and market share liability, which emerged in the personal injury context, also may affect litigation with regard to groundwater contamination. It should be noted that common law reme- dies for environmental damage caused by hazardous substances are not well developed in Florida or other states. Furthermore, it is possible a State or Federal court would hold that federal law preempts state common law actions.” To date, however, no other reported decision has been found in which state common law remedies were held to be preempted under CER- 194 FLORIDA SCIENTIST [Vol. 52 CLA. To the contrary, the cases seem to at least implicitly recognize the via- bility of common law actions.*! Negligence. There are five basic elements of common law negligence: (1) conduct, act or failure to act; (2) a duty or an obligation recognized by the law; (3) a breach of that duty; (4) causation - a reasonably close causal connection between the conduct or omission and the resulting injury; and (5) factual damage. In Florida, as in many states, negligence has been defined as the failure to observe such care and diligence as the circumstances justly demand, or as the omission by a responsible person to use that degree of care, diligence and skill that it was his legal duty to assume to protect another person from injury.” Because effects of toxic substances may not manifest themselves for an extended period of time, the plaintiff typically must plead and prove actual loss or injury, as “the threat of future harm not yet realized is not enough.”* A majority of jurisdictions still adhere to this rule.* In granting the defendant’s motion to dismiss, the court stated “plaintiff has not pleaded injury to any- body, only risk of injury.” Slip op. at 8 (emphasis in original). Other state courts have held that actual injury is essential to recovery in tort.® The law of negligence developed in the personal injury context. Actions for property damage historically arose in the context of trespass and/or nui- sance. The personal injury negligence claims generally allege: 1) failure to warn; 2) defective product design; and 3) failure to adequately test products which have been marketed. However, recent litigation in Florida and other states demonstrates the novel application of common law negligence in the context of environmental damage. In 1986, the State of Florida and the FDER brought an unprece- dented suit against a pesticide manufacturer alleging, inter alia, that it knew or should have known that the injection of its pesticide products into sandy, permeable soils in Florida would result in groundwater contamination; that it failed to conduct environmental fate tests for use of the pesticide in Florida; and that it failed to provide any warnings on its labels regarding potential contamination of groundwater by its pesticide. The gravamen of the State’s claim is that the company failed to exercise the requisite degree of care in manufacturing and selling its pesticides, exposing it to liability in tort to the State for damages to its property, the groundwater. Furthermore, a Florida municipality has brought suit against the manufacturer upon the same ground for damages to its public water supply. Also, private citizens have brought similar actions against the manufacturers arising out of alleged per- sonal injuries and property damage caused by the pesticide contamination of groundwater underlying their properties. The manufacture, distribution and labeling of toxic chemicals is a highly regulated and monitored commercial activity. Pesticides, for example, are regulated under the Federal Insecticide, Fungicide, and Rodenticide Act, 7 No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 195 U.S.C. Section 136 et seq., and the Florida Pesticide Law, Chapter 487, Florida Statutes (1987). In this context, the law of negligence seems inappro- priate as a cause of action to remedy groundwater contamination. Although the courts often find that violation of a statute or regulation is admissible as evidence of negligence, and that the standards of care can be established by other means than regulatory standards, compliance with the standard of care imposed by statute or regulation is at times all the law requires.” Since the production and distribution of most toxic substances is so highly regulated, compliance with legislative regulatory standards may defeat liability, unless it is pleaded and proved that the manufacturer should have taken additional precautions beyond the ones imposed by the regulation or statute. Furthermore, it is questionable whether a manufacturer can owe a legal duty of care in the manufacture and distribution of chemicals to a state or municipality, as such a duty generally runs only to users or consumers.” Also, as mentioned above, where, as in the Florida suit, the manufacturer com- piled with all federal and state requirements for registration and labeling of the product, it is questionable whether a breach of the legal standard of care has occurred. This is particularly true in such areas as pesticides and other regulated toxics, where the federal and state agencies are charged with the responsibility of determining whether such products posed an unreasonable risk of harm to man or the environment prior to their sale and use in the State. Strict Product Liability Traditionally, the imposition of strict liability for ultrahazardous activities comes from the “non-natural uses of land” theory of the landmark English decision.* The rule of Rylands provides for the imposition of liability for damages proximately caused by a defendant’s dangerous, non-natural use of land regardless of the standard of care the defendant may have used in con- ducting a particular activity. There are two types of strict liability which may be asserted in an envi- ronmental contamination case. Section 402(A), The Restatement of Torts 2d, establishes strict liability for the sale of a product in a “defective condition unreasonably dangerous to the user or consumer.” Within this category are actions based on failure to warn, breach of implied warranty, innocent mis- representation [Section 402(B) Restatement of Torts 2d], defective design, and defective manufacture. The Florida Supreme Court has expressly adopted this type of strict liability.” The second type of strict liability concerns abnormally dangerous activity. Sections 519, 520, Restatement of Torts 2d. Courts have recently held that strict liability should be applied to claims based upon radiation damage." Furthermore, at least one court has established that “the creation, location, operation, and closure of the toxic chemical dumpsite by defendant was and is an inherently and abnormally dangerous activity.” In Cities Service Company v. State, 312 So. 2d 799 (Fla. 2d DCA 1975), a Florida court found the owner of a phosphate operation strictly liable when 196 FLORIDA SCIENTIST [Vol. 52 one of its reservoirs broke and discharged phosphate slime into the Peace River. In reaching this conclusion, the court weighed the following six factors listed in Restatement (Second) Torts Section 520 (1977): (a) Whether the activity involves a high degree of risk or some harm to the person, land or chattels of others; (b) Whether the harm which may result from it is likely to be great; (c) Whether the risk cannot be eliminated by the exercise of reasonable care; (d) Whether the activity is not a matter of common usage; (e) Whether the activity is inappropriate to the place where it is carried on; and (f) The value of the activity to the community. In other jurisdictions, there is a split of authority on the question of whether the generation and disposal of hazardous waste is an abnormally dangerous or ultrahazardous activity.” Also, in a case in which a plaintiff was attempting to recover CERCLA “response costs” under strict liability, the court, in dictum, found that such costs “are not the type of harm for which strict liability generally attaches.’ However, it is generally recognized that the standard of liability under CERCLA is strict liability.” It should be noted that even if strict liability evolves as the standard in groundwater contamination cases, a threshold requirement for maintaining an action on this ground (as well as other common law grounds) is that the plaintiff identify the manufacturer or other product seller responsible for placing the product into the stream of commerce. Stated another way, the plaintiff must establish the defendant’s instrumentality in bringing about the injury.” Claims based upon strict product liability were advanced by the State of Florida municipality in recent litigation against a private pesticide manufac- turer. The deficiency in those claims is that in West, supra, the Florida Su- preme Court limited standing to maintain strict liability actions to “foresee- able bystanders.’ Also, the deficiency in these claims, as in negligence, in effect constitutes estoppel. Throughout the period during which the pesticide was sold and distributed in Florida, the manufacturer complied with all state regulatory/statutory requirements. In fact, the State itself approved and used the product. It can be argued that the State, by accepting the manufacturer’s adherence to regulation, made a determination that the products were nei- ther defective nor abnormally dangerous. As Section 402A (Restatement of Torts 2d) and Florida law clearly state, this is an essential element for recov- ering on a strict liability claim.” This situation differs from the case of a typical injured third party claimant, who has no role in establishing the standards by which the product is sold, or labeled, or used. Thus, strict prod- uct liability, like negligence, appears ill-suited to provide a remedy for groundwater contamination. No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 197 Nuisance A cause of action for private nuisance arises when a person commits an act which either annoys, injures or endangers the comfort, health or safety of another, or which unlawfully interferes with or tends to obstruct, or in any way render unsafe, other persons in life or in the use of their property.” To prove nuisance in a private action the plaintiff must show that the interfer- ence with his interest is either intentional and unreasonable, reckless or the result of an abnormally dangerous activity. Although a nuisance generally implies a continuity of action over a substantial period of time, a single act may establish a nuisance if the degree of harm is severe.” Where a nuisance has been abated, the injured party’s recovery is mea- sured by the damages resulting from the loss of his property during the period of the nuisance.” In addition, the injured party is also entitled to his reason- able expenses in preventing, reducing or abating the results of the defendant’s wrongful acts.” In cases involving an invasion of one’s land by a noxious substance from the land of another, in the context of an ultrahazardous activity, nuisance and strict liability are closely related and recovery may be had under either the- ory. Proof of negligence is not required.® In essence, both nuisance and strict liability can be utilized to circumvent the need to prove the lack of due care, if the interference is caused by an abnormally dangerous activity.™ Trespass Trespass is a physical invasion and interference with an individual’s exclu- sive right to possession of his property.” Trespass has frequently been alleged in environmental contamination cases by plaintiffs seeking property and/or personal damages from the owners and operations of hazardous waste dump- ing and treatment facilities. Trespass is particularly useful in property dam- age cases where toxins have migrated off-site and onto an adjoining property owner’s land. In cases where there is seepage or leakage of oil or gas onto plaintiff's land caused by conditions created by a defendant on his land, courts have used both a trespass and nuisance approach.” Commentators have suggested that where the leakage is onto the surface of the land, the action lies in trespass; and where the leakage is a subsurface invasion, the action lies in nuisance.* In North Dade Water Co. v. Adken Land Co., 130 So. 2d 894 (Fla. 3d DCA 1961), defendant’s sewage disposal plant discharged effluent into its own lake and then through a conduit to lakes on plaintiff's property. The court en- joined defendant’s conduct as both a nuisance and continuing trespass. Generally, the proper measure of damages in an action for trespass to land is the difference in value of the property before and after the trespass.® In Clark v. J.W. Conner & Sons, Inc., 441 So. 2d 674 (Fla. 2d DCA 1983), however, the court recognized that where reduction in market value is an inadequate measure, recovery may be allowed for losses personal to the plain- tiff. For example, where a trespass is created by a person’s refusal to remove his property from the land of another, the costs of removal may be recov- ered.” 198 FLORIDA SCIENTIST [Vol. 52 Difficulty may arise in a trespass action due to the running of the statute of limitations. In order to obviate any difficulty with the statue of limita- tions, an allegation of continuing trespass can be made. Under the theory of continuing trespass, the invasion is continued by a failure to abate the offen- sive activity. An actionable wrong continues as long as the offensive activity continues.*' Some courts, rather than recognize a continuing trespass, will label the migration of toxic chemicals from a hazardous waste site as a contin- uing nuisance.” The trespass/nuisance distinction is founded upon the pri- vate/public nature of the property which is damaged and the collateral ef- fects that a particular course of conduct may have upon the public welfare. Emerging Theories—Enterprise Liability—The theory of enterprise lia- bility has emerged in the context of personal injury liability; however, its applicability in the environmental damage context is apparent. Plaintiffs may be able to state a cause of action under a theory of enterprise liability when a particular defendant-manufacturer cannot be identified and where “the existence of industry wide standards or practices could support a finding of joint control of risk.”® The theory was considered but not applied in Mor- ton v. Abbott Laboratories, 538 F.Supp. 593 (M.D. Fla. 1982). In Hall, plaintiffs were injured by defective blasting caps. Since it was obviously im- possible to identify the particular blasting cap (and hence its manufacturer) plaintiffs joined the entire American blasting cap industry. In allowing join- der of an entire industry, the court noted that very few companies made up the entire industry, that the companies were jointly aware of the risk that their products posed, and that they possessed the joint capacity to affect those risks. The court shifted the burden of proof to each manufacturer to show that its cap had not caused the injury. Because no defendants could meet the burden, all defendants were held jointly and severally liable. In the context of environmental contamination such as groundwater pollu- tion, it may be difficult if not impossible to identify a particular manufacturer’s products as having actually caused the damage. Further, typically a small number of manufacturers have manufactured a particular product. Thus, en- terprise theory may be appropriate in this context. However, the applicability of enterprise theory in environmental claims in Florida is uncertain. Market Share Liability—Liability may attach under the market share theory where a small number of defendants manufacture a significant amount of a particular product. The plaintiff must join all defendants repre- senting a substantial share of the particular market. Liability will be prorated to the defendants’ market share.“ In Sindell, the California Supreme Court proposed, but did not accept, the proration of liability related to DES problems based on the market share of each potentially liable DES manufacturer, at the time that the plaintiff consumed the drug. The Court shifted the burden of proof to the defendants based on several factors: 1) plaintiff had joined 95% of the DES manufactur- ers; 2) all DES manufacturers would be equally culpable, if DES was proved No. 3, 1989] GUILDAY AND DEMEO—LEGAL ISSUES 199 to be inherently dangerous or defective; and 3) since the manifestation of injury could be extremely slow and the defendants knew more about the drug and its testing, the burden of proof was appropriately shifted. Sindell left unanswered several questions such as defining a substantial market share, the appropriate burden of proof, and the impositions of joint and several liability. The use of market share theory has been rejected in the asbestos injury context.® Florida has also rejected market share theory in the asbestos cases.” The application of market share theory to environmental contamination claims in Florida is unclear. Damages At common law, damages were compensatory in nature. In the damage to property context, it would logically follow that the appropriate measure of damages for a common law claim is the diminution in the value of the dam- aged property.” However, where the alleged damage is contamination of the state’s groundwater, traditional damages theories do not fit. For example, in the Florida case brought by the municipality, the municipality sought both cleanup of its drinking water supply and diminution in the value of its prop- erty. There are a number of problems with these claims for damages. To begin with, the State of Florida owns the groundwater, and any suit for cleanup of such groundwater is more properly brought by the FDER. Furthermore, these claims are duplicative: if the groundwater is cleaned up, no diminution in property value has occurred. The courts have applied an ad hoc cost of cleanup damages measure out- side the scope of traditional common law damages in environmental contam- ination cases. Damage to the environment is often considered “property dam- age.’ The measure of such damage is the cost of cleanup.® As discussed above, cleanup costs are elements of damages as generally provided for by environmental statutes or regulations. Perhaps the confusion created by the issue of damages as outlined above is the key reason why common law causes of action appear to be incongruent with groundwater contamination litigation. These and similar difficulties will be faced by courts which permit plaintiffs, particularly governments, to bring common law causes of action against defendant manufacturers of toxic substances for groundwater contamination. Defenses In addition to the problems with stating a claim under the common law causes of action summarized above, a number of other defenses are applica- ble to damages claims for environmental contamination brought by state and local governments and other parties. Standing—State Agencies—It is well-settled that state administrative agencies, such as pollution control agencies, are creations of statute and can operate only in the areas and manner provided for by the enabling statute.” Administrative agencies have only those powers which are legally conferred upon them by statute.’ Furthermore, where the state legislature provides 200 FLORIDA SCIENTIST [Vol. 52 that an agency’s authority should be exercised in a certain way, this precludes the exercise of that authority in any other way.” The scope of an agency’s statutory authority is to be strictly construed. If there is a reasonable doubt as to the lawful exercise of agency powers, the doubt should be resolved against the agency’s exercise of power in the manner challenged. Cape Coral, supra. Inasmuch as environmental statutes gener- ally confer upon the agencies only certain enumerated duties and powers, to the exclusion of common law powers, it may be argued that they are unau- thorized to bring a common law cause of action. State Standing—It may be argued that states have retained no common law powers to remedy environmental contamination inasmuch as they have delegated the task of environmental cleanup to state agencies through explicit and narrow statutory provisions. In Florida, for example, the legislature has delegated the authority to instigate civil actions for damage to the environ- ment to an environmental agency, as discussed above. Criminal prosecutions for the willful or negligent causing of pollution have also been delegated to a state agency. Since legislative intent was to delegate authority to bring civil actions and criminal actions to these agencies, the state arguably has dele- gated all of its authority to litigate environmental claims for damages, and retains no authority to assert any common law claims against manufacturers. Preemption—Preemption has emerged as a viable defense to tort claims involving regulated hazardous substances which cause environmental con- tamination. The preemption doctrine is based upon the principle that federal law may preempt state regulation in a specific area.” Preemption occurs even though federal law has not wholly superseded state regulation in a specific area, but does not to the extent that state law “actually conflicts” with federal law.“ Actual conflict exists where state law “stands as an obstacle to the accomplishment and execution of the full purposes and objectives of Con- gress.” In the absence of contravening evidence, it is generally presumed that Congress did not intend to preempt state law.” An appropriate example of federal preemption in a toxic substance context is that of the Federal Insecticide, Fungicide, Rodenticide Act (“FIFRA”), 7 U.S.C. Section 136(v). Under 7 U.S.C. Section 136(v) (b), Congress deter- mined that exclusive federal regulation of pesticide labeling would best pro- mote the interest of protecting public health and the interest of pesticide manu- facturers in operating under uniform regulations. Permitting the pursuit of common law claims against manufacturers of pesticides for insufficient warn- ings on labels would contravene congressional intent. The U.S. Supreme Court has, on several occasions, recognized the regulatory effect of state law damage claims and their potential for frustrating congressional objectives.” The decision of the Third Circuit highlights the preemption defense.” In Cipollone, plaintiffs brought an action against several cigarette manufactur- ers alleging, in part, that Defendants were subject to liability on the basis of negligence and strict product liability for their failure to warn of the hazards of cigarette smoking. Id. 184. Defendants contended that Plaintiffs claims No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 201 were preempted by the congressionally mandated warning provided in the Federal Cigarette Labeling and Advertising Act, 15 U.S.C. Sections 1331, Fas5 (1982), The third Circuit held that although the Act did not expressly prohibit Plaintiffs’ eight common law claims, these claims were subject to preemption insofar as they challenged the adequacy of the warning on Defendants’ ciga- rette packages. Id. at 185 through 187. The Third Circuit found that Plain- tiffs’ state common law claims were in actual conflict with the Act’s prohibi- tion of state labeling requirements and thus were preempted.” At least two courts have expressly held that state common law claims have been preempted by FIFRA.® Other courts have found no preemption of com- mon law claims against manufacturers. A court upheld a private tort action against the pesticide manufacturer’s argument that the claim was preempted by FIFRA." Deciding the case almost two years prior to Cipollone, the Fere- bee court was persuaded that prohibition of state pesticide labeling require- ments under Section 136(v) (b) did not prohibit a private tort action challeng- ing the adequacy of the Defendant’s paraquat label. Commerce Clause— Another viable defense to groundwater contamina- tion claims by government is the Commerce Clause. Article I, Section 8, Clause 3 of the United States Constitution gives Congress exclusive power to regulate interstate commerce. This Clause, even without implementing legis- lation by Congress, is a limitation upon the power of the states.” This clause focuses on undue burdens on interstate commerce and is an affirmative grant of power to Congress and an implied restriction on the power of States.* The test of whether state legislation is in violation of this clause is whether laws are unduly burdensome and unreasonable in scope.™ An “undue or un- reasonable burden on interstate commerce” is defined as one which materi- ally effects interstate commerce where uniformity of regulation is necessary.* In determining whether a rule of law imposed by a court would constitute an unreasonable restraint of commerce, these same tests would be applied as is applicable to legislative enactments.* Inasmuch as toxic substances are highly regulated, intervention by common law claims by state governments and state agencies should be closely scrutinized. Any toxic substance which is regulated by the federal government may be amenable to a defense based upon an undue or unreasonable burden on interstate commerce. Statute of Limitations—Historically, statutes of limitations were given literal application to completely bar any action. In the toxic tort context, many of the injuries are latent and take years to manifest themselves. In this regard, personal injury cases and property damage cases are coextensive. The discovery rule, wherein the statute of limitations is in effect tolled until damage or injury manifests itself, has significantly reduced the effec- tiveness of defenses based on statute of limitations.*’ In addition to the discov- ery rule, the inherent complexities of detection and causation of toxic spills and contaminations further reduce the ability to successfully assert a statute of limitations defense as a bar to a toxic tort case. Furthermore, under the 202 FLORIDA SCIENTIST [Vol. 52 Superfund Amendments and Reauthorization Act of 1986, amending CER- CLA, 42 U.S.C., Section 9601 et seq., all claims for damage resulting from environmental contamination are subject to a federal commencement date, which is the date the plaintiff knew or reasonably should have known that the damages were caused by or contributed to by the hazardous substance. Section 309(a), CERCLA. State of the Art—The state of the art defense is one of the most important defenses in environmental litigation. As technology quickly outstrips itself, the establishment of a state of the art defense is, by its nature, factually based. It is important to distinguish between the state of the art and an industry standard. The state of the art refers to the level of scientific and technological knowledge at a given point in time. Industry standards or customs refers to methods which are generally accepted within an industry with respect to a certain manufac- turing process or product. Section 402 (A) of the Restatement of Torts 2nd indicates that some prod- ucts are inherently dangerous. However, if a certain product is made as safely as it can be made then it can be said to be the state of the art. The state of the art doctrine has developed primarily in conjunction with a failure to warn action where a defendant manufacturer can not be held to a duty to warn if it did not know and should not have had reason to know about the hazardous nature of the product or byproducts.’ While the state of the art defense can be successfully employed in a negligence action, courts differ whether it is applicable in strict liability actions.” The Beshada case is a personal injury case based upon a failure to warn of the harmful side effects of asbestos dust fibers. The court found that the state of the art defense was not applicable to strict liability cases based on a failure to warn.” The state of the art defense was also rejected by the District Court for the Western District of Tennessee. *! In Sterling, the Defendant asserted the state of the art defense against Plaintiffs’ claims for injuries from a chemi- cal dump site contending that Velsicol exceeded the then known state of the art in the selection and operation of a hazardous waste disposal site. It should be noted, however, that the viability of the state of the art defense has not been resolved in the environmental damage context. Miscellaneous Defenses—In addition to the above, other defenses are ap- propriate to resist damages claims by government for environmental contam- ination. For example, the state government may be negligent itself and breached duties arising by: being in a position of equal or superior knowledge to the manufacturers regarding the environmental characteristics and the potential for products to result in contamination, but may have failed to test and review such products prior to approval for use; failing to develop proce- dures to monitor the environment for the presence of contamination; or oth- erwise failing to carry out its statutory responsibilities. Also, the state may be estopped from or may have waived its right of claiming the product to be hazardous by virtue of having approved it for use in the state, in the case of | regulated substances. Further, the defendant may be able to show that regu- No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 203 lations which provide what constitutes “pollutants” or contamination are ar- bitrary and capricious or otherwise do not meaningfully describe the alleged contamination. ConcLusions—It should be apparent from the above that the law is un- dergoing a mutation in the area of traditional common law actions brought by state and local governments and private parties against toxic substance manufacturers for environmental damages. The courts, if they permit these actions against private entities, may have circumvented the intent and will of both the state legislatures and Congress, as well as misapplied principles of law ill-suited for their new roles. The state and federal governments have provided comprehensive statu- tory/regulatory schemes for the redress of environmental contamination. The structure of the common law is ill equipped to deal with the complexities and intricacies of groundwater contamination and subsequent litigation. Applica- tion of common law principles by the courts will result in ineffective, piece- meal and inconsistent legal solutions to a situation which is more appropri- ately and adequately dealt with by specific well-articulated statutory provisions and the regulatory process equipped to handle the same. Given the broad scope of the problem of groundwater contamination which is as yet ill- defined, states should work with the federal government to define more care- fully the nature of the problem and arrive at more meaningful legislative, as opposed to judicial, solutions. FOOTNOTES “The term ‘public water system’ means a system for the provision to the public of piped water for human consumption, if such a system has at least fifteen service connections or regu- larly serves at least twenty-five individuals.” 42 U.S.C. Section 300f(4) (emphasis added). The SDWA applies to all public water systems except those which consist of only distribution and storage facilities without collection and treatment facilities; which obtain all water from but are not owned or operated by a public water system; which do not sell water to any person; and which are not carriers which convey passengers in interstate commerce. 42 U.S.C. Section 300g. In an opinion of a U.S. Environmental Protection Agency (“EPA”) Regional Counsel, it was determined that since the SDWA does not define the term “human consumption,” the definition of that term should be no broader than its common meaning: to eat, drink, or ingest. Thus, a system providing water only for nondrinking purposes does not fall within the scope of the SDWA. USEPA RCO (Region 10) October 25, 1978. See, 42 U.S.C. Section 300h (1974). 3The EPA Administrator “shall by rule establish recommended maximum contaminant levels for each contaminant which, in his judgment based on the report on the study conducted pursu- ant to subsection (e), may have an adverse effect on the health of persons.” 42 U.S.C. Section 300g-1(b) (1) (B) (1974) (emphasis added). Conversely, if the Administrator determined that health risks were minimal, or speculative, no regulations need be issued. 4The authorization for appropriations expired on September 30, 1982. The Congressional Conference Committee that worked out the details of the reauthorization did not approve the final measure until March of 1986, a span of some four years. >The standard setting schedules and deadlines, located at 42 U.S.C. Section 300g-1(b) (1986), mandate that the Administrator “shall publish maximum contaminant level goals and promul- gate national primary drinking water regulations” for approximately 80 listed contaminants over a three year span from the date of enactment of the Amendments. The listed contaminants are published “in volume 47, Federal Register, page 9352, and in volume 48, Federal Register. page 45502.” Id. 204 FLORIDA SCIENTIST [Vol. 52 6See, 42 U.S.C. Section 300¢-3(g) (1986). 7See, 42 U.S.C. Section 300g-1(b) (4-7) (1986) (setting the standards for contaminants based upon the best available technology to remove that contaminant). 8See, 42 U.S.C. Section 300g-1(b) (3) (1986). Following this three year review cycle, the Administrator must list contaminants which require regulation. Within 24 months of listing, proposed regulations must be issued, and within another 12 months final regulations must be promulgated. 942 U.S.C. Section 300g-1(b) (5) (1986). 10The Amendments were enacted June 19, 1986. See, 42 U.S.C. Section 300g-z (1986). 12Florida Administrative Code Annotated R. 17-22.101 (Nov. 1987). 13Id., l4See, supra note 5. 5See, supra notes 8-9. 16In assessing civil penalties, the court must “[take] into account the seriousness of the viola- tion, the population at risk, and other appropriate factors.” 42 U.S.C. Section 300g-3(b) (1986). 17See, 42 U.S.C. Section 300g-3(b) (1974). 1842 U.S.C. Section 300g-3(g) (1986). 1942 U.S.C. Section 300g-3(g) (2) (1986). 2042 U.S.C. Section 300g-3(g) (3) (B) (1986). 2142 U.S.C. Section 300g-3(g) (3) (C) (1986). 2242 U.S.C. Section 300j-8 (1974). 2342 U.S.C. Section 300j-8(a) (1974). This provision also allows suit to be brought against the Administrator for failure to perform a non-discretionary duty. Id. 2442 U.S.C. Section 300j-8(b) (1974). 25“Nothing in this section shall restrict any right which any person (or class of persons) may have under any statute or common law to seek enforcement of any requirement prescribed by or under this title or to seek any other relief.” 42 U.S.C. Section 300j-8(e) (1974) (emphasis added). 26See, United States v. Ottati & Goss, Inc., 630 F.Supp. 1361, 1383 (D. N.H. 1985). 27See, Wickland Oil Terminals v. Asarco, Inc., 792 F.2d 887 (9th Cir. 1986). Bulk Distribution Centers v. Monsanto Co., 589 F.Supp. 1437 (S.D. Fla. 1984); Artesian Water Co. v. Government of New Castle County, 659 F. Supp. 1269 (D. Del. 1987); State of New York v. Shore Realty Corp., 648 F. Supp. 255 (E.D. N.Y. 1986). 29See, Village of Tequesta v. Jupiter Inlet Corporation, 371 So2d 663 (Fla. 1979), cert. denied, 441 U.S, 965 (1979). 30See, City of Philadelphia v. Stepan Chemical Co., 544 F. Supp. 1135 (E.D. Pa. 1982); United States v. Price, 523 F. Supp. 1055 (D. N.J. 1981), aff'd., 688 F.2d 204 (3d Cir. 1982) (RCRA and CERCLA preempt federal common law of nuisance in the area of hazardous waste disposal). 31See, e.g., City of Philadelphia v. Stepan Chemical Company, 544 F. Supp. 1135, 1152-53 (E.D. Penn. 1982). 32DeWald v. Quarnstrom, 60 So.2d 919 (Fla. 1952); Smith v. Hinkley, 123 So. 564 (Fla. 1929); Lake Parker Mall v. Carson, 327 So.2d 121 (Fla. 2d DCA 1976), cert. denied, 344 So.2d 325 (Fla. 1977): 33Prosser and Keeton on the Law of Torts Section 30 at 165 (W. Keeton 5th Ed. 1984). 34See, Rheingold v. E.R. Squibb and Sons, Inc., No. 74-3420 (S.D. N.J. Oct. 8, 1975) (wherein plaintiff sought to establish a class action claiming defendants should create a fund to compensate girls whose mothers had ingested DES during pregnancy.) No. 3, 1989] GUILDAY AND DEMEO— LEGAL ISSUES 205 35[d. See also, Mink v. University of Chicago, 460 F. Supp. 713 (N.D. Ill. 1978); Morrissy v. Eli Lilly & Co., 394 N.E. 2d 1369, 76 Ill. App. 3d 752 (1979). (The mere ‘risk’ of developing a future injury or disease is an insufficient basis for recovery in tort). 36See, Prosser and Keeton on Torts, at 233. 37See, Johnson v. Murph Metals, 562 F.Supp. 246 (N.D. Tex. 1983). 38Rylands v. Fletcher, 3 HL 330 (1868). 39West v. Caterpillar Tractor Co., 336 So.2d 80 (Fla. 1976). 40See, Bennett v. Mallincrockett, Inc., 698 S.W. 2d 854, 867-869 (Mo. Ct. App. 1985), cert. denied, 106 S. Ct. 2903 (1986). 41Sterling v. Velsicol Chemical Corp., 647 F.Supp. 303, 315 (W.D. Tenn. 1986). 42See, generally, Note, Strict Liability for Generators, Transporters, and Disposers of Hazard- ous Wastes, 64 Minn. L. Rev. 949, 967-85 (1980). 43Pinole Point Properties, Inc. v. Bethlehem Steel Corp., 596 F. Supp. 283, 292 n.5 (N.D. Cal. 1984). 4442 U.S.C. Sections 9601(32); New York v. Shore Realty Corp., 759 F.2d 1032 (2d Cir. 1985). 45See, e.g., Gray v. U.S., 445 F.Supp. 337 (S.D. Tex. 1978). 46 West, 336 So.2d at 92. 47See, West, 336 So.2d at 80. 48Jones v. Trawick, 75 So.2d 785 (Fla. 1954). 49Restatement (Second) of Torts Section 822 (1977). 5038 Fla. Jur.2d Nuisances Section 16 (1982). 51Ford v. Dania Lumber & Supply Co., 7 So. 2d 594 (Fla. 1942). 52Nitram Chemicals Inc. v. Parker, 200 So.2d 220 (Fla. 2d DCA), cert. denied, 204 2d 330 (Fla. 1967) (citing Prosser, Torts Section 89 (5th Ed. 1964)); Bousquet v. Commonwealth, 372 N.E.2d 257 (1978); McCormick, Damages Section 127 (1985). 3Bunyak v. Clyde J. Yancey & Sons Dairy, Inc., 438 So. 2d 891 (Fla. 2d DCA 1983), pet. for rev. denied, 447 So. 2d 885 (Fla. 1984). >4But see Prosser and Keeton, Torts, Section 89 (5th ed. 1984) (nuisance should require an intentional interference). See, Guin v. Riviera Beach, 338 So.2d 604 (Fla 4th DCA 1980). 56See, Maryland Heights Leasing, Inc. v. Mallinckrodt, Inc., 706 S.W.2d 218 (Mo. Ct. App. 1985). s7F.g., compare, Burr v. Adam Eidemiller, Inc., 126 A.2d 403 (Pa. 1956 (nuisance approach) with Phillips v. Sun Oil Co., 121 N.E. 2d 249 (N.Y. 1954) (trespass approach). 58Prosser and Keeton, Torts Section 13 (5th ed. 1984). °Kirchoff v. Moulder Bros., Inc., 391 So. 2d 347 (Fla. 5th DCA 1980). 6Anchorage Yacht Haven, Inc. v. Robertson, 264 So.2d 57 (Fla. 4th DCA 1972). 61Prosser and Keeton on Torts, Section 13 at 71. 62See, State v. Schenectady Chemicals, Inc., 117 Misc.2d 960, 459 N.Y.S. 2d 971 (N.Y. S.Ct. 1983), modified 103 A.2d 33 (1984). 63See, Hallv. E.I. DuPont de Nemours & Co., 345 F.Supp. 353 (E.D. N.Y. 1972). See, Sindell v. Abbott Labs, 607 P.2d 924, 26 Cal.3d 588, 163 Cal.Rptr. 132, cert. denied, 449 U.S. 912 (1980). 6See Thompson v. Johns-Manville Sales Corp., 714 F.2d 581 (5th Cir. 1983), cert. denied, 104 S.Ct. 159 (1984). ®Celotex Corp. v. Copeland, 471 So.2d 553 (Fla. 1985). 206 FLORIDA SCIENTIST [Vol. 52 67A tlantic Coastline R.R. Co. v. Saffold, 178 So. 288 (Fla. 1938). 68See Lansco, Inc. v. Department of Environmental Protection, 350 A.2d 520 (N.J. Super. 1975). “United States Aviex Company v. Travelers Insurance Co., 336 N.W.2d 838 (C.O.A. Mich. 1983); Chemical Application Co., Inc. v. Home Indemnity Co., 425 F.Supp. 777 (D. Mass. 1977). Department of Health and Rehabilitative Service v. Florida Psychiatric Society, 382 So.2d 1280 (Fla. lst DCA 1980). Cape Coral v. GAC Utilities, Inc., 281 So.2d 493 (Fla. 1973). Kirk v. Publix Super Markets, 185 So.2d 161 (Fla. 1966). 3Silkwood v. Kerr-McGee, 464 U.S. 238 (1984). “Pacific Gas & Electric Company v. Energy Resources Conservation & Development Comm’n., 461 U.S. 190, 204 (1982). Silkwood, 464 U.S. at 248. Maryland v. Louisiana, 451 U.S. 725, 746 (1981). 7E.g., Fidelity Federal Savings and Loan Ass'n v. De la Cuesta, 458 U.S. 141, 156-159 (1982); Chicago & North Western Transport Co. v. Kalo Brick & Tile Co., 450 U.S. 311, 324-25 (1981). 8Cipollone v. Liggett Group, Inc., 789 F.2d 181 (3rd Cir. 1986). Cipollone, 789 F.2d at 187. 80See, Fitzgerald v. Mallinckrodt, Inc., Civ. Action No. 86-2598 (E.D.M. 1987); Stiltjes v. Ridco Exterminating Company, Inc., 343 S.E. 2d 715 (Ga. App. 1988). See also, 2 Toxics Law Reporter 1104 (BNA March 9, 1988). 51Ferebee v. Chevron Chemical Company, 736 F.2d 1529 (DC Cir.), cert. denied, 469 U.S. 1062 (1984). 82Great Atlantic and Pacific Tea Company, Inc. v. Cottrell, 424 U.S. 366, 96 S. Ct. 923 (ISTO): 83. aborers Local Union #374 v. Felton Constr. Co., 654 P.2d 67, 98 Wash. 121 (1982). ‘4Interstate Motor Freight System v. Service Terminals, Inc., 455 N.Y.S. 2d 910, 116 Misc. 2d 517 (1982). Union Pacific Railroad Company v. Woodahl, 308 F. Supp. 1002 (D.C. Mont. 1970). 86 West v. Broderick & Bascom Rope Company, 197 N.W. 2d 202 (Iowa 1972). ‘See e.g. In re Agent Orange Product Liability Litigation, 597 F.Supp. 740 (E.D. N.Y. 1984), summary judgment granted, 603 F.Supp. 239 (E.D. N.Y. 1985). 88See, Basko v. Sterling Drug, Inc., 416 F.2d 417 (2nd Cir. 1969). 89Cf. Beshada v. Johns-Manville Products Corporation, 447 A.2d 539 (1982). %Compare Beshada with Karjala v. Johns-Manville Products Corporation, 523 F.2d 155 (8th Cir. 1975), in which the 8th Circuit Court of Appeals reached the opposite conclusion—that the state of the art defense was inapplicable—based on virtually the same set of facts. Sterling v. Velsicol Chemical Corporation, 647 F.Supp. 303 (W.D. Tenn. 1986). Florida Sci. 52(3):183-206. 1989. Accepted: August 19, 1988. Environmental Chemistry LABORATORY MODELS FOR ASSESSING THE FATE OF GROUNDWATER CONTAMINANTS JosEPH J. DELFINO, Patricia V. CLINE, AND CARL J. MILEs' Department of Environmental Engineering Sciences, A.P. Black Hall, University of Florida, Gainesville, Florida 32611 Asstract: Groundwater in Florida has been contaminated by serveral classes of chemicals. Regulatory interest in these chemicals has focused on impacts such as the loss of potable water supplies as well as cleanup and reclamation strategies. Also of interest, but often overlooked, are chemical transformation processes that occur in the subsurface aquatic environment. Using labo- ratory microcosms as model groundwater systems, we studied: (a) hydrolysis reactions and re- lated processes involving aldicarb, a carbamate pesticide, including possible changes that could occur upon interaction with surface waters; (b) the degradation of industrial and petroleum derived organic chemicals; and (c) the abiotic conversion of 1,1,1,-trichloroethane to 1,1-dichlo- roethene. The application of these laboratory model results to field sites can provide useful inter- pretive information about the fate of these classes of chemicals in groundwater. GROUNDWATER is an especially valuable resource in Florida as it provides approximately 92% of the annual potable water needs of the state’s citizens (Fernald and Patton, 1984). As a consequence of Florida’s geology and hy- drology, coupled with a rapidly growing population and increasing urbaniza- tion, groundwater contamination is becoming a more common occurrence (Delfino, 1986). It is difficult to perform complete studies of chemical and microbiological interactions involving contaminants in situ due to the necessity of multiple sampling sites. This would necessitate construction of an extensive network of monitoring wells, something that is generally infeasible due to construction, monitoring and sample analysis costs. An alternative to performing in situ studies is the use of laboratory model systems (often called microcosms) that are based on conditions existing at contaminated field sites. Generally, there are sufficient monitoring well data available once a contamination incident has been discovered that will allow identification of the nature of the contaminant(s), concentrations, and associ- ated physical and chemical parameters. The results of chemical analyses per- formed on groundwater samples drawn from the monitoring wells can be used to design the matrices for the laboratory experiments. While laboratory experiments cannot be expected to completely replicate conditions in situ, they do offer the opportunity to elucidate reaction mechanisms under con- trolled conditions, something that is difficult to accomplish under field condi- tions. 1Present Address: Pesticide Residue Research Laboratory, Henke Hall, University of Hawaii, Honolulu, Ha- waii 96822. 208 FLORIDA SCIENTIST [Vol. 52 The types of contaminants generally found in Florida’s groundwater rep- resent several chemical classes including pesticides, industrial and petroleum- based compounds, and halogenated solvents (Delfino, 1986). We designed laboratory model systems to study the behavior of chemicals from each of these classes. The compound aldicarb represented carbamate pesticides; ben- zene, naphthalene, p-cresol and methy-ethyl-ketone represented the indus- trial and petroleum derived chemicals; and 1,1,1-trichloroethane was the halocarbon solvent representative. MetuHops— Reactions of aldicarb in various groundwater (natural and sterilized) matrices were conducted in glass containers with Teflon coated crimp-seals under both aerobic and anaer- obic conditions, in the presence and absence of crushed and sieved limestone (Miles and Delfino, 1985). Analyses of aldicarb and its oxidation and hydrolysis products were performed by high pressure liquid chromatography (Miles and Delfino, 1984). Degradation studies of the industrial and petroleum-derived compounds were similarly performed under aerobic and anaerobic condi- tions using both natural and sterilized groundwater in the absence of limestone; analyses of these compounds were performed by gas chromatography with mass spectrometry detection (Delfino and Miles, 1985). The degradation experiments involving 1,1,1-trichloroethane (TCA) were per- formed in heat-sealed glass ampules that were incubated at various temperatures. Analyses of TCA and its principal degradation product, 1,1-dichloroethene (DCE), were performed by gas chromatography with a flame ionization detector (Cline et al., 1986). RESULTS AND Discuss1on—Our laboratory experiments were designed to provide data to interpret observations of chemical behavior under field con- ditions. Aldicarb [2-methyl-2-(methylthio)-propionaldehyde 0-(methyl-carba- moyl)oxime], a carbamate pesticide, is applied to the soil root zone in granu- lar form to minimize air drift and applicator exposure (Fig. 1). While this is an effective mode of application, it can cause environmental problems espe- cially where sandy soils predominate. Florida is a classic example of this situation. There are nine different chemical species of concern that can result from the initial application of the parent aldicarb compound. These include aldi- carb sulfoxide and sulfone (which, together with aldicarb, are collectively known as total toxic residue-TTR) and their corresponding oximes and ni- triles (Moye and Miles, 1988). If aldicarb is not completely degraded in soils, infiltration to groundwater or runoff to surface waters can occur (Fig. 1). There have been extensive aldicarb degradation studies performed in various soils under many conditions, and a variety of soil-specific chemical and mi- crobial processes and reaction rates have been observed. These have been reviewed and summarized by Moye and Miles (1988) and will not be treated here. Our work emphasized the hydrolysis of aldicarb in groundwater micro- cosms under both aerobic and anaerobic conditions, with and without crushed limestone that represented matrix material found in the Floridan aquifer (Miles and Delfino, 1985). Aldicarb had an average first order alka- line hydrolysis half life of 63 days under aerobic groundwater conditions but this rate extended to over 600 days under anaerobic conditions. Degradation No. 3, 1989] DELFINO, ET AL. MODELS FOR CONTAMINANTS 209 ALDICARB [TEMIK ®] 2 - methyl - 2 (methylthio) - propionaldehyde O - (methyl carbamoyl) oxime APPLICATION IN ROOT ZONE (granular, 5 - 20% 4a.i.) SOIL ® RELEASE FROM GRANULE ° INFILTRATION BY MOISTURE © RUNOFF GROUND WATER SURFACE WATER © TOXICITY TO TARGET PESTS * PLANT UPTAKE ° HYDROLYSIS ° DEGRADATION ¢ TOXICITY TO NON- Hydrolysis © PHOTOLYSIS Fic. 1. Aldicarb reaction pathways in the environment. rates for the sulfoxide and sulfone were more rapid but once again, average half-lives under anaerobic conditions were longer than under aerobic condi- tions (Table 1). We determined, as have others (Moye and Miles, 1988), that the typically less toxic oximes were the predominant hydrolysis products of the three TTR compounds that are generally found in soil. We also found that crushed limestone retarded the rate of alkaline hydrolysis but that the lime- stone had minimal sorptive capacity for aldicarb. TABLE 1. Average first order Half-Lives for the alkaline hydrolysis of Aldicarb compounds in Floridan groundwater microcosms. Compounds Half-Life (days) Aerobic Anaerobic Aldicarb 63 635 Aldicarb sulfoxide 11 26 Aldicarb sulfone 6 26 We discovered that aldicarb sulfoxide could be reduced back to the parent aldicarb compound under anaerobic conditions (such as occur in the deeper Floridan aquifer) in the presence of limestone and added organic matter (Miles and Delfino, 1985). This finding was confirmed by Lightfoot and co- workers (1987) who also suggested that laboratory studies could augment but not replace field studies. However, Moye and Miles (1988) observed consider- able variability in degradation rates among the many field sites reported in the literature. They also indicated that laboratory studies were quite useful in understanding the environmental fate of pesticides such as aldicarb. 210 FLORIDA SCIENTIST [Vol. 52 The TTR compounds can reach surface waters via runoff or infiltration (Fig. 1). They have been observed in streams and rivers that drain citrus agricultural areas (Foran et al., 1986). Because of these findings, we investi- gated the response of these TTR compounds toward photolysis degradation processes. Earlier, aldicarb and the sulfoxide were found to be relatively toxic to a zooplankton species (Foran et al., 1985) although the test concentrations were considerably higher than those reported in surface waters. Aldicarb sulfone was the least toxic of the TTR compounds under acute bioassay condi- tions. Aqueous solutions of the aldicarb TTR compounds were exposed to sunlight and aldicarb and the sulfoxide degraded relatively quickly over a two week period while aldicarb sulfone resisted photodegradation during the same period (Fig. 2, adapted from Germuska, 1985). Fortunately, the photol- ysis and acute bioassay experiments both showed that the more toxic aldicarb and sulfoxide compounds were most susceptible to photolysis in surface wa- ters (and thus were least stable) whereas the least acutely toxic compound, aldicarb sulfone, was the most persistent TTR compound in surface waters. The presence of colored humic matter in lake water had only a slight inhibi- tory effect on the rate of aldicarb degradation but did not affect the photoly- sis rates of the other two compounds (Fig. 2). 2.0 ¢ Aldicarb Sulfone ES EE— LO 0 i —S ZN {\ A \ a @xS - re) [S Aldicarb Sulfoxide e fe) Y\ —_ @ — NB = 1.0 (= Y\ o oO 0 /\ Oo Aldicarb 2 O {\ f\ 0 & Lake Water O Distilled 0 Water 0 0 2 4 6 8 10 12 14 Time (Days) Fic. 2. Photolysis of Aldicarb compounds in lake and distilled water under spring sunlight conditions. No. 3, 1989] DELFINO, ET AL. MODELS FOR CONTAMINANTS 211 Besides pesticides, Florida’s groundwaters have been affected by various classes of non-agricultural chemicals. These include phenols and cresols; oxy- genated and halogenated solvents; and aromatic and polynuclear aromatic compounds. Four compounds chosen to represent these classes in biodegrada- tion experiments were, respectively: p-cresol; methyl ethyl ketone; and ben- zene and naphthalene. Degradation studies were performed under aerobic and anaerobic conditions using actual groundwaters that contained indige- nous bacteria. Additional studies involving the abiotic degradation of 1,1,1- trichloroethane are discussed later. Surficial groundwater (pH=5.3) was used for the aerobic degradation studies. All four of the target compounds degraded completely under these aerobic conditions with rates ranging from eight days (p-cresol and naphtha- lene) to 16 days (benzene) (Delfino and Miles, 1985). These findings were generally consistent with the results of related experiments published in the literature. Our data, though, did not always agree with observations made in the field at the locations of spills, tank leaks, etc. This is not too surprising since disagreement between laboratory and field data can be caused by dif- ferent microbial populations, varying pH, redox and nutrient conditions, and excessively high chemical concentrations that might be toxic to the natural microbial populations. However, researchers involved in groundwater reme- diation activities need to know, or be able to predict, degradation rates so that appropriate cleanup strategies can be designed and implemented. Tech- niques for assessing in situ biodegradation rates are still in the experimental stages, thus resulting in a need for laboratory derived degradation rate data which can be employed in computational models that complement ground- water remediation actions. The anaerobic biodegradation studies employed Floridan aquifer water (pH 7 to 8). The literature suggested that biodegradation rates would be considerably slower under anaerobic conditions (Delfino and Miles, 1985) and our results supported such predictions. The biodegradation of p-cresol, having taken eight days under aerobic conditions, took about 45 days under anaerobic conditions. Given the predominance of phenol and cresol domi- nated waste sites in Florida (related to the wood treatment industry), the longer degradation rate was not welcome information. These laboratory results, though, were consistent with the continuing presence of phenols and cresols at a Superfund site in Gainesville, FL (McCreary et al., 1983). Even more troubling was the relative stability of naphthalene and ben- zene under anaerobic conditions. While these compounds degraded within one to two weeks under aerobic conditions, they resisted anaerobic biodegra- dation for a period of at least three months after which the experiments were terminated (Delfino and Miles, 1985). These latter two compounds fre- quently enter surficial aquifers following petroleum fuel leakage or spillage. Movement into subsurface anaerobic zones could considerably increase the longevity of these aromatic compounds and thereby inhibit strategies de- signed to enhance natural biodegradation processes. 22, FLORIDA SCIENTIST [Vol. 52 An extension of the non-agricultural chemical studies led to research in- volving the abiotic degradation of the solvent 1,1,1-trichloroethane (TCA) and its degradation product 1,1-dichloroethene (DCE). A review of data col- lected at several contaminated sites indicated the possibility that abiotic deg- radation of TCA might be occuring, even in preference to biodegradation (Cline et al., 1986). Observations in Florida were supported by reports of laboratory studies that indicated TCA could degrade under abiotic condi- tions (Dilling et al., 1975; Vogel and McCarty, 1987; Haag and Mill, 1988). The chlorinated solvent 1,1,1-trichloroethane (TCA) is a widely used chemical and is frequently found as a groundwater contaminant. The com- pound has been observed to undergo both bio-mediated and abiotic degrada- tion. Anaerobic biodegradation results primarily in the formation of 1,1- dichloroethane (1,1-DCA) and cis-(1,2)-dichloroethene (cis-DCE) with 1,1-DCE being a minor product. Under field conditions that favor abiotic rather than bio-degradation, however, the primary degradation product is 1,1-DCE. This phenomemon, whereby the 1,1-DCE product is formed from TCA by an elimination mechanism (loss of HC1), has been observed at vari- ous contamination sites throughout the U.S. (Cline et al., 1986). Abiotic degradation experiments performed in ampules incubated in the laboratory under various temperatures showed that TCA followed first order kinetics as it degraded to 1,1-DCE. Arrhenius plots were used to extrapolate reaction rates at temperatures that closely approximated ambient conditions. The extrapolated first order half-lives for TCA at different temperatures were: 10.2 months (25 C); 2.0 years (20 C); and 4.5 years (15 C) (Cline and Delfino, 1988). At any temperature, the reaction rates were found to be inde- pendent of pH over the range pH =4.5 to 8.5. Matrix effects appeared to be minimal since reaction rates in groundwater and in buffer solutions were similar at a given temperature, although a seawater matrix did increase the degradation rate by 10-14%, perhaps due to catalysis or ionic strength influ- ences. The stability of the TCA abiotic degradation product, 1,1-DCE, was as- sessed in experiments lasting over one year, at which time no significant deg- radation had been seen (Cline and Delfino, 1988). The conversion of TCA to 1,1-DCE, coupled with the latter compound’s considerable stability, helps to explain the frequent observations of 1,1-DCE in water supplies. The rate of transformation of TCA to 1,1-DCE appears to be influenced predominantly by temperature. Conc.Lusions— Degradation studies of groundwater contaminants per- formed in laboratory microcosms can provide kinetic data that will assist in understanding fate and transport observations made under field conditions. The results of such laboratory studies can be widely applied in descriptive or interpretive models whereas field studies performed at specific sites run the risk of poor transferability due to unique conditions that might be present at a given site. The combined use of both laboratory and field data, recognizing potential limitations, represents a good approach in groundwater contamina- tion studies. No. 3, 1989] DELFINO, ET AL. MODELS FOR CONTAMINANTS Daye ACKNOWLEDGMENTS— Funding for these studies was provided by the Florida Department of Environmental Regulation, the Florida State University System STAR Program and the Univer- sity of Florida Engineering and Industrial Experiment Station. William M. Davis provided ana- lytical assistance, including GC/MS identification and confirmation analyses for the TCA experi- ments. LITERATURE CITED Cun, P. V., J. J. DELFINO, AND W. J. Cooper. 1986. Hydrolysis of 1,1,1-trichloroethane; forma- tion of 1,1-dichloroethene. Pp. 239-248. Proc., NWWA/API Conference on Petroleum Hydrocarbons and Organic Chemicals in Ground Water-Prevention, Detection and Res- toration, NWWA, Dublin, Ohio. , AND J. J. DELFINO. 1988. 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J. DELFINO. 1985. Acute toxicity of aldicarb, aldicarb sulfoxide and aldicarb sulfone to Daphnia laevis. Bull. Environ. Contam. Toxicol. 35:546- 500. , W. L. Mriiuer, S. Doyan, anp M. Krrausu. 1986. Temik contamination in surface water and potential effect on a daphnid species in Florida. Environ. Poll. 40, 369-380. GermuskA, P. J. 1985. Photolysis of aldicarb, aldicarb sulfoxide and aldicarb sulfone in surface waters. Unpublished M. S. Report, Dept. Environ. Engr. Sciences, Univ. Florida, Gainesville, Florida. Haac, W. R., anv T. Mix. 1988. Effect of a subsurface sediment on hydrolysis of haloalkanes and epoxides. Environ. Sci. Technol. 22:658-663. Licutroot, E. N., P. S. THorne, R. L. Jones, J. L. HANSEN, AND R. R. Romine. 1987. Labora- tory studies on mechanisms for the degradation of aldicarb, aldicarb sulfoxide and aldi- carb sulfone. Environ. Toxicol. Chem. 6:377-394. McCreary, J. J., J. JACKSON, AND J. ZOLTEK, JR. 1983. Toxic chemicals in an abandoned phenolic waste site. Chemosphere. 12:1619-1632. Mites, C. J., AnD J. J. Detrino. 1984. Determination of aldicarb and its derivaties in ground- water by high-performance liquid chromatography with uv detection., J. Chromatogr. 299: 275-280. , AND J. J. DELFINO. 1985. Fate of aldicarb, aldicarb sulfoxide and aldicarb sulfone in Floridan groundwater. J. Agric. Food Chem. 33:455-460. Moye, H. A., and C. J. Mixes. 1988. Aldicarb contamination of groundwater. Reviews Environ. Contam. and Toxicol. 105, in press. VocEL, T. M., AND P. L. McCarry. 1987. Rate of abiotic formation of 1,1-dichloroethylene from 1,1,1-trichloroethane in groundwater. J. Contam. Hydrol. 1:199-308. Florida Sci. 52(3):207-213. 1989. Accepted: August 17, 1988. Biological Sciences GROUND-WATER CONTAMINATION PROGRAMS OF THE U.S. GEOLOGICAL SURVEY IN FLORIDA IRWIN H. KANTROWITZ U.S. Geological Survey, WRD, Suite 3015, 227 North Bronough Street, Tallahassee, Florida 32301 Asstract: The U.S. Geological Survey has the principal responsibility within the Federal Government of providing the hydrologic information and understanding needed to achieve the best use and management of the Nation’s water resources. The Survey is involved in studies of ground-water contamination as part of its federally funded research program, in support of the programs of other Federal agencies, and in its cooperative water-resources program with State and local agencies. Research investigations currently (1988) active in Florida include studies of ground-water contamination from agricultural practices, landfills and hazardous waste sites, wastewater disposal, and seawater intrusion. These and other research activities, and the opera- tion of an extensive data-collection network, provide much of the scientific basis for the ground- water management programs of about 70 State, regional, and local agencies in Florida. THE U.S. GEOLOGICAL SuRvEY is a bureau within the Department of the Interior and has the principal responsibility within the Federal Government of providing the hydrologic information and understanding needed to achieve the best use and management of the Nation’s water resources. It is noteworthy that “management” refers to the actions of other agencies, and that “regulation” is not within the mission of the Survey. It is the Survey’s role to provide the hydrologic information and understanding to those agencies, both Federal and State, who are the managers and regulators. Ground water is the principal source of freshwater for public-supply, ru- ral, industrial, and irrigation demands in Florida (Fig. 1). More than 4 bil- lion gallons per day of fresh ground water were withdrawn, on average, in 1985 for these uses (Solley et al., 1988, table 24). The continuing rapid growth in Florida’s population is increasing the demand for high-quality po- table water. Florida’s aquifers consist of surficial sands and carbonate rocks; the distri- bution of the principal aquifers is shown (Fig. 2). Although generally abun- dant statewide, supplies in some areas where the surficial and intermediate aquifers are used may be somewhat limited. All the aquifers are subject to seawater intrusion in coastal areas. Over large parts of the State, the aquifers occur at land surface or are overlain by less than 10 feet of fine-grained sediments (Fig. 3) and are therefore vulnerable to contamination infiltrating from activities and structures at the land surface. These potential sources of contamination include 6,000 surface impoundments for waste disposal, 9,500 drainage wells, 60,000 underground storage tanks, 800 municipal landfills, more than 400 industrial waste sites, hundreds of thousands of septic tanks, and the widespread and intensive use of some 9,500 pesticides (Irwin and Bonds, 1987). Because of this vulnerability and the statewide dependency on No. 3, 1989] KANTROWITZ— USGS SURVEYS OF CONTAMINANTS 215 Rural/Domestic O 20 40 60 80 100 PERCENTAGE OF WATER USE DERIVED FROM GROUND-WATER SOURCES Fic. 1. Percentage of total water withdrawals in Florida derived from ground-water sources, by water-use category, 1985 (data from Solley et al., 1988). ground water, much of the Geological Survey’s data-collection and research efforts in Florida are directed toward studies of ground-water contamina- tion. GROUND- WATER CONTAMINATION PROGRAMS—Ground-water contamina- tion is an issue of national as well as State concern and the Survey is address- ing this problem in each of its three major authorized program areas: the Federal Program, the Other Federal Agencies Program, and the Federal- State Cooperative Program. These three programs differ in the sources of their funding, in the role the Survey plays, and in the objectives of the work. In each of the programs, however, the Geological Survey is involved in all scientific aspects of ground-water contamination studies—from basic re- search to resource appraisal. Federal Program: Funds are appropriated to the Survey by the U.S. Con- gress to support several high priority topical programs; the two programs concerned with ground-water contamination are the Toxic Waste—Ground- Water Contamination Program and the National Water-Quality Assessment (NAWQA). Although national in scope, significant research components of both programs are ongoing in Florida. The purpose of the Toxic Waste— Ground-Water Contamination Program is to provide the hydrologic understanding needed to improve waste-disposal practices and mitigate or prevent further contamination of water resources by toxic substances (Cardin et al., 1986). There are two research components of the program. One component consists of intensive interdisciplinary field 216 FLORIDA SCIENTIST [Vol. 52 87° 86° 85° 84° 83° 82° 81° 80° 31° 30° 29° 28° EXPLANATION BISCAYNE AQUIFER 27° SAND-AND-GRAVEL AQUIFER soc) UNNAMED SURFICIAL AQUIFERS AND INTERMEDIATE AQUIFERS, UNDIFFERENTIATED zeeF = [_]sFLORIDAN AQUIFER SYSTEM 26° 100 MILES SRSARRAREEAEEE 100 KILOMETERS Fic. 2. Approximate areal extent over which the principal aquifers are the primary source of supply (Vecchioli and Foose, 1985). investigations to apply theoretical and analytical methods in real-world situa- tions; to improve monitoring, sampling, and predictive procedures; and to develop new ways of mitigating ground-water contamination. One of six sites being investigated is at an abandoned wood-treatment plant overlying the surficial aquifer in western Florida. The research emphasis at this site is on hydrologic, geochemical, and microbial processes affecting the distribution, movement, and fate of creosote and other related chemicals introduced into the surficial aquifer from waste-disposal ponds (Franks, 1987). The second research component of the Toxic Waste—Ground-Water Contamination Pro- gram seeks to relate land-use and associated activities to the distribution and magnitude of selected manmade chemicals. One of the six areas being studied nationally is in central Florida where water quality beneath urban, citrus- No. 3, 1989] KANTROWITZ— USGS SURVEYS OF CONTAMINANTS Zp Fh 87° 86° 85° 84° 83° 82° 81° 80° SIG 30° 29° 28° 27° EXPLANATION 26° <5] PRINCIPAL AQUIFERS--occur at land surface or are overlain by less than 10 feet of fine-grained sediments 25° 50 100 MILES 100 KILOMETERS Fic. 3. Areas where the principal aquifers are most vulnerable to contamination from surface activities (modified from Healy and Hunn, 1984; and Miller, 1986, pl. 25). growing, phosphate ore mining and processing, and forested (natural) areas is being described and compared (Rutledge, 1987). NAW OA is the other Federal program concerned with ground-water con- tamination. The goals of the NAWQA Program are to provide a consistent description of current water-quality conditions for a large part of the Nation’s water resources, to define long-term trends in water quality, and to identify and describe the relations of current conditions and trends in water quality to natural and human factors (Hirsch and others, 1988). This program is still in its formative stage. Research is currently (1988) being conducted in Florida to evaluate different network-design strategies that may be used to collect perti- nent water-quality information. The research uses a large EDB (pesticide) data base for central Florida to test different sampling strategies. Subsets of 218 FLORIDA SCIENTIST [Vol. 52 the entire data base will be evaluated for their ability to describe the distribu- tion of EDB, first regionally and then in relation to various hydrogeological factors. Other Federal Agencies Program: The Survey is commonly requested by other Federal agencies (OFA) to assist them by supplying hydrologic informa- tion or expertise pertinent to their needs. About 60 agencies participate in this program on a cost-reimbursement basis. Nationally, the OFA program in- cludes considerable research involving various aspects of ground-water con- tamination; some examples are the development of a solute-transport model- ing technique (Department of Defense), research related to pesticide migration in the unsaturated zone (Department of Agriculture), and a study of contamination by trace metals associated with irrigation-drainage projects (Bureau of Reclamation). In Florida, ground-water contamination aspects of the OFA program have been limited to an advisory role in support of the Environmental Protection Agency’s toxic-waste cleanup actions, and a site evaluation for the Department of Energy. Federal-State Cooperative Program: Federal funds are appropriated by Congress for use by the Survey to match, on a dollar-for-dollar basis, funds offered by State or other tax-supported agencies. The Cooperative Program covers a variety of data-collection and research activities in which the Survey represents national responsibilities and the cooperating agencies represent State and local interests. In Florida, about 70 cooperating agencies work together with the Survey in determining the nature of the program; this part- nership approach ensures that the needs and priorities of all parties are met. Of the 45 cooperative-program investigations currently (1988) active in Florida, 23 are related to ground-water contamination issues. Of these 23 investigations, 2 deal with contamination from agricultural practices, 3 with landfills and hazardous waste sites, 6 with wastewater and highway-runoff disposal, 3 with saltwater intrusion, and 9 with general issues of ground- water quality management. For the most part, these investigations are con- cerned with local problems, but they also enable the Survey to develop new methods of study or better understand the movement and fate of contami- nants in the natural environment. As an example, the modeling of the growth and subsequent decay of a chloride plume from a flowing well will assist a local government in locating future ground-water supplies and will also result in an improved understanding of dispersion and dilution phenomena with expected transfer value to studies of contamination abatement. In addition to research activities, the Cooperative Program also supports a statewide hydrologic-data collection network. As part of this network, wa- ter-quality data are collected at more than 700 wells and springs. These data provide much of the basis for water-management decisions made by State and local agencies and are the scientific foundation for the research activities of the Survey and others in Florida. ADDITIONAL INFORMATION-—It is the policy of the Survey that, wherever possible, the results of its programs are made available to the pub- No. 3, 1989] KANTROWITZ—USGS SURVEYS OF CONTAMINANTS 219 lic, generally in the form of published reports. More than 350 reports on the ground-water resources of Florida have been published (Claiborne and oth- ers, 1987), and almost all contain some data on water quality. Investigations currently (1988) underway are described by Glenn (1988). Data collected at network sites or as part of investigations are published in annual data reports and are also generally available in machine-readable form. LITERATURE CITED Carpin, W. C., J. E. Moore, AND J. M. Rusin. 1986. Water Resources Division in the 1980's. U.S. Geological Survey Circular 1005. CLarBorNgE, M., T. L. Emsry, N. D. Hoy, D. H. WeLDon, ANnp T. D. Witson. 1987. Bibliography of U.S. Geological Survey reports on the water resources of Florida, 1886-1986, ed. 4, U.S. Geological Survey Open-File Report 85-424. Franks, B. J. 1987. Movement and fate of creosote waste in ground water near an abandoned wood-preserving plant near Pensacola, Florida. Pp A3-A9. In: Franks, B. J. (ed.), U.S. Geological Survey program on toxic waste. U.S. Geological Survey Open-File Report 87- 109. GLENN, M. (ed.) 1988. U.S. Geological Survey water resources activities in Florida, 1987-88. U.S. Geological Survey Open-File Report 88-199. Hearty, H. G., anv J. D. Hunn. 1984. Occurrence of beds of low hydraulic conductivity in surficial deposits of Florida. U.S. Geological Survey Water-Resources Investigations Re- port 84-4210. Hirscu, R. M., W. M. ALLEy, AND W. G. Wiser. 1988. A summary of the U.S. Geological Survey National Water-Quality Assessment Program. U.S. Geological Survey Open-File Report 88-95. IRwIN, G. A., AND J. L. Bonps. 1987. Florida ground-water quality, U.S. Geological Survey Open-File Report 87-0719. Miter, J. A. 1986. Hydrologic framework of the Floridan aquifer system in Florida and in parts of Georgia, Alabama, and South Carolina. U.S. Geological Survey Professional Paper 1403-B. Rut.ence, A. T. 1987. Effects of land use on ground-water quality in central Florida—prelimi- nary results. U.S. Geological Survey Water-Resources Investigations Report 86-4163. Sou_ey, W. B., C. F. Merk, AND R. R. Pierce. 1988. Estimated use of water in the United States in 1985. U.S. Geological Survey Circular 1004. VECCHIOLI, J., AND D. W. Foose. 1985. Florida ground-water resources. Pp. 173-178. In: U.S. Geological Survey, National water summary 1984, U.S. Geological Survey Water-Supply Paper 2275. Florida Sci 52(3):214-219. 1989. Accepted: September 23, 1988. Environmental Chemistry PESTICIDES AND GROUND WATER PROTECTION CHARLES C. ALLER Bureau of Ground Water Protection, Florida Department of Environmental Regulation, 2600 Blair Stone Road, Tallahassee, Florida 32399-2400 Axpstract: The development of the Department of Environmental Regulation (DER) pesti- cide program, based on a preventative strategy of ground water protection, is reviewed. Included is the current status of program activities in ground water protection, interagency pesticide activ- ities, field studies and monitoring, and remediation provided through the statewide ethylene dibromide (EDB) clean-up. Future areas of emphasis for ground water protection through man- agement of agricultural chemicals are reviewed. As the State lead agency for water quality, which includes the establish- ment of standards for all chemicals in ground water, the Florida DER has a major role in protecting water resources from pesticide contamination. To understand how the present ground water rules relate to pesticides, a review of the water quality and permitting procedures is necessary. Chapter 17-3 is the DER rule for Water Quality Standards (Chapter 17-3, Florida Administrative Code (FAC)). It has two important features— ground water classification and water quality standards. All ground waters in the state are classified by total dissolved solids (TDS) and degree of confinement into potable and non-potable uses (Swihart et al., 1984). Potable ground waters are classes G-I and G-II which have less than 10,000 mg/l TDS. AI- though there are presently no G-I ground waters, G-II probably covers more than 90% of the readily available ground water in the state. Class G-III and G-IV are non-potable classes, both having greater than 10,000 mg/l TDS with G-III being unconfined and G-IV confined. G-III ground waters may be found in coastal areas and the Florida Keys while G-IV ground waters underlay the entire state at depth (Conover et al., 1984). The second part of Chapter 17-3 establishes water quality standards in three parts: minimum criteria, primary drinking water standards, and sec- ondary drinking water standards. These standards are applied to those activi- ties which discharge to ground water and are designed to provide a high degree of protection to the resource that supplies over 90% of our drinking water, much of it used directly from private wells without treatment. The minimum criteria, also known as “free-froms” are those substances (in concentrations) which are carcinogenic, mutagenic, teratogenic, toxic to humans, or create a nuisance. No numerical standards are set for these com- pounds, although health advisories or other EPA or state guidelines are fre- quently used in applying these criteria. Most pesticides fall into this category. The primary drinking water standards are adopted by the federal govern- ment and the State and are applied as ground water standards. They include the inorganics, the organics or older pesticides, the volatile organics which No. 3, 1989] ALLER—GROUND WATER PROTECTION 22M include some gasoline constituents and industrial solvents, radionuclides, and turbidity, microbiological, and total trihalomethanes. The secondary drinking water standards are not adopted by the federal government, but rather are provided to the states as non-enforceable guide- lines. Florida, however, has adopted them as drinking water standards for certain larger public systems and does use them as ground water standards. For these standards certain older installations are exempt provided they do not cause violations in drinking water supplies. The secondary standard con- stituents’ effects are primarily aesthetic, although at high levels adverse health impacts are possible. The Chapter 17-3 standards for classification and water quality criteria are applied to ground water discharges through permitting procedures which are contained in Chapter 17-4. The basis of this is that installations which discharge to the ground water should be allowed a portion of the aquifer within which the water quality standards may be exceeded, known as the zone of discharge (ZOD), but at some point removed from the discharge which will vary with the criteria and the type of installation, the standards must be met. Only in unusual circumstances, such as an exemption, will this go beyond the dischargers’ property boundary. For permitted discharges, monitoring wells are required to assure compliance. The dimensions of this zone of discharge will vary, depending on the water quality criteria (Chapter 17-4, FAC). There are many activities which can create a discharge to ground water that do not require a permit or site specific monitoring. Among these are agricultural fields which may, through the application of pesticides and fer- tilizers, affect either the minimum criteria or, for nitrates, the primary drink- ing water standards. Agricultural fields are given, by the rules, a zone of discharge of 100 feet or to the property boundary. This allows the standards to be exceeded beneath a field, but requires that they be met off site where another person’s ground water (drinking water) may be affected. A clear provision of the rules, and one which is not widely understood is a provision for permitting agricultural operations if water quality standards are threat- ened. Prior to 1983, the Department did not have a program or capability spe- cific to pesticides. With the passage of the Water Quality Assurance Act, the Legislature provided positions in the Department with scientific expertise to review the environmental effects of pesticides. These include experts in the disciplines of hydrogeology, entomology, biology, chemistry, toxicology, and data management. Since the establishment of the Department’s pesticide program in 1983, DER scientists have worked closely with scientists at the Florida Department of Agriculture and Consumer Services to assess the environmental effects of pesticide use patterns. Considering these effects, such as the potential to con- taminate ground water, means that pesticide management strategies in lieu of use prohibition can be employed on a product or chemical specific basis. 222 FLORIDA SCIENTIST [Vol. 52 Applied during the process of registering pesticides for use in the State, such techniques as application timing, rate of application, different formulations, or restriction of use on sensitive soils can be employed as preventatives to ground water contamination. A prime example of this approach, and one which is likely to be more heavily employed in future years, is the case of aldicarb. On certain soil types there are seasonal, formulation, and well set- back limits specified for the use of the chemical (Chapter 5E-2, FAC). CurrRENT Status—In the early 1980s the reported presence of aldicarb (Swihart et al., 1984) in Florida ground water publicly raised the issue of pesticides entering ground water in a broad sense. It has been, however, EDB and other similar soil fumigants applied to control nematodes which have created the largest impact on ground water used as drinking water. To date, EDB has been found in about 1600 drinking water wells, and the state has, at a cost of millions of dollars provided over 1000 remedies in the form of GAC filters, new wells, or connections to existing utility systems. The statewide distribution of EDB contamination reflects its use in agriculture such as citrus and row crops, and turf applications such as golf courses. EDB has been detected in drinking water wells in 24 counties. In Polk County, the known area of application, principally along the ridge where citrus is grown, closely reflects the pattern of positive detections. Both of these patterns correlate highly with agricultural land use, which demonstrates a principal tool of the statewide Ambient Ground Water Moni- toring Network. Using such land use information in relation to hydrogeologic vulnerability will assist in siting monitor wells around the state to survey for other pesticides which may have entered ground water (Alexander and Miller, 1986). In the early phases of EDB sampling a review of the use patterns, method of application, and physical chemical properties of several other chemicals suggested that they might also be found in ground water. Because of this, analysis for EDB also began to include a scan for eight other pesticides or degradation products. At present there have been detections for dibromo- chloropropane, 1, 2-dichloropropane, and 1, 3, dichloropropene. The geo- graphic distribution of these chemicals reflects the sampling that has been done through the EDB program. Most of the wells containing these pesticides also contain EDB, however, a few do not. Sample results on finished water demonstrates that the GAC filters also remove these chemicals to below de- tection limits. The widespread detection of EDB in Florida ground water and reports of detections of other pesticides in other states provided impetus for a more scientific means of evaluating pesticides for their potential to leach to ground water. Following more than a year of staff work on the part of the Florida Department of Agriculture and Consumer Services (DACS) and DER, the Florida Pesticide Review Council, an interagency scientific advisory council, No. 3, 1989] ALLER— GROUND WATER PROTECTION upg published the Pesticide Assessment Procedure (Florida Pesticide Review Council, 1986). This procedure provides a means of combining pesticide and hydrogeolo- gic factors to develop a ranking for decisions on pesticide field studies. Pesti- cides are rated by sorption coefficient, half life, application method, and toxicological factors. The hydrogeologic setting in which a given pesticide is used may be ranked by proximity to the source aquifer, confinement, sinkhole probability, recharge capacity, and surface runoff. Although, as with most ranking systems, some subjectivity is introduced in assigning points to the various categories, the procedure establishes a means of relating critical pesti- cide characteristics to hydrogeologic sensitivity. The state has, in fact, used this or similar ranking to set priorities for the scientific evaluation of pesticides. Currently there are 62 products tentatively assigned to the product review program which is run by DACS. Assisting DACS in this review, the Department will be evaluating these materials as potential ground water contaminants. Within this group, twelve chemicals are under active review, and eight have been or will be studied in the field through carefully controlled ground water monitoring programs. Such stud- ies are always performed in cooperation with DACS and, usually, the regis- trant of the product. Four other compounds are likely candidates for future field studies. A final note on field studies should include mention of the Collier County regional study now in progress. The first phase of this effort involved well installation and ground water sampling for a variety of crops in a 60 square mile area in Collier County for 22 pesticides. No detections were reported during two seasonal sampling runs. Working with EPA, the Water Manage- ment Districts, IFAS, the Soil Conservation Service, DACS, and others, fu- ture work is planned to assess the impact of regional agricultural develop- ment on water quality in this area. CoNncLusion— Data collected during the last several years and compiled by the Department demonstrates that 16 pesticides have been found in ground water as a likely consequence of normal use. When loading and dis- posal operations are considered, another 20 chemicals are known to have been found in ground water. Most of these detections represent localized events, dependent on hydro- geologic and climatologic factors, patterns of chemical use, cropping and irrigation practices. The single example of EDB, however, demonstrates that without proper regard for careful pesticide management, based upon chemi- cal and hydrogeologic properties, pesticides are capable of widespread ground water contamination. The state now has in place a preventative mechanism to evaluate the environmental effects of pesticides. The Department supports this preventa- tive approach with scientific expertise in data review, hydrogeologic field studies, and remedial action. In the future, as more information is available, management strategies based upon local soil and hydrogeology, application 224 FLORIDA SCIENTIST [Vol. 52 rates, formulations, or timing of applications will be used for certain chemi- cals. Such an approach will be essential to protect our ground water resources and maintain the continued use of some chemicals needed for agricultural production. LITERATURE CITED ALEXANDER, J. F., AND W. L. Miter. 1986. An Information System to Locate Potential Threats to Ground Water Resources. Contract No. 59-3-0209K for the Florida Department of Envi- ronmental Regulation. Univ. Florida, Gainesville. CuHapTer 17-3, Florida Administrative Code. CHAPTER 17-4, Florida Administrative Code. CHAPTER 5E-2, Florida Administrative Code. Conover, C. S., J. J. GeRATHY, AND G. G. ParKER, Sr. 1984. Ground Water, Pp. 36-53 In: FERNALD E. A., AND D. J. PaTTon (eds.). Water Resources Atlas of Florida. Florida State Univ., Tallahassee. FLorRIDA PESTICIDE REvIEW CouncIu. 1986. Pesticide Assessment Procedure. Tallahassee. SwiHaert, T., J. HAND, D. Barker, L. BELL, J. CARNES, C. Cosper, R. DEUERLING, C. GLUCKMAN, W. HInktey, R. Leins, E. Livincston, AND D. York. 1984. Water Quality. Pp. In: FERNALD, E. A., AND D. J. Patron (eds). Water Resources Atlas of Florida. Florida State Univ., Tallahassee. Florida Sci. 52(3):220-224. 1989. Accepted: October 11, 1988 INSTRUCTIONS TO AUTHORS Individuals who publish in the Florida Scientist must be active members in the Florida Academy of Sciences. Submit a typewritten original and two copies of the text, illustrations, and tables. All type- written material—including the abstract, literature citations, footnotes, tables, and figure legends — shall be double-spaced. Use one side of 82 x 11 inch (21% cm X 28 cm) good quality bond paper for the original; the copy may be xeroxed. Margins should be at least 3 cm all around. Number the pages through the Literature Cited section. Avoid footnotes and do not use mimeo, slick, erasable, or ruled paper. Use metric units for all measurements. Assistance with production costs will be negotiated directly with authors of papers which exceed 10 printed pages. 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Florida's Estuaries —-Management or Mismanagement? — Academy Symposium FLoripA SCIENTIST 37(4) — $5.00 Land Spreading of Secondary Effluent —Academy Symposium FLORIDA SCIENTIST 38(4) — $5.00 Solar Energy — Academy Symposium FLORIDA SCIENTIST 39(3) — $5.00 (includes do-it-yourself instructions) Anthropology — Academy Symposium FLoripa SCIENTIST 43(3) — $7.50 Shark Biology —Academy Symposium FLoripa ScieNTIsT 45(1) — $8.00 Future of the Indian River System — Academy Symposium FLoripa SCIENTIST 46(3/4) — $15.00 Individual orders should be sent with payment. A statement will be sent in re- sponse to a bona fide purchase order over $10.00 from a recognized institution. Ad- dress all orders to: The Florida Academy of Sciences, Inc. c/o The Orlando Science Center 810 East Rollins Street Orlando, Florida 32803 Phone: (305) 896-7151 ISSN: 0098-4590 Scientis Volume 52 Autumn, 1989 nnv 1 3 1989 _ LIBRARIES) CONTENTS Symposium: “Groundwater Contamination and Protection “rh FLOTIG 2. GOTTTECG (2) sneer meg eae ar aay 225 Prevention and Cleanup of Petroleum Contamination of Grounadwater—Florida’s Super Act... 2... 1... oe eee eee eee Craig Ash, Connie Garrett, and Susan Gray 225 The Status of Superfund and State-Funded Cleanup Sites 2 [ELCIRIGIEL +o-5 oto Soleo kee ere John M. Ruddell 230 Impact of Groundwater Contamination on Public OSS? (SNP OTO) TSS eG aan ae ce J. Edward Singley 240 Regular Contributions: The Neural Derivative, OCOS and Motion Detection ........... David Lawson 244 maeopous Enaemic to Florida Scrub .... 6.6.0.0... cee eee eee. Mark Deyrup 254 Productivity of Departments of Chemistry at Florida ese Ake PUSPIUUILIONS) ae ies ce se cd RA a eae Se thee ee 8 John C. Follman and Dean F. Martin a Infestation and Epidemiology of Head Lice in Elementary Sensolsin Hillsborough County, Florida .............06.0..000-- W. Wayne Price and Amparo Benitez 278 An Investigation of the Meteorological Conditions Affecting Pespersion of Ozone in the lampa Bay Region ................-. Dewey M. Stowers, Jr. and Neva Duncan Tabb 289 EE re ie ie he) eee ys ca eae. oe ete eae tes wD EES 299 ITE or esclcen en a gr 300 STROM ABUTS IO ts 5 ee os oie os Soe be Cate evele ols 8 BAS ars 301 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE F'LORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1989 Editor: Dr. DEAN F. MartTIN Co-Editor: Mrs. BARBARA B. MARTIN Institute for Environmental Studies Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoripDA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. 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Martin, Editor BARBARA B. Martin, Co-Editor Volume 52 Autumn 1989 Number 4 Geological Chemistry PREVENTION AND CLEANUP OF PETROLEUM CONTAMINATION OF GROUND WATER— FLORIDAS SUPER ACT Craic ASH, CONNIE GARRETT!, AND SUSAN GRAY? Florida Department of Environmental Regulation, Tallahassee, Florida 32399-2400 Asstract: In an effort to protect public drinking water supplies, the Florida Legislature enacted the “State Underground Petroleum Environmental Response Act” of 1986 (SUPER Act). This bill created a trust fund which provides monies for the state to conduct site rehabilitation projects or for reimbursement to persons who have voluntarily or through negotiated enforcement cleaned up their sites. Participation in the program is encouraged by the establishment of a grace period (July 1, 1986-December 31, 1988) during which owners or operators of petroleum storage systems that report suspected contamination will not be held liable for the costs of restoring their sites. More than 9,000 applications were received during the 30 month grace period. Rules covering the order of site cleanup and reimbursement, establishing cleanup guidelines, and providing for reimbursement of reasonable costs have been adopted by the Environmental Regulation Commission. GROUND WATER is such a critically important resource in the United States that half the country’s residents rely on it as their primary source of drinking water (Environmental Protection Agency, 1984). Florida’s ground water de- pendency is even greater. Approximately 92 percent of the state’s residents use ground water for their potable supply, and 20 percent of those obtain their drinking water from untreated and unregulated private wells (Florida Depart- ment of Environmental Regulation, 1985). Ground water dependency has been increasing with Florida’s relentless growth. Since 1980, the population has increased by an average of 893 persons per day (Zwick, 1986). Florida’s concern for leaking petroleum storage tanks has been on the rise over the past several years. Increased awareness coupled with a growing de- pendency on ground water resources prompted the state’s commitment to ground water protection. Florida’s comprehensive program for protection from leaking petroleum storage tanks began with the passage of the 1983 Water Quality Assurance Act. This act led to the promulgation of the Stationary Tanks Rule, Chapter 17-61, Florida Administrative Code (FAC) which became effective in May, 1984. The Stationary Tanks Rule requires that owners and operators of vehicular fuel tanks with a capacity greater than 550 gallons register their tanks with the 1Florida Department of Health & Rehabilitative Services— Toxicology & Hazard Assessment, Tallahassee, Florida 2Dames & Moore, Boca Raton, Florida 226 FLORIDA SCIENTIST [Vol. 52 Florida Department of Environmental Regulation (FDER). The rule also re- quires maintenance of inventory records, installation and use of monitoring systems, a schedule for replacement or retrofitting of old tanks and proper abandonment of tanks that are no longer in use. An important requirement of the Stationary Tanks Rule is that the FDER be notified of significant petroleum product discharges. Following such noti- fication, the owner or operator found responsible for environmental contami- nation resulting from a leaking petroleum storage system would be held liable for the costs associated with cleanup. Because of the potentially devastating economic consequences of a discharge and subsequent cleanup, many known leaks were not reported. A combination of highly permeable soils and shallow water tables makes Florida’s ground water extremely vulnerable to contamination from leading petroleum storage tanks. There are more than 76,000 petroleum storage tanks at an estimated 28,000 registered facilities. Many of these facilities are gaso- line stations that contain original bare steel tanks, which are now at leak- developing stages. Studies done in Florida by the American Petroleum Insti- tute indicate an average product loss of 2,300 gallons in leaks that last for more than one day (Florida Petroleum Council, 1986). Improperly aban- doned tanks are also a major concern. As they begin to rust and deteriorate, conditions exist for any remaining product to leak into the surrounding soils and eventually into the ground water. Discussion—SUPER Act Early Detection Incentive and Reimbursement Programs— The number of reported ground water contamination incidences involving petroleum products rose from several in 1983 to 695 by July 1, 1986. In response to the increasing number of confirmed contamination problems and the reluctance of facility owners and operators to report a leak or discharge, Florida enacted the State Underground Petroleum Environ- mental Response Act of 1986 (SUPER Act). This act was established to pro- tect the state’s drinking water supplies by easing the regulatory and financial burdens placed on owners and operators of leaking petroleum storage sys- tems. A series of funding mechanisms, incentives for early leak reporting, and allowances for reimbursement of costs for voluntary cleanups were devel- oped. Perhaps the most important provisions of the SUPER Act are the Early Detection Incentive and Reimbursement Programs. Storage facilities that are eligible for these programs are not held liable for remediation costs associated with petroleum contamination reported to the FDER. Restoration of a site may be accomplished by doing a voluntary cleanup, requesting a state-ad- ministered cleanup, or a combination of the two programs. The SUPER Act, as amended during the 1988 legislative session, set forth a 30 month “grace” period beginning July 1, 1986 and ending December 31, 1988. This grace period is known as the Early Detection Incentive (EDI) Program. A storage facility which has a discharge reported for the first time No. 4, 1989] ASH ET AL.—FLORIDA’S SUPERACT Hapag| during the EDI period may apply for either state cleanup or reimbursement. Special provisions of the Reimbursement Program entitle site owners or oper- ators to apply for reimbursement of voluntary cleanup costs incurred on or after January 1, 1985. Other eligibility requirements for the programs in- clude: 1) the facility cannot be federally owned or operated; 2) the owner or operator cannot have denied site access to the FDER; and 3) the owner or operator must comply with the registration, inventory, monitoring, and ret- rofitting requirements of the Stationary Tanks Rule, Chapter 17-61, FAC. In addition, the discharge must not have been caused by gross negligence or intentional tampering on the part of the owner or operator. Intent to conceal a discharge also disqualifies a site. Additional eligibility requirements for the Reimbursement Program include: 1) submittal of documentation of site con- ditions prior to cleanup; 2) no previous discharge violations without appro- priate cleanup actions; and 3) filing a notice of intent to apply for reimburse- ment to the FDER. The order in which the state will cleanup or reimburse owners or opera- tors of eligible sites is determined by the Site Priority Ranking Rule, Chapter 17-71, FAC. This rule provides for the assignment of numerical scores to various environmental and public health considerations for each eligible pe- troleum contamination site. Factors that are taken into consideration for cal- culating each site’s score include fire or explosion hazard, threat to uncon- taminated drinking water supplies, migration potential of the contaminant, and environmental setting. All eligible sites are then ranked according to their environmental threat score on a quarterly basis. The sites with the high- est ranks are given priority for the assignment of funds. The number of sites funded quarterly depends upon the available balance in the trust fund. One of the most critical issues of site rehabilitation is the question, “How clean is clean?”. The Petroleum Contamination Site Cleanup Criteria Rule, Chapter 17-70, FAC, became effective November 1, 1987. This rule considers such factors as: the degree to which human health, safety or welfare may be affected by exposure to the contamination; the present and future uses of the affected aquifer or surface waters; the effect of the contamination on the environment; the individual site characteristics; and, the application of state water quality standards. Procedures and program tasks to be completed dur- ing site rehabilitation are described in the rule. SUPER Act Funding and Reimbursement Guidelines—Both the Early Detection Incentive and Reimbursement Programs receive funding from the Inland Protection Trust Fund which was created by the SUPER Act. The Inland Protection Trust Fund is financed by a tax on pollutants and petro- leum products that are imported into or produced in Florida. Taxing provi- sions in the Act ensure the fund will remain in effect until 1997. Under SU- PER Act, this trust fund provides monies for costs associated with contaminated soil removal, petroleum product recovery and ground water restoration. Funds are not to be used for costs associated with tank testing, replacement or repair, or litigation fees. 228 FLORIDA SCIENTIST [Vol. 52 Reasonable cost guidelines for reimbursable expenses are addressed by the FDER in the Reimbursement for Contamination Site Cleanup Rule, Chapter 17-73, FAC, which became effective in May, 1988. This rule establishes pro- cedures to be used and documentation required to receive reimbursement for site rehabilitation costs. SUPER Act Preventative Programs—Other preventative provisions con- tained in the SUPER Act are the Pollutant Storage System Specialty Contrac- tor Program (PSSSC), the Registration Program and the Compliance Verifi- cation Program. These programs are designed to prevent leaks associated with petroleum storage tanks. The PSSSC program is a joint effort between the state’s Department of Professional Regulation and FDER. An examina- tion and certification program for tank installers and removers has been de- veloped. By establishing consistent criteria for these contractors, the number of leaks due to improper installation or removal of tanks is being reduced. The Registration Program is important for identifying all storage tanks within Florida, including not-in-service or abandoned tanks. A registration fee is assessed to all storage tanks with a capacity greater than 550 gallons that store vehicular fuel. Any change in ownership or tank status, including tank replacement, must be reported to the FDER. The FDER has registered ap- proximately 76,000 tanks at over 28,000 facilities to date. The 1987 amend- ments to the SUPER Act extended the registration program to all tanks re- gardless of size, except for residential heating oil tanks. The development of a Compliance Verification Program to ensure proper tank construction, monitoring and maintenance was required by SUPER Act to be in effect by December 31, 1987. The priority for facility inspections places emphasis on inspecting sites in wellfield protection areas, areas with private potable wells, new installations and abandoned tanks, The FDER is contracting with local governments to perform the inspection program. Involvement of State Agencies and Counties in the SUPER Act—State agencies other than FDER have responsibilities mandated by the SUPER Act. These agencies include the Department of Professional Regulation, the De- partment of Agriculture and Consumer Services, the Department of Health and Rehabilitative Services and the Department of Revenue. The Florida Department of Professional Regulation worked with FDER to develop requirements for certification of petroleum storage tank installers and removers. The Department of Agriculture and Consumer Services (DACS) checks for proof of valid tank registration during its regular inspec- tions of retail facilities with pollutant storage systems. Unregistered facilities are reported to the FDER. In addition, as of July 15, 1987 it was illegal for gasoline distributors to fill unregistered tanks. This action is being monitored and regulated by the DACS. The Department of Health and Rehabilitative Services provides the FDER with well location information, and toxicologic © and public health information for potable wells in the proximity of reported x No. 4, 1989] ASH ET AL.—FLORIDA’S SUPERACT 229 petroleum contamination sites. This information is collected by county health agents who have access to detailed well location information. ConcLusion— The SUPER Act is an important part of the comprehensive legislation passed by the Florida government to protect the state’s ground water. The SUPER Act and the Stationary Tanks Rule together provide for inspection of new petroleum storage tank installations, maintenance and monitoring of tanks that are in operation, and incentives for reporting dis- charges in the form of state funding for site cleanup or cleanup reimburse- ment. The specific intent of the SUPER Act is to protect Florida’s critical ground water resources. By minimizing the number of petroleum contamina- tion sites, the health risks associated with contamination sites are lessened. Public safety and welfare are also protected in cases of petroleum contamina- tion by having established procedures for dealing with the repair or replace- ment of contaminated potable water systems. Response to the SUPER Act’s incentive providing programs was encouraging: More than 10,500 applica- tions were received during the grace reporting period. By mid-year 1989, it is projected that assessment and cleanup activities will be initiated at approxi- mately 1,500 petroleum contamination sites. LITERATURE CITED Anonymovs. 1984. Protecting ground water, the hidden resource. U.S. Environ. Prot. Agency Jl. August, 1984. FLoRIDA DEPARTMENT OF ENVIRONMENTAL REGULATION. 1985. Fact Sheet 1, July. Tallahassee, Florida. FLoRIDA PETROLEUM CouncIL. 1986. Benzene in Florida ground water, an assessment of the significance to human health. American Petroleum Institute. Washington, D.C. October, 1986. Zwick, C. 1986. Speech delivered to the Florida Council of One-hundred. The Breakers, Palm Beach, Florida. November 7. Florida Sci. 52(4):225-229. 1989. Accepted: January 20, 1989. Geological Sciences THE STATUS OF SUPERFUND AND STATE-FUNDED CLEANUP SITES IN FLORIDA JoHN M. RupDDELL Bureau of Waste Cleanup, Florida Department of Environmental Regulation, 2600 Blairstone Road, Tallahassee, Florida 32399-2400 AssTract: Of the 469 Florida sites with potential contamination identified as of May i988, 39 are National Priorities List (NPL) sites and 23 are State Action sites—making them eligible for either the federal Superfund program or the state funded cleanup program. Thirteen of the Superfund sites are under state lead and 26 are managed by the U.S. Environmental Protection Agency (EPA). Forty one percent of Superfund sites are in the design or construction steps. Interim Remedial Measures (IRM) have been completed or are underway at 25 Superfund sites. Three of the Superfund sites have been proposed for deletion from the NPL—the cleanup com- pleted or long term treatment installed and operating. Fifty two percent of the State Action Sites are in design or construction. IRM’s have been completed or are underway at 7 State Action Sites. Remedial actions have been completed at 5 State Action Sites. Progress from site discovery to cleanup for all Florida sites is discussed along with the technical and institutional issues that affect the time it takes to complete the various stages of a cleanup. HAZARDOUS waste sites are a potential source of contamination of Florida’s groundwater. With 92% of the state’s drinking water coming from ground- water, hazardous waste contamination poses a significant public health and economic threat, in addition to the environmental problems it causes (Lewis et. al., 1986). In 1980 Congress passed Public Law 96-510, the Comprehen- sive Environmental Response Compensation and Liability Act (CERCLA), commonly known as “Superfund”’. The law was amended in 1986 by passage of Public Law 99-499, the Superfund Amendments and Reauthorization Act (SARA). CERCLA, and the subsequent SARA amendments, established the programs, authorities, and funding to allow the U.S. Environmental Protec- tion Agency (EPA), in conjunction with the various states, to investigate and clean up abandoned or uncontrolled hazardous waste sites throughout the country. In 1983 the Florida Legislature passed Chapter 83-310, Laws of Florida, the Water Quality Assurance Act (WQA). This law provided specific authori- ties and a source of funds to the Florida Department of Environmental Regu- lation (FDER) for the development of the state’s hazardous waste response program. The WQA made it possible for Florida to participate in Superfund by providing money for the required state matching funds. It also provided the state with the resources to investigate and cleanup sites that would not qual- ify for the federal program. It is these two laws, and the subsequent programs developed from them, that form the basis of hazardous waste cleanup in Florida. With Superfund nearly 8 years old and the state program in place almost 5 years, it is an appropriate point to assess the status of hazardous waste cleanup in Florida. No. 4, 1989] RUDDELL— STATUS OF SUPERFUND 231 MerHops— All data for this paper are summarized from EPA and FDER publications as well as unpublished FDER records and files. Data are arranged to show the progressive stages in both the Superfund and state programs. The stages include site discovery, site verification, site investi- gation, and site remediation. Various elements of each stage are discussed and areas that cause delays are identified. RESULTS AND Discussion—Both the federal Superfund and the state cleanup programs follow similar procedures in moving from the discovery of a potential contamination site to its eventual cleanup. The two programs differ only in detail and the relative emphasis placed on each stage. In many cases, the procedures in the federal program are more formal with less flexi- bility than those in the state program. Both programs begin with site discov- ery. Site Discovery Stage—Federal—EPA’s Comprehensive Environmental Response, Compensation, and Liability Information System (CERCLIS) in- ventory of potential hazardous waste sites lists approximately 27,000 sites nationally. The U.S. General Accounting Office (GAO) considers this list to represent only a small fraction of the potential sites throughout the country (GAO, 1987). The GAO (1987) estimates range from 130,300 to 425,400 potential sites. CERCLIS, however, is a listing of only those sites reported to EPA. Sites are discovered by EPA as a result of a variety of statutory reporting requirements. All past and present owners of hazardous waste facilities or facilities where hazardous substances were stored were required to report to EPA by June of 1981 in accordance with Section 103(c) of CERCLA. Section 103(a) also requires that releases of hazardous materials must be reported to EPA. The various states also report sites they discover. In addition EPA main- tains a “hot line” where citizens can report suspected hazardous waste sites. These and other sources have provided the basis of CERCLIS. There are currently 855 Florida sites on CERCLIS (FDER, unpublished data). State— Florida discovers sites in a variety of ways. Sites are discovered by inspections in one of FDER’s other programs; e.g. Industrial Waste or Solid Waste permitting. Sites are discovered by other state agencies and local gov- ernments; e.g. the Florida Department of Transportation through highway right-of-way acquisitions. Sites are identified through state or federally spon- sored regional sampling programs. EPA will report sites to the state from CERCLIS. Citizens report sites directly to FDER or through the Department of Natural Resources’ Resource Alert Hotline. The FDER list of hazardous waste sites is known as “The Sites List” (FDER, 1988b). Unlike EPA’s CERCLIS, sites are added to the state’s list only after there has been some verification of contamination, and sites are removed from the list once cleanup has been completed. There are currently 469 sites on “The Sites List” (FDER, 1988b). Site Verification Stage—Federal—The primary site verification efforts of EPA are carried out through the CERCLA Site Screening Program. Either 232 FLORIDA SCIENTIST [Vol. 52 EPA, or the states under contract to EPA, will verify through a three-step process that sites are contaminated. The first step is a Preliminary Assessment (PA) where all available information is gathered about a particular site. Based on this information a site is ranked “high”, “medium”, or “low” prior- ity for the next step—a Site Investigation (SI). In the SI step EPA, or state- hired contractors, will sample the various sites according to their priority in order to confirm contamination. Once contamination is confirmed the site is scored through EPA’s Hazard Ranking System (HRS). Any site scoring 28.5 points or above, out of a possible 100 points, qualifies for inclusion on the National Priorities List (NPL) (EPA, 1986). NPL listed sites are the true Su- perfund sites and are eligible for federal cleanup money. Table 1 summarizes the status of PA’s and SI’s for Florida sites on CER- CLIS and distinguishes between the number of PA’s and SI’s done by the state and those done by EPA. To date, 829 of the 855 Florida sites have had PA’s completed and SI’s have been done for 244 sites (FDER, unpublished data). The pace in completing PA’s and SI’s has been controlled primarily by available EPA funds. There was a marked slow down in the number of sites examined during the year long period that corresponded to Superfund reauthorization. One other significant area of delay, primarily in the SI step, can be in gaining site access. If the owner of a site does not voluntarily allow the government to sample, then relief must be sought through the courts. This can be a lengthy process which can delay investigations from several months to several years. TABLE 1. Preliminary assessments and site investigations completed (May, 1988). Activity FDER EPA Total Preliminary Assessments (PA) ols 316 829 Site Investigations (SI) 109 135 244 State—Site verification is done on the state level partially through the CERCLA Site Screening Program and partially through FDER staff investi- gations. FDER enforcement in a number of different programs will also result in investigations that find and confirm contaminated sites. Many of these investigations are ordered by FDER and then carried out by the owners of the sites (Responsible Parties). Once contamination is confirmed the site is added to “The Sites List”’. Table 2 is a summary of the categories of sites on “The Sites List’. It includes the Superfund sites in the state as well as sites that are part of the state cleanup program (State Action Sites). Of the 469 sites on the state list, 332 of them fall to responsible parties for further investigation and subse- quent cleanup. Table 2 also breaks down the type of contamination found at the sites with confirmed contamination. Groundwater contamination occurs No. 4, 1989] RUDDELL—STATUS OF SUPERFUND Dapeke) TABLE 2. “The Sites List” summary (May, 1988). Total Sites on List 469 Superfund Sites 39 State Action Sites 23 Responsible Party Sites 332 Sites Requiring Investigation 75 Total Sites with Confirmed Contamination 295 Groundwater Contamination 262* Soil Contamination aa Surface Water Contamination Sly *A site may have more than one type of contamination. at 89% of the sites. This confirms that hazardous waste sites in a state like Florida, with its high groundwater table and porous soils, do pose a signifi- cant threat to the groundwater. The discrepancy in total sites on “The Sites List” and the number of sites with confirmed contamination comes from the way the original list was com- piled in 1983 (FDER, 1983). The first “Sites List’ was a combination of all state and federal lists in effect at the time. A number of the lists contained only suspected sites. FDER is systematically working through these old sites with unconfirmed contamination in order to verify their status. Site Investigation/Site Remediation Stages— Federal—Once a site is eligi- ble for the National Priorities List (NPL) the lead agency to oversee site inves- tigation and cleanup is determined by EPA in consultation with the state. The lead can fall either to the state or to EPA. For all NPL sites it is preferable for Potentially Responsible Parties (PRP’s) to conduct site remedial activities with government oversight. When the PRP is unable or unwilling to conduct the investigation and cleanup then the lead government agency will do the work using federal Superfund money and seek cost recovery. There are 888 sites nationwide either on the NPL or officially proposed for addition to the NPL (EPA, 1986). Table 3 lists the 39 Florida sites on or proposed for the NPL (32 on the list, 7 proposed) and their locations. The lead is indicated on Table 3 as is the party responsible for doing the work (FDER, 1988a). In the federal Superfund Program the site investigation and site remedia- tion stages are divided into several very specific steps. The steps that must be followed at a Superfund site include: a Remedial Investigation (RI) that char- acterizes the nature and extent of contamination; a Feasibility Study (FS) (including a Risk Assessment) that sets the cleanup targets, and identifies and evaluates the alternatives available for cleanup; a Record of Decision (ROD) that establishes the selected and approved remedy as part of the official EPA 234 FLORIDA SCIENTIST [Vol. 52 record; a Design (DES) that will provide for detailed engineering of the se- lected remedy and develop specifications for its construction; and finally Construction (CON) of the remedy itself. Once the remedy is in place, any on-going treatment facilities, such as groundwater recovery systems, are op- erated and the site is monitored until cleanup target levels are reached. In the case of groundwater treatment this last step may go on for many years. Figure 1 represents the number of Superfund sites in each stage of cleanup. This figure reflects the “pipeline” nature of the cleanup process. The large number of sites initially on the NPL in 1982 have moved their way through the steps of a cleanup with newer sites being added to the front end. In comparing the sites in each category, it is apparent that a significant num- ber of Florida’s sites are through the initial study steps with 16 of 39 sites (41%) having reached or completed the design or construction stages. Three sites have had remedial actions completed and these are pending EPA delist- ing from the NPL. Normally, federal construction money will not be spent at a site unless all steps have been completed. The one exception to this rule is the Initial Reme- dial Measure (IRM) step. IRM’s are primarily source control measures de- signed to eliminate or abate a continuing source of contamination while the studies and design are underway. Since the study and design steps can take several years to complete, sources that continue to leach contaminants can pose a continuing threat. The IRM step is designed not only to eliminate an environmental threat, but also to prevent the size of the problem at a site from becoming larger and the final cleanup more expensive. IRM’s are lim- ited by law (CERCLA Section 104(c)(1) as amended) to $2 million and 1 year to complete. They are intended to be limited and interim in nature. Figure 1 shows that IRM’s are done or underway at 25 Florida sites. NUMBER OF SITES 40 COMPLETED Be UNDERWAY TOTAL 30 20 = 1e,) oO Be NS $255 —_ (je) (xX) Sd 5504 © O05 OX SSC £282 oe) Ma O o, , OM , ©, O,, 6252 WG RQ AA? SS , SAA? OY OY 2% \ SOS2 ) oO 7 ] lj ] j l Z G j V l; CMO NE we oteres oe, x 05 LA} SO ne, IN x» ore; ree se FS ROD DES CON IRM a Fic. 1. Status of Florida Superfund sites. (Total number of sites is 39, and remedial actions have been completed at 3 sites. Note: A site may be in more than one category). No. 4, 1989] TABLE 3. Florida Superfund sites (May, 1988). Name Alpha Chemical American Creosote Biscayne Aquifer Brown Wood Preserving Cabot Carbon/Koppers City Chemical-Forsyth Road! Coleman Evans Davie Landfill Dubose Oil Products Florida Steel Co. Gold Coast Oil Harris Corporation! Hipps Road Landfill Hollingsworth Solderless Terminal Kassouf/Kimmerling Montco Research Products! Munisport Landfill Northwest 58th St. Landfill Parramore Surplus® Peak Oil/Bay Drum Pepper’s Steel & Alloys Petroleum Products! Picketville Road Landfill Pioneer Sand Piper Aircraft Corp/Vero Beach Water & Sewer Dept.! Pratt & Whitney Aircraft! Reeves Southeast Galvanizing Sapp Battery Salvage Schuylkill Metals Sherwood Medical 62nd Street Dump Sydney Mine! Taylor Road Landfill Tower Chemical Tri City Oil Conservationist® Varsol Spill Whitehouse Oil Pits Yellow Water Road Zellwood Groundwater/Drum Services 1Sites proposed for NPL. County Polk Escambia Dade Suwanee Alachua Orange Duval Broward Escambia Martin Dade Brevard Duval Broward Hillsborough Putnam Dade Dade Gadsden Hillsborough Dade Broward Duval Escambia Indian River Palm Beach Hillsborough Jackson Hillsborough Volusia Hillsborough Hillsborough Hillsborough Lake Hillsborough Dade Duval Duval Orange RUDDELL—STATUS OF SUPERFUND HRS Score 38.95 56.27 58.04 45.51 45.51 50.92 59.14 59.14 34.18 45.92 58.14 35.57 40.80 51.26 41.43 19.70% 32.37 79.85 34.84 58.15 53.68 40.11 44.00 47.71 42.80 51.18 2Sites where cleanup costs are to be shared by EPA and county government. 3Site has been rescored with new information since original proposal to add to list. 4Score in dispute. 5Remedial actions completed. Lead State EPA EPA EPA EPA State EPA EPA State 235 Person Responsible PRE GOV PRP/GOV?2 PRP GOV PRP GOV PRP/GOV? PRP PRP PRP PRP PRP GOV PRP PRP PRP PRP PRP PRP PRP GOV PRP PRP PRP PRP PRP GOV PRE. PRP GOV PRP PRP GOV GOV PRP/GOV? GOV PRP GOV 236 FLORIDA SCIENTIST [Vol. 52 State—The criteria for sites to become State Action Sites and eligible for state cleanup money are different than for sites that are on the National Priorities List. In the state program, all enforcement steps necessary to get a responsible party to do the work must be exhausted before state funds can be spent. That usually means that the state cleanup program is dealing with sites where the responsible party does not have the resources to do the necessary work. The only exception to that rule is when enforcement has become stalled in the court system to the extent that the delay will cause a threat to public health or the environment. All State Action Sites have the government re- sponsible for carrying out the work (FDER, 1988a). Table 4 lists the State Action Sites as of May 1988. In the state cleanup program, the steps that are followed in the investiga- tion and remediation stages are similar to those in the federal Superfund program. The Remedial Investigation step is referred to as a Contamination Assessment (CA) in the state program and there is no formal Record of Deci- sion as in the federal program. The purpose of each step is the same in both programs; however, there is generally more latitude in the state program to deviate from the prescribed procedures if site conditions warrant (e.g., not all sites will require a formal design before the construction is done). NUMBER OF SITES Za Py COMPLETED BS UNDERWAY Z, Y TOTAL 20 . 4 b dq see KX? Fic. 2. Status of Florida state action sites. (Total number of sites is 23, and remedial actions have been completed at 5 sites. Note: A site may be in more than one category). Figure 2 shows the progress to date in the state funded cleanup program. Progress in the state program compares favorably with that in the federal Superfund program in spite of the fact that the federal program is three years older. Twelve of the 23 State Action Sites (52%) have reached the design or construction steps. There have been 7 IRM’s started or completed on State Action Sites and final remedial actions have been completed at 5 sites. No. 4, 1989] RUDDELL— STATUS OF SUPERFUND 037 TaBLe 4. Florida state action sites (May, 1988). Name County HRS Score Party Responsible Baseline A.G. Industries! Marion 18.40 GOV Belleview Gasoline Marion 27295 GOV Camview Builders! Marion 18.42 GOV Citra Gasoline! Marion Qaas GOV City Chemical-University Blvd. Orange 25.70 GOV Cocoa Beach Brevard D200 GOV Control Products St. Johns 31.76 GOV Edmonds Salvage Yard Dixie ene GOV Emerson Electric Seminole UPR GOV Florida Peach Corp.! Marion 10.65 GOV Jorge Leon! Dade 34.86 GOV K&K Grocery Holmes eae GOV Lake Butler! Union 28.10 GOV Miguel’s Auto Service Dade 31.53 GOV Old 441 Gasoline Marion 26.04 GOV Omni-Vest Landfill Escambia 42.49 GOV Peach Corp. of Florida! Marion 14.25 GOV Silvex St. Johns 13.38 GOV Southern Crop Services Palm Beach 28.56 GOV Sparr Gasoline Marion 17.11 GOV Town & Country Cleaners Clay 10.39 GOV Vroom Polk 31.83 GOV Wacissa Groundwater Jefferson 24.80 GOV 1Remedial actions completed. The final component of the state cleanup program is a category of sites being handled by responsible parties. The steps to be followed are similar to those followed in government-funded cleanup. However, since private funds are being used to investigate and cleanup the sites not all steps are required in responsible party cleanups. In many cases the Feasibility Study is not done, although it is encouraged by state regulators. When a Feasibility Study is not done, the Risk Assesssment required to set cleanup targets is done as part of a Contamination Assessment. In addition, since Designs are not required as separate documents under state enforcement actions, separate information on the number of Designs is not kept by the state. The numbers of Responsible Party Sites in each of the steps of investiga- tion and cleanup are summarized in Figure 3. Progress on non-Superfund responsible party cleanups is slower than for the two government funded programs, with only 43 of the 332 Responsible Party Sites (13%) in the De- sign/Construction step. The low percentage is a reflection of; 1) the large number of sites that are in this category when compared to FDER’s enforce- ment resources, and 2) the inevitable delays that are encountered either in negotiating cleanups with responsible parties, or in working through the court system when negotiations fail. In addition, many of the sites in this category are owned by small businesses. With the costs of investigations and cleanups often running into the millions of dollars for each site, cash flow for 238 FLORIDA SCIENTIST [Vol. 52 NUMBER OF SITES 100 80 60 40 Ju 5050 o, ?, OOO0) 5252505 >, \7 20 OOS eee, << xX 6252505 nee CA ES DES/CON IRM je) ox Fic. 3. Florida responsible-party sites. (Total number of sites is 332; and remedial actions have been completed at 13. (Note: a site may be in more than one category). the responsible party plays a significant factor in the pace of cleanup. As Figure 3 indicates, 57 IRM’s have been done at Responsible Party Sites and final cleanups have been completed at 13 sites. ConcLusions—Superfund is now 8 years old and the state program is 5 years old. Initial progress in both programs was slow. The pace of cleanup was primarily controlled by the difficulties of setting up complex new pro- grams and an unanticipated series of unfamiliar legal, administrative and technical challenges that had to be met. For both programs the initial opti- mistic expectations of how quickly things could be accomplished and what it would cost, fast ran up against reality. Everyone—state and federal officials, consultants, cleanup contractors, past and present owners of sites, and virtu- ally anyone else who became involved—soon realized that they were dealing with something with which they had very little experience. The programs have matured. Procedures have been developed and are being refined. As a result, the rate of progress has picked up considerably in the last two to three years. Sites are now moving through the pipelines of both state and federal cleanup programs. Continued improvement in the pro- grams will continue to increase the pace of cleanup and allow new sites to be addressed more quickly. How much can these programs be speeded up? There are two areas that will limit the rate of cleanup in the near future. The first is the time it takes to resolve legal issues at sites. The second is the availability of experienced pro- fessionals to carry out the programs, both in government and in the private sector, and the availability of laboratory capacity to do all of the work that is necessary. The first area is a fundamental question of a person’s right to due process. The time it takes to guarantee an individual’s rights is up to the No. 4, 1989] RUDDELL— STATUS OF SUPERFUND 239 courts. It is an area that will more than likely continue to be an uncontrolla- ble factor in the time it takes to complete a site cleanup. The second area is a function of the free market and the educational system. It may take some time to resolve. Market forces will dictate the availability of consulting firms working in the area of hazardous waste cleanup and the availability of labo- ratory capacity. It is the establishment of programs in the colleges and univer- sity systems that will determine how soon new groundwater hydrologists, geologists, engineers, chemists, and other graduates with specialized techni- cal skills will be available to staff the consulting firms, man the laboratories, and fill the government jobs that oversee and carry out the cleanup of hazard- ous waste sites. ACKNOWLEDGMENTS— The author acknowledges the help of Joe McGarrity, Susan Gray, Jona- than Rakestraw, Dan DiDomenico and Ruth Gray in the preparation of this paper. LITERATURE CITED GAO (U.S. General Accounting Office). 1987. Superfund— Extent of Nation’s Potential Hazard- ous Waste Problem Still Unknown. GAO/RCED 88-44, Dec. 1987. Washington, D.C. AOp. EPA (U.S. Environmental Protection Agency). 1986. National Priorities List Fact Book. EPA, Office of Emergency and Remedial Response. Washington, D.C. 94p. FDER (Florida Department of Environmental Regulation). 1983. The Sites List—Summary Sta- tus Report. FDER, Bureau of Operations. Tallahassee, FL. n.p. (Florida Department of Environmental Regulation). 1988a. NPL/State Sites Summary Tracking Report— May 1988. FDER, Bureau of Waste Cleanup. Tallahassee, FL. 19p. (Florida Department of Environmental Regulation). 1988b. The Sites List—Summary Status Report. FDER, Bureau of Waste Cleanup. Tallahassee, FL. n.p. Lewis, J., W. STEVENS, S. STUART, AND K. CANANAUGH. 1986. Florida—State of the Environ- ment. Florida Department of Environmental Regulation, Office of Public Information. Wilderness Graphics. Tallahassee, Florida. 48p. Florida Sci. 52(4):230-239. 1989. Accepted: August 26, 1988. Geological Sciences IMPACT OF GROUNDWATER CONTAMINATION ON PUBLIC WATER SUPPLIES J. EDwWarp SINGLEY James M. Montgomery, Consulting Engineers Inc., 1020 N.W. 23rd Avenue, Suite D, Gainesville, FL 32609 Asstract: The presence of a wide variety of organic contaminants in groundwater has re- quired that additional treatment processes be added to the present treatment train in potable water treatment plants. The relevant drinking water quality standards have been summarized as well as the estimated risks some of these contaminants pose to the public. CONTAMINATION of groundwater resources utilized by public water sup- plies has been observed in every state in the country. The problem is particu- larly acute in Florida since 92 percent of the population uses groundwater for consumption and 20 percent use untreated groundwater. Among the many contaminants identified, the most concern has been focused on those of industrial origin, the synthetic organic compounds. These most commonly detected in groundwater in various national and state sur- veys were trichloroethylene, tetrachloroethylene, carbon tetrachloride, TABLE 1. Estimated cancer risks (EPA, 1974) Concentration Resulting in 10-6 Risk Contaminant (ug/l) Vinyl Chloride 0.015 Trichloroethylene 2.6 Carbon Tetrachloride 0.27 1,2-Dichloroethane 0.38 Benzene 1.3 para-Dichlorobenzene 2.08 Toxaphene 0.03 Chlordane 0.0218 Alachlor 0.15 Epichlorohydrin 3.54 Dioxin 22, 104 PAHs 2.8 PCBs 0.008 Phthalates B Acrythamide 0.01 DBCP 0.025 EDB 5 x 10-4 Trihalomethanes ° Heptachlor Epoxide and Heptachlor 0.00065 aDraft potency estimate. bUnder review by the Carcinogen Assessment Group (CAG). cCurrent estimate for systems serving 10,000 or more people, having a concentration of 100 ug/l or more. May be different if CAG risk estimate for chloroform, currently under review, changes. Population Exposed (millions) Cases Per Year 40 1 12 No. 4, 1989] SINGLEY— IMPACT OF GROUNDWATER CONTAMINATION 24] 1,1,1-trichloroethane, 1,2-dichloroethane, vinyl chloride, methylene chlo- ride, benzene, chlorobenzene, dichlorobenzenes, trichlorobenzenes, 1,1- dichloroethylene, cis-1,2-dichloroethylene, and trans-1,2-dichloroethylene in relative descending order as listed. Several hundred others have been identi- fied. Trichloroethylene, a common degreasing solvent, has been found at levels above 30 mg/l and 1,1,1-trichloroethane, also a degreasing solvent, at concentrations above 100 mg/l. There are many inadvertent sources of this contamination ranging from leaking underground storage tanks to industrial waste disposal on-site. There are those resulting from pest control, such as ethylene dibromide (EDB) and dibromochloropropane (DBCP), which were used as nematocides. EDB has shown up in over 100 wells in Florida alone, and DBCP and EDB in wells in California and Hawaii. When many of these are found in the drinking water, they are estimated to present a risk to human health. Some of those that are, or may be, carcino- genic have been evaluated by the Environmental Protection Agency’s (EPA's) Carcinogen Assessment Group, as shown in Table 1. The Safe Drinking Water Act (PL 93-523) requires that these be regulated. Potential or known carcino- gens must have a maximum contaminant level goal (MCLG) set at zero, with the maximum contaminant level (MCL) set as close to zero as feasible (tech- nologically and economically). Florida, in 1984, established regulations for control of some of the more common of the volatile organic contaminants (VOCs) under FAC 17- 22.104(1)(g) as shown in Table 2. Florida also requires monitoring of each source (well) for 118 additional synthetic organic compounds in order to build a database on such contaminants. TABLE 2. Florida primary drinking water maximum contaminant levels for volatile organic contaminants Level, Micrograms Contaminant Per Liter Trichloroethylene 3 Tetrachloroethylene 3 Carbon Tetrachloride 3 Vinyl Chloride 1 1,1,1-Trichloroethane 200 1,2-Dichloroethane 3 Benzene 1 Ethylene Dibromide 0 EPA promulgated MCLGs for eight VOCs in 1985 and MCLs in June 1987. These are shown in Table 3. Although there are some differences in the lists, they will be brought into agreement in the near future with the national values prevailing, since they are based upon more recent risk data. The EPA also proposed MCLGs for several synthetic organic contaminants (SOCs) as shown in Table 4. 242 FLORIDA SCIENTIST [Vol. 52 TABLE 3. EPA volatile organic contaminants, maximum contaminant level goals, and maxi- mum contaminant levels Final Final MCLG: MCL (mg/1) (mg/1) Trichloroethylene 0 0.005 Carbon Tetrachloride 0 0.005 Vinyl Chloride 0 0.002 1,2-Dichloroethane 0 0.005 Benzene 0 0.005 para-Dichlorobenzene 0.075 0.075 1,1-Dichloroethylene 0.007 0.007 1,1,1-Trichloroethane 0.2 On aFinal MCLGs were published November 13, 1985. The MCLG and MCL for para-dichlorobenzene were reproposed at 0 and 0.005 mg/I on April 17, 1987; comment was requested on levels of 0.075 mg/l and 0.075 mg/l, respectively. TABLE 4. EPA proposed maximum contaminant level goals for synthetic organic contaminants (SOC) Existing Proposed SOCs NPDWR: (mg/l) MCLG (mg/l) Acrylamide -- 0 Alachlor — 0 Aldicarb, aldicarb sulfoxide and aldicarb sulfone -- 0.009 Carbofuran —- 0.036 Chlordane aa 0 cis-1,2-Dichloroethylene — 0.07 DBCP — 0 1,2-Dichloropropane _ 0.006 o-Dichlorobenzene — 0.62 2,4-D Om 0.07 EDB — 0 Epichlorohydrin — 0 Ethylbenzene -- 0.68 Heptachlor _ 0 Heptachlor epoxide o- 0 Lindane 0.004 0.0002 Methoxychlor O 0.34 Monochlorobenzene — 0.06 PCBs a 0 Pentachlorophenol os 022 Styrene -— 0.14 Toluene a= 2.0 2,4,5-TP 0.01 0.052 Toxaphene 0.005 0 trans-1,2-Dichloroethylene — 0.07 Xylene — 0.44 «National Primary Drinking Water Regulations No. 4, 1989] SINGLEY— IMPACT OF GROUNDWATER CONTAMINATION 243 Discussion—One immediate impact upon all public water supplies is the expense of the analyses required. This has been estimated to be from $1,000 to $1,500 per well—a not insignificant sum to a small water supply. This becomes important when it is recognized that there are over 2500 systems in Florida serving fewer than 3300 persons. The major impacts upon the public water supply industry will result from the finding of these contaminants in their well supply. Unless another supply is available, additional treatment will be required. This treatment can easily double the cost of water to the consumer under conditions where severe con- tamination has occurred. In those cases where one of the volatile organics is present, the cost of treatment may increase by $0.05 to $0.10 per thousand gallons. The average cost of water to the customer is approximately $1.25 per thousand gallons. Several major supplies in Florida have already had to respond to such a problem. North Miami Beach was the first, when approximately 80 different organic compounds were found in their new East Drive Wellfield, located down gradient from an industrial area. by using activated carbon, they were able to control the levels and keep the wellfield in service for several years. It, and the Sunny Isles Water Treatment Plant it served, were later removed from service and water was purchased. The investment in wells and plants was several million dollars. Treatment technology has advanced to the point that any contaminant can be removed but the cost varies dramatically. The American public has said that it is willing to pay more for better water and it will soon have to do sO. LITERATURE CITED FLorRIDA DEPARTMENT OF ENVIRONMENTAL REGULATION. 1984. FAC 17-22, Public Drinking Water Systems— Volatile Organics. ENVIRONMENTAL PROTECTION AGENCY. 1987. Unfinished Business: A Comparative Assessment of Environmental Problems. Appendix I: Report of the Cancer Risk Work Group. Appendix II: Non-Cancer Work Group. . 1978. National Organics Monitoring Survey (NOMS). Technical Support Division, Office of Drinking Water. NATIONAL REVISED PRIMARY DRINKING WATER REGULATIONS. 1985. Volatile Synthetic Organic Chemicals in Drinking Water. Federal Register, 50:46880. Syms, J.M., T.A. Betrars, J.K. CARSwELL, G.G. Rosecx, K.L. Kropp, D.R. SEEGER, C.J. SLocuM, B.L. SmirH, AND A.A. STEVENS. 1975. National Organics Reconnaissance Survey for Halogenated Organics. J. Am. Wat. Works Ass. 67:621-635. Florida Sci. 52(4):240-243. 1989 Accepted: February 1, 1989. Computer Sciences THE NEURAL DERIVATIVE, OCOS AND MOTION DETECTION Davip LAWSON Department of Mathematics and Computer Sciences, Stetson University, DeLand, Florida 32724 ABsTRACT: This paper presents a computational neural mechanism, a neural derivative, and confirms through the use of Schiller’s (1982) experiments that this theoretical structure behaves in exactly the same manner as do motion detectors observed in the rhesus monkey. This strongly suggests that the neural derivative presented is a fundamental functional unit of the early visual system. These experiments indicate that the on-center off-surround architecture can operate as a neural derivative, and can thus be one of the motion detection mechanisms employed in vivo. The Veto mechanism, another motion detector, may or may not be a neural derivative, depend- ing on the structure of its dendritic trees. NeEuRAL modelers often work at two distinct levels. One is the develop- ment of computational neural elements capable of specific tasks. The other is the attempt to discover the structure and function of anatomically identifi- able groups of cells. Both elements can be found in the following paper which presents an ideal element, a neural derivative, and then demonstrates that this derivative is part of the early vision system. The neural derivative dis- played is of special interest because it reacts in accordance with recent experi- ments involving motion detection, recordings made in the lateral geniculate nucleus (LGN) and the striate cortex of the rhesus monkey. Motion detection plays a central role in many biological systems, and many animals have special purpose motion detection mechanisms. The frog has detection mechanisms capable of responding selectively to small dark objects moving in its visual field (Lettvin, Maturana, McCulloch and Pitts, 1959). The housefly can track moving objects even when the object and its background are indistinguishable in the absence of motion (Reichardt and Poggio, 1980). Some species, including the rabbit can perform rudimentary motion analysis at the retinal level. There are other examples of this nature, examples of motion detection mechanisms that are clearly associated with special behavioral needs of the organism. Thus, neural mechanisms for motion detection may be as varied as the behavior of the animals that employ these mechanisms. As an example, direc- tion sensitive cells have been found in the retina of the grey squirrel (Cooper and Robson, 1966), the turtle (Jenson and Devoe, 1983), and the rabbit (Bar- low and Levick, 1965). Evidently, each of these animals can safely ignore objects or animals that are moving away. On the other hand, the neural mechanisms may not be so varied. Perhaps, as one might expect of a function of the central nervous system, the same mechanisms are widespread, essentially identical, and are variations on a central theme. Perhaps we will find special adaptations for special purposes No. 4, 1989] LAWSON— NEURAL DERIVATIVE 945 in some animals and none in others. No direction sensitive motion detectors have yet been found in the monkey’s retina (Lennie, 1980). Primates, how- ever, tend to depend on their ability to organize information within the brain and react to it, rather than relying on special purpose neural anatomy. It is almost as if the primate solves in software what other animals solve by em- ploying special purpose hardware. Our major point is that the neural derivative which we will introduce below is capable of motion detection, and that neurons that react in the same fashion as the neural derivative can be found in the early visual system. Spe- cifically, the neural derivative is an implementation of the gradient method of motion detection. This neural mechanism is also an odd (and rather inter- esting) mathematical entity—a geometic, or graph theoretic, definition of the derivative. Motion Detection—Two standard theoretical approaches to motion de- tection are the Veto mechanism and the gradient method (Ullman, 1983). I shall discuss both, show the strengths and weakness of each, and demonstrate that our proposed neural derivative is an implementation of the gradient method. I will demonstrate that the neural derivative is preferable to the Veto mechanism if one wishes to detect motion over a wide range of velocities and directions. Delayed Comparison and the Spaghetti Nightmare— First, I shall present an early scheme proposed by Barlow and Levick (1965) (Fig. 1). The delayed comparison method is a unidirectional computational scheme which will detect motion from right to left (or left to right) but not vice-versa. This is in keeping with experimental results for both cats (Hubel and Wiesel, 1962) and monkeys (Hubel and Weisel, 1968), which show that there are neurons which respond to stimuli in one direction but not the other. The delayed-comparison mechanism works in the following fashion (Fig. 1): Neuron C will fire if it receives inputs from neurons A and B at the same time. If an object moves from left to right, neuron A will fire prior to neuron B. The delay will retard the signal from B. Thus, the signal from A will arrive before the signal from B, and C will not fire. If an object moves from right to left B will fire, then A. The delay will cause the signals from A and B to both arrive at C at approximately the same time. C will then fire. There is a fundamental flaw in the delayed comparison mechanism. A stationary object which can fire both A and B will fire C as well. To avoid this problem, Barlow and Levick proposed the Veto mechanism (Fig. 2). Neuron B inhibits neuron C (Fig. 2). Thus if B fires first, then C will not fire. In other words, input to B will veto input to A. In addition if B fires and continues to fire, then C will not fire. Thus, a stationary object over A and B cannot fire C. A great deal of progress is being made toward understanding the chemi- cal, electrical and structural underpinings of the Veto mechanism (Torre and Poggio, 1978; Jensen and DeVoe, 1983; Poggio, 1983; Poggio and Koch, 1987). 246 FLORIDA SCIENTIST [Vol. 52 (<--) A B <-- CELL BODY <--- AXON DELAY <--- SYNAPTIC KNOB DELAYED COMPARISON Fic. 1. Motion detection from right to left. C will fire if the signals from A & B arrive at the same time. (Se) DELAY THE VETO MECHANISM Fic. 2. Motion detection from left to right. The darkened synaptic knob is inhibitory. If an object moves from B to A, then the signal from B will cancel the signal from A and C will not fire. No. 4, 1989] LAWSON— NEURAL DERIVATIVE 247 THE GRADIENT METHOD Fic. 3. This is a light edge moving to the left. As a light spot passes in front of P the intensity I will rise. The gradient is the slope of the curve I, the rate at which I rises. ot THE NEURAL DERIVATIVE Fic. 4. B(t) is the rate at which neuron B fires at time t. Neuron B fires at time t at the same rate that neuron A fired the time step before. This means B(t) = A(t-1). In addition, B is inhibitory and thus C(t) = A(t-1)-B(t-1). Thus, C(t) = A(t-1)-A(t-2) =delta A=delta I, the change in inten- sity. C(t) is therefore the time derivative of A. 248 FLORIDA SCIENTIST [Vol. 52 A problem exists with the Veto mechanism, the Spaghetti Nightmare. The Veto mechanism is a special purpose device. It senses motion in only one direction. If this information is of value to the organism then the Veto mecha- nism is a marvelous device, for it reduces the two step process of discovering motion and determining the direction to just one step. If, however, one has use for motion information in many directions, using the veto mechanism would require replication of this arrangement in all directions. For an electri- cal engineer, interested in building a sensing device, this becomes a spaghetti nightmare of wires running needlessly in all directions. The Gradiant Method and the Neural Derivative—The neural derivative will solve this problem. But first we must discuss the gradient method of motion detection (Fig. 3). If P is a point on the retina, and if the intensity I increases significantly at point P, then the gradient (the change in I) is increasing. As a light object moves across P the intensity at P will increase. If P is connected to neuron C, then C will fire. If the intensity I increases rapidly, then C will fire at a faster rate. In other words, C responds to the slope of I, firing at a faster rate for a steeper positive slope. In addition, if the intensity I does not change then C will not fire at all. A geometric derivative—The operation of the neural derivative is a result of its geometry, the connections between A, B, and C. The neural derivative (Fig. 4) is an implementation of the gradient method. It is a derivative because the output of the neural derivative is a measure of the rate of change of the intensity of the light that it receives. As an example, the neural derivative will not fire if the intensity I remains constant. This is because the rate of firing at neuron A is matched by the rate at B. The two inputs then meet at C, with the inhibitory input from B cancel- ing the excitatory input from A. This, if the input does not change the output will be zero. This is exactly the same response given by the standard mathe- matically defined derivative. In addition, if the intensity I increases, then the input from A will be larger than the input from B, and C, the neural deriva- tive’s output, will fire (Fig. 5). Further, if the intensity increases more quickly, then the output of C will be greater. In other words, the output C of the neural derivative tracks the change of the intensity of the light which it receives (e.g., the output measures the rate at which the change in intensity occurs. If the intensity changes quickly then the output C will be high, if the intensity changes gradually, then the output of C will be low). There is one flaw. The neural derivative proposed above will not work correctly for a negative slope. If the intensity I is falling C will merely refuse to fire. This problem is not difficult to solve. We mention two different meth- ods. First, if C has a resting firing rate which is greater than zero, then the offset from the resting rate will be the derivative. In this case, if the slope of I is negative then B will merely suppress the firing of C below the resting rate. A second method takes advantage of the ON and OFF channels of early No. 4, 1989] LAWSON— NEURAL DERIVATIVE 249 THE OPERATION OF THE NEURAL DERIVATIVE Fic. 5. Att=0 A fires at the rate of 4 times a second. At t=1 A & B both fire 4 times a second, and C doesn’t fire. At t=2 A fires 10 times a second, B fires 4 times a second, and C fires 6 times a second. (eg. C fires at the rate of A - B, which is 10 - 4). vision. If A is an ON cell the neural derivative will measure positive slopes, if A is an OFF cell, then changes in the firing rate of A will correspond to negative slopes. This is because an OFF cell has a resting rate which increases as the illumination I decreases. As a result, in the latter case, the output of C would correspond to the absolute value of the slope of I, whenever the slope was negative. Neurons which exhibit this second method have been found in the early visual system of the rhesus monkey (Schiller, 1982). Properties of the neural derivative—A light edge is the leading edge of a light figure moving across a dark background. Similarly, a dark edge is the leading edge of a dark figure moving across a light background. Fact 1. The ON-center neural derivative presented above will respond to a moving light edge only. Fact 2. An OFF-center model of the neural derivative will respond to a moving dark edge only. Neural recordings of neural derivative activity in the rhesus monkey— Neural recordings made in the lateral geniculate nuclei (LGN) of the rhesus monkey showed that retinal ganglion cells react in exactly the same fashion as does the neural derivative above. (Schiller, 1982) These recordings indicate the presence of a neural derivative composed of ON cells. Schiller used DL-2-amino-4-phosphonobutyric acid (APB) to reversibly block the ON responses in the retina. ON bipolar, ON amacrine and ON ganglion cells become unresponsive. Receptors, horizontal cells, OFF bipo- 250 FLORIDA SCIENTIST [Vol. 52 lars, OFF amacrine, and OFF ganglion cells were not affected. Recordings were then made in the LGN while the retinal ON responses were reversibly blocked. Schiller discovered that the response of ON cells was surpressed, while OFF cells were unaffected by APB infusion. The same experiment was repeated with the recordings of complex cells in the striate cortex. Because ON cells can be surpressed, Schiller was able to determine whether or not cells in the LGN and striate cortex would react in the same manner as the neural derivative introduced above. Schiller presented the monkey with a moving light bar. Under normal conditions the cell responded to the light edge of the bar. After retinal infu- sion of APB the cell no longer responded to the light edge. In other words the cell reacted as though it was an ON cell implementation of a neural deriva- tive. The same cell responded to a dark edge. Its response was what one would have suspected if the neuron was meant to be the absolute value of the derivative. Schiller found cells in the striate cortex which reacted in a complex man- ner of a different sort. One cell was found which would respond to the mov- ing light edge and not a dark edge prior to APB infusion, but would reverse its response following APB infusion. In the later case, the cell would respond to a moving dark edge, but not to a moving light edge. K/ DENDRITIC TREE Fic. 6. The Veto mechanism with overlapping dendritic trees. No. 4, 1989] LAWSON— NEURAL DERIVATIVE Dial Thus, Schiller’s work shows that the ON and OFF cell implementations of the neural derivative are distinct entities at the level of the LGN, but within the striate cortex they are subunits of much more complex cell assemblies. The Veto mechanism and the neural derivative—a comparison—Therte is a striking similarity between the structure of the Veto mechanism (Fig. 2.) and the neural derivative (Fig. 4). If the dendritic trees of neurons A and B of the Veto mechanism overlap significantly (Fig. 6.), then we could represent the network presented in Figure 6 with the functionally equivalent network of Figure 7. Notice that the essential feature of the neural derivative, represented alge- bracically by the equation A(t) = B(t-1), is satisfied by figures 6 and 7. Thus, the Veto mechanism with largely overlapping dendritic trees is functionally equivalent to a neural derivative. One can say that the Veto mechanism is simply a neural derivative, but with disjoint dendritic trees. In other words, depending on the nature of the dendritic trees, the Veto mechanism can function as a either a directional motion detector or as a neural derivative. This could explain the evolutionary development of the Veto mechanism from the neural derivative (or vice- versa). DELAY Fic. 7. The Veto mechanism with disjoint neuron C in place of Neurons A and B. 252 FLORIDA SCIENTIST [Vol. 52 Discuss1on— Animals respond to change in their environment. The deriv- ative is a measure of change. Thus, it makes sense that a derivative would have a fundamental place within the early vision system, and Schiller’s (1982) experiment demonstrates that the neural derivative is a part of that system. The development of theoretical neural structures can help to guide our search for order within the brain. Thus, such structures receive a great deal of attention. Examples abound. It has long been recognized that the on-center off-surround architecture can report intensity changes and thus determine the edges of an object. Practitioners of signal processing clearly recognize that this is an implementation of the derivative. ON and OFF channels working in conjunction can even produce the Laplacian, a second-order differential op- erator (Marr and Hildreth, 1980; Marr and Ullman, 1981). Theoretical constructions can guide our intuition and our investigations. The identification of such a unit may lead to other discoveries. As an exam- ple, Lennie (1980) has conjectured that the Veto mechanism proposed by Barlow and Levick (1965) during their investigation of the rabbit retina is an adaptation of the on-center off-surround architecture. The evolutionary trail of the development of motion detection mechanisms has yet to be forged, but it may be that the neural derivative is the unit of motion detection from which other more specialized and sophisticated mechanisms have sprung. The theoretical neural derivative we have presented could alter our con- ception of the early vision system. Edge detection is not a derivative, but instead an application of the derivative. Motion detection is another applica- tion of the derivative. Both are fundamental functions of the nervous system, but of a different order than the neural derivative. Viewed in this fashion it even seems possible that edge detection and then shape formation could be an emergent property arising from early motion detection mechanisms. ACKNOWLEDGMENTS— This research was supported by the Air Force Office of Scientific Re- search/AFSC, United States Air Force, under Contract No. F49620-85-C-0013, and Contract No. F49620-85-C-0013/SB5851-0360. The United States Government is authorized to reproduce and distribute reprints for governmental purpose notwithstanding any copyright notation hereon. No. 4, 1989] LAWSON— NEURAL DERIVATIVE 253 LITERATURE CITED Bartow, H. B. anp W. R. Levick. 1965. The Mechanism of Directionally Selective Units in the Rabbit’s Retina. J. Physiol. 178:477-504. Cooper, F. G. AND J. G. Rosson. 1966. Directional selective movement detectors in the retina of the squirrel. J. Physiol. 186:116P-117P. Huse, D. H. anp T. N. Wiesev. 1962. Receptive fields, binocular interaction and functional architecture in the cat’s visual cortex. J. Physiol. 160:105-154. . 1968. Receptive fields and functional architecture of monkey striate cortex. J. Phy- siol. 195:215-243. JENSEN, R. J. AND R. D. DeVoe. 1983. Comparisons of Directionally Selective with other Gan- glion Cells of the Turtle Retina: Intercellular Recording and Staining. J. Comp. Neurol- ogy 217:271-287. LENNIE, P. 1980. Parallel Visual Pathways: A Review. Vision Res. 20:561-594. Lettvin, J. Y., H. Maturana, W. S. McCu.ttocu, anv W. H. Pitts. 1959. What the frogs eye tells the frogs brain. Proc. Inst. Radio Engineers 47: 1940-1951. Marr, D. anp E. Hivpretu. 1980. Theory of edge detection. Proc. Royal Soc. London B. 207:187-217. AND S. ULLMAN. 1981. Directional selectivity and its use in early visual processing. Proc. Royal Soc. London B. 211:151-180. Poccio, T. 1983. Visual Algorithms. Pp. 128-175. In: Brappicx, O. J. AND A. J. SLEIGH (eds.) Physical and biological processing of images. Springer-Verlag. Berlin. AND C. Kocn. 1987. Synapses That Compute Motion. Scientific American 256, (5):46- 52. REICHARDT, W. AND T. Poccio. 1980. Biological Cybernetics 35:81-100. ScHILLER, P. H. 1982. Central connections of the retinal ON and OFF pathways. Nature. 297:580-583. Torre, V. AND T. Poccio. 1978. A synaptic mechanism possibly underlying directional selectivity to motion. Proc. Royal Soc. London B. 202:409-416. ULLMAN, S. 1983. The measurement of visual motion. Trends in Neurosciences 6: 177-179. Florida Sci. 52(4):244-253. 1989. Accepted: November 29, 1988. Biological Sciences ARTHROPODS ENDEMIC TO FLORIDA SCRUB Mark Dryrup Archbold Biological Station, P.O. Box 2057, Lake Placid, FL 33852 Asstract: Florida sand pine scrub is a community type found on well-drained ridges and dunes. Some of these uplands escaped the periodic flooding of the Florida Peninsula. Organisms of Florida scrub are adapted to sterile sand, drought, and fire. Some species appear to be relicts. An annotated list identified 46 species of scrub endemic arthropods. Species are concentrated on the central ridges, especially the Lakes Wales Ridge. There appear to be cases of divergent evolu- tion on different scrub islands. The high proportion of flightless species in the list may not reflect adaptation for island life. Detailed studies of biogeography of scrub endemic arthropods may be thwarted by rapid eradication of many scrub sites. A LakcE proportion of the species unique to the Florida peninsula is con- centrated on a series of islands far from the tropical Florida keys. These islands are ancient inland sand ridges and dune systems, that served as biotic refuges during Miocene, Pliocene or Pleistocene inundations (Neill 1957). These islands were also sites for evolutionary divergence of isolated popula- tions. After the retreat of the sea from Florida’s lowlands, the ridges and dunes retained some of their insular character because their deep, sterile, excessively drained sands are unsuitable for many lowland species, and the lowlands are likewise unsuitable for many ridge species. Florida scrub is a characteristic community of these ridges, and is easily identified by its vegeta- tion of low, evergreen, sclerophyllous oaks, often with sand pine (Pinus clausa) or Florida rosemary (Ceratiola ericoides). Patches of bare sand are frequent, and the low sparse nature of the vegetation is maintained in part by fire (Abrahamson et al. 1984). Scrubs occur along both coasts of Florida, in the extensive Marion Uplands and several smaller ridge areas in the north- central part of the Peninsula, and along the Lake Wales Ridge, which extends south from Lake Apopka about 100 miles, ending in southern Highlands County (Figs. 1, 2). The only general discussion of arthropods of Florida scrub is that of Hub- bell (1961), who mentioned a number of genera that include scrub endemics. Hubbell also noted the western affinities of some species, and hypothesized that the patterns of variation seen in certain species were the result of rapid evolutionary change in small insular populations. In a much earlier paper (1932), Hubbell analyzed the biogeography of a group of short-winged grass- hoppers inhabiting Florida’s xeric upland habitats. He concluded that this group may have entered during the Miocene, spread from central Florida, and radiated as populations became isolated by changes in climate and sea level, persisting to the present in the most stable islands of xeric habitat. In the 56 years since Hubbell pointed out that Florida has its own archipelago for the study of evolutionary patterns, there have been few publications on No. 4, 1989] DEYRUP— FLORIDA SCRUB ARTHROPODS Zp Fic. 1. Location of Florida scrub (redrawn from Davis, 1967) 256 FLORIDA SCIENTIST [Vol. 52 1. Brooksville Ridge 2. Mount Dora Ridge 3. Lake Wales Ridge 4. Crescent City Ridge 5. Deland Ridge 6. Atiantic Coastal Ridge 7. Lakeland Ridge 8. Bombing Range Ridge Fic. 2. Location of ridges (modified from White, 1970) No. 4, 1989] DEYRUP— FLORIDA SCRUB ARTHROPODS 257 the biogeography of scrub arthropods. Endemism in scrub arthropods has been mentioned with respect to scarab beetles (Woodruff 1973, 1982), spiders (Edwards 1982), and ants (Deyrup and Trager 1986). The term “scrub en- demic” is applied here to species that apparently evolved in, and are re- stricted to, scrub habitat. This does not mean these species were always re- stricted to the exact geographic ranges they presently occupy. The purpose of this paper is to summarize published and unpublished research on arthropods endemic to Florida scrub, with a view to stimulating more research by entomological colleagues. Current information is fragmen- tary, but nonetheless useful in a variety of ways. The distribution of scrub arthropods is of interest as it sheds further light on the biogeography of Flor- ida scrub; this initial list of invertebrates already includes more species than the number of scrub endemic plants, and thus provides many new data points to be used in analyses of scrub biogeography. Identification of scrub arthro- pods is also of interest as it relates to the biogeography and systematics of arthropods groups. An analysis of the characteristics of scrub arthropods should give some new ideas of general adaptations of scrub endemics. The presence of endemic scrub arthropods may even be useful in identifying patches of scrub worthy of preservation. MeEtTHops— Distribution records were obtained from published literature and from personal collecting records. Collections of scrub arthropods are maintained at the Archbold Biological Station (ABS) as a component of the long-term ABS projects on the organisms of Florida scrub. During September, 1987 through April, 1988 a series of pitfall traps were set in scrubs along the Lake Wales Ridge. These traps were at the Hendrie Ranch (Venus, Highlands Co.), the former YMCA Camp Florida (4 km s. Lake Placid, Highlands Co.), Virginia Avenue in Highlands Park Estates (Lake Placid, Highlands Co.), Flamingo Villas (Sebring, Highlands Co.), Lake Arbuckle Management Area (Frostproof, Polk Co.), Flaming Arrow Scout Camp (Lake Wales, Polk Co.). This work was supported by a grant (GF 184-010) to S. Christman from the Nongame Wildlife Program. The ABS (9 kms. Lake Placid, Highlands Co.) also has an independent survey of scrub arthropods, concentrating on species found in scrub habitats of the ABS. Several problems beset any large-scale arthropod survey of a particular habitat. One problem is the definition of the habitat. Florida scrub shares many species with Florida sandhill habitat (a habitat dominated by scattered pines and a thick ground cover of wiregrass) (Abrahamson et al. 1984), especially sandhills that have been protected from fire. Where the water table is higher there may be an intermediate habitat classified as scrubby flatwoods (Abrahamson et al. 1984). I have tried to confine the list of species to those restricted to scrub, but some of these species also occur to some extent in sandhill or scrubby flatwood habitats. A second problem is the very large number of arthropod species involved, and a third is identification of arthropods. Since many of the species are restricted to small areas, some of them have not been previously collected, and there is a relatively high percentage of undescribed species. Finally, the distribution of many Florida arthropods is poorly known. Not only is our knowledge of the distribution of many scrub endemics obviously incomplete, but some species known only from a single scrub site may not be scrub endemics. The alleculid beetle Onychomira floridensis Campbell, for example, represents a monotypic genus known only from the ABS, and is likely to be scrub endemic, but all specimens were taken in light traps, and may have originated in some habitat other than the scrub. I am conservative in designation of scrub endemics, selecting species from taxonomic groups that have been adequately surveyed, or species whose known ecological requirements suggest that they are probably restricted to scrub. Geographic references are taken from White (1970) and Davis (1967). 258 FLORIDA SCIENTIST [Vol. 52 ANNOTATED List oF SpEcIES—Class Diplopoda, Order Spirobolida, Spriobolidae—Floridobolus penneri Causey. A large, dark gray millipede, distinguishing characters provided by Causey (1957). Known only from southern Lake Wales Ridge from Lake Placid to Venus (Highlands Co.). Found under sand at base of plants, active on surface of sand at night. Read- ily taken in pitfall traps in Highlands Co., but not taken in similar traps in Polk Co. Class Arachnida, Order Aranae, Theridiidae—Latrodectus bishopi Kas- ton, Red Widow Spider. Red legs and cephalothorax distinguish this species from other widow spiders in Florida (color illustration in McCrone and Stone, 1965). Known from the southern Lake Wales Ridge (Highlands Co.), Marion Uplands (Putnam, Marion, Lake, Orange Co.) and Atlantic Coastal Ridge (Martin, Palm Beach Co.) (Edwards 1982). This species is probabiy more widespread in Florida scrubs than presently recognized, as it has marked fluctuations in abundance and is difficult to find in certain years. The webs are usually in palmetto leaves, especially the leaves of scrub pal- metto (Sabal etonia) which always remain partially folded, providing an en- closed site for the web. This has been recognized as a scrub endemic since 1964 (McCrone and Levi). Salticidae—Phidippus xeros Edwards. A large jumping spider with a di- agnostic color pattern, illustrated by Edwards (1978, 1982). Known from scrubs in the Marion Uplands (Marion, Alachua, Putnam, Lake, Orange Co.), from a Gulf Coast scrub in Escambia Co. (Edwards 1982), and from the southern Lake Wales Ridge (Highlands Co.). Gnaphosidae—Zelotes ocala Platnick and Shadab. A fast-moving black spider in leaf litter scrub. This species is distinguished primarily by genitalic characters, illustrated by Platnick and Shadab in the revision of the genus Zelotes (1983). This revision covers the distribution of Zelotes in considerable detail, and it seems likely that the two species collected only in scrub are confined to that habitat. Zelotes species are likely to depend on a particular type of ground litter. Z. ocala is known from the Marion Uplands (Marion, Alachua, Putnam Co.) and from the Southern Lake Wales Ridge (Highlands Co.). Gnapshosidae—Zelotes florodes Platnick and Shadab. Similar to the pre- ceding species; diagnostic characters provided by Platnick and Shadab (1983). Known from the Lake Wales Ridge (Polk, Highlands Co.). Lycosidae— Geolycosa xera McCrone, McCrone’s Burrowing Wolf Spi- der. This species can be distinguished from sympatric species by the whitish pubescence covering the upper surface of the body; additional characters are provided by McCrone (1963). There are two subspecies. G. x. xera is known from the Marion Uplands (Lake, Orange, Seminole, Volusia Co.) and most of the Lake Wales Ridge (Polk, Highlands Co.) (McCrone 1963). G. x. archboldi occurs on the southern Lake Wales Ridge (Highlands Co.) (McCrone 1963). The habitat is usually bare white sand. The burrows remain open and are easily found. No. 4, 1989] DEYRUP—FLORIDA SCRUB ARTHROPODS 259 Lycosidae—Lycosa ceratiola Gertsch and Wallace. This species is distin- guished from the similar species L. lenta Hentz by reduced spines beneath the first and second tibiae, and by genitalic characters of the female (Gertsch and Wallace 1935) and the male (Gertsch and Wallace 1937). It is known from scrub areas in Mount Dora Ridge (Lake Co.), the Atlantic Coast Ridge (Mar- tin Co., Palm Beach Co.) (Gertsch and Wallace 1937), and the Lake Wales Ridge (Highlands Co.). Lycosidae—Lycosa pseudoceratiola Wallace. This species is distinguished primarily by genitalic characters (Wallace 1982). It appears to be confined to the Atlantic Coast Ridge (Dade, Palm Beach, Martin, Indian River Co.); it is not a beach species (Wallace 1982). Lycosidae—Lycosa ericeticola Wallace, Rosemary Wolf Spider. This is another species recognized by distinctive male genitalia (Wallace 1982). It is known from rosemary scrubs in the Marion Uplands (Putnam Co.) (Wallace 1982, Edwards, 1982, Reiskind 1987). Lycosidae—Sosippus placidus Brady, Lake Placid Funnel Wolf Spider. This large, web-building wolf spider is distinguished by its apricot-colored ventral pubescence (Brady 1972). It is known from the southern Lake Wales Ridge, from Lake Placid to Venus (Highlands Co.). Like the sympatric S. floridanus Simon, which was studied by Brach (1976), the female S. placidus builds a large web where the young remain until about a third grown, feed- ing on prey caught by their mother. Class Insecta, Order Orthoptera, Polyphagidae—Arenivaga floridensis Caudell, Florida Sand Cockroach. The body of this distinctive species is al- most circular in outline when viewed from above, and is white with black flecks. Known from coastal scrub in Pinellas Co., from the Lakeland Ridge (Polk Co.) (Blatchley 1920), and from the Lake Wales Ridge (Polk, Highlands Co.). This species is an isolated eastern representative of a genus of south- western sand roaches. The adult male is winged and occasionally comes to lights; the female is wingless and remains under the sand. Acrididae—Schistocerca ceratiola Hubbell and Walker, Rosemary Bird Grasshopper. S. ceratiola is the smallest (24-36 mm) and slenderest of eastern bird grasshoppers, and is also distinguished by its host plant, as it is the only bird grasshopper on Ceratiola ericoides (Hubbell and Walker 1928). It is known from the Orlando Ridge (Orange Co.), the Mount Dora Ridge (Lake, Orange Co.) (Hubbell and Walker 1928), the Atlantic Coastal Ridge (Martin Co.), and southern Lake Wales Ridge (Highlands Co.). This species is exclu- sively nocturnal. Acrididae—Melanoplus insignis Hubbell. This species is recognized by the characteristics of the male genitalia (Hubbell 1932). It appears to be closely related to M. forcipatus, and may actually be a distinctive isolated population of that species (Hubbell 1932). M. insignis is known from the Atlantic Coastal Ridge (Palm Beach Co.) (Hubbell 1932). This species be- longs to a group of small flightless grasshoppers that are usually restricted to xeric upland habitats. 260 FLORIDA SCIENTIST [Vol. 52 Acridadae—Melanoplus forcipatus Hubbell. The most distinctive fea- tures of this species are characteristics of the male genitalia. It is known from the Orlando and Mount Dora Ridges (Orange Co.) and the southern Lake Wales Ridge (Highlands Co.) (Hubbell 1932). The habitat descriptions pro- vided by Hubbell, as well as our own collecting, suggest that this species is primarily a scrub inhabitant, although it also occurs in sandhill areas. The elimination or modification of sandhill habitats through much of the range of this species makes its habitat preferences difficult to analyze. Acrididae— Melanoplus tequestae Hubbell. The comments on the identi- fication of M. forcipatus also apply to this sympatric species. It is known from the Orlando Ridge (Orange, Seminole Co.), Mount Dora Ridge (Orange Co.), and the southern Lake Wales Ridge (Highlands Co.) (Hubbell 1932). Class Insecta, Order Mallophaga, Philopteridae—Bruelia deficiens (Piaget). This bird louse is host specific on the scrub jay (Aphelocoma caeru- lescens), and therefore indirectly confined to scrub. The relationship between this louse and the louse found on western populations of the scrub jay is currently under investigation (Fitzpatrick 1988). Class Insecta, Order Coleoptera, Cicindelidae—Cicindela highlandensis Choate. This small black tiger beetle is distinguished from similar species by its smooth elytra and hairless ventral surface (Choate 1984). It is known from two small scrub areas (both recently destroyed) on the southern Lake Wales Ridge (Highlands Co.). Tiger beetles have been intensively collected in Flor- ida, and it is not likely that this species has an extensive range that has been overlooked. Cicindelidae—Cicindela scabrosa Schaupp. This small black species is distinguished from similar species by its heavily punctate elytra and hairy ventral area (Choate 1984). It is widely distributed in Florida scrubs, though the only records on hand are from the Crescent City Ridge (Putnam Co.) (Choate 1984), the Atlantic Coastal Ridge (Broward Co.) and the Lake Wales Ridge (Highlands Co.). Like other members of its genus, C. scabrosa is seen running rapidly over the ground in open areas, readily taking flight when alarmed. Histeridae— Terapus sp. An undescribed species of Terapus inhabits the nests of the ant Pheidole morrisi in scrub at the ABS. This beetle is conspicu- ous in the nests of its host, and has also been taken in Malaise traps and window traps. It seems likely, therefore, that this species is not widespread, or it would have been collected elsewhere. Terapus is a southwestern genus (Hinton 1934), and the Florida species may be another example of an isolated Florida scrub representative of an otherwise southwestern group. Scarabaeidae—Note on the genera Phyllophaga, Anomala, and Serica: Larval scarabs of these genera live in soil and presumably feed on the roots of living plants. Such species are not only sensitive to the species of available plants, but are also sensitive to the soil type and the depth of the water table. Members of these genera which are common in scrub habitats and unknown No. 4, 1989] DEYRUP—FLORIDA SCRUB ARTHROPODS 261 elsewhere may be provisionally considered scrub endemics, even though the host plant of the larvae are unknown. Scarabaeidae—Serica frosti Dawson. This species is recognized by its iri- descent sheen and by characters of the male genitalia (Dawson 1967). It is very common at the ABS, unknown from elsewhere. Scarabaeidae—Anomala eximia Potts. This species is distinguished from sympatric species by the lack of a tooth on the outer edge of the protibia, and by the irregularity of the rows of elytral punctures (Potts 1976). This species is common in scrub habitats at the ABS, but is known from no other site. It is a diurnal species and is readily taken in Malaise traps. Scarabaeidae—Phyllophaga elizoria Saylor. This species is distinguished from other June beetles by its unusual hairiness and by the shape of the male genitalia. It is known from the Atlantic Coastal Ridge (Indian River Co.) and the Lake Wales Ridge (Highlands Co.) (Woodruff 1982). Scarabaeidae—Phyllophaga elongata (Linell). This species is best recog- nized by the shape of the male genitalia. It appears to be a “relictual endemic of the central Florida sand ridges” (Woodruff 1982). It is known from the Lake Wales Ridge (Polk, Highlands Co.), the Sumter Uplands (Marion Co.), Polk Uplands (Hillsborough Co.), and unspecified sites in Lake, Levy, and Marion Co. (Woodruff 1982). Scarabaeidae— Phyllophaga panorpa Sanderson. This species is best rec- ognized by the shape of the male genitalia. It is known from the Lake Wales Ridge (Polk, Highlands Co.) (Woodruff 1982). Scarabaeidae—Phyllophaga okeechobeea Robinson. This is an unusual small, hairy, diurnal species. It is known from the Lake Wales Ridge (High- lands Co.) and from Okeechobee (Woodruff 1982). Woodruff (1982) con- siders that this species is an endemic of the south-central sand ridges, which agrees with our observations. This would seem to cast doubt on the accuracy of the type locality of Okeechobee. Scarabaeidae— Trigonopeltastes floridana (Casey). This small, brightly colored diurnal species has a conspicuous yellow “V” on the pronotum. It is known from the Lake Wales Ridge (Highlands Co.), the Orlando Ridge (Orange Co.), the Atlantic Coastal Ridge (Indian River Co.), the Mount Dora Ridge (Marion Co.) and the Northern Highlands (Alachua Co.) (Wood- ruff 1960). Adults are often found on flowers of scrub palmetto (Sabal eto- nia), less frequently on the earlier blooming saw palmetto (Serenoa repens). Scarabaeidae—Mycotrupes pedester Howden. This is the only flightless geotrupine scarab in its range. Specimens have been collected from islands of scrub in DeSoto, Charlotte, and Lee counties (Woodruff 1973). This is the only scrub arthropod known only from southwestern Florida scrub areas. Scarabaeidae—Peltotrupes youngi Howden. This species may be only subspecifically distinct from the more widespread species Peltotrupes profun- dus Howden (Woodruff 1973). It is known from scrub areas of the Mount Dora Ridge (Putnam, Marion Co.) (Woodruff 1973). 262 FLORIDA SCIENTIST [Vol. 52 Scarabaeidae—Onthophagus aciculatulus Blatchley. Woodruff (1973) provides characters for distinguishing this species from similar small black species of Onthophagus. It is very common in scrub at the Archbold Biologi- cal Station. The only other known collection site is the type locality of Dune- din, a locality which includes some scrub (it is also the type locality of the Florida sand cockroach) where Blatchley collected. Scarabaeidae—Ataenius saramari Cartwright. Woodruff (1973) provides characters for distinguishing this species from a large number of similar spe- cies. It is known from the Mount Dora Ridge (Marion Co.), the Sumter Up- lands (Marion Co.), the Atlantic Coastal Ridge (St. Lucie, Martin Co.), scrub areas near St. Cloud (Osceola Co.) (Woodruff 1973) and the Lake Wales Ridge (Highlands Co.). This species is usually collected in sand pine needle litter (Woodruff 1973) but can be collected by sifting in a variety of scrub habitats. The wings are short and may be non-functional; this would explain why specimens are never taken at lights. Scarabaeidae—Psammodius sp. Specimens of this undescribed small wingless scarab have been collected by sifting sand in scrub at the ABS. It appears to be closely related to P. hydropicus Horn, another wingless sand- dwelling species. P. hydropicus occurs on beaches in the southeastern U.S., and there are wingless beach species in Oregon and California (Cartwright 1955). The ABS species is the first inland wingless arenicolus species. It is possible that this or similar species are widely distributed in Florida scrubs as they are not likely to be found by normal collecting methods. Cerambycidae—Aethecerinus hornii (Lacordaire). This reddish brown cerambycid with pale elytral lines is known from a scrub area in Lee County (Woodruff 1973) and from the Lake Wales Ridge (Highlands Co.). I have taken an adult in its pupal cell and several larvae in dead stems of Quercus inopina in a rosemary scrub area. The two other members of this genus are southwestern (Linsley 1962). Anthicidae— Mecynotarsus sp. An undescribed species of mottled black and white anthicid occurs in scrub on the Lake Wales Ridge (Highlands, Polk Co.). This species is related to species found on beaches on both coasts of Florida (Chandler 1986). Polk Co. specimens are marked differently from Highlands Co. specimens. A species of Mecynotarsus in Nevada provides a parallel example of an endemic of relict inland dune systems (Rust 1986). Lampyridae—Pleotomodes needhami (Green). Characters for identifying this small sluggish firefly are given by Geisthardt (1986). This species is known only from the ABS, where it is abundant. Adults, pupae, and larvae have recently been found in the fungus gardens of the ant Trachymymex septentrionalis (McCook) (Beshars, 1988, Sivinsky, 1988). Lampyridae—Lucidota luteicollis LeConte. This black species with a bright orange pronotum is the only day-flying firefly in Florida scrub. I have seen specimens from the Lake Wales Ridge (Highlands, Polk Co.), the Mount Dora Ridge (Marion Co.) and the Brooksville Ridge (Levy Co.). The female is wingless. No. 4, 1989] DEYRUP—FLORIDA SCRUB ARTHROPODS 263 Chrysomelidae—An unidentified species of Pachybrachys, near P. confor- mis Suffrian, occurs on Ceratiola ericoides on the Lake Wales Ridge. Consid- ering the idiosyncracies of the host plant, and the host specificity of most Pachybrachys it is extremely unlikely that this species has any other hosts. Class Insecta, Order Diptera, Mydidae—Nemomydas melanopogon Stey- skal. Characters for distinguishing eastern species of these small mydas flies are given by Steyskal (1956). This species is known from the Mount Dora Ridge (Lake Co.), the Crescent City Ridge (Putnam Co.) (Steyskal 1956) and the Lake Wales Ridge (Highlands, Polk Co.). Mydidae—Nemomydas lara Steyskal. This species is known from the Mount Dora Ridge (Marion, Lake Co.) and the Orlando Ridge (Orange Co.) (Steyskal 1956). Larvae of this genus are probably predators inhabiting sandy areas. There are 7 western species (Cole 1969) and 3 eastern, all the latter in Florida. Class Insecta, Order Siphonaptera, Rhopalopsyllidae—Polygenis flori- danus Johnson and Layne. Technical characters for identification of this flea are provided by Johnson and Layne (1961). It is restricted to the scrub en- demic mouse Podomys floridanus (Johnson and Layne 1961), and is likely to share the wide distribution of that species in scrubs throughout Florida. The closest relatives of this flea are on South and Central American Podomys; both flea and host appear to be further examples of Neotropical scrub- adapted lineages isolated in Florida scrub (Johnson and Layne 1961). Class Insecta, Order Lepidoptera, Geometridae—Nemouria outina Ferguson. This species is one of a large group of green geometrid moths re- vised by Ferguson (1969). It is known from the Lake Wales Ridge (Highlands Co.), the Brooksville Ridge (Hernando Co.), the Atlantic Coastal Ridge (Martin, Dade Co.), and the Orlando Ridge (Orange Co.) (Ferguson 1969). I have reared this species from Ceratiola ericoides; it is highly unlikely that there are other hosts. Class Insecta, Order Hymenoptera, Mutillidae—Dasymutilla archboldi Schmidt and Mickel. This species is distinguished from a number of similar species by characters provided by Schmidt and Mickel (1979) (description of female) and by Manley (1983, 1984) (description of male). It is known only from the Lake Wales Ridge (Highlands, Polk Co.), where it is often the most abundant species in open scrub. Mutillidae—Photomorphus archboldi Manley and Deyrup. This species may be distinguished from other Photomorphus by the characters given by Manley and Deyrup (1987). It is a nocturnal species known from specimens taken in Malaise traps on the Lake Wales Ridge (Highlands Co.), the Sumter Upland (Marion Co.), and the Mount Dora Ridge (Marion Co.) (Manley and Deyrup 1987). Formicidae—Conomyrma elegans Trager. This ant is distinguished from its congeners by its elongate body form and appendages (Trager 1988). It is known from three scrub sites on the Lake Wales Ridge (Highlands Co.); these sites are unusual in that they have yellow rather than white sand. 264 FLORIDA SCIENTIST [Vol. 52 Formicidae—Conomyrma flavopectus (M. R. Smith). The name flavo- pectus has been applied to several species of Conomyrma but should be re- served for a distinctive black and orange species found in scrub habitats (Tra- ger 1988). It is known from the Lake Wales Ridge (Highlands Co.) and the Mount Dora Ridge (Lake, Marion Co.). Formicidae—Odontomachus clarus Roger. This species is distinguished from the two other Florida Odontomachus by its smooth petiolar scale and sparsely pubescent gaster (Deyrup et al. 1985). It is known from the Lake Wales Ridge (Highlands, Polk Co.). It appears that this species is replaced by the less ecologically restricted species O. brunneus (Patton) in the scrubs of Marion Co. O. clarus is also known from Mexico and the southwestern U.S. (Brown 1976). Assuming that the Florida population is really the same species as the western clarus, this is the first arthropod species known to have dis- junct populations in scrub and in the Southwest, like those of the scrub jay. Halictidae—Dialictus placidensis Mitchell. A description of this species and keys for identifying members of the large genus Dialictus are provided by Mitchell (1960). This bee was described from two specimens, one from Lake Placid (Highlands Co.), the other from Oneco (Manatee Co.); we do not know the habitat at the latter site. We include this species as a scrub species because of its great abundance in scrub at the ABS (it is the most common halictid) and absence of specimens from elsewhere, with the exception of the Oneco specimen. Mitchell looked at great numbers of Dialictus from Florida, and one would expect placidensis to appear from other sites if it were widely distributed. Discuss1on—Status of Annotated List—This list is necessarily prelimi- nary because our knowledge of many groups of Florida arthropods is rudi- mentary. There are whole groups of arenophilous arthropods common in scrub whose distribution is too poorly known to allow us to single out the scrub endemics. In some cases the classification is also unresolved. In the family Bombyliidae, for example, there are 6 undescribed species of a genus of tiny flies whose larvae are in ant nests. These flies are likely to be sensitive to soil and temperature conditions, and some species are probably confined to Flor- ida scrub. Since these flies require specialized collecting methods, and since they are known only from a single site (ABS), it is impossible to draw any conclusions at present. I would certainly expect to find scrub endemics among arenophilous members of the families Linyphiidae, Carabidae, Staphylinidae, Elateridae, Alleculidae, Tenebrionidae, Bombyliidae, Asili- dae, Therevidae, Sphecidae (s. 1.), and Pompilidae. There should be addi- tional species in some groups already known to include scrub endemics, espe- cially the families Scarabaeidae, Lycosidae, Halictidae, and Mydidae. Soil mites and parasitic mites constitute an enormous group of poorly known ar- thropods and probably include as many scrub specialists as all other arthro- pods combined. No. 4, 1989] DEYRUP—FLORIDA SCRUB ARTHROPODS 265 Species that depend on scrub endemic plants are even less understood than species that are directly dependent on the physical environment of scrub. Ceratiola ericoides, for example is confined to scrub in Florida, and appears to have its own specialized herbivores, including unidentified species of Hemiptera, Homoptera, and Lepidoptera. Carya floridana and Pinus clausa are attacked by large numbers of herbivorous insects, but the host relationships of these insects are unclear, as there are other species of Carya and Pinus that might be potential hosts. It is highly probable that some her- bivores on scrub hickory and scrub pine are host specific, but cannot be rec- ognized at present. Finally, nobody has investigated whether specialized her- bivorous insects are associated with herbaceous plants endemic to scrub. Origin and Biogeography of Endemic Scrub Arthropods—Even though the arthropod list is incomplete and the geographic ranges of most of the species poorly known, it is possible to discern some large scale biogeographic patterns. As in the case of a few reptiles (Auffenburg 1982), the mouse Podo- mys floridanus (Johnson and Layne 1961), and the scrub jay, certain arthro- pods have close relatives in the southwestern U.S., Mexico, and even Central America. Most of these species show poor long-range mobility and, like the reptiles, provide evidence of an ancient continuous austral band of xeric habi- tat. Examples of these species with strong western affinities are Arenivaga floridensis, Terapus sp., Nemomydas melanopogon, N. lara, Polygenus flori- danus, and Odontomachus clarus (references in annotated list). The term “relic” would be applied to these arthropods if they were considered at the genus level, though the species (except for O. clarus) are likely to have evolved in Florida and are in that sense autochthonous. There are no scrub endemic arthropods that appear to be West Indian in origin. Within Florida, relict and autochthonous lineages can be considered at the species level. Species presently found in widely disjunct scrubs are proba- bly relic populations left from a time when a drier climate occurred (Neill 1957). Examples of such species are Latrodectus bishopi, Arenivaga floriden- sis, Schistocerca ceratiola, Cicindela scabrosa, Trigonopeltastes floridanus and Ataenius saramari. As in the vertebrates (Neill 1957, Christman 1980a, Auffenburg 1982), there are cases where species complexes or subspecies ap- pear to have diverged on various scrub islands. Examples are provided by Melanoplus spp., the subspecies of Geolycosa xera, the Cicindela abdominalis complex (includes C. highlandensis and C. scabrosa), Peltotrupes youngi and most of the endemic Phyllophaga. Many more examples would undoubtedly be revealed by analyses of variation in scrub arthropods. In the ant Conomy- rma bossuta Trager for example, there appears to be a distinctive pale-colored population in scrubs on the southern Lake Wales Ridge. There is a current aversion to formally describing or (in the case cited above) even mentioning geographic races of arthropods in descriptions of new species. This makes taxonomy simpler, but makes biogeographic information much more difficult to extract from the literature. 266 FLORIDA SCIENTIST [Vol. 52 The origins of some scrub endemics are difficult to interpret even when information on their ranges and relationships is available. The flightless are- nicolous beetles Psammodius sp. and Mecynotarsus sp. have relatives in the Southwest and also on Florida beaches, and can be considered either isolates from the former austral arid zone or beach species abandoned by the sea. In such cases chemical techniques would be appropriate for determining rela- tionships. The southern Lake Wales Ridge has one organism that has no close relatives anywhere. This is the millipede Floridobolus penneri (Causey 1957, Keeton 1959). There are upland species more widely distributed in Florida that present the same problem. These include the two species of the firefly genus Pleotomodes (Geisthardt 1986); scrub rosemary, Ceratiola ericoides (Moore et al. 1970); and the reptiles Neoseps reynoldsi and Stilosoma ex- tenuatum (Auffenberg 1982). These could all be either relict or autochtho- nous lineages at the generic level. Analysis of the distribution of scrub endemic arthropods shows some pat- terns too pervasive to be due to collecting bias. The majority of species occur on the central ridges (Mount Dora, Orlando, Lake Wales), which are more or less contiguous. Of the 46 species listed above, 43 occur on the central ridges, with 20 apparently restricted to the central ridges. The central ridges, espe- cially the Lake Wales Ridge, are also the center of distribution of scrub en- demic plants (S. Christman, personal communication). There are two spe- cies, a wolf spider (L. pseudoceratiola) and a grasshopper (M. insignis), known only from the Atlantic Coast Ridge. There are a few plants restricted to this ridge, such as Dicerandra immaculata (Lakela 1963) and Asimina tetramera (Austin and Tatje 1979). One species, a scarab (M. pedester), is known only from scrubs in southwest Florida. Woodruff (1973) contends that this is a distinct species, but it would be possible to argue that it is a distinc- tive allopatric population of M. cartwrighti Olson and Hubbell. Its distribu- tion is apparently unique among scrub organisms, though there is a gray- throated form of the green anole (Anolis carolinensis) whose distribution is similar to that of M. pedester (Christman 1980b). Collecting may have been concentrated on the scrubs of the central ridges, but this sampling bias could not be strong enough to affect the overwhelming evidence that the central ridges are the center of distribution of scrub endemic arthropods. The Ecology of Scrub Endemic Arthropods—There is no analysis of com- munities of arthropods of Florida scrub, as there has been for arthropods of isolated dunes in the West (Rust 1986). The results of such analysis would clearly depend on the type of scrub. An open Ceratiola stand would probably have the lowest total diversity of species and the highest proportion of endem- ics, considering the harshness of the habitat and the relatively high propor- tion of scrub endemic plants. A more mesic mature sand pine stand with oak understory would probably have a much higher species diversity and a lower proportion of scrub endemic arthropods. When all scrub habitats at the ABS are combined for analysis, the proportion of scrub endemics appears very No. 4, 1989] DEYRUP—FLORIDA SCRUB ARTHROPODS 74 SS9 low. Among the ants (Formicidae) there are 59 species found in scrub, and only 3 scrub endemics. There are 35 species of velvet ants (Mutillidae) in scrub, with only 3 endemics, about 70 bees (Apoidea) with one endemic, about 80 scarab beetles (Scarabaeidae) with 9 endemics, 45 bark and ambro- sia beetles (Scolytidae) with no endemics. Since Florida scrub is found on former islands, and still occurs as islands of highly distinctive habitat, one might expect scrub arthropods to show some of the characteristics associated with island populations. One of these charac- teristics is an ecological “release”: island species or groups of species may have broader ecological roles than equivalent mainland lineages as the island pop- ulations have evolved to take advantage of underexploited resources (MacAr- thur and Wilson 1967). This characteristic, if it ever occurred, may be sup- pressed in scrub endemics because the islands of habitat are less stringently isolated than when surrounded by water, and there may have been considera- ble movement and dispersion of upland species during drier periods. The endemic velvet ant Dasymutilla archboldi is a possible example of ecological release: it shows an extraordinary size variation that has been explained as possibly the result of a broad ecological role on former islands (Deyrup and Manley 1986). A more easily observed adaptation of island populations is a reduction in dispersal ability (MacArthur and Wilson 1967). Of the 36 species of insects listed above (parasites of vertebrates excluded), 12 are flightless, at least in the case of the female. This is certainly a high proportion of flightless species, but most of these belong to groups that are always flightless. Only 5 species (Melanoplus spp., L. luteicollis, Psammodius sp.) belong to genera that in- clude flying species. In the case of Melanoplus and Psammodius there are flightless species that occur outside of scrub or island situations, so the ances- tors of the scrub species may well have been flightless. This situation would strongly suggest that flightlessness leads to endemism on scrub islands, rather than the reverse. The simplest explanation for this counterintuitive conclu- sion is that large numbers of species that were endemics when the sea level was high never lost their powers of dispersal. Since these species subsequently spread through a number of xeric habitats in a broad geographic area, they are not regarded as endemics. Flightless species, including the relatively large number of spiders, remain marooned as recognized endemics. Scrub endemic arthropods that are host specific on endemic plants in- clude no known examples of flightless species. The need to move readily from one host plant to another may have counterbalanced any tendency toward loss of dispersal ability, if such a tendency even existed. Retrospection reminds the biogeographer that views of endemism can change dramatically over relatively short intervals of geologic time, even when the same organisms and land areas are involved. Many scrub species that are common in other upland habitats in Florida and adjacent states may have been restricted to scrub habitats during inundations of the Florida pe- ninsula. Hypothetical biologists present during these times of flooding would 268 FLORIDA SCIENTIST [Vol. 52 probably have found the island biota remarkably distinct from that of the mainland, and would have urged the (hypothetical) federal government to set aside portions of the archipelago as biotic reserves. ConcLus1on—The arthropods of Florida scrub are a rich and relatively unexploited source of biogeographical and ecological information. The 46 endemic species discussed here are probably only a fraction of the endemics present. The exact ranges of the known arthropods are poorly known, but enough is known to show that these arthropods reveal a fascinating history of the evolution of isolated populations and the impact of climatic and geologic changes. It is frustrating to know that many islands of Florida scrub habitat have already been destroyed by human activity, and many more will disap- pear before their arthropods have been studied. The efforts of conservation- ists have generally been focused on particular endangered species of scrub plants and vertebrates. Arthropods lack such general appeal, and it is easy to look beyond the individual species to the evolutionary principles they em- body. From this viewpoint it becomes imperative to salvage what we can, in the form of information if not the actual habitats, of the entire inland archi- pelago, in order to understand the biogeography of Florida. It is sad to think of North American biogeographers and evolutionary biologists preparing large scale grant proposals to study the biotic patterns of distant island chains, while their opportunities in Florida slip away. ACKNOWLEDGMENTS—I gratefully acknowledge the assistance of the following individuals. Dr. Steven Christman (Florida State Museum), initiated scrub surveys on the Lake Wales Ridge, and reviewed this paper. Paige Martin (ABS), and James Cronin (Florida State University), collected arthropods from pitfall traps. Dr. Eric Menges, Dr. Shirley Denton, Dr. James Layne, Dr. James Wolfe, and Fred Lohrer (ABS) reviewed the manuscript. Nancy Deyrup and James Cronin assisted in computer compilation of data from the pitfall traps. Dorothy Carter typed the manuscript. LITERATURE CITED ABRAHAMSON, W. G., A. F. Jounson, J. N. LAYNE, AND P. A. PERoNI. 1984. Vegetation of the Archbold Biological Station, Florida: an example of the southern Lake Wales Ridge. 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Vol. I. N.C. Agr. Exper. Stat. Tech. Bull. 141:1-538. Moore, D. M., J. B. HARBoRNE, AND C. A. WituiaMs. 1970. Chemotaxonomy, variation, and geographical distribution of the Empetraceae. Bot. J. Linn. Soc. 63:277-293. NeILi, W. T. 1957. Historical biogeography of present-day Florida. Bull. Fla. State Mus. 2:175- 220. Piatnick, N. I. anDM. U. SHapas. 1983. A revision of the American spiders of the genus Zelotes . (Araneae, Gnaphosidae). Bull. Amer. Mus. Nat. Hist. 174:97-192. Potts, W. L. 1976. New Species of American Anomala (Scarabaeidae: Anomalinae). PAN-PAC. Ent. 52:220-226, REISKIND, J. 1987. Status of the rosemary wolf spider in Florida. Univ. Fla. Coop. Fish Wildl. Res. Unit Tech. Rep. 28:1-13. Rust, R. W. 1986. Seasonal distribution, trophic structure and origin of sand obligate insect communities in the Great Basin. Pan-Pac. Ent. 62:44-52. ScHMupT, J. O. AND C. E. Micke. 1979. A new species of Dasymutilla from Florida (Hymenop- tera: Mutillidae). Proc. Ent. Soc. Wash. 81:576-579. Sivinsky, J. 1988. University of Florida, personal communication. STEYSKAL, G. C. 1956. The eastern species of Nemomydas Curran (Diptera:Mydaidae). Occ. Pap. Mus. Zool. Univ. Mich. 573:1-5. TRAGER, J. C. 1988. A revision of Conomyrma (Hymenoptera: Formicidae) from the southeastern United States, especially Florida, with keys to the species. Fla. Entomol. 71:11-29. Wa..ace, H. K. 1982. Order Araneae, species accounts. Pp. 120-129. In: Franz, R. (ed.). Rare and Endangered Biota of Florida. Vol. 6. Invertebrates. University Presses of Florida, Gainesville. 131 pp. Wuire, W. A. 1970. The geomorphology of the Florida peninsula. Fla. Dept. Nat. Res. Geol. Bull. 51:164 pp. Wooprurr, R. E. 1960. Suppression of the genus Roplisa Casey with notes on the United States species of Trigonopeltastes Burmeister (Coleoptera:Scarabaeidae). Fla. Ent. 43:139-145. 1973. The scarab beetles of Florida (Coleoptera:Scarabaeidae), part 1. The Laparos- ticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae). Florida Dept. Agric., Div. Plant Indust. Arthropods of Florida and Neighboring Land Areas 8: 1-220. 1982. Family Scarabaeidae, species accounts. Pp. 84-102. In: Franz, R. (ed.). Rare and Endangered Biota of Florida. Vol. 6. Invertebrates. Univ. Presses of Florida, Gaines- ville. 131 pp. Florida Sci. 52(4):254-270. 1989. Accepted: November 22. 1988. Science Teaching PRODUCTIVITY OF DEPARTMENTS OF CHEMISTRY AT FLORIDA GRADUATE INSTITUTIONS JOHN C. FOLLMAN! AND DEAN F. MarrTIN? (1) Department of Psychological and Social Foundations, College of Education, University of South Florida, Tampa, FL 33620 and (Chemical and Environmental Management Services (CHEMS) Center, Department of Chemistry, University of South Florida, Tampa, FL 33620 USA Asstract: Data comparing Departments of Chemistry at six Florida universities (number of faculty, postdoctoral appointments, degrees granted at the masters and doctoral levels) were gathered. These were compared with faculty productivity, including total number of publica- tions for a two-year period (January 1, 1985 through December 31, 1986), mean number of publications per faculty member, percentage of those faculty who did not publish during that period, and the maximum number of publications for one faculty member in the department. “PUBLISH-OR-PERISH’ has been a convenient catch-phrase to describe one of the traditions of academic life in the latter half of the twentieth century. The expression implies that rewards in higher education are determined by publications and research/creative activities, rather than by teaching and/or service activities. Evidence for the accuracy of the publish-or-perish premise for rewards is at best diffuse. There is considerable inductive, anecdotal evidence, and much rhetoric. For example, one of us can recall as a young faculty member being told that the sun rises and sets on research at the University of X, a major research institution. The rhetoric is that pay, promotion and tenure decisions are based on a faculty member’s publication, rather than on teach- ing activities (Lindsey, 1979; Mahoney, 1979). It is noteworthy that six books on how to publish were released in a recent five-year period prior to 1983 (Smith, 1983). If the reward system is based on publication then the basic question be- comes: how prolific are faculty members in the penultimate decade of the twentieth century? In particular, what is the veridicality, the perceived valid- ity of Lotka’s (1926) Law of Literary Productivity which postulates there is an inverse relationship between scholarly productivity and the number of producers, i.e., 1/n’. This empirical law was developed first for chemists, and then for physicists. Consequently, a rule of thumb of faculty publication is that about one-third of the faculty produce nothing, 50% produce 15% or fewer of the publications, about 10-15% produce about half of the publica- tions, and about 2-3% produce about 15% of the total. The present study examines the productivity of chemists at six Florida institutions with graduate programs in chemistry. LA FLORIDA SCIENTIST [Vol. 52 MeEtHops—Data on demographic characteristics of schools reporting graduate programs (Table 1) to the American Chemical Society (ACS) Committee on Professional Training (CPT) were obtained from the Directory of Graduate Research (CPT, 1987). This source was also used to find the number of specific publications for faculty members at each of the six Florida universi- ties studied (CPT, 1987). The total number of publications for each department was divided by the number of listed faculty members to compute the mean publications per faculty member per department (Table 2). In addition, the percentage of faculty who had not listed a publication or for whom no publications had been retrieved through the ACS Chemical Abstracts Service was calculated. Finally, we recorded the number of publications for the most productive faculty member in a given department (Table 2). The listings of all Florida chemistry department degree programs and faculty members were obtained in ACS (1986). Correlation coefficients and regres- sion equations were computed. RESULTS AND Discuss1oN—Some limitations in our data should be noted. The number of research publications for the 10 chemistry faculty at Florida International University and six faculty at Florida A & M (ACS, 1986) were not available in the Directory of Graduate Research (CPT, 1987). In addi- tion, the total number of publications for a given department could be over- estimated because of collaboration. Finally, books, textbooks and mono- graphs were not counted as publications in CPT (1987), even though these represent scholarly activities. For example, colleagues at the University of South Florida (USF) have written over twenty monographs and textbooks including two organic chemistry textbooks (Solomons, 1986; Raber and Ra- ber, 1988); also colleagues at UCF have written a definitive textbook on in- dustrial chemistry (Clausen and Mattson, 1978). Other examples of scholarly activity could be cited. In addition, the number of faculty members reported for a given depart- ment may not match the number listed in the Directory of Graduate Re- search (CPT, 1987). In all instances, the overall number listed exceeded the number reported in the statistical summary. The difference is perhaps linked to senior-level research associates who have a limited teaching function, or in some instances linked to associates in chemistry who have a higher-than- typical teaching assignment and publish. In some instances, though appar- ently not the ones we covered, the number of faculty listed in full with biog- raphies and cited publications is significantly less than the reported number of full-time faculty. A prime example is CUNY Brooklyn College, for which 20 faculty members were listed in the biographical/bibliographical section while 27 were reported in the statistical section. The difference represents the disparity between those faculty recognized as committed to graduate educa- tion and research versus those committed to undergraduate education and other forms of scholarly endeavor other than traditional research papers (Ziegler, personal communication, 1985). Data Interpretation—The Department of Chemistry at The University of Florida (UF) and Florida State University (FSU) are larger than those at the other four institutions for most of the criteria (Table 1). Both UF and FSU lead in publication productivity. Table 2 indicates that mean faculty publica- tions are fairly comparable across five of the departments, except for the University of Central Florida (UCF), the youngest department, which has a 273 FOLLMAN AND MARTIN—CHEMISTRY DEPARTMENTS No. 4, 1989] ‘sydeisououw pur syooq jou pure sjeuinol peyUel1o-Ajsturayo ur paysiqnd saporsze o1fQUaros 0} Sajal _SUOT}BOI[GNZ,,x 9G eV ce L6 Bployy YyNos cI GG VY 6L Ture ly] 89 jail 9°8 C6E epltoly IZ 6 gg Cc6I 9781S BPHOLy VP cc CPG 09 onUueNYy epliop.y él SL aol LI Bpllopy [ere UMNUTXBUT A}{NOVF O[BUIS peyst{qnd jou yueo18g Aynoey iad uvayy s[eq0], HSIOATU A) esuoTyeol|qng a ee ee SS ee SONISIOATUN BPLIOLy 72 SpusuZIEdap AIysTUAYO UT AY[NIeJ JO (QQET “G86T) suoneorqnd jo Areuiwins *% aTavy, ‘U9ATS ST [BAQ] STU] 3B BaIZap OU Sa}eOIpUI Y/N ‘potsad sty} Bulinp Aue aals jou pip ynq aeisep ay} Soars JUsWIedap 9} SozOIPUT (q ‘98/L JO See € I oS tl Vv 86 BpHoyy YNosg I 6 0 € g 0 bP OI g 0 LI TUIRTYY cI 0Z II € PSI OF OL € oP epHoly Ol ra 9 9 96 GZ 13 0 9¢ 9781S BPLOL A V/N V/N I 0 G I I I Or oNUBPY epHopy V/N aV/N a0 0 0 0 0 0 II BPHOLy [eID 986T C861 986I C861 [P3707 IBOA-ISITF SeoTUlOe ce WETS ATS SLANG CS6I VS6I CS6I VS6I JUSUW][OIUS oyeNpeID e[B1OJOP sod vO UIT} -}18 J vO UT} -[[N pepleMmy sooid9q SOMISIOATUN BPO] UT SyUsWAIedap AI}STUIIYO JO SONST}VIS P9}DI]VS * | ATAVI, 274 FLORIDA SCIENTIST [Vol. 52 M.S. in Industrial Chemistry. Noteworthy are the remarkable single faculty two-year outputs of 44 at Florida Atlantic University (FAU), and 68 at UF, respectively. Several correlations were computed to get an indication of factors affect- ing publication prolificity for a given chemistry department (Table 3). For the six departments, the total number of publications was a linear function of the total number of faculty, i.e., the degree of association (r’) is 0.80, which can be interpreted that 80% of the variation in the total number of publica- tions for a department could be accounted for simply by the number of fac- ulty members. These results are surprising in the light of academic traditions and expectations that some faculty members are considerably more produc- tive than others (the range at UF, for example, was 0-68). The relative stand- ard deviation (standard deviation divided by mean) was sizeable for smaller departments, e.g., 242% (USF), 240% (FAU), and generally less for larger ones, 180% (USF), 120% University of Miami (UM), 87% (FSU) and 147% (UF). We believe that this reflects the greater productivity of the larger de- partments and that the most prolific faculty members have a relatively smaller impact on the overall record. We assumed that the number of publications should be related to the activity of the graduate students working closely with faculty members. There are significant (p <0.01) correlations of 0.94 between the total number of publications and the number of doctorates awarded during a two year period, and 0.94 between the total number of graduate degrees (PhD + MS; six schools). Although there is no difference between the correlation coeffi- cients, the defining equation has less variation for the total number of de- grees. The Graduate Directory of Research contains not only complete citations of faculty publications, but it also lists theses and dissertations resulting from a faculty member’s service as advisor. There is commonly a time lag between appearance of the thesis/dissertation and appearance of articles generated as a result of the collaborative research that led to the thesis. Allison (1980), for example, provides an extended treatment of the random variation by scien- tists, including chemists, in their publishing activities over a two-year period. Nevertheless, there may be a common pattern: that those faculty who are advising thesis/dissertation students will continue to do so over several years and their papers and thesis/dissertation lists will continue to be out-of-phase, but that continued productivity in advising graduate students will be re- flected in continued productivity in publishing in traditional chemistry jour- nals. We also examined the relationships between total publications and total graduate enrollment (as of September 1986) and found a direct, significant (p <0.001) correlation of 0.975. The importance in productivity by postdoctoral associates also should not be underestimated, and one should not overlook the advantages of advising postdoctoral research associates who spend one or two years with a research 2795 FOLLMAN AND MARTIN—CHEMISTRY DEPARTMENTS No. 4, 1989] ‘(€861) SVN Wor eyep Aprerprure qq ‘(QS ¥ UvaUI ale sonjeA q pue ve) ¥q+te=e 986I1- F861 syuouryurodde 100°0> ve 9S76 0 SOFS Cl CS ZL S8G= 9 suoHeorqnd [e10}90p-ysod [BIO], 98/6 2 PRISE | OAKS) 9861 -F86I 100°0> SSil VL86 0 91 0*86T Vt Bescil 9 ANS AS) aN G, suoneorqnd [230], Aynoey juswj1edap ED O861-6L61 Sd0°0-S0'0 el 66 0 £0'0+¢61'0 Say FORE 4 gAH IETF suoneorqnd [eq], Aynoeyj juowj1edap 9} Jo 9861 -F86I SN OL 0< G LLVL’O 1€0'0 + L670'°0 LL 8€ V qAH TPT [FUe suoneorqnd [eo], 986I-FS6I 986I-FS6T suoneorqnd popieme 10°0> 8S VLI6 0 98°0+09'9 IGS TE 9 [e3en ‘SWt ddd 986IT-P861 suoneorqnd 100°0-10°0 aT L688°0 6*+66 COO 62> 9 [eIeL Aynoey [BIOL 986I-F86T 9861 -FS86I 10: 0> LG SVL6°0 89° 1 +89'6 CE +P 66 Vv SUSE ON GIN Jepertreny VIE Mol d ad I =4 =e N x X euoTyenboe sululjaqd Apnjs sty} 107 suonenbs Sululjap pur SUONReIIO pa}Oa]9S *¢ ATAV], 276 FLORIDA SCIENTIST [Vol. 52 group as part of an initiation into an academic career in chemistry. The initiative and productivity of the experienced associates can be significantly greater than that of the typical graduate student. Some associates on the other hand, have suffered from a short-lived post-dissertation burnout, and are less productive than anticipated. For example, Reskin (1977) showed em- pirically that the influence of the advisor was primarily on early productivity of advisees, but only in high-caliber chemistry departments. The productivity (total publications) at six Florida institutions was directly related (r=0.95, p=0.001) to the total number of postdoctoral associates present in a chemis- try department. Surprisingly, total number of publications was not related (p= >0.05) to the reported (NAS, 1983) perceived quality of the faculty or to the perceived familiarity (r=0.75; p>0.1) of the faculty members in four doctoral institu- tions. The data used for these perceived values were obtained from a NAS (1983) evaluation of about 600 doctoral programs in mathematics and physi- cal sciences. A standardized scale was used in the NAS study (1983) for fac- ulty quality and familiarity of the faculty to those surveyed, where the aver- age score of all rated departments was 50. On this scale, UF and FSU had virtually the same degree of familiarity, 55, and 58, respectively, as did USF and UM, 41, and 39, respectively. There was no difference in faculty quality for FSU and UF (both 57) nor between USF and UM (both 37). Perhaps this observation should not be unexpected because Hagstrom (1971) demon- strated empirically that while an aggregation of chemistry department char- acteristics accounted for large amounts (71%, 21% respectively) of depart- mental prestige, no single variable accounted for much variance across both of the two regression equations. Probably, if there is a relationship between familiarity and number of publications, data would be needed for several years before 1983. Selecting the correct number of years is a singularly arbitrary choice, but we selected the number of publications for the period January 1, 1979 through December 31, 1980 (CPT, 198) with the thought that these papers might have had a recent impact on those being surveyed. The correlation between the total number of papers and the familiarity was not only higher than when 1984-86 publications were used (r=0.93), but it was also statistically significant (p<0.05, Table 3). While we admit that total number of publications is an imperfect crite- rion of faculty/department quality, there is an empirical basis for its use. Blume and Sinclair (1973), in a sample of British universities, found correla- tions of 0.63, 0.66, and 0.75, between productivity (number of papers) from a university and quality (peer evaluation). We believe that our results point to a problem that was raised in 1958 and in 1986. Caplow and McGee (1958) popularized the phrase “publish or per- ish” as a result of their survey of faculties at ten major universities. They called attention to a problem, but did not provide a solution. The authors noted a reality of the teaching profession, “a real strain in the academic role No. 4, 1989] FOLLMAN AND MARTIN—CHEMISTRY DEPARTMENTS DHT arises from the fact that [faculty members] are, in essence, paid to do one job [teach] whereas the worth of their services is evaluated on how well they do > . another [research].”’ While one might question this, our study calls attention to the correlation between perception of chemistry faculty quality at six Flor- ida universities and the total number of publications published by the faculty in the respective departments of chemistry. The perception is insidious when we recall the conclusion of Bowen and Schuster (1986) that the most critical influence on higher education is the caliber of its faculties. If the perceived caliber is determined solely by the quantity of the faculty publication, then we might very well find that we could publish and perish. ACKNOWLEDGMENTS— We are grateful to Walter K. Taylor for serving as consulting editor. LITERATURE CITED Auuison, P. D. 1980. Inequality and scientific productivity. Social Stud. Sci. 10:163-179. AMERICAN CHEMICAL Society. 1986. College Chemistry Faculties, ACS, Washington, DC. BiumgE, S. S. AND R. Sincuair. 1973. Chemists in British universities: A study of the reward system in science. Am. Sociolog. Rev. 38:126-138. Bowen, H. R. anv J. H. Scuuster. 1986. American Professors: A National Resource Imperiled. Oxford University Press. New York. Captow, T. AnD R. J. McGee. 1958. The Academic Marketplace. Crausen, C. A. IIT AND G. Mattson. 1978. Principles of Industrial Chemistry. Wiley, New York. CoMMITTEE ON PROFESSIONAL TRAINING. 1987. Directory of Graduate Research. American Chem- ical Society, Washington, DC. . 1980. Directory of Graduate Research. American Chemical Society, Washington, DC. Hacstrom, W. O. 1971. Inputs, outputs, and the prestige of university departments. Sociolog, Educ. 44:75-397. Linpsey, D. 1979. Editorial vision and journal product. Am. Sociologist. 14:41-242. Lorka, A. J. 1926. The frequency distribution of scientific productivity. J. Wash. Acad. Sci. 16:317-323. Manuoney, M. J. 1979. Psychology of the scientist: an evaluative review. Soc. Stud. Sci. 9:349- 375. NATIONAL ACADEMY OF SCIENCES. 1983. An Assessment of Research-Doctorate Programs in the United States: Mathematics and Physical Sciences. National Academy Press, Washington, DC. Raber, D. J. ANDN. K. Raper. 1988. Organic Chemistry. West. New York. RESKIN, B. F. 1977. Scientific productivity and the reward structure of science. Am. Sociolog. Rev. 42:491-504. SmitH, V. H. 1983. A dry but informative look at writing for publication. Book review of Getting Published in Education Journals. Phi Delta Kappan. 64:666. SoLomons, T. W. G. 1986. Organic Chemistry 4th ed. Wiley. New York. ZIEGLER, H. E. 1985. Pers. Comm., City University of New York, Brooklyn College, Brooklyn, New York. Florida Sci. 52(4):271-277. 1989. Accepted: November 21, 1988. Biological Chemistry INFESTATION AND EPIDEMIOLOGY OF HEAD LICE IN ELEMENTARY SCHOOLS IN HILLSBOROUGH COUNTY, FLORIDA W. WAYNE PRICE AND AMPARO BENITEZ Department of Biology, University of Tampa, Tampa, FL 33606-1490 Asstract: An investigation of the prevalence and epidemiology of human head lice (Pedicu- lus humanus capitis) was conducted in the elementary schools of Hillsborough County, Florida during the academic years of 1985-86 and 1986-87. A questionnaire survey of county schools showed that 6.7% of the students (excluding Blacks) were infested in 1985-86. Nearly half of the cases were due to reinfestations. Students in one school were examined 4 times, at approximately 3 month intervals in 1986-87, and answered questionnaires. Ninety-seven (6.4%) of the 1515 White, Hispanic, and Asian students were infested, but none of the 436 Blacks was infested. The distribution of pediculosis in non-black students was significantly influenced by season, sex, hair length, race, family size, crowding in the home, infestation of other family members, socioeco- nomic level, and mode of transportation to and from school. Classroom activities of students associated with increased lice infestation included using headsets, crowding at tables, and sitting on the floor. THE human head louse, Pediculus humanus capitis, is a blood sucking parasite that is found on the scalp and hair. Recent studies indicate that its prevalence is high in both developed and developing countries (Gratz, 1985). Although there is no nationwide reporting system for head lice infestations in the United States, at least 3 million households were infested in 1984 (Altschuler and Kenney, 1986). Pediculosis is particularly prevalent among children in elementary schools where it is a major health problem (Robinson and Shepherd, 1980). Within the United States the few recent studies docu- menting the prevalence of pediculosis have been conducted in elementary and junior high schools. Infestation levels were reported to be 4.3% in Tuc- son, Arizona (Lang, 1975), 3% in Barrow County, Georgia (Slonka et al., 1976), and 7.2% in Buffalo, New York (Slonka et al., 1977). Sampling of schoolchildren in New York, Georgia, Pennsylvania, and Florida showed an infestation rate of 10.4% (Juranek, 1985). This study was conducted to eval- uate the magnitude of the pediculosis problem among elementary school chil- dren in Hillsborough County, Florida, and to determine factors involved in the transmission of the disease. MATERIALS AND METHOps—At the beginning of the study in April, 1986, students were en- rolled in 132 schools in Hillsborough County. There were 92 elementary schools with 60,447 students, 27 junior highs with 31,134 and 13 high schools with 23,404. The study consisted of two parts. First, all elementary schools (Kindergarten-Grade 5 or 6) in Hillsborough County were surveyed by questionnaire to determine the prevalence and seasonality of head lice infestation reported in the academic year 1985-86. Second, the student population at a representative Hills- borough County Elementary school was examined for head lice four times: April 1986, Septem- ber 1986, January 1987, and May 1987) by county health personnel and trained volunteers. Infestation was diagnosed by direct inspection of the hair and scalp for the presence of adult head lice, nymphs, or nits (eggs). The hair was parted systematically using wooden applicator sticks. No. 4, 1989] PRICE AND BENITEZ— HEAD LICE 279 Lice and nits were usually visible to the naked eye. Samples of suspected lice or nits were con- firmed by microscopic examination. Infested children were sent home and not readmitted to school until they were free of lice. Epidemiologic data at the elementary school were obtained using questionnaires completed by the students at the time of the examination or sent home to parents and returned by the students. The following information was requested in the questionnaires: 1) grade; 2) sex; 3) race; 4)hair length; 5) family size; 6)number of bedrooms per household; 7) number of preschool children in household; 8) type of housing (single or multiple family dwelling); 9) frequency of hair washing; 10) transportation method to and from school; 11) use of daycare facilities; 12) previous pediculosis cases of child and other household members. At the conclusion of the study, each of the 21 homeroom teachers at the elementary school filled out a questionnaire which requested information concerning various classroom activities and characteristics. Data were analyzed using standard statistical tests including Chi-square, Student’s t-test, and the Kruskal-Wallis test. Values were considered statistically significant at p<0.05. Resutts—Lice Infestation of Elementary Schools in Hillsborough County —All of the 36 schools responding to the questionnaire reported cases of head lice infestation in the academic year 1985-86. A total of 2460 infesta- tions were reported ranging from 6 to 295 cases per school (mean 70.3). The prevalence for all schools was 6.7% and ranged from 0.8-38.9% . High attack rates were uncommon with 3/4 of the schools reporting rates less than 7.4%. Prevalences were calculated after excluding Blacks from the school popula- tions. Blacks have been shown to be refractory to lice infestation in all other U.S. studies (Juranek, 1985) and the present one. Nearly half (49.5%) of the total number of infestations were due to rein- festations, with rates ranging from 4 to 95% per school. More than 3/4 of the schools reported rates of <50%. Table 1 shows the number of lice infestations in each of the four 9 week periods for the elementary schools. Although the greatest number of cases occurred during the late fall, winter, and early spring months of periods II and III, there were no significant differences among the four time periods (Kruskal-Wallis test, p >0.05). Epidemiology of Lice Infestation at one Elementary School—Enrollment from kindergarten through grade 5 in the elementary school chosen by county health officials varied from 525 to 535 from spring, 1986 to spring, 1987. The racial composition of the students for the study was as follows: White-40.0% , Hispanic-36.4%, Black-21.2%, and Asian-2.4%. Twenty-one classrooms were utilized as homerooms. Generally, students spent more time in their homerooms than in any other setting during a school day. TaBLeE 1. Number of lice infestations per 9-week period in Hillsborough County Elementary Schools (K-5 or 6) for the academic year 1985-86. Nine-week period Number of infestations I (Aug.-Oct.) 417 II (Oct.-Dec.) 730 III (Jan.-March) 690 IV (March-May) 538 280 FLORIDA SCIENTIST [Vol. 52 For the entire study, 97 (6.4%) of the 1515 White, Hispanic, and Asian students were infested with head lice, but none of the 436 Blacks was in- fested. Because of the lack of infestation in Blacks, they were excluded from further analyses. Prevalence of lice infestation by students by grade and sex for the four sampling periods is shown in Table 2. In January, 1986 a lice check by school and county health personnel found 30 (7.8%) of 383 non-black students in- fested. The attack rate decreased to about 4% in the spring and fall of 1986, and then increased steadily to 10.4% the following spring. Significant differ- ences existed among the five sampling periods (X*= 17.12, p=0.002). TABLE 2. Prevalence (% ) of lice infestation of children at the elementary school by grade and sex for 4 sampling periods and entire study. Grade Sampling Period K K-1 1-2 2-3 3-4 4-5 5 Total April, 1986 Female = As 6.7 2.6 8.0 32 0.0 3.9 Male — 370 10.0 0.0 6.5 BA 0.0 4.0 Total — Si) 9.1 LS Hes One 0.0 3.9 Sepi. Jo Female 18.8 9.4 5.3 0.0 16.0 27. — ee Male 0.0 0.0 4.8 3.4 0.0 ey — 2.0 Total (5 BoD 5.0 6 6.3 3.9 — 4.4 Jan, 1987 Female eS 11.4 ASS 3.4 9.1 15.8 — 9.8 Male 4.0 1.6 8.0 a 6.7 50) — Ane Total T.23 5.2 6.3 oe4 7.9 10.3 — 6.8 May, 1987 Female 29.4 13.5 8.7 Oxi. 16.0 12-2 — 13.8 Male 8.0 1253 3.6 8.6 8.7 2.4 — AAG Total 16.7 1278 5.9 9.1 12.5 72 = 10.4 Entire Study Female 90.4 10-2 6.3 3.8 12.0 8.8 0.0 8.7 Male Ant 4.4 TW 3.8 4.9 3.9 0.0 4.5 Total 10.6 6.6 6.7 3.8 8.5 6.4 0.0 6.4 Mean daily temperatures and relative humidities for the winter-spring periods of 1986 and 1987 were obtained from the U.S. Weather Bureau Of- fice, Tampa International Airport. Although temperatures were not signifi- cantly different for these two time periods (1986-19.4° C; 1987-18.9° C), the relative humidity in 1987 (71.3%) was significantly greater than in 1986 (66.7%) (t=4.30, p=0.0001). Although the highest attack rates generally occurred in kindergarten chil- dren, there was no association between overall prevalence and grade (X’ test, p=0.15). Females had a higher prevalence of infestation than males in three of four sampling periods (Table 2), and had a significantly higher overall attack rate (8.7% vs. 4.5%) than males (X’?= 11.20, p=0.0008). Hair length of students was classified as short (above collar and ears), medium (touching ears and/or to collar), or long (touching collar) (Table 3). The prevalence of infestation for boys was significantly greater for those with No. 4, 1989] PRICE AND BENITEZ— HEAD LICE 281 medium hair (11.1%) than with short hair (3.8%) (X’?=8.64, p=0.003). Of girls examined with long hair, 12.7% were infested, compared to 4.0% with medium hair and 0.0% with short hair. These differences in attack rates were significant (X?= 10.40, p=0.005). TaBLeE 3. Lice infestation of children at the elementary school by hair length and sex. Prevalence (% ) Hair Length Male Female Total Short 3.8 0.0 3.8 Medium Js bell 4.0 6.3 Long — ail OT, Of the 814 White children examined for head lice in the study 58 (7.1%) were infested as were 25 (4.1%) of the 617 Hispanic children and 2 (4.8%) of the 40 Asian children. The differences in attack rates between whites and the other races combined were significant (X?=6.07, p=0.014). Family size of students was placed in one of three categories (Table 4). Prevalence of infestation increased with increasing family size. Families with four or fewer members had the lowest attack rate (5.4%) and families with seven or more members had the greatest (18.8%). These differences in preva- lence of students based on family size were significant (X*?= 18.50, p=0.0001). TABLE 4. Lice infestation of children at the elementary school by sex and family size. Prevalence (%) Number of family members Male Female Total <4 AV ead, 5.4 5-6 ao 582 LEO >7 20.0 17.6 18.8 For each student a crowding index was determined by the number of persons in the household per bedroom and placed into one of three categories (Table 5). Students in households with the lowest crowding index had the lowest attack rate (6.1%). Attack rates increased with increasing crowding indexes to a high of 15.8%. These differences in attack rates were significant (X?=9.49, p=0.009). TABLE 5. Lice infestation of children at the elementary school by sex and by number of persons in the household per bedroom. Prevalence (%) Number of persons/ bedroom Male Female Total <1.5 3.8 9.2 6.1 Des 6.0 Weal 8.5 >2.5 20.0 13.7 15.8 282 FLORIDA SCIENTIST [Vol. 52 Data concerning infestations of other family members are shown in Table 6. In spring, 1986, 12 students reported that another household member had been treated for head lice during the academic year 1985-86. Four (33.3%) of these students were infested. Of the 219 students who reported that no house- hold member had been treated previously, 5 (2.3%) of this group were in- fested. Seventy-one students reported that another household member had been treated for head lice during the academic year 1986-87; 23 (32.4%) of this group were infested. Of 313 students who reported no household member had been treated, 15 (4.8%) were infested. For both school years, the preva- lence for infested students who had another household member treated for head lice was significantly higher than for those who had no household mem- ber treated (X’ = 29.29, p=0.0001 and X*= 49.45, p=0.0001, respectively). TABLE 6. Lice infestation of children at the elementary school by whether another household member was treated previously. Prevalence (% ) Previous Treatment 1985-86 1986-87 Yes eines go4 No 2.3 4.8 Fifty-nine families had at least one child with head lice during the aca- demic year 1986-87. Thirteen (22% ) of these families reported multiple infes- tations and accounted for 41.3% of the cases recorded during the academic year. Of the 95 infestations recorded during the four sampling periods, 24 (25.3%) were from 12 children who were infested twice and 71 were due to single infestations. During the spring, 1986 investigation, all 8 of the infested students who responded to the questionnaire reported that they had been treated previously for head lice that school year. Of the 219 responding non- infested students, 29 (13.2%) had been treated previously. For the academic year, 1986-87, 72.4% (42 of 58) of the students who were found infested during a sampling period reported having been treated previously for head lice during that year. For the academic year 1986-87, students were divided into two socioeco- nomic levels: low and high. Students whose families qualified for free meals in the federal government’s meal program were placed in the low socioeco- nomic level, whereas all other students were placed in the high socioeco- nomic level. Infestation levels were significantly greater for children in the low level (7.7 % ) as compared to the high level (4.5%) (X?=4.70, p=0.0301). The mode of transportation to and from school was determined for the students: 30% rode buses, 3% rode in car pools, and 67% rode in family cars, bicycles, or walked. The prevalence of infestation in bus riders (14.6%) was significantly greater than the 5.5% for students using all other types of transportation combined (X?= 22.50, p=0.0001). No statistical association was found between infestation and the following factors: frequency of hair washing, type of housing, use of after-school day- care facilities, and number of preschool children per household. No. 4, 1989] PRICE AND BENITEZ— HEAD LICE 283 TABLE 7. Lice infestation of children at the elementary school by classroom. Number of cases Number of Classrooms O22 a 3-4 a 5-6 5 7+ y) Epidemiological evidence suggesting both person-to-person and fomite transmission of head lice was obtained from teacher questionnaires by com- paring characteristics of classrooms having three or more cases with class- rooms having two or fewer cases (Table 7). Due to the small number of class- rooms involved, statistical analysis of the data was not possible. However, some trends were evident (Table 8). Children sat and/or played on the floor in 38.5% of 13 classrooms having three or more cases compared with the 14.2% of seven classrooms having two or fewer cases. Tables were used for student seating to about the same extent in both classroom categories. However, in 90% of the classrooms having three or more cases, four or more students were seated per table compared with 33.3% in classrooms having less than three cases. Headsets were used in 58.3% of the 12 classrooms having three or more cases, but were not used in any of the seven classrooms having less than three cases. No trends were found between the number of classroom cases and presence of carpeted floors, pillows, stuffed toys, combs, hats, blankets, tow- els and costumes in classrooms, and method of clothes storage. Garments were placed under or on the backs of students’ chairs in 17 of the 21 class- rooms. TABLE 8. Characteristics of classrooms associated with lice infestations of children at the elementary school. Classrooms with 3+ cases Classrooms with <3 cases Number classrooms with Number classrooms with char./no. classrooms char./no. classrooms Characteristic examined % examined % Sit/play on floors 5/13 38.5 eit 14.2 Tables in classroom 10/13 76.9 6/7 Sonal 2-3 students/table 1/10 10.0 4/6 66.7 4+ students/table 9/10 90.0 2/6 SS} Headsets used HD 58.3 0/7 0.0 Discuss1on—Prevalences of 6.7% for Hillsborough County elementary schools surveyed by questionnaire and 6.4% for the representative elemen- tary school were similar to results obtained from other studies of schoolchil- dren in the United States (see Introduction). Most of the previous studies were performed during epidemic situations (Slonka et al., 1976; Slonka et al., 1977; Juranek, 1985), whereas the present study was not. These results indi- cate that the magnitude of the pediculosis problem is substantial among the elementary schoolchildren in Hillsborough County. 284 FLORIDA SCIENTIST [Vol. 52 There was no clear-cut seasonal pattern for the infestation rates recorded at the elementary school. During the winter-spring of 1986 prevalence de- creased markedly, but the opposite occurred during the winter-spring of 1987. The higher humidity in 1987 may have increased survivability of eggs, nymphs, or adults off the host and thus facilitated transmission of lice. At 31° and 36° C, Lang (1975) found more hatching of head lice eggs at 70-83% relative humidity than at 50%. Previous studies provide conflicting evidence with respect to patterns of seasonal abundance of head lice. Some found no seasonal fluctuations, espe- cially in equatorial areas (Buxton 1936, 1938; Lang, 1975; Petrelli et al., 1980). Buxton (1938) found higher infestations in Kenya and Jerusalem in dry months as compared to wet months, but recorded opposite findings for a Nigerian population. Although climate is acknowledged as a contributing cause of these fluctuations, seasonal differences in human activities are gener- ally considered to play the major role in temporal changes of head lice popu- lations (Buxton, 1946; Lang, 1975; Busvine, 1985). Long term studies must be performed before the relative effects of these two factors can be deter- mined accurately. No significant differences in prevalence were found among the various elementary grades. Similar findings were noted in some studies (Grainger, 1980; Petrelli et al., 1980; Ewasechko, 1981), but several investigators (Mel- lanby, 1942a; Jalayer, 1967; Slonka et al., 1976; Slonka et al., 1977; Juranek, 1985) found a greater prevalence of head lice among younger than older children. Juranek (1985) suggested that this relationship was due to a greater frequency of head-to-head contact in and out of the classroom among youn- ger children. However, lower infestation rates were reported in younger than older children in Ghana (Kwaku-Kpikpi, 1982) and Nigeria (Ogunrinade and Oyejide, 1984). This condition was attributed to shorter hair and better hy- giene in the younger age groups, and to the practice of plaiting hair (in girls) in the older age groups. After being plaited the hair was not washed or combed for a week or more, providing a stable environment for head lice. The higher infestation rates of females in this study were similar to the findings of most investigators (Sobel, 1913; Mellanby, 1941; Jalayer, 1967; Hopper, 1971; Maguire and McNally, 1972; Donaldson, 1976; Grainger, 1980; Petrelli et al., 1980; Kwaku-Kpikpi, 1982; Sinniah et al., 1983; Ogunrinade and Oyejide, 1984; Juranek, 1985). However, in studies in Geor- gia (Slonka et al., 1976), Buffalo, New York (Slonka et al., 1977), and Al- berta, Canada (Ewasechko, 1981) no significant differences were found be- tween males and females in at least parts of the investigations. Longer hair lengths and elaborate hair styles of females have been the principal factors used to explain the sex difference in prevalence of head lice (Mellanby, 1941; Jalayer, 1967; Orkin et al., 1976; Kawaku-Kpikpi, 1982; Ogunrinade and Oyejide, 1984). The prevalence of infestation for boys and girls increased with increasing hair length. Similar observations were reported in Pennsylvania (Juranek, No. 4, 1989] PRICE AND BENITEZ— HEAD LICE 285 1985), Ghana (Kwaku-Kpikpi, 1982), Nigeria (Ogunrinade and Oyejide, 1984), and Malaysia (Busvine and Reid, 1949; Sinniah et al., 1983). Buxton (1938) found a positive correlation between hair weight and lice infestation in a tribe of Nigerians. In contrast, several studies (Slonka et al., 1976; Slonka et al., 1977; Petrelli et al., 1980; Ewasechko, 1981; Juranek, 1985) found no significant differences in attack rates based on hair length for either sex. Differences in attack rates based on hair length probably have no physiologi- cal basis, but may be due to the increased difficulty in grooming and treating long hair. Blacks were not infested with head lice even though they were at equal risk of infestation with other races. Previous studies in the U.S. found no or low frequencies of infestation in black children (Green, 1898; Sobel, 1913; Slonka et al., 1976; Slonka et al., 1977; Billstein and Laone, 1979; Juranek, 1985). Low attack rates in Blacks have been attributed to various physical characteristics of the hair (Ashcroft, 1969; Rasmussen, 1984), grooming methods (Sobel, 1913; Nuttall, 1918), hair length or weight, (Buxton, 1946), and cultural phenomena (Buxton, 1946; Gratz, 1985). It is interesting to note that several studies in Africa have found moderate to high prevalences among Blacks (Buxton, 1936, 1938; Iwuala and Onyeka, 1977; Kwaku-Kpikpi, 1982; Ogunrinade and Oyejide, 1984). White children had higher infestation rates than either Hispanic or Asian children in this study. Slonka and co-workers (1977) found that the attack rate of white children (22%) was twice that of other races (Puerto Ricans, Orientals, American Indians) in a city-wide survey of third to eighth graders in Buffalo, New York. Head lice were significantly more common among American Indians than Whites in a survey of elementary schools and junior high schools in rural Central Alberta, Canada (Ewasechko, 1981). In a pedic- ulicide study involving infested children in California (Billstein and Laone, 1979), the percentage of Hispanics was much greater than their representa- tion in the total population, while Whites were underrepresented. Higher prevalences were found among Malays and Indians than Chinese in two stud- ies in Malaysia (Busvine and Reid, 1949; Sinniah et al., 1983). These differ- ences were attributed to lower socioeconomic level and longer hair of Malays and Indians. The attack rate of head lice was significantly greater in larger than smaller families. Similar findings have been reported in the literature (Mel- lanby, 1942b; Slonka et al., 1976; Slonka et al., 1977; Petrelli et al., 1980: Kwaku-Kpikpi, 1982; Sinniah et al., 1983). Often, large family size is an indication of household crowding. The prevalence of pediculosis increased with increasing crowding index. This relationship has been found in other studies (Slonka et al., 1976; Slonka et al., 1977; Sinniah et al., 1983) and suggests that there is a correlation between attack rate and the number of family members per bedroom. In Malaysia (Sinniah et al., 1983) higher prevalences were recorded for children who shared a bed with another person than for those who slept alone. 286 FLORIDA SCIENTIST [Vol. 52 The data on household crowding suggest that infested elementary school children should be more likely to have other infested family members than noninfested students. This hypothesis was substantiated by findings from this and previous studies (Maguire and McNally, 1972; Slonka et al., 1976; Nitzkin, 1977; Slonka et al., 1977; Juranek, 1985). Almost 100% infestation rates for siblings of infested children were found in Dade County, Florida (Nitzkin, 1977). This phenomenon was attributed to failure to treat all family members on discovery of the initial infection. Using epidemiological informa- tion from four states, Juranek (1985) reported that 59% of infested children had at least one other infested household member as compared to 2% for uninfested children. Data from academic year 1986-87 indicated that a small number of fami- lies with infested schoolchildren accounted for over 40 % of the cases reported at the elementary school. Slonka and co-workers (1977) found that 9.1% of the families represented at an elementary school in Buffalo, New York were responsible for 62% of all cases recorded at the school. These findings strongly suggest that the family is a primary source of lice transmission out- side of the school. Results from this study suggested that a sizable proportion of the infested children had a history of repeated infestation. Slonka and co-workers (1977) reported that 31% of the infested children at an elementary school in Buf- falo, New York had a history of continuous or repeated infestation. Similar results were found for 89% of infested school children in Armagh, Ireland (Maguire and McNally, 1972). Infestation rates were significantly greater for students from the low so- cioeconomic level than for those from the high level. An increased prevalence of pediculosis in lower income groups has been reported in the literature (Slonka et al., 1976; Slonka et al., 1977; Billstein and Laone, 1979; Ewa- sechko, 1981; Sinniah et al., 1983). Juranek (1977) pointed out that the socio- economic level is not correlated with susceptibility, but is more a reflection of a person’s ability to deal with the problem once a family becomes infested. People from lower income groups may be unable to pay for proper medical care or have appliances (washers, dryers) that help to reduce infestations. Inability to provide individual toilet articles (combs, brushes, towels) to fam- ily members could increase fomite transmission of head lice. Children from lower income levels are not necessarily the source of infes- tations for higher levels. Infestation problems have been noted in public, private, and parochial schools at all socioeconomic levels, and can occur in children from upper-class families in the absence of children from lower in- come families (Rasmussen, 1984). The attack rate for students who rode buses was significantly greater than for students using all other types of transportation combined. Juranek (1985) presented similar findings for Pennsylvannia schools during an outbreak of head lice. He suggested that crowded seating conditions and frequent seat changes by children provided ample opportunities for head-to-head transmis- No. 4, 1989] PRICE AND BENITEZ— HEAD LICE 287 sion of lice. These same riding conditions exist in Hillsborough County buses. Indirect transmission of lice from infested seats is possible throughout the school year in west central Florida since freezing temperatures are rare in this area. Adult lice may survive nearly 50 hours at 4° C (Lang, 1975). Three classroom activities were linked to increased head lice infestation: children sitting and/or playing on floors; four or more children using a table; children using headsets. The first two activities suggest person-to-person transmission and probably are a result of head and body contact between students. Greater opportunities exist for student contact when they sit on the floor or crowd around tables than when they sit at conventional desks. Simi- lar results were reported for these activities in schools in Pennsylvania (Juranek, 1985) and Ghana (Kwaku-Kpikpi, 1982). The use of headsets was the only evidence suggesting transmission by a fomite. Since head lice can survive off the host for 48-55 hours at room temperature (22° C) (Juranek, 1985), transmission by headsets could be accomplished if used by different students during any 1-2 day period. The method of clothes storage at schools has been implicated in the trans- mission of head lice in previous studies. Sharing lockers (Hopper, 1971; Juranek, 1985), using unassigned wall hooks (Juranek, 1985), and piling hats and coats together on the floor (Nitzkin, 1977) have been suggested as major factors in the spread of head lice. In this study most students placed their garments under or on the backs of their chairs. This method seemed to sepa- rate clothing adequately and is one suggested by Juranek (1977) as a means of decreasing lice transmission. The results of this study indicate a considerable pediculosis problem among elementary schoolchildren in Hillsborough County. Coordinated ef- forts between school officials and parents involving prevention, screening, education, and treatment programs are necessary to control this annoying and costly public health problem. ACKNOWLEDGMENTS—We gratefully acknowledge the assistance of Mary Ellen Gillette, Elaine Diaz, and her administrative and teaching staff of the Hillsborough County Public Schools; Barbara Kinnee, Ruth Wyatt, and health support aides from the Florida Department of Health and Rehabilitative Services, Hillsborough County Public Health Unit; and volunteer stu- dent helpers from the University of Tampa. This work was supported by a University of Tampa Faculty Development Grant. LITERATURE CITED ALTSCHULER, D. Z., AND L. R. KENNEY. 1986. Pediculicide performance, profit, and the public health. Arch. Dermatol. 122:259-261. AsucrortT, M. T. 1969. Racial difference in Pediculus h. capitis infestation in Guyana, Trans. R. Soc. Trop. Med. Hyg. 63:547. BILLSTEIN, S., AND P. Laone. 1979. Demographic study of head lice infestations in Sacramento County school children. Int. J. Dermatol. 18:301-304. BusvineE, J. R. 1985. Pediculosis: treatments on the horizon. Pp. 231-237. In: Orkin, M., AND H. Maisacu (eds.) Cutaneous Infections and Insect Bites. Marcel Dekker, Inc., New York, New York. ., ANDJ. A. Rew. 1949. A simple remedy for head lice. Malaysia Med. J. 3:232-235. 288 FLORIDA SCIENTIST [Vol. 52 Buxton, P. A. 1936. Studies on populations of headlice (Pediculus humanus capitis: Anoplura). I. Parasitology 28:92-97. . 1938. Studies on populations of head-lice. (Pediculus humanus capitis: Anoplura). II. Parasitology 30:85-110. . 1946. The Louse, an Account of the Lice Which Infest Man, Their Medical Impor- tance and Control. Williams and Wilkins, Baltimore, Maryland. Dona .pson, R. J. 1976. The head louse in England: prevalence amongst schoolchildren. J. R. Soc. Health 96:55-57. Ewasecuko, C. A. 1981. Prevalence of head lice (Pediculus capitis |DeGeer]) among children in a rural, Central Alberta school. Can. J. Pub. Health 72:249-252. GRAINGER, C. R. 1980. Pediculus humanus capitis on children in Mahe, Seychelles. Trans. R. Soc. Trop. Med. Hyg. 74:296-299. Gratz, N. 1985. Epidemiology of louse infestations. Pg. 187-198. In: Orkin, M., AND H. MAILBACH (eds.), Cutaneous Infections and Insect Bites. Marcel Dekker, Inc., New York, New York. GreEEN, E. M. 1898. Pediculosis in Boston’s public schools. Boston Med. Surg. J. 138:70-71. Hopper, J. M. H. 1971. An epidemic of nits. Can. J. Public Health 62:159-160. Iwuaa, M., AND J. OnyekKA. 1977. The incidence and distribution of head lice Pediculus hu- manus var capitis (Insecta, Anoplura) in primary and post-primary school pupils in Nsukka, East Central State, Nigeria. Niger. Med. J. 7:174-283. JALAYER, T. 1967. Head louse infestation in villages of Shiraz, Iran. J. Parasitol. 53:216. JuRANEK, D. D. 1977. Epidemiology of lice. J. Sch. Health 47:352-355. . 1985. Pediculus capitis in school children. Epidemiologic trends, risk factors, and recommendations for control. Pp. 199-211. In: Orkin, M., AND H. Marsacu (eds.), Cuta- neous Infections and Insect Bites. Marcel Dekker, Inc., New York, New York. Kwaku-Kpixp1, J. E. 1982. The incidence of the head louse (Pediculus humanus capitis) among pupils of two schools in Accra. Trans. R. Soc. Trop. Med. Hyg. 76:378-381. Lanc, J. 1975. Biology and control of the head louse, Pediculus humanus capitis (Anoplura: Pediculidae), in a semi-arid urban area. Ph.D. thesis. University of Arizona. MaculrE, J., AND A. McNALLy. 1972. Head infestation in school children: extent of problem and treatment. Community Med. 128:374-375. MELLANBY, K. 1941. The incidence of head lice in England. Med. Officer 65:39-43. . 1942a. Natural population of the head louse (Pediculus humanus capitis: Anoplura) on infected children in England. Parasitology 34:180-184. . 1942b. Relation between size of family and incidence of head lice. Public Health 56:31-32. Nitzkin, J. L. 1977. Pediculosis capitis. J.A.M.A. 237:530. NutTAa.u, G. H. F. 1918. The biology of Pediculus humanus. Parasitology 10:80-185. OcUNRINADE, A., AND C. Oyeyine. 1984. Pediculosis capitis among rural and urban schoolchil- dren in Nigeria. Trans. R. Soc. Trop. Med. Hyg. 78:590-592. Orkin, M., E. Epstern, AND H. I. Marpacu. 1976. Treatment of todays scabies and pediculosis. J. Am. Med. Ass. 236:1136-1139. PETRELLI, G., G. Majorit, M. Macinni, F. Tacci, AND M. Maro.ut. 1980. The head louse in Italy: an epidemiological study among school children. J. R. Soc. Health 100:64-66. RASMUSSEN, J. E. 1984. Pediculosis and the pediatrician. Pediatr. Dermatol. 2:74-79. RoBInson, D., AND D. SHEPHERD. 1980. Control of head lice in schoolchildren. Curr. Ther. Res. 27:1-6. SINNIAH, B., D. SINNIAH, AND B. Rayeswari. 1983. Epidemiology and control of human head louse in Malaysia. Trop. Geog. Med. 35:337-342. StonKA, G. F., T. W. McKInLey, J. E. McCroan, S. P. Sinciair, M. G. ScHuLTz, F. Hicks, AND M. Hix. 1976. Epidemiology of an outbreak of head lice in Georgia. Am. J. Trop. Med. Hyg. 25:739-743. ., M. L. FLEISSNER, J. BERLIN, J. PuLEo, E. K. Harrop, AnD M. G. Scuuttz. 1977. An epidemic of Pediculosis capitis. J. Parasitol. 63:377-383. SoBEL, J. 1913. Pediculosis capitis among school children. N. Y. Med. J. 98:656-664. Florida Sci. 52(4):278-288. 1989. Accepted: December 27, 1988. Atmospheric Sciences AN INVESTIGATION OF THE METEOROLOGICAL CONDITIONS AFFECTING DISPERSION OF OZONE IN THE TAMPA BAY REGION Dewey M. STowers, JR. AND NEVA DUNCAN TABB Department of Geography, University of South Florida, Tampa, FL 33620-8100 Asstract: During the last decade, the ozone levels of both Hillsborough and Pinellas counties have significantly increased. Although each county is primarily urban and in close proximity, Hillsborough County has traditionally experienced a greater number of exceedences of the Fed- eral standard (120 ppb). An investigation was undertaken to determine the effects meteorological elements, principally wind velocity, surface temperatures, and the seabreeze phenomenon, have on the mass transference of ozone in the region. In addition, the continual expansion of urban areas and the resultant increase of vehicular traffic were also considered. The results of this research indicate that the impending ozone crisis is a regional rather than an individual county problem. A GRADUAL decline in the levels of harmful pollutants in the air over major urban areas has been occurring since the passage of the original Clean Air Act more than two decades ago. Among these pollutants, ozone remains one of the most difficult to control, primarily because it is formed in a complex series of reactions involving both weather and emission sources. Although ozone has declined 10% nationally, the Environmental Protection Agency estimates at least 80 million Americans, approximately one third of the United States population, are still being exposed to levels exceeding the Federal air quality standard of 120 ppb. EPA monitoring of 84 major metropolitan areas in 1982 revealed 32 areas, including Tampa Bay, failing to meet the Federal standard (Raloff, 1986). Despite a five year extension period, many regions, including Tampa Bay, were still unable to bring their ozone levels down to the standard by the end of 1987. The combination of optimum weather conditions and abundant emission sources in the Tampa Bay area illustrates the difficulty many regions face in trying to control ozone levels. Ozone is formed by a series of reactions involving nitrogen oxides and hydrocarbons in the presence of sunlight. The major source of these two chemicals can be traced to the heavy vehicular exhaust generated in rapidly- growing urban populations. In addition to vehicular exhaust, nitrogen oxides are also emitted by electric utility and industrial boilers. Hydrocarbons enter the atmosphere from gasoline stations, handlers, and transporters, and by paint and dry-cleaning solvents (DEM, 1980). The simplified reactions involved in the formation of ozone are illustrated (Fig. 1). Atmospheric nitrogen and oxygen disassociate during combustion and form nitric oxide (NO). Nitric oxide then gradually interacts with oxygen to form nitrogen dioxide (NO,). In the presence of high solar insolation, nitro- gen dioxide breaks down to form nitric oxide and atomic oxygen, the latter of 290 FLORIDA SCIENTIST [Vol. 52 which can combine with the atmospheric oxygen to form ozone. Ozone usu- ally requires from 15 minutes to three hours to form. Normally, ozone would interact with any nitric oxide but if hydrocarbons are present they will com- pete with the ozone for the nitric oxide. This allows the ozone concentration to build (DER, 1985). Highly industrialized subtropical areas where stagnant air masses can prevail offer an ideal environment for ozone formation due to maximum solar insolation and minimal mixing of the atmosphere. FORMATION OF OZONE N+ 0,-NO +O O + N,—-NO + N 2NO + 0, ~2NO, NO, = NO + O Fic. 1.Chemical formation of ozone. Analysis of the chemical reactions which generate ozone reveals the im- portance of ultraviolet radiation in instigating the formation of atomic oxy- gen which, when exposed to molecular oxygen present in the atmosphere, produces ozone. Both latitudinal location and the presence of high-pressure systems influence the amount of solar radiation reaching the surface. High pressure areas are usually accompanied by clear skies, allowing for greater absorption of ultraviolet radiation. In addition, high pressure areas often create stagnant atmospheric conditions, particularly in autumn and spring when convection is not as well developed as in summer. These stagnant air conditions are also conducive to increased levels of ozone production because of reduction in wind speed. Lower wind speeds decrease atmospheric dilu- tion, allowing for greater accumulation of nitrogen oxides and hydrocarbons. No. 4, 1989] STOWERS AND TABB— OZONE 291 500mb HEIGHT CONTOURS APRIL 24, 1987 7:00 A.M. SCALE OQ 200 300400 KILOMETERS Fic. 2. A 500 mb map showing location of a typical double high pressure pattern, April 24, 1987. Recent research (Chu, 1986) has shown the role of a double high-pressure system weather pattern in influencing ozone concentrations in Florida (Fig. 2). The combination of a stalled high pressure system at 500 mb in the Gulf and a seasonal oceanic high in the Atlantic creates an extremely stable air layer between the two highs, leading to the formation of a temperature inver- sion at the surface and resulting in a significant increase in ozone concentra- tions. Research indicates 87% of the observed exceedences in Florida be- tween January, 1981 and June, 1986 occurred in the presence of these double high-pressure systems, most commonly in the spring and autumn. The Tampa Bay area occupies a favorable location for the formation of high concentrations of ozone. Over the five year period 1981-1986, only this region in the state showed a recognizable increase of 10-50% in the monthly mean concentration of ozone (Chu, 1986). In addition to the subtropical location (latitude 28° N), the proximity of this area to the Gulf and Tampa Bay encourages greater ozone production due to the high reflectivity of water and the resultant increase in ultraviolet radiation over the water (Hessling, 1987). This rapidly-growing area contains numerous emission sources al- though the heavy vehicular traffic is considered the principal polluter. The study area shows a strong seasonal cycle of ozone formation, peaking in the spring, and to a lesser degree, the fall, when high pressure systems dominate. Despite the increased ultraviolet radiation in the summer months, strong 292 FLORIDA SCIENTIST [Vol. 52 convective activity in conjunction with increasing cloudiness and rain tended to dampen ozone formation. However, it is the westerly sea breezes and their effect upon ozone exceedences in the study area that will be most closely examined in this study. MetHops— The Tampa Bay area airshed is the focus for this study. The Federal government considers Pinellas and Hillsborough counties to share this airshed, a region in which emissions are shared and measured collectively, although separate agencies oversee each county. Six sites in the study area, three in Pinellas and three in Hillsborough, continually monitor ozone concentrations (Fig. 3). The three Pinellas sites are Azalea Park, St. Petersburg Junior College (Clearwater campus), and Brooker Creek Park. The three Hillsborough sites are located at Beach Park, Davis Island, and Simmons Park. BAY AREA OZONE MONITORING NETWORK PASCO CQ. __ o g feamen! eBrooker Creek \ Batis Island e Beach) Park Isles : | \ - ae e Simmons’ Park | 0 aes iefaseeuxn cael oe ws 2 if — 2 —_.. Mls, Oo = < Babat! = ade: oO Us a = © a \¢ # Q ol ao tela ope aaa SCALE Oo 3 (km) Fic. 3. Location of ozone monitoring sites in Hillsborough and Pinellas counties. No. 4, 1989] STOWERS AND TABB—OZONE 293 In recording ozone concentrations, Pinellas County utilizes an infrared electronic analyzer, whereas Hillsborough County relies on ultraviolet photometry. Both methods are EPA-approved and allow for continuous monitoring, and data are recorded as a series of one-hour average concentration values. The Tampa Bay area is in nonattainment of the Federal standard if read- ings are reported exceeding the standard (120 ppb) an average of more than once per year over three years. In addition to ozone readings, weather data on the days of the ozone exceedences were also examined. Temperature, humidity, barometric pressure, and rainfall readings were recorded at the University of South Florida weather station. Wind data were collected at Brooker Creek Park in Pinellas County and the University of South Florida in Hillsborough County. In 1987, an unusually high number of exceedences was recorded in the Tampa Bay region compared to previous years. Exceedences were reported on six days: April 23, 29, and 30; July 24 and 25; and August 7 (Novak, 1987). All of these exceedences occurred late in the work week after several days of heavy vehicular traffic. Hillsborough County recorded exceedences at one or more sites on each of the six days, whereas Pinellas County only recorded one exceedence during this time period, on July 24. An analysis of the wind data for these six days, however, reveals a closer relationship might exist than has previously been suspected. Discusslon—The mass transference of ozone by light surface winds is a recognized occurrence today by the researchers of air pollution (Altshuller, 1986). Although ozone can deteriorate readily during the transfer process, meteorological conditions such as subsidence aloft, temperature inversions, and the presence of large bodies of water tend to retard the process. There exists a complex formula to evaluate this breakdown which involves the use of tethered ballons to observe wind velocity above the surface. The use of these balloons in the Tampa Bay region is prohibited by the F.F.A. due to the close proximity of five heavily-used airports. Therefore, a reasonable percentage of the transfer of ozone by winds under 10 km/hr had to be established. After a discussion with Dr. Hans Neuberger, a recognized authority in air pollution studies and former chairperson of the Department of Meteorology at Pennsyl- vania State University, a figure of 25% was deemed appropriate for this study. During the five year period 1981-1986, 24 exceedences were recorded in Hillsborough County. The highest reading in Hillsborough County (145 ppb) occurred on July 15, 1983, at the Davis Island site. The remaining 19 ex- ceedences during this period ranged from 125 ppb to 135 ppb and occurred predominately during the months of April, May, June, and September. In 1987, the highest exceedences of the period occurred when four exceedences in excess of 150 ppb were recorded. A contributing factor to this situation was a shift in the usual summer wind pattern over the region (Stowers and Tabb, 1987). This shift, first observed in the late 1970s, replaced the usual easterly winds with a southwesterly flow, allowing for the mass transference of ozone across Tampa Bay. FLORIDA SCIENTIST [Vol. 52 294 “L861 ‘WAC ‘L861 “Oda = 991N0Se oOVE ~ 08 VO = 16 oar a = VOD vol Vv a Ere lL Leh | L8-L°8 dAOQ® SB UIeS dAOQR SB OWLS ,Gran = GO16 Qc] I yileg yorog 009 BBE Chips € 006 SC F'8 ILT I 212d SUC TCS L8-VG-L aA0qe Sv JUIeS dAOq” Sv oUIeS 406... = SEO Sal P yrleg yorog o0GE ~— 96 98 = & o0€ . =—I¢ € Lol Vv Spe SS rG o0GE ~— S96 901 - F ov = IGE Tél 9 TE SUC L8-€6-L 0866 — S96 ne ae) 0096 — =O9'T col I [28 (suo taarsS L8-0&-P oFlE ~ 068 (AC ae ae | ofll€ ~— OOT [St G 2 keke | EATON SS) L8-66-P ocLG ~ S08 699 = ~cl oL96 ~— SVY 661 EOE? at L8-E6-V qdd ‘urd qdd ‘urd sooisoq IY /wWy SUIpeay ‘OUIL], sdoisoq 1Y4/Un suIpeoy ‘OUT ], (Yoe1D 19x001g) (y1eg volezy) Ayuno’) Ayunor) Ayunor) JOUIIINIIO AJUNOT) SeTOUTg AJUNOT) SP][OUTY ysno10qgsy [iH ysno10qgs| [iH ysno10gs| tH jo :A}OOTAA, PUTA, :AYOOTAA, PUTA, :UOlB00'T a3eq e(L86T nsny-[ldy) varie Avg edWey, a4} Ul S9OUIPIdOXa JUOZO IO} BILP [BOIBO]OIODJOJ ‘| ATAV], No. 4, 1989] STOWERS AND TABB—OZONE 295 A total of six ozone exceedences was recorded in the Tampa Bay region during 1987 (Table 1). The first exceedence was recorded on Thursday, April 23, at the Beach Park station in Hillsborough County. The highest reading, 129 ppb, occurred at 1 p.m. under a 6.5 km/hr wind from 267 degrees. One hour earlier, a reading of 69 ppb was recorded at Azalea Park in Pinellas County under a wind of 8.1 km/hr from 272 degrees. Using the transfer factor of 25%, a figure of 17.25 ppb is calculated. This figure subtracted from 129 ppb recorded at Beach Park would result in a non-exceedence read- ing of 111.75 ppb at Beach Park without the mass transference from Pinellas County. This procedure is duplicated for the remaining exceedences. On April 29, 1987, a record exceedence of 151 ppb was recorded at Sim- mons Park, Hillsborough County, at 2 p.m. under a 1.6 km/hr wind from 311 degrees. At the Azalea Park site a recorded reading of 114 ppb during a 8.9 km/hr wind from 314 degrees was observed. This results in a mass transfer- ence factor of 28.5 ppb. Utilizing this percentage of transfer, the resultant figure at Simmons Park is 122.5 ppb. Although this reading is high, it is still slightly below the exceedence standard. A third exceedence occurred at Simmons Park, Hillsborough County, on April 30, 1987 at 1 p.m. with a reading of 125 ppb under wind velocity 1.6 km/hr at 260 degrees. While this exceedence met Federal standards, the transfer factor from Pinellas County makes the actual reading at Simmons Park much lower. On this date, Azalea Park recorded a reading of 77 ppb with a wind velocity of 9.7 km/hr from 298 degrees. Applying the transfer factor, the resultant figure is 19.25 ppb; therefore, the actual site reading is 105.75 ppb, significantly below the Federal standards. On July 23, 1987, exceedences were recorded at all three sites in Hillsbo- rough County (Simmons Park, Davis Island, and Beach Park). The highest reading of 142 ppb occurred at 4 p.m. at Beach Park with a wind velocity 6.5 km/hr from 20 degrees. Correspondingly, the Pinellas County site recorded 86 ppb at 3 p.m. under a wind of 9.7 km/hr from 320 degrees. The mass transference at this time would be 21.5 ppb. This reduces the on-site reading at Beach Park to 120.5 ppb which, although again a high reading, does not exceed the standard. The situation reversed on July 24, 1987 at 1 p.m. with an exceedence occurring at Azalea Park in Pinellas County. The reading at this site was 142 ppb with a wind velocity of 3.22 km/hr from 60 degrees. Two exceedences were recorded in Hillsborough County at 12 p.m. on the same date (Simmons Park and Beach Park). The highest reading of 140 ppb occurred at Simmons Park with a wind velocity of 8.0 km/hr from 20 degrees. This NNE wind provided a transfer factor of 35 ppb from Simmons Park and a factor of 31.75 ppb from Beach Park to Azalea Park in Pinellas County. Applying these trans- fer numbers to Azalea Park results in non-exceedence readings of 107 ppb and 110.25 ppb respectively. Unlike the previous dates of exceedence, Hillsbo- rough County did experience an on-site exceedence. Also in the previous dis- cussions the transfer was from Pinellas County to Hillsborough County. On this date, the transfer was reversed. 296 FLORIDA SCIENTIST [Vol. 52 The last exceedence to occur in the Tampa Bay area in 1987 was recorded at the Beach Park site on August 8 at 4 p.m. At this time Beach Park recorded a reading of 164 ppb with a wind velocity of 6.0 km/hr from 45 degrees. At Azalea Park at 2 p.m. a reading of 94 ppb with a wind velocity of 8.0 km/hr from 340 degrees was recorded. Applying the transference factor of 23.5 ppb still leaves Beach Park in exceedence with an adjusted reading of 140.5 ppb. Consequently, of the six ozone exceedences which occurred in Hillsbo- rough County, four appeared to be the result of mass transference of ozone by light west-southwest winds from Pinellas County. The two exceedences which did occur would have been far less severe had transference not oc- curred. Conversely, the only ozone exceedence recorded in Pinellas County (July 24, 1987) appeared to be the direct result of transference by northeast winds from Hillsborough County. An examination of the data for several days both before and after the six exceedence days offers additional support for the argument of mass transfer- ence of ozone. For example, on Friday, April 24, maximum ozone readings in Hillsborough County ranged from 78 ppb at Davis Island to 98 ppb at Sim- mons Park, with a maximum of 80 ppb at Azalea Park. A cold front which had passed through earlier in the day was most likely responsible for bringing in cleaner air and lowering ozone readings in comparison to the highs of the previous day (Fig. 4). However, the highest reading in Hillsborough County BBEAN SEA xs > cARI SURFACE PRESSURE CHART APRIL 24 1987 7:00 A.M. Z SCALE et pace © 200 300400 wee Vie ———————— = 60 75 KILOMETERS jl 70 Fic. 4. The passing of a cold front, April 24, 1987. No. 4, 1989] STOWERS AND TABB—OZONE 297 (Simmons Park) occurred shortly after high readings in Pinellas and under a northwesterly breeze. Azalea Park is located northwest of the Simmons Park station. Following two days of exceedences, highest readings on Saturday, April 25, occurred in the early morning hours, an unusual phenomenon. In Hillsborough County, the highest reading was recorded at Simmons Park, 97 ppb. An hour earlier, Azalea Park reached its highest reading of 82 ppb. Again, a northwesterly wind was present. Another example can be seen on July 22, when highs in Hillsborough County ranged from 55 ppb at Simmons Park to 111 ppb at Beach Park anda maximum of 97 ppb at Azalea Park. Although the high readings in both counties occurred at approximately the same time, the ozone levels at Beach Park had started to rise an hour earlier and a northeasterly wind transported ozone to the southwest from Beach Park to Azalea Park. Following two days of ozone exceedences, highest readings on July 25 were 76 ppb (12 p.m.) in Pinellas County and 89 ppb in Hillsborough County at Simmons Park (11 a.m.). The southeasterly breeze present at that time transported ozone across the bay to Azalea Park. ConcLusions— Ozone is considered to be one of the most damaging air pollutants threatening the urban air over American cities today. Unfortu- nately, it is also considered to be one of the most difficult to control. Although much blame may be attached to the prevalence of combustion engines, cer- tain meteorological conditions, such as subsidence aloft resulting from dou- ble-highs, can intensify the concentration levels. In addition, documentation exists for the mass transference of ozone from one region to another under proper wind conditions (Altshuller, 1986). Therefore, all of these factors must be considered when determining why an exceedence of the ozone standard occurred in a particular region. Traditionally, air quality in both Pinellas and Hillsborough Counties has been handled by separate local agencies: the Pinellas County Department of Environmental Management in Pinellas County and the Environmental Pro- tection Commission in Hillsborough County. A careful examination of the meteorological conditions on the six days on which exceedences occurred in 1987 provides evidence for transference of ozone across the bay. In conjunc- tion with the low wind speed and an advantageous direction, the mass trans- ference that resulted was sufficient to elevate opposing stations to exceedence levels. This research indicates ozone pollution in west central Florida cannot be simply considered a county problem but a regional one. Therefore, the most beneficial action for those living in this airshed would be: 1) to encourage cooperation between Hillsborough and Pinellas counties; 2) to consider form- ing a regional EPA to service both areas; and 3) to ensure that extensive records are kept on both ozone levels and meteorological conditions in both areas. This type of cooperation would provide the greatest opportunity for averting a major ozone crisis in this area. 298 FLORIDA SCIENTIST [Vol. 52 LITERATURE CITED ALTSHULLER, A. 1986. The role of nitrogen oxides in nonurban ozone formation in the planetary boundary layer over N. America, W. Europe, and adjacent areas of the ocean. Atmos. Sci. 20:245-268. Cuu, S.-H. 1986. Couplings of High Pressure Systems and Outbreaks of High Surface Ozone Concentrations. Bureau of Air Quality Management, Department of Environmental Reg- ulation. Tallahassee, Florida. DEPARTMENT OF ENVIRONMENTAL MANAGEMENT. 1980. Air Quality 1979: Annual Report. Clear- water, Florida. DEPARTMENT OF ENVIRONMENTAL MANAGEMENT. 1987. Hourly ozone readings and hourly wind tables, 1987. Division of Air Quality. Clearwater, Florida. DEPARTMENT OF ENVIRONMENTAL REGULATION. 1986. Ambient Air Quality in Florida 1985. Bu- reau of Air Quality Management. Tallahassee, Florida. ENVIRONMENTAL PROTECTION ComMIssION. 1987. Hourly ozone readings and hourly wind tables. Hillsborough County, Florida. HeEss.inc, P. 1987. Division of Air Quality, Clearwater, Florida. Interview, 30 March. Novak, L. 1987. Environmental Protection Commission, Tampa, Florida, Interview, 19 March. Ravorr, J. 1986. Smog’s Ozone: EPA wants more controls. Science News, 129:405. Stowers, D. M., AND N. D. Tass. 1987. An investigation of the variances from the traditional summer precipitation in the west-central Florida region (1978-1985). Florida Sci. 50:177-183. Florida Sci. 52(4):289-298. 1989. Accepted: January 30, 1989. No. 4, 1989] AWARDS 299 Outstanding Student Paper Awards, Awardees Fifty-Third Annual Meeting of the Florida Academy of Sciences Florida Community College at Jacksonville— March 30-April 1, 1989. AGRICULTURAL SCIENCE—Mr. J.F. Holderbaum, Department of Agronomy, University of Flor- ida, “Nutrient composition of stock-piled bermudagrass as affected by fertilizer nitrogen rate and method of application.” ANTHROPOLOGICAL SCIENCE— Ms. Anna M. Estes, Department of Anthropology, Florida State University, “Assessment of cortical thickness by computer tomography in an early Archaic skele- tal sample from Windover (8Br246).” ATMOSPHERIC AND OCEANOGRAPHIC SCIENCES— Ms. Haejung An, Department of Food Science and Human Nutrition, University of Florida, “Immunological identification of marine species using the rock shrimp (Sicyonia brevirostris) as a model.” BIOLOGICAL SCIENCE—Mr. Brian Bowman, Division of Science, University of Tampa, “Isola- tion and characterization of the enzyme glutathione S-transferase from two rotifer species,’ Un- dergraduate Award. Mr. Chase C. Smith, Natural Sciences, Eckerd College, “A SEM study of pollen exine devel- opment in Hibiscus tiliaceus,” Undergraduate Honorable Mention. Ms. Karen R. Lips, Archbold Biological Station, “Vertebrates associated with gopher tortoise (Gopherus polyphemus) burrows in four habitats in south-central Florida,’ Graduate Award. Ms. Mary Fahning Brooks, Department of Biological Sciences, Florida Atlantic University, “Notes on hippolytid cave shrimps in the Bahamas (Crustacea: Hippolytidae) ,’ Graduate Honor- able Mention, also receiving The Explorers Club Award. COMPUTER SCIENCE AND MATHEMATICS—Mr. Jay Stryker, Department of Mathematics/Com- puter Science, Stetson University, “A new geometrical method of generating prime numbers,” Undergraduate Award, also receiving the AAAS Award. Ms. Lynn K. Stevens, Operations Research Program, Florida Institute of Technology, “Models for course scheduling,’ Graduate Award. ENVIRONMENTAL CHEMISTRY—Ms. Karen E. Wright, Millar Wilson Laboratory for Chemical Research, Jacksonville University, “Determination of formaldehyde in urban atmosphere,” Un- dergraduate Award, also receiving the AAAS Award. Ms. M. Christine Flynn, Department of Chemistry, University of South Florida, “Inhibition of the red tide organism, Ptychodiscus brevis, by the green alga, Nannochloris oculata, Graduate Award. PHYSICAL AND SPACE SCIENCES—Mr. Ragaiy Zidan, Department of Physics and Space Sci- ences, Florida Institute of Technology, “Parameters affecting the electromigration phenomenon.” AMERICAN ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE AWARD— Mr. Jay Stryker (see Com- puter Science and Mathematics above). Ms. Karen E. Wright (see Environmental Chemistry above). UNIVERSITY OF FLORIDA CHAPTER, SIGMA X1 AwARD—Ms. Haejung An (see Atmospheric and Oceanographic Sciences above). CENTRAL FLORIDA CHAPTER, THE ExpLoRERS CLusB AwARD—Ms. Mary Fahning Brooks (see Biological Science above). THE Vice ADMIRAL WILLIAM W. BEHRENS, JR./FLORIDA INSTITUTE OF OCEANOGRAPHY AWARD— Mr. Robert A. Frease, Department of Oceanography and Ocean Engineering, Florida Institute of Technology, “Polycyclic aromatic hydrocarbons associated with an artificial reef constructed of coal and oil ash waste products,’ Atmospheric and Oceanographic Sciences Section. We are grateful to Dr. Richard L. Turner, Chairman, Florida Academy of Sciences Awards Committee, for compiling this information. ERRATUM Through a remarkable oversight, an error was made in: D. R. Richardson. 1988. The sand pine scrub community: An annotated bibliography. Florida Scientist. 52(2):65-95. Paragraphs 4-6 on p. 66 were repeated on pp. 67-68. The Editor assumes full responsibility and extends a personal and a profes- sional apology to the author and to the readers. 300 FLORIDA SCIENTIST [Vol. 52 REVIEW Lawrence H. Keith and Douglas B. Walters (Editors). Compendium of Safety Data Sheets for Research and Industrial Chemicals (volumes IV, V, VI) VCH Publishers, Deerfield, Florida; New York, 1987 pp. 1863-3559, $295 This series continues the useful series that was reviewed previously [Flor- ida Scientist, 49(4):191-192 (1986)]. Material Safety Data Sheets (MSDS) pro- vide the range of useful information that those affected by worker-right-to- know must have, either as employer or employee. All of us who work with substances or who are affected by them in our daily lives will find this infor- mation useful, and, in some instances, absolutely essential. How useful and how valuable will, of course, depend upon the individual. This information, it must be added, is available from other sources and in other forms. A well known chemical firm provides a collection of MSDS on PC-compatible com- puter disc for $1,300, and while there may be more substances, the selection may be greater than most of us would need, or care to pay for. Another disc is available from the Canadian Centre for Occupational Health and Safety; it is updated quarterly for $100, but the information is rather more specialized (herbicides are heavily emphasized). And in both instances, it is given that appropriate computers are available. The Compendium has several features that are attractive. Thoroughness of coverage, general and specific references, error-free copy, thorough defini- tion of terms, remarkably useful index to entries in all six volumes, by prefer- red name, by synonym, by Chemical Abstracts Registry Number, and by molecular formula. The price is reasonable, considering what is available. A total of 1607 substances is listed (740 in this set, 867 in the previous three volumes) and the listings are carefully prepared and thorough. Volume VI contains a complete index; each volume contains the fifty references that were used to compile the information, thus providing useful background for those who wish to add additional compounds. The choice of compounds is always a matter of potential disagreement: Many substances on the EPA List of Extremely Hazardous Substances will be found here, some will not be. Most of the substances selected have been se- lected for study by the National Toxiocology Program, and that may mean that they are suspected of being potentially hazardous. That surely does not apply to all, however, including agar, which is a fairly common material that is usefully included. The current three volumes of the Compendium maintain the high stand- ards previously established, and the information provided is notably useful. These volumes are recommended to all who have need of this significant information—Dean F. Martin, University of South Florida, Tampa Florida Scientist QUARTERLY JOURNAL of the FLORIDA ACADEMY OF SCIENCES VOLUME 52 DEAN F. MARTIN Editor BARBARA B. MarTIN Co-Editor Published by the FLORIDA ACADEMY OF SCIENCES, INC. Orlando, Florida 1989 The Florida Scientist continues the series formerly issued as the Quarterly Journal of the Florida Academy of Sciences. The Annual Program Issue is published independently of the journal and is issued as a separately paged Supplement. Copyright© by the FLoripa ACApEmy OF SCIENCES, INc. 1989 CONTENTS OF VOLUME 52 NuMBER 1 Response of Laying Hens to Various Grains and Pease; S Moo lemneNtatlOM eae mie. atocacnes oats dorel + Mh aeivtoge se ges 48 ncere R. D. Miles, J. E. Marion, R.D. Barnett, N. Ruiz, and R. H. Harms First Record of Pawpaw Consumption by the Florida Mouse ............. Cheri A. Jones Atlantic White Cedar (Chamaecyparis thyoides) in the Southern SUDUSS 6 ated beech ic ca Daniel B. Ward and Andre F. Clewell Water Quality Efficiency of an Urban Commercial Wet Detention Stormwater Management System at the Boynton Mall in Sw telecine peach: COUMLYOE LOG a. § sare sepey- or cy open tvatiy hs Rave cuca cucoant i hhaee & apo Jeffrey Dee Holler Allocation of Energy Resources in the Freshwater Angiosperms Vallisneria americana Michx. and Potamogeton pectinatus L. in Florida ........... Clinton J. Dawes and John M. Lawrence pekmomuledomentsOl WeViCWEIS 2... 26 ce ee ce eee tase cence NuMBER 2 The Sand Pine Scrub Community: An Annotated Bibliography ........... Donald R. Richardson Calculated X-Ray Data Aid in Collecting High Quality X-Ray Power Data—Oxytetracycline Dihydrate, C,,H,,N,O,¢2H,O), asan Example... Frank N. Blanchard Bostrichobranchus diagonas: Confirmation of its Presence in the SASL GLE LUC O aig eh Oe ee eee a Gerald E. Walsh Community Waste to Energy System Technologies (CWEST) ............. Alex E. S. Green ine istmbution of the Tiustles of Floridas. 2. ya oe ka ee ee ee John B. Iverson and Cory R. Etchberger NUMBER 3 Boosting Science Careers for the Physically Eindchicap pea StiGent.....k 2c doce wuss «4 kewe sone ys Luretha F. Luckey Fate of Satellite-Tracked Buoys and Drift Cards of the Southeastern Atlantic Coast of the United States..................-.-- George A. Maul and Nicholas J. Bravo The Abundance of Aeromonas hydrophila L. at Lake Harney on the St. Johns River with Respect to Red Sore Disease in Sempped Mullet(iMusil cephalus 1G. \ ne ees hcs es ce eee ct eee ees John A. Osborne, Gerald E. Fensch, and Julius F. Charba The Occurrence of the Crayfish Fallicambarus (Creaserinus) fodiens SPMLSIEIC le ey ews... ee eee eters, Wel pk tes h5 aid 3 Barry W. Mansell What is the Impact of Engineering on Florida’s Ecomony— TAG AS? B57! Wop woratonaes ob oath, oo GOL 4 ao eee ee Betty Preece Symposium: “Groundwater Contamination and Protection in ouiel EEE rt i ee a a AS ae ah Pain yore daw al» S Emerging Legal Issues in Groundwater Contamination Cases ............ Thomas J. Guilday and Ralph A. DeMeo 48 58 65 94 100 104 LS, 145 154 171 lad Lae 182 183 Laboratory Models for Assessing the Fate of Groundwater Contaminants. 0c rere ree Joseph J. Delfino, Patricia V. Cline, and Carl J. Miles Ground-Water Contamination Programs of the U.S. Geological Survey in Florida... 0.06.66 6 cues ob Oe Onl 6 oe Gee eer Irwin H. Kantrowitz Pesticides and Ground Water Protection ................ Charles C. Aller NUMBER 4 Symposium: “Groundwater Contamination and Protection in Florida (cont’d.)” 6.80 ae ee ee Prevention and Cleanup of Petroleum Contamination of Groundwater—Florida’s Super Act .............:.5..50:)) sepeeeee Craig Ash, Connie Garrett, and Susan Gray The Status of Superfund and State-Funded Cleanup Sites in Florida 2.00. eb SY Aas So ee oe ore ee Impact of Groundwater Contamination on Public Water Supplies « .2.. sse-eees source eee sea ee ae ee J. Edward Singley Regular Contributions: The Neural Derivative, OCOS and Motion Detection ................. Arthropods Endemic to Florida Scrub. « ..< 0 os.) «4505s le Productivity of Departments of Chemistry at Florida Graduate Institutions, ss. s..aehed ia as eee a idles So ee eee John C. Follman and Dean F. Martin Infestation and Epidemiology of Head Lice in Elementary schools:in Hillsborough County, Florida... .. 25... .2-2225. 2 eee W. Wayne Price and Amparo Benitez An Investigation of the Meteorological Conditions Affecting Dispersion of Ozone in the Tampa Bay Region ................4 sce Dewey M. Stowers, Jr. and Neva Duncan Tabb ROEVICW hase kd aa 8 le wore Hale. DAE ee 207 214 220 225 225 230 240 244 254 271 278 289 299 300 301 INSTRUCTIONS TO AUTHORS Individuals who publish in the Florida Scientist must be active members in the Florida Academy of Sciences. Submit a typewritten original and two copies of the text, illustrations, and tables. All type- written material—including the abstract, literature citations, footnotes, tables, and figure legends — shall be double-spaced. Use one side of 8’ x 11 inch (21% cm X 28 cm) good quality bond paper for the original; the copy may be xeroxed. Margins should be at least 3 cm all around. Number the pages through the Literature Cited section. Avoid footnotes and do not use mimeo, slick, erasable, or ruled paper. Use metric units for all measurements. 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Florida's Estuaries — Management or Mismanagement? — Academy Symposium FLoripa SCIENTIST 37(4) — $5.00 Land Spreading of Secondary Effluent —Academy Symposium FLORIDA SCIENTIST 38(4) — $5.00 Solar Energy — Academy Symposium FLORIDA SCIENTIST 39(3) — $5.00 (includes do-it-yourself instructions) Anthropology —Academy Symposium FLORIDA SCIENTIST 43(3) — $7.50 Shark Biology — Academy Symposium FLoripa SCIENTIST 45(1) — $8.00 Future of the Indian River System — Academy Symposium FLORIDA SCIENTIST 46(3/4) — $15.00 Individual orders should be sent with payment. A statement will be sent in re- sponse to a bona fide purchase order over $10.00 from a recognized institution. Ad- dress all orders to: The Florida Academy of Sciences, Inc. c/o The Orlando Science Center 810 East Rollins Street Orlando, Florida 32803 Phone: (305) 896-7151 ISSN: 0098-4590 ‘Florida Scientist Volume 53 Winter, 1990 Number 1 CONTENTS Monitoring Florida’s Riverine Fish Communities................. D. Gray Bass, Jr. 1 Vegetation on the Florida Atlantic University Ecological SICS 22-05 coo Obi Gonna eS neg Daniel F. Austin 11 Nonpoint Source Phosphorus Control By a Combination Wet Detention/Filtration Facility in Kissimmee, Florida ............ Jeffrey Dee Holler 28 Survival of Florida Bay Fish Tagged with Internally Anchored APC IAS ee oe ogee Se oe Bee Gerald M. Ludwig, Jorgan E. Skjeveland, and Nicholas A. Funicelli 38 Postures Associated with Immobile Woodland Salamanders, CHBMISICINOGON 0h hoon ec es ek sans C. Kenneth Dodd, Jr. 43 Emewmonedoment Of REVIEWETS ... 2... 5. clee cc eee ee eet ens ~ 49 Note on the feeding behavior of the Common Atlantic Marginella Prunum Apicinum (Gastrododa, Marginellidae) .............. Thomas M. Baugh 50 SLL? LESTE ea A Clinton J. Dawes 51 Steven P. Christman and Walter S. Judd a2 Landfills—A Thing of the Past? .......... Richard C. Johnson, Sr. 74 y MAITHSONI 4, ~ APA 0 2 1990 LIBRARIES_/ j QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1990 Editor: Dr. DEAN F. MarTIN Co-Editor: Mrs. BARBARA B. MARTIN Institute for Environmental Studies Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoripDA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. Applica- tions may be obtained from the Executive Secretary. Both individual and institutional members receive a subscription to the FLoripa ScIENTIST. Direct subscription is avail- able at $20.00 per calendar year. Original articles containing new knowledge, or new interpretation of knowledge, are welcomed in any field of Science as represented by the sections of the Academy, viz., Biological Sciences, Conservation, Earth and Planetary Sciences, Medical Sci- ences, Physical Sciences, Science Teaching, and Social Sciences. Also, contributions will be considered which present new applications of scientific knowledge to practical problems within fields of interest to the Academy. Articles must not duplicate in any substantial way material that is published elsewhere. Contributions are accepted only from members of the Academy and so papers submitted by non-members will be accepted only after the authors join the Academy. Instructions for preparation of manuscripts are inside the back cover. Officers for 1989-90 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. ERNEstT D. ESTEVEZ Treasurery: Dr. ANTHONY F. WALSH Mote Marine Laboratory 5636 Satel Drive 1600 City Island Park Orlando, Florida 32810 Sarasota, Florida 33577 Executive Secretary: Dr. ALEXANDER DICKISON President-Elect: Dr. FRED BuUONI Department of Physical Sciences Operations Research Program Seminole Community College Florida Institute of Technology Sanford, Florida 32771 Melbourne, FL 32901 Program Chair: Dr. PAULA THOMPSON Division of Natural Science Secretary: Dr. Patrick J. GLEASON Florida Community College 1131 North Parkway 11901 Beach Blvd. Lake Worth, Florida 33460 Jacksonville, FL 32216 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BarBaARA B. Martin, Co-Editor Volume 53 Winter 1990 Number 1 Biological Sciences MONITORING FLORIDAS RIVERINE FISH COMMUNITIES D. Gray Bass, Jr. Blackwater Fisheries Research and Development Center, Florida Game and Fresh Water Fish Commission, Route 1, Box 79F, Holt, FL 32564 Asstract— Fishes of eleven Florida rivers were sampled annually to determine relative abun- dance, biomass and species richness. The goal of this program was to create a statewide catalog of riverine fish resources and to provide a mechanism for detecting long-term trends in fish popula- tions and communities. Both increases and declines of abundance and biomass were observed in some rivers and baseline data for continued monitoring were established. However, preliminary data do not suggest overall, statewide, declines in riverine fish resources. Rivers of Florida support commercial and sport fisheries of considerable economic and recreational value. Commercial fisheries in the St. Johns River alone yielded more than two million dollars from fish and blue crab landings during the 1986-87 reporting year (Hale et al. 1987). Freshwater sport fishing in lakes and streams, provides in excess of 39 million days of recreation annu- ally and freshwater anglers generate more than one billion dollars annually within the state (Brantly 1988). Approximately 24% of all licensed resident freshwater sport fishermen in Florida prefer to fish in rivers (Rayburn and Posnansky 1987). Reviewing the aquatic resources of Florida, Seaman (1985) observed many of the environmental qualities of the state which are attractive to resi- dents, tourists and immigrants are the same features often damaged by in- creased human populations. Currently Florida’s population is growing at a phenomenal rate. Between 1960 and 1980 alone the state gained almost 4.8 million people, and by the turn of the century may contain 15.2 million or more (Shoemyen, Floyd and Drexel 1987). Maintenance of high quality river ecosystems and stream fisheries is a major challenge for a rapidly growing state. Bass and Cox (1985) listed many problems affecting Florida river sys- tems, including channelization, impoundment, dredging and spoil disposal practices, floodplain encroachment, destruction of wetlands, excessive nutri- ent wastes, riparian land development, flood control projects, phosphate mine wastes, and alteration of hydroperiods. As the number of anglers ex- pands along with the overall population, fishing effort will also become an increasingly important factor to consider in fishery management. Consider- ing these adverse environmental forces it has become desirable to monitor the ecological well-being of Florida rivers for detection of long-term trends in fish resources. 2 FLORIDA SCIENTIST [Vol. 53 In addition to the need to maintain optimal abundances of exploited sport and commercial species, fish as a group possess advantages as biological indi- cators of overall ecosystem health. Karr and co-workers (1986) discussed ra- tionales for utilizing fish for biological monitoring; including the role of fish assemblages in integrating environmental conditions within a stream system. Recognizing the need to identify long-term changes in fish communities, the Florida Game and Fresh Water Fish Commission established a statewide river monitoring program in 1983. Seven rivers were surveyed during the initial year, with subsequent expansion of the program to eleven others by 1987. Rivers monitored are the Escambia, Choctawhatchee, Chipola, Apa- lachicola, Ochlockonee, Suwannee, Withlacoochee, Oklawaha, Peace, and lower St. Johns Rivers and Holmes Creek. This paper summarizes data col- lected through 1987. METHOps— Two to six permanent stations were selected on main-channel segments of each river and each station sampled once per year, during the fall low-water season, except the Withlacoochee River, which was sampled in the spring (Fig. 1). Standardized electrofishing boats were employed to collect fish during night sampling sessions. All sizes and species present were collected. cc cw tate oe |. daipven foxes scence Escambia R waste R Chipola R. Holmes C. ea ie) chtioekonee R Apalachicola R. Suwannee R Withlacoochee R. Fic. 1. Locations of Florida’s riverine fish monitoring stations, 1983-1987. No. 1, 1990] BASS—RIVERINE FISH COMMUNITIES 3 RESULTS AND Discuss1on— Results were expressed in terms of electrofish- ing catch per unit effort (CPUE). Standard sampling procedures dictated a 30-minute electrofishing session per station. CPUE represents number or bio- mass of fish collected per minute of actual electrofishing time. In addition, species richness was denoted by the total number of species collected per river. CPUE data were analyzed for (1) the entire fish community, (2) sport and commercial species, and (3) largemouth bass. Summaries for each river were compiled by combining electrofishing data from all stations into a com- posite sample and calculating mean CPUE. Although five years is insufficient time for demonstration of long-term trends in fish populations or communities, current results demonstrate how the monitoring program functions, what information it yields, and provide baseline data for future surveys. Throughout the following discussion I have not drawn conclusions from rivers monitored less than four years. Where possible I have attempted to identify trends in abundance, biomass and spe- cies richness. However, these conclusions should be considered tentative. Rivers in which there has been a general, though not necessarily uniform, increase in overall fish abundance over at least a four-year period are the Escambia, Apalachicola, and Ochlockonee Rivers (Fig. 2). It is of interest to 70 & £ Ss ri a E ay ZL LJ = fae O ( > SUW WIT Rivers of Florida WAY) > 4983 ICN) 1984 ZA 1985 AX 1986 DX] 1987 Fic. 2. Electrofishing catch per unit effort (CPUE), for all species combined, by number, 1983-1987. (ESC = Escambia R., CHO = Choctawhatchee R., HOL= Holmes C., CHI=Chipola R., APA = Apalachicola R., OCH = Ochlockonee R., SUW = Suwannee R., WIT = Withlacoochee R., LSJ =St. Johns R.,. OKL = Oklawaha R., PEA = Peace R.). 4 FLORIDA SCIENTIST [Vol. 53 note, however, that numbers of fish declined in seven of the eleven rivers during 1987, compared to 1986 abundances, including the Escambia, Apa- lachicola and Suwannee Rivers. Future monitoring will determine whether these recent reductions represent downward trends. The only river showing a clear downward trend in overall fish abundance over the monitoring period is the Choctawhatchee River. With regard to relative density, the Peace River contains low numbers of fish, especially when compared to other fertile rivers on the peninsula, such as the Withlacoochee, lower St. Johns and Oklawaha Rivers. Fish biomass generally increased in the Escambia River (Fig. 3). Definite downward trends in biomass are not evident for any river, although biomass reductions are apparent for the Choctawhatchee and Peace Rivers. In the former, overall fish weight was much greater during the first two years of sampling than the past three seasons. Peace River biomass increased over the first three years and declined substantially the past two years. 3 7 mal AS N 6b= Q : N a yi ant F WH NN N ag 4 U NN A € NN Nd an q ea Fa 18 - \ NN A ON Y q WAN ) fe eo Ne we Q v hd g a h N oo ENG? Ska Pin Wo ey) ONAN ON ere PROMS vue x WS RN Le g AN : NI) A Ad x WAN xe Y we ot WAY ON NA ON OTN ON ONAN INE : HN OM ON NOT NOAH EA WA SRN] ON) WANT Th TY AY AAG TNT MAS ANG rN N vi IN Re Ue ) AN TAM NAS if Ne aN Ne IN ) d rh Ne IN ON axe VN > Ne NOME UGG DEE NOMEN AZNOINGUL NOMNONZNOIEO RNIN NOMIONGNOIANZNG Wi NAY ORY NANT INN TA TA Nj NN Nd ONS Abd iA‘ NY Ane aN Abd Vb x] SSN O MIG NOMMIUCOMMGNINOIMANANOINPIASBe WANG) yc it NL RARE FST AAS URRY PRESS AN RAS TAA TRS ON ENN INN INAS ER NAS EO CHO HOL CHI APA OCH SUW WIT. ES OKL PEA YY) 1983 [IN 1984 aA ae. TER 1986 [XX] 1987 Fic. 3. Electrofishing catch per unit effort (CPUE), for all species combined, by weight, 1983-1987. (ESC = Escambia R., CHO = Choctawhatchee R., HOL=Holmes C., CHI =Chipola R., APA = Apalachicola R., OCH = Ochlockonee R., SUW = Suwannee R., WIT = Withlacoochee R., LSJ =St. Johns R., OKL= Oklawaha R., PEA = Peace R.). No. 1, 1990] BASS—RIVERINE FISH COMMUNITIES 5 24 22 N . N] N 6 20 N N AG NN N is NON Ne NN NM S 16 IN OUND E ON N UN IN N Ne S 14 ON NAR we ? A eo I ee N N WAY LAS ea. Ma ai NIN SGN WOES , : PANG NU NAN NAN z » INE ers NAW INN oS ON ONAL TAA NE | NonmN NOBUO NNO NO EEN INNO IUZNO SANS IAN Se NOME UZNOIICUINORT 12 NOUN ZNO IIANZNS SINAN IN A Ran ON ONAN TAA NORA TAA AN CAN RA OND NAD TAN AA NORA OA + TYANT NAS NAT oN OYA NN A ONAN TS as YN NA RSS NAN ENB NAS HEN NA LINAS AN TNA NAN PR RA TAN AA TN NA EAA oo 2-1 ENA NAH A NA SY ENA HEN NAINA oe 5 LNASLINAS NAY AT NAS NY NA EN NNN BSG CHO HOL CHI APA OCH SUW WIT LSJ OKL PEA VA AI8S ICN) 1984 We a 1986 XX] 1987 Fic. 4. Electrofishing catch per unit effort (CPUE), for sport and commercial species, by number, 1983-1987. (ESC = Escambia R., CHO = Choctawhatchee R., HOL= Holmes C., CHI= Chipola R., APA=Apalachicola R., OCH=Ochlockonee R., SUW=Suwannee R., WIT = Withlacoochee R., LSJ =St. Johns R., OKL= Oklawaha R., PEA = Peace R.). Sport and commercial fishes—Both relative abundance and biomass of sport and commercial species, as a group, increased geographically in a west to east direction, from rivers of the panhandle to those of the peninsula (Figs. 4 and 5). A notable exception to this geographic production pattern was the Peace River, which yielded low numbers and weights of fish compared to other peninsular streams. This general progression of biological productivity, from the northwest to the peninsula was previously noted by Kautz (1981) and Bass and Cox (1985) and attributed to increased dissolved solids and increased nutrient concentrations in peninsula streams. Numerical abun- dances of sport and commercial fish exhibited generally upward trends over the monitoring period in the Suwannee and Peace Rivers. Data collected for the past three years also demonstrated numerical increases in the Escambia, Choctawhatchee and Oklawaha Rivers. Declines in abundance were not evi- dent for any stream. Electrofishing CPUE, by weight, exhibited generally upward trends in the Escambia and Suwannee Rivers, while declines have occurred in the Apa- lachicola and Peace Rivers (Fig. 5). Largemouth bass—Populations of largemouth bass increased continu- ously in the Escambia River, and also over the first four years in the Peace River (Fig. 6). Overall declines in abundance of bass are suggested for the 6 FLORIDA SCIENTIST [Vol. 53 [A/T = a N N cS N 7 YAN 14 Vl 13,4 NANG . VN pond) vis € | NY Q Aes RZNOEREN ae NZNOMAN NS eee n NY Be) al SL N NES aa peek K NIV 6 nN NIN : ia ih 8 OW NY NY WA N NY LANG LYN UN NY Ne UN > Nw MA x RNA) A YS) a TA AN WN NY UN NY TAM DAN ON WAT Ih SmI SHIVA 1 c Ly : aN Na CHO HOL CHI APA OCH SUW WIT 1983 1984 GN eee OS 1986 XX] 1987 Fic. 5. Electrofishing catch per unit effort (CPUE), for sport and commercial species, by weight, 1983-1987. (ESC = Escambia R., CHO = Choctawhatchee R., HOL = Holmes C., CHI = Chipola R., APA=Apalachicola R., OCH=Ochlockonee R., SUW=Suwannee R., WIT = Withlacoochee R., LSJ =St. Johns R., OKL = Oklawaha R., PEA = Peace R.). Apalachicola, Ochlockonee and Suwannee Rivers. However, the cycles in numerical abundances of bass may be due to the presence of especially strong year-classes which dominate the population for several years. Clear upward trends in largemouth bass biomass are not evident for any river (Fig. 7). Obvious trends in bass weight are not evident, except in the Ochlockonee River, where biomass was much higher in 1983-84 than during 1986 and HOST. Species richness— Over the course of the survey, 132 species were collected from rivers of Florida. The number of species collected per river, per year, ranged from 19 (Peace River, 1984) to 56 (Escambia River, 1985, 1987). No obvious trends are evident, except for Holmes Creek, where the number of species reported increased over the past three years (Fig. 8). Increases in Holmes Creek species numbers may be a sampling artifact as only two sta- tions are maintained on this small stream. In other streams, species richness appears relatively stable. Current data provides a baseline from which to measure future changes in species richness in main-channel segments of Flor- ida rivers. As a matter of note, it should not be inferred, on the basis of species richness alone, that streams of the northwest, such as the Escambia and Apa- lachicola Rivers, ecologically are more viable than peninsular watercourses. Ichthyofauna of the panhandle is much richer in species than the peninsula, No. 1, 1990] BASS—RIVERINE FISH COMMUNITIES i 1.4 q ‘ 13 \ N N N y N 1.2 N y N ) y N 11 g N ms h 4 \ A) & S U y N | vie . : A ON ONG S 0.9 <7 LN Ny ih mn at Nf a ay BN eS INUND : eee ie IN INE BS NBS EL IINGN8 oN A PY REE BN NAN AN € A on ONAN 7 WN NN LAO INAS Me oe WANN Pe Nn NE Ne NIRS 2 os NN NAY OW OOM ON ORAS r MeL IANISGE WANGh INL aIN@NGP phere TpASOb UL ZN mANANIANANGI mS 0.4 INA IN AY NAY ONY NAY INA NAY GN ONAN NAN AA INN NAD OPN RAN TAA NORA Ao PNA PNY NAY AY NAY INS NAS IN ONAN NA ON 03 INN YN NN EN NA IN AY NA HEN ONAN NAO ao INA TAA NAY RE ORAS TN SE TRANS HES NAN TAS A 2 TN TAN NA A NA TAS TAA LN NA TA 01 INA AAS NAY NAY NAY ENA TAN AA ENA FAY > LNA NAN RNA INI NA TANNA NYA YAS ESC CHO HOL CHI APA OCH SUW WIT LSJ OKL PEA Vea 1983 INSNIEF i934 a eee OOS 1986 XX] 1987 Fic. 6. Electrofishing catch per unit effort (CPUE), for largemouth bass by number, 1983- 1987. (ESC = Escambia R., CHO =Choctawhatchee R., HOL=Holmes C., CHI=Chipola R., APA = Apalachicola R., OCH = Ochlockonee R., SUW = Suwannee R., WIT = Withlacoochee R.., LSJ =St. Johns R., OKL =Oklawaha R., PEA = Peace R.). due to the past geological events (Neill 1957, Bass and Cox 1985, Swift et al. 1986, Gilbert 1987). Changes in species number should be employed to evalu- ate ecological changes only where the total number of species in the stream under investigation is known, or where historical records demonstrate species occurrence during previous undisturbed conditions. Even addition of new species to a fish assemblage does not necessarily denote ecological improve- ment, especially if the new forms are exotics, such as walking catfish (Clarias batrachus) or blue tilapia (Oreochromis aureus) both of which have invaded peninsular Florida. Conversely, extirpation of even one, or a few, native spe- cies may signal severe environmental degradation, as these species may be markedly sensitive to pollution or habitat alteration. Conc.usions— Riverine ecosystems are dynamic. Rise and fall of popula- tions and the composition of communities are influenced by random physical forces (e.g. floods, droughts) and biological interactions (e.g. interspecific competition, predation). Whether a population or community is ever stable, or reaches a state of equilibrium, is a subject of much current ecological investigation. The importance of random physical forces, interspecific com- petition, or both, in determining the fate of communities is likewise a lively 8 FLORIDA SCIENTIST [Vol. 53 Zg N ON N ON N x h n- N we a N 7 | UN N ¢ N CeO] |) NN t Hn 6 OTN ON NN . q N : HAN NN ¢ N N rH 6h SCOUT ON OA : " A TN ONO A OMA © N Ng go | NNN NNO NANO UZN u N Re gee a YN oA LN TAA Oe 5 Nein Asad NAG FN NAS NAY NAN in mR . A AH VEN = NA NA INA eae VA g ph VAM ( ZX NAN WA WN Tr WN OU COCO ONO Nn VND ONY we TD TN OO es yn VN) NO RNR NN ON a y PN A DN TNO ts os RY NAT OAS NAY NN A EN ONANS T Ny NA OR NAY INAS YA HN NAS A ONS Ny NAN FRE NAW ENB ENA EN NA ESE SSS Nd _NAN FAS NAN IN TNA AN NAS NA OIA ESC CHO HOL CHI APA OCH SUW WIT LSJ OKL PEA ZA, W983 IAN) 1984 A eee. Bw 1986 XX] 1987 Fic. 7. Electrofishing catch per unit effort (CPUE), for largemouth bass by weight, 1983- 1987. (ESC = Escambia R., CHO =Choctawhatchee R., HOL=Holmes C., CHI=Chipola R., APA = Apalachicola R., OCH = Ochlockonee R., SUW = Suwannee R., WIT = Withlacoochee R., LSJ =St. Johns R., OKL=Oklawaha R., PEA = Peace R.). subject of present-day debate (see Schoener 1987). Disregarding the outcome of the current academic brouhaha in ecology, one thing is certain: Numbers and biomass of stream fishes cannot be expected to remain precisely the same between contiguous years. Community stability may also be less in some sections of rivers than others. Tidal zone assemblages, for example, may be more unstable than upstream assemblages. However, long-term monitoring should reveal profound changes despite effects of between-year or between- site variations. The goal of a monitoring system is to detect and quantify long-term ecological perturbations and to stimulate technological advances in fish management. With the present program, we have a beginning and some suggestions of current trends in lotic fish assemblages. In addition to routine monitoring, this program also provides data which may be employed to measure ecological stability and persistence in fish com- munities and to study community dynamics (see Connell and Sousa 1983, Schoener 1987). These materials may also be used to develop simpler methods of environmental analysis such as the Index of Biotic Integrity (Karr et al. 1986). Finally, monitoring of fishes will track abundance and biomass of important sport and commercial fishes. BASS—RIVERINE FISH COMMUNITIES No. 1, 1990] 60 CK CK EK TELLS TIPLE ES LTT MTT ETLTETLT ETA ISS SASS ASSET VSI IDOE EE GE GNA SENG BOO DPPWDOPaass Lat Ltt ltt litle: KQQAAAWTAAT WA LILI LID ED AY CY VERTED OS LET UT LS LA SDTUNGENGENS EN SNES LOX SOKO XG ULL 22 ie L LLL Le LLL LLL aaa LLL GQAQNNN AVA a SEDI LILES FET “DD LE EG UD DED LEDS UG LEG (Zp A (CLL L Le Lee LE LLL“ LLL La RQQNNAYSSOo.||eE_EA Dw A Md FEF EBEE DGG SESE ARNG SEG) ROBB SDOS DBD ODGa CULL 222i ZL Le LeeLee LLL LLL LLL PQQ SASS SASS ASS ASS SIS ASISASSASSASSASSSSN ED LED VD AUS DD UPD AD ay EB LEG ESL SSS SYNE OAT ASE SEN EEE NG WG EGG QOD OLEB DDD BODE (LLL LLL LLL LLL LLL L LLL LL LLL LLL LLL (GL LG LE (FED ED ULL EEG LL DUES (BF CSS|’WES EN ENE ENE ONE NENG] QO2OOOSEE BSD BDBDDDODPDS OLA ATT YA AP AD AY AL AD A A A A AAD AAD A A AA AV AD AA LA AL AA A ALLA A RQRNQNRHACAAAAAAVUAM GSMA _-*E"FEEF*EE8BE Ba Oo LIL GLI ESUE GL UF GET UG OT EL ED UES OED ED OEE EES SOS DOD SSG COLUM UME ALE LLL MLM LM zee ls GQINNNSO OO O_O oO 2x Dea Ss CLD DISS PLATO TIT SLSLTIT SLT PQS SSSSASSSSAS SSS ASSSDAS SSA CLL LES GILG DES TG BS ETL ESE BDS OOD DBPDBOSE LLL tyyyiltytltlilltill lll AQRRVKQAQAHARHA NMA sd DELLA EDD OD ELT EES EDAD ALG UL ED UG ETL ENE NNN SSE ERNE ENE SENG] DOQQ QOOXQDODBDDIID DD DPDDDLSS ALIS UU TUTITEPUT TESTIS TTL TITUTLMTITLTLLLT UD ET AA NAM QQ aa aaa PLETE GT ag ED 0D UP TE DG A TD TOBE LTD SDN EEN {e) (o) oO (e) oO jo) wo se ne) N v5 seiseds jo uequinn CHO HOL CHI APA OCH SUW WIT LSJ OKL PEA ESC Rivers of Florida 1985 1987 KX 1986 SN VLLA 1984 [Soy 1983 VA, a (se (ae Cg w ml [= Qo SEouS Ge) oles) oA SY ed Man l 3, ll OtR A + 3 Ba aes 3 Se -- (oe) er 2 a fem te ou 8 23 § Toes > 213 =i wp, EOS 5-5 Be & En & Oo 0 EES (o) Bis ol 3s aA eren Ome ll SaNH (Sh Wigs oO cect Be Oca 6 Es .4 3.379 2 8) ora! om x NGS Sil ie (@)jae ca] 10) OS) Peace R.). Oklawaha R., PEA In sum, our current, albeit short-term, data do not suggest overall state- wide declines in Florida’s riverine fish communities. It is also possible, even certain in some cases, that changes in riverine fish populations and communi- ties occurred prior to the current monitoring program. However, current results do point out present (Peace River) or potential (Choctawhatchee River) trouble spots throughout the state. Several more years of survey will be term population cycles or represent real long-term directions in fish abun- necessary to determine whether current trends are merely artifacts of short- dance, biomass or species richness. OKL ACKNOWLEDGMENTS—I would like to express my appreciation to the many fishery biologists and laboratory technicians throughout the state who collected fishes from streams of their re- gions. This study was partially funded by Wallop-Breaux Federal Aid Project F-36. LITERATURE CITED SEAMAN, W. (ed.) Florida Aquatic Habitat and Fishery Resources. Florida Chapter, and freshwater fish 1988-1993. 3rd Ed. Florida Game and Fresh Water Fish Commission. American Fisheries Society, Kissimmee, FL. Tallahassee. 80 pp. Bass, D. G. anno D. T. Cox. 1985. River habitat and fishery resources of Florida. Pp. 121-187. In: BRANTLY, R. M. 1988. A strategic plan for the comprehensive management of Florida’s wildlife 10 FLORIDA SCIENTIST [Vol. 53 ConNneELL, J. H. AnD W. P. Sousa. 1983. On the evidence needed to judge ecological stability or persistence. Am. Nat. 121(6):789-824. GiLBerT, C. R. 1987. Zoogeography of the freshwater fish fauna of southern Georgia and penin- sular Florida. Brimleyana No. 13:25-54. Hate, M. M., J. E. Crumpron, Aanp D. J. RENFrro. 1987. Florida Game and Fresh Water Fish Commission 1986-87 commercial fisheries investigations report. Florida Game and Fresh Water Fish Commission, Tallahassee, FL Karr, J. R., K. D. Fauscn, P. L. ANGERMEIER, P. R. YANT, AND I. J. SCHLOssER. 1986. Assessing biological integrity in running waters. A method and its rationale. Ill. Nat. Hist. Surv., Spec. Publ. No. 5:1-28. Kautz, R. S. 1981. Fish populations and water quality in north Florida rivers. Proc. Ann. Conf. Southeast. Assoc. Fish Wildl. Agen. 35:495-507. NEILL, W. T. 1957. Historical biogeography of present-day Florida. Bull. Fla. St. Mus., Biol. Sci. 27) 175-220: Raysurn, J. D., HI anp G. Posnansky. 1987. A survey of attitudes and preferences of licensed fresh water fishermen in Florida: An update. Communications Research Center, Florida State Univ., Tallahassee, FL. 61 pp. SEAMAN, W. 1985. Introduction to the aquatic resources of Florida. Pp. 1-19. In: SEAMaAn, W. (ed.) Florida Aquatic Habitat and Fishery Resources. Florida Chapter, American Fish- eries Society, Kimmissee, FL. SHOEMYEN, A. H., S. S. FLoyp anp L. L. Drexeu. 1987. 1987 Florida Statistical Abstract. Univ. Presses of Florida, Gainesville, FL. 706 pp. SCHOENER, T. W. 1987. Axes of controversy in community ecology. Pp. 8-16. In: MaTTHEws, W. J. AND D. C. Hens (eds.). Community and Evolutionary Ecology of North American Stream Fishes. Univ. of Oklahoma Press, Norman, OK. Swirt, C. C., C. R. Grupert, S. A. BorToNE, G. H. Burcess AND R. W. YeERGER. 1986. Zoogeogra- phy of the freshwater fishes of the southeastern United States: Savannah River to Lake Ponchartrain. Pp. 213-265. In: Hocutt, C. H. ano E. O. Winey. (eds.). The Zoogeogra- phy of North American Freshwater Fishes. John Wiley and Sons, New York. Florida Sci. 53(1):1-10. 1990. Accepted: January 23, 1989. Biological Sciences VEGETATION ON THE FLORIDA ATLANTIC UNIVERSITY ECOLOGICAL SITE DANIEL F. AUSTIN Department of Biological Sciences, Florida Atlantic University, Boca Raton, FL 33431 Asstract: A discussion of historical and modern habitats and animals is given along with the recent history of the site. Plants known from the area are listed and their frequency, endanger- ment and introduced or native status given. In May of 1984 the administration of Florida Atlantic University (FAU) declared a 91.6 acre (37 ha.) parcel on the northern edge of the campus as a “Natural Preserve” for continued study as an ecological site. This action for- malized the preservation of land that had been studied by the Department of Biological Sciences since 1970. Since the area is preserved and contains both endangered plants and animals, it was decided to record some of the results of the studies (Austin, et al, 1987). The following discussion presents more detail on the ecological site. History—Prior to World War II the only major change in the vegetation on the FAU ecological site was the construction of a city airport (Curl, 1976). This early airport, one of the few on the southeastern coast at the time, was built between 1936 and 1937 and covered about 185 acres. The northwestern corner of the city airport occupied the southeastern corner of the present FAU ecological site. The overlap of the airport with the ecological site was in T 47 S, R 42 E, Sect. 13, SE 1/2 of the SE 1/4 of the NE 1/4 and T 47S, R 43S, Sect. 18, SW 1/2 of the SW 1/4 of the NW 1/4. The overlap area was “cleared of trees, stumps and palmetto grubbed” as of 31 July 1936 (Curl, 1976). Other parts of the modern ecological site were left undisturbed. The military inspected the site in 1941 and by 15 Oct. 1942 an Air Force base was officially open (Brown, 1985). The military base enlarged consider- ably the area formerly used by the Boca Raton City Airport, and much of the area of the modern campus was cleared of vegetation. Aerial photography from the 1940s shows the marked alteration of the native vegetation during the construction of the runways, aprons for tying down grounded planes beside the runways, buildings, and other facilities. The land now in the FAU ecological site was bulldozed, but apparently never cleared as drastically as most of the remaining land. Oak trees that show in the 1945 and 1947 aerial photography are still extant today in several places within the site. Cores taken from the largest of these trees in 1970 showed that they ranged in age from 25 to 30 years. Every tree in this age range grows in clusters and the clumps are thought to be stump sprouts from the plants cut in the late 1930s and early 1940s. There is a hiatus in the 12 FLORIDA SCIENTIST [Vol. 53 historical record from the end of the military usage until the land became the site for Florida Atlantic University. Presumably, no major changes in vegeta- tion occurred. Florida Atlantic University, officially established in 1961, at first used the remaining military buildings for offices while the university buildings were being constructed (Brown, 1985). It was not until July 1964 that the majority of the present buildings were finished, and classes began in September of 1964 (T. T. Sturrock, 1988). Some time about then the Grounds Department began systematic mowing, not only of the cultivated lawns and other grass- lands, but also of the triangle that was to become the FAU ecological site. The parcel was mowed at least once annually, and in most years more often, except where vegetation was so large by the time this practice began that the mower (bush-hog) used could not remove it. Oak hammocks, a few pine trees, saw palmetto clumps and isolated cabbage palms were the dominant arborescent vegetation in 1970 when this study began. By 1973 the Depart- ment of Biological Sciences was able to convince the university that mowing the triangle was not only cost ineffective but detrimental to the teaching of classes. After cessation of mowing, the vegetation began recovery. The plot has now been in undisturbed secondary succession for fifteen years. ANIMALS—No complete survey of the animals has been attempted, but Burch & Marsh (1987) have compiled potential or probable lists for the entire FAU campus. Observations over the years have given more specific partial lists of species known to use the ecological site. Only vertebrates will be listed for this report. The most important animal on the site in terms of the state Game and Freshwater Fish Commission criteria is the gopher tortoise (Gopherus po- lyphemus). Shortly before the parcel was designated an ecological site, poachers almost completely extirpated this legally protected reptile. Since designation as an ecological site, efforts at guarding against this problem have been more successful. From a low population of an unknown number of animals, the latest estimate of gopher tortoises is 200-300 individuals (T. Mann, 1988). This density by far exceeds those reported by Auffenberg and Franz (1982) and by Richardson et al. (1986) for similar but not identical habitats. Over the past two years, observations suggest an increased number of hatchlings and pre-reproductive individuals. Numbers of breeding age adults (10 yrs or older, fide Franz and Tonnessen, 1983; Jackson, 1985; Go- pher Tortoise Council, n.d.) seem to be more or less constant. These data indicate that this is a viable, reproductive population of gopher tortoises, one of the few in southeastern Florida. Comparatively few of the gopher tortoises that now inhabit the ecological site were purposely introduced. One FAU student, Ms. Chris Collins, moved in about 20 individuals from nearby construction sites at the request of the city. Only three of these marked individuals have been recaptured. The re- maining individuals arrived on the ecological site without any known assis- No. 1, 1990] AUSTIN—FAU VEGETATION 13 tance from man. Presumably, those individuals that made it to the site with- out assistance came from disappearing nearby habitats that were lost to them through urbanization, although this is debatable. Gopher tortoises have been seen eating mostly grasses of various kinds, but primarily the introduced bahia (Paspalum notatum) and Digitaria longiflora. Fruits of gopher apple (Licania michauxii), prickly pear (Opuntia compressa) and saw palmetto (Serenoa repens) have either been seen being consumed or identified in scat. No detailed study of their food habitats in this southern marginal extreme of their range has been attempted. Other reptiles that have been found on the site are the rough green snake (Opheodrys aestivus), eastern coachwhip (Masticophis flagellum), eastern diamondback rattlesnake (Crotalus ademanteus), coral snake (Micrurus ful- vius), six-lined racerunner (Cnemidophorus sexlineatus), and the eastern glass lizard (Ophisaurus ventralis). The only amphibians recorded to date on the site were the green tree frog (Hyla cinerea), oak toad (Bufo guercicus), the southern toad (Bufo terrestris). Greenhouse frogs, Cuban tree frogs, Gi- ant toads, and narrow-mouth toads are known from nearby on the campus, but have not been recorded on the ecological site. Some of the birds seen on the site include mourning dove, ground dove, mockingbird, catbird, ani, shrike, chuck-wills-widow, great-horned owl, burrowing owl, red-tailed hawk, kestrel, sharp-shinned hawk, harrier, fish crow, turkey vulture, cardinal, blue jay, boat-tailed grackle, starling, towhee, and a variety of migrants such as warblers. Examination of burrowing owl pellets indicates that the cotton mouse (Peromyscus gossypinus) and cotton rat (Sigmodon hispidus) are on the site or nearby (Tatje, 1979). Other mammals recorded recently are spotted skunk (Spilogale putorius), armadillo (Dasypus novemcinctus), racoon (Procyon lo- tor), opossum (Didelphis virginiana), mole (Scalopus aquaticus), cottontail rabbit (Sylvilagus floridanus), grey fox (Urocyon cinereoragenteus), and feral housecats (Felis catus). The feral cats are thought to be a major factor in the lack of some other animals that would be expected on the site. For example, no records exist for the Florida mouse (Peromyscus floridanus) or for the gopher frog (Rana areolata). At least the mouse is quickly extirpated when feral cats are present (J. Stout, 1986). Studies of the nearby Yamato tract failed to find the gopher frog and some factor other than feral cats may be responsible for its absence (Richardson et al., 1986). There were formerly abundant brown anoles (Ano- lis sagrei) on campus, yet this prolific introduced lizard has not been seen since the feral cat population reached its current level in the 1980s. Cats are also implicated by circumstantial evidence in the destruction of burrowing owls since tracks have been found associated with feathers and the disappear- ance of young before fledging. 14 FLORIDA SCIENTIST [Vol. 53 E| WET PRAIRIE LOW HAMMOCK DRY PRAIRIE SAND PINE SCRUB === 4 = OVE Ee 0 SF OO 3 0,8 , PONDED WET PRAIRIE S=225 Oe? ‘ tT, ee ——— mw Ps zJ S =a SSeS erat a — =F i r 2 CEI R LG a ee Me eS = = “te oc Ps rc Le <& Yor f= \. c Fae ee eg Runway. eA sb-a0 Ai roort © 00° Oo ° ° ° © 000 0 oo =o o 0°90 ° ooo © 00° 0 ov. 00° 0 600 0 Fic. 1. Vegetation patterns of the Florida Atlantic University Ecological Site as they appeared prior to 1940. Runways and ditches prepared by the military are overlain on the vegetation to indicate modern localities. VEGETATION— The FAU ecological site now occupies parts of the south- eastern triangle of the NE 1/4 of section 13, and the western half of the NW 1/4 of section 18, Palm Beach County. Vegetation of the FAU ecological site has been studied on aerial photography taken from 1940 to the present. Since all of the original parts of the site are recognizable in the early and more recent phography, it is possible to make an accurate map of the vegetation in its pre-1940 condition. Habitats on the ecological site in 1940 consisted of Wet Prairie, Dry Prai- rie, Low Hammocks and Scrub Pinelands (Fig. 1). These habitats have been described in terms of species elsewhere (Austin, et al., 1978). The eastern part of the ecological site was a complex of Wet and Dry Prairie. Wet Prairie dominated on the eastern fringe near the El] Rio Canal, and Dry Prairie occupied the area immediately to its west. The N-S runway No. 1, 1990] AUSTIN—FAU VEGETATION LS E] WET PRAIRIE S|] DRY PRAIRIE PONDED WET PRAIRIE LOW HAMMOCK SAND PINE SCRUB SCALE ead ©) 200° Ft: > ¥) - a CNY. fa a. — a BAG OO Wen ee Be ee ee ~Mowed Area Airport Runway ees me ee Fic. 2. Vegetation patterns of the Florida Atlantic University Ecological Site as they appear in 1988. (now Palm Beach Avenue) was built near the interface between these two habitats. Another Wet and Dry Prairie system occupied the western part of the triangle. The north-central part of the ecological site was a Scrub Pinelands in the 1940s. This habitat has already been reported on in terms of how the site compares with other Scrub Pinelands in the Boca Raton vicinity (Austin, et al., 1987). Surrounding the Scrub was an ecotonal Dry Prairie. Low Ham- mocks, dominated by several oak species, were scattered through the Scrub Pineland and Dry Prairies. The same habitats are represented on the modern ecological site that oc- curred in the 1940s (Fig. 2). They have, however, been altered by long-term mowing and a regional drop in the water table (Thomas, 1974). Moreover, the usage of the fringe areas of the ecological site during the years of military occupation resulted in an increase in the number of individuals and species characteristic of disturbed sites. 16 FLORIDA SCIENTIST [Vol. 53 Wet Prairies have been altered to the extent that only soils and species analysis will show clearly their original limits. Some of the species remaining that show the former existence of Wet Prairies under wetter conditions are cord grass (Spartina bakeri), blue maidencane (Amphicarpum muhlenber- gianum) and button bush (Cephalanthus occidentalis). Soils where the Dry Prairies occurred are Basinger fine sands or Pompano fine sands (U.S.D.A., 1978). The Basinger is a poorly drained, nearly level, deep sandy soil where the water table is within ten inches of the surface for two to six months of the year. Pompano is almost identical in drainage, topog- raphy and water; they differ chiefly in the thickness of their upper layers. Elevations of the Wet Prairies are from the 12-foot contour down, with the lowest areas near the nine foot level. Dry Prairie species remain in the areas where they were in the 1940s, and have partly invaded the margins of the now-dry Wet Prairies. Typical species of the Dry Prairies are saw palmetto (Serenoa repens), gallberry (Ilex glabra), runner oak (Quercus minima) and fetterbush (Lyonia lucida). Five un- planted slash pines (Pinus elliottii) and several seedlings remain in the Dry Prairie, although there was an unsuccessful attempt to establish this species in much of the habitat in the late 1960s. From the planted slash pines two adults and two seedlings remain. Soils in the Dry Prairies are Immokalee fine sands (U.S.D.A., 1978). While similar to the Basinger sands, these soils have a dark colored layer below thirty inches which is weakly cemented with organics. The water table is within ten inches of the surface for two to four months. Elevations in the Dry Prairies are in the 12-13 foot range. Scrub Pinelands, in greatly altered form, occupy part of their original extent. Two ditches were constructed by the military and formerly connected with a ditch system that took excess water from the entire military base into nearby wetlands (Brooks, 1988). One of these ditches extended the length of the highest ridge where the Scrub Pinelands dominated. This alteration among others has decreased the area of the habitat, although many of its distinctive species remain precariously on the ecological site (Austin, Posin & Burch, 1987). Sand pines (Pinus clausa) are represented by two remaining trees and two small seedlings (Ritchey, 1988). Two other sand pines were formerly present (1970-1979), but these have died for unknown reasons. Soils for the scrub area were listed in the country survey as Immokalee fine sands (U.S.D.A., 1978). These soils are typically associated with Pine Flatwoods and Dry Prairies and the report must be in error. Since the soil maps were made from interpretations of aerial photography, small local pockets of soils are not always correctly labeled. The soils on the part occu- pied by Scrub appear to be typical St. Lucie fine sands. Elevations on the campus survey show the Scrub areas as being 13 feet or above. The highest point surveyed was 13.5 feet. Hammocks are now occupied by oaks, with the four species occurring in different combinations at different sites. The dominant species in the area is No. 1, 1990] AUSTIN—FAU VEGETATION IL9f the myrtle oak (Quercus myrtifolia), although live oak (Quercus virginiana), chapman’s oak (Quercus chapmanii) and sand live oak (Quercus geminata) are also present. These species are not confined to the hammocks and also may be found in the Dry Prairies and Scrub sites. Soils under the Hammocks are not differentiated from the other types mentioned, being either St. Lucie or Immokalee. Elevations in the hammocks are above the 12 foot contour. The majority of disturbed site plants have declined in frequency since 1970 (Conroy, 1970). For example, in 1970 the herb layer was dominated by Natal grass (Rhynchelytrum repens) and Para grass (Panicum purpurascens). Para grass is said to have been planted as camouflage by the military. Broom grass (Andropogon virginicus) was almost as common as the other two grasses as was the herb flat-top goldenrod (Euthamia tenuifolia). All of these have declined in frequency. In the 1970 sample, Calusa grape (Vitis shuttle- worthii) was found to be more frequent than the Muscadine (Vitis rotundifo- lia). They have subsequently reversed in frequency, with the Muscadine now being about twice as common as the Calusa. Puncture vine (Tribulus cis- toides) is now frequent on the FAU campus and elsewhere in southern Florida (Austin, 1978). This species was probably introduced during World War II on airplane tires (D. Porter, 1970). One species that continues to gain in frequency is the pepper tree (Schinus terebinthifolius). Since the termination of mowing in 1973, there has been a marked increase in the number of individuals of this species. Moreover, indi- vidual plants of this tree are expanding the area they cover and out-compet- ing or at least shading the native species. The ultimate impact this species will have on the ecological site is unknown. Another exotic tree which has invaded the ecological site is the ear-leaf acacia (Acacia auriculiformis), widely cultivated in southern Florida as an ornamental. This short-lived tree was first noted on the site in 1979. Since that time three other individuals about the same age (in the 15-20 foot size class) and several smaller plants have been found. The species is becoming more widespread in southern Florida, and little is known about its impact on the native environment. Fifty-six families and 180 species have been identified on the site. Al- though the species may occur in more than one habitat, the following per- centages illustrate the tendencies. Species are counted in a particular habitat where they are most abundant. There are some 73 species that indicate dis- turbance or about 40.5% of the total. This high figure largely reflects the impact of man since the early 1940s. Scrub Pinelands are next in richness with 55 species or about 30.5% of the total. This figure does not include ten species (e.g., Quercus virginiana and Sabal palmetto) reported in Scrub by Austin and co-workers (1987) because they are found more frequently in other habi- tats. The next highest is the Dry Prairie with 27 of the total, or about 15%. Although Dry Prairie species represent a small proportion of the total species, they are represented by large numbers of individuals. The remaining habitats 18 FLORIDA SCIENTIST [Vol. 53 have 17 Wet Prairie species (9.4%) and eight Low Hammock (4.4%) species. The Low Hammocks are expected to have few species because they are young, but the Wet Prairies show marked erosion of species richness (Goo- drick, 1974). This loss of species in the Wet Prairies is undoubtedly linked to the drop in the water table. PLANT List—Names used are those that seem to be most appropriate for the species. If synonyms are given, they represent names used by Wunderlin (1982) or Wunderlin and co-workers (1985, 1988) where a difference in opin- ion exists as to generic or specific limits, or, in rare cases, where there has been a long usage of another name. Endangered status is summarized by Wood (1985). Introduced status is based on Austin (1978); unless otherwise noted, the remaining species are presumed to be native. Frequency notes: Abundant=found every few yards; common = found every few tens of yards; occasional=sometimes found; infrequent = more than a half dozen or so individuals known, but still uncommon; rare=a half dozen or fewer individuals known from the site. Data are based, except where noted, on the period from 1980-1988; before this period (1970-1979) there were major shifts in frequency due to termination of annual or more frequent mowing. ACANTHACEAE Dyschoriste oblongifolia (Michx.) Kuntze.-TWINFLOWER. Rare in dry prairie. AIZOACEAE Mollugo verticillata L.-_INDIAN CHICKWEED. Rare on disturbed mar- gins. Introduced. AMARANTHACEAE Froelichia floridana (Nutt.) Moqg.-COTTONTOP. Common on disturbed scrub & prairie. Gomphrena serrata L. (=Gomphrena decumbens Jacq.)-Common on disturbed margins. Iresine diffusa Humb. & Bonpl. ex Willd.-IRESINE. Occasional on dis- turbed margins. ANACARDIACEAE Schinus terebinthifolius Raddi-PEPPER TREE. Occasional in disturbed sites throughout. Introduced. ANNONACEAE Asimina reticulata Chapman—PAWPAW. Occasional on scrub ridge. APIACEAE Eryngium aromaticum Baldw. ex Ell.-BUTTON SNAKEROOT. Occa- sional on scrub ridge. No. 1, 1990] AUSTIN—FAU VEGETATION 19 APOCYNACEAE Catharanthus roseus (L.) G. Don-PERIWINKLE. Common in disturbed margins. Introduced. ARALIACEAE Schefflera actinophylla (Endl.) Harms (=Brassaia actinophylla Endl.)- UMBRELLA TREE. Infrequent as seedling trees in hammocks. Introduced. ASCLEPIADACEAE Cynanchum scoparium Nutt.-HAIRNET VINE. Rare on disturbed mar- gins. ARECACEAE Sabal palmetto (Walt.) Lodd. ex Schultes-CABBAGE PALMETTO. Oc- casional throughout the prairies; rare in scrub. Serenoa repens (Bartr.) Small-SAW PALMETTO. Common throughout. AQUIFOLIACEAE Ilex cassine L.-DAHOON HOLLY. Rare in prairies. Ilex glabra (L.) Gray-GALLBERRY. Common in prairies and scrub. ASTERACEAE Ambrosia artemisiifolia L.__RAGWEED. Common in disturbed margins. Aster dumosus L.-ASTER. Rare in prairies. Baccharis halimifolia L._SALTBUSH. Rare in prairies. Balduinia angustifolia (Pursh) Robins.-BALDUINIA. Rare in scrub. Bidens alba (L.) DC.-SPANISH NEEDLES. Common in disturbed sites throughout. Carphephorus corymbosus (Nutt.) Torrey & Gray-FALSE BLAZING STAR. Infrequent in scrub and dry prairies. Centaurea cyanus L._BATCHELOR’S BUTTON. Not seen since 1971. Introduced. Conyza canadensis (L.) Cronquist-HORSETAIL. Common in disturbed prairies. Elephantopus elatus Bertol._ELEPHANT’S FOOT. Rare in scrub-dry prairie ecotone. Emelia fosbergii Nicolson-TASSLE FLOWER. Occasional in disturbed margins. Erechtites hieracifolia (L.) Raf. ex DC.-FIREWEED. Rare on disturbed margins. Erigeron strigosus Muhl.-DAISY FLEABANE. Occasional in disturbed margins. Eupatorium capillifolium (Lam.) Small-DOG FENNEL. Common in prairies. Euthamia tenuifolia (Pursh) Greene (=E. minor (Michx.) Greene, Soli- dago microcephala (Greene) Bush)-FLAT-TOP GOLDENROD. Common in prairies. Heterotheca graminifolia (Michx.) Shinners—(=Pityopsis graminifolia (Mich.) Nutt.)-GOLDEN ASTER. Common in prairies; occasional in scrub. Heterotheca subaxillaris (Lam.) Britt. -GOLDEN ASTER. Common in prairies; occasional in scrub. 20 FLORIDA SCIENTIST [Vol. 53 Lactuca graminifolia Michx.-WILD LETTUCE. Occasional in prairies. Liatris chapmanii Torr. & Gray-BLAZING STAR. Occasional in dry prairies. Liatris tenuifolia Nutt. BLAZING STAR. Occasional in scrub and dry prairies. Pluchea rosea Godfrey-CAMPHORWEED. Rare in prairies. Pterocaulon pycnostachyum (Michx.) Ell. (=P. virgatum sensu auct., non (L.) DC.)-BLACKROOT. Occasional in scrub. Palafoxia feayi Gray-PALAFOXIA. Occasional in scrub. Palafoxia integrifolia (Nutt.) Torrey & Gray-PALAFOXIA. Occasional in scrub. Pectis prostrata Cav.-PECTIS. Occasional in disturbed margins. Solidago odora Ait. var. chapmanii (Gray) Cronq. (=S. chapmanii Torr. & Gray)-GOLDENROD. Occasional in prairies. Solidago stricta Ait.-GOLDENROD. Infrequent in wet prairies. Tridax procumbens L.-TRIDAX. Occasional on disturbed margins. In- troduced. Vernonia cinerea (L.) Less.-IRONWEED. Rare in disturbed margins. Introduced. BLECHNACEAE Blechnum serrulatum L.C. Rich.-SWAMP FERN. Rare in wet prairies. BORAGINACEAE Heliotropium polyphyllum Lehm.-PINELAND HELIOTROPE. Occa- sional in dry prairies. BRASSICACEAE Lepidium virginicium L.-PEPPER GRASS. Occasional in disturbed mar- gins. Introduced. BROMELIACEAE Tillandsia balbisiana Schult.-RECURVED WILD PINE. Rare in ham- mocks, Tillandsia fasciculata Sw.-CARDINAL WILD PINE. Rare in ham- mocks. Considered threatened by the Florida Department of Agriculture. Tillandsia recurvata (L.) L.-BALL MOSS. Common in hammocks. Tillandsia usneoides (L.) L.-SPANISH MOSS. Occasional in hammocks. CACTACEAE Opuntia compressa (Salisb.) Macbr. (=Opuntia humifusa Raf.) Raf.)- PRICKLY PEAR. Common in disturbed prairies. Considered threatened by the Florida Department of Agriculture. Opuntia stricta Haw.-PRICKLY PEAR. Occasional in scrub and dry prairies. Considered threatened by the Florida Department of Agriculture. CAPPARACEAE Aldenella tenuifolia (Torrey & Gray) Greene( = Polanisia tenuifolia Torrey & Gray). Occasional in scrub. CARYOPHYLLACEAE Stipulicida setacea Michx. Common in scrub; occasional elsewhere. No. 1, 1990] AUSTIN—FAU VEGETATION ZA CHENOPODIACEAE Chenopodium ambrosioides L._MEXICAN TEA, EPIZOTE. Occasional in disturbed margins. Introduced. CHRYSOBALANACEAE Licania michauxii Prance-GOPHER APPLE. Abundant in scrub; com- mon in dry prairies. CISTACEAE Helianthemum corymbosum Michx. Occasional in scrub. Helianthemum nashii Britt. Common in scrub. Lechea cernua Small. Rare in scrub. An endangered species under review by the U.S. Fish & Wildlife Service. Lechea deckertii Small. Rare in scrub. Lechea divaricata Shuttlew. ex Britt. Occasional in scrub. COMMELINACEAE Commelina erecta L._DAYFLOWER. Infrequent in scrub and dry prai- ries. Cuthbertia ornata Small-ROSELING. Rare in scrub. CONVOLVULACEAE Ipomoea sagittata Poir-MARSH MORNING GLORY. Rare in wet prai- ries. CYCADACEAE Zamia pumila L.-COONTIE. Rare in scrub-hammock margin. Consid- ered threatened by commercial exploitation by the Florida Department of Agriculture. CYPERACEAE Bulbostylis ciliatifolia (Ell.) Fern. HAIR SEDGE. Common in scrub. Cyperus nashii Britt. (=Cyperus retrorsus Chapm.)-SCRUB NUT- GRASS. Common in scrub. Cyperus compressus L. Occasional in disturbed margins. Rhynchospora megalocarpa Gray-SCRUB SEDGE. Rare in scrub. Scleria ciliata Michx. Rare in scrub. Scleria reticularis Michx. Rare in wet prairies. ERICACEAE Befaria racemosa Vent.-TARFLOWER. Common in dry prairies; rare in scrub. Lyonia ferruginea (Walt.) Nutt. (=Lyonia fruticosa (Michx.) Torr.)- STAGGERBUSH. Common in dry prairies and scrub. Lyonia lucida (Lam.) D. Don-FETTERBUSH. Common in dry prairies. Vaccinium myrsinites Lam.-SHINY BLUEBERRY. Common in dry prai- ries and scrub. EUPHORBIACEAE Chamaesyce hirta (L.) Millsp.-SPURGE. Occasional on disturbed mar- gins. Chamaesyce hyssopifolia (L.) Small-SPURGE. Occasional on disturbed margins. 99 FLORIDA SCIENTIST [Vol. 53 Chamaesyce mendezii (Boissier) Millsp.-SPURGE. Occasional on dis- turbed margins. Cnidoscolus stimulosus (Michx.) Engelm. & Gray-STINGING NETTLE. Rare in scrub and dry prairies. Croton glandulosus L.-CROTON. Occasional in dry prairies and scrub. Euphorbia polyphylla Engelm.-SCRUB SPURGE. Common in scrub. Phyllanthus abnormis Baillon. Occasional in scrub. Phyllanthus amarus Schumm. & Thonn. Rare in disturbed margins. Poinsettia cyathophora (Murr.) Kl]. & Gke.-POINSETTIA. Occasional in disturbed margins. Stillingia sylvatica L.-QUEEN’S DELIGHT. Rare in scrub. Under re- view by the U.S. Fish & Wildlife Service as threatened. FABACEAE Abrus precatorius L.-PRECATORY PEA. Occasional in disturbed sites throughout. Introduced. Acacia auriculiformis A. Cunn. ex Benth.-EARLEAF ACACIA. Rare in disturbed margins. Introduced; first found in 1979. Cassia chamaecrista L. (= Cassia fasciculata Michx., Chamaecrista fasci- culata (Michx.) Greene)-PARTRIDGE PEA. Occasional in scrub and dry prairies. Cassia aspera Muhl. ex Ell. (=Cassia nicitans L. var. aspera (Muhl. ex Ell.) Torrey & Gray, Chamaecrista nicitans (L.) Moench var. aspera (Muhl. ex Ell.) Irwin & Barneby)-PARTRIDGE PEA. Infrequent in disturbed mar- gins. Cassia obtusifolia L. (=Senna obtusifolia (L.) Irwin & Barnaby)-SICK- LEPOD. Rare in disturbed margins. Introduced. Cassia pilosa L. (=Chamaecrista pilosa (L.) Greene) Infrequent on paths. Introduced. Crotalaria rotundifolia (Walt.) Gmelin (=Crotalaria maritima Chap- man)—-RATTLEBOX. Occasional in scrub and dry prairies. Desmodium incanum DC.-BEGGAR TICKS. Common in disturbed margins. Galactia elliottii Nutt.-MILK PEA. Common in dry prairies. Galactia volubilis (L.) Britt.-MILK PEA. Rare in scrub. Macroptilium lathyroides (L.) Urban (= Phaseolus lathyroides L.)WILD BEAN. Rare in disturbed margins. Introduced. Vigna luteola (Jacq.) Benth._COW PEA. Rare in disturbed margins. FAGACEAE Quercus chapmanii Sarg.-CHAPMAN’S OAK. Rare in hammocks and scrub. Quercus geminata Small-SAND LIVE OAK. Rare in dry prairie. Quercus minima (Sarg.) Small-RUNNER OAK. Common in scrub and dry prairies. Quercus myrtifolia Willd.-MYRTLE OAK. Abundant in hammocks. Quercus virginiana Mill.-LIVE OAK. Occasional in hammocks. No. 1, 1990] AUSTIN—FAU VEGETATION 23 IRIDACEAE Sisyrinchium solstitiale Bickn. (=S. xerophyllum sensu auct., non Bickn.)—-BLUE-EYED GRASS. Rare in scrub. LAMIACEAE Satureja rigida Bartr. ex Benth. (=Piloblephis rigida (Bartr. ex Benth.) Raf.)-PENNYROYAL. Rare in scrub and dry prairie. Trichostema dichotomum L.-BLUECURLS. Rare in scrub. LAURACEAE Cassytha filiformis L._DODDER. Occasional wherever shrubs grow. Persea borbonia L.-RED BAY. Rare in dry prairies. LOGANIACEAE Polypremum procumbens L.-RUSTWEED. Occasional in disturbed margins; common in scrub. MALVACEAE Sida acuta Burm. f.-INDIAN TEA. Occasional in disturbed margins. Sida cordifolia L._TEAWEED. Common in disturbed margins. Intro- duced. Urena lobata L.-CAESAR’S WEED. Rare in disturbed margins. Intro- duced. MORACEAE Ficus aurea Nutt.-STRANGLER FIG. Rare in low hammocks; a single epiphytic specimen on a cabbage palm. MYRICACEAE Myrica cerifera L. (=M. pusilla Raf.)-WAX MYRTLE. Occasional in dry prairies. OLACACEAE Ximenia americana L.-TALLOW WOOD. Rare in scrub. ONAGRACEAE Gaura angustifolia Michx.-GAURA. Common in disturbed margins. Ludwigia maritima Harper. Occasional in prairies. Ludwigia suffruticosa Walt. Occasional in wet prairies. PHYTOLACCACEAE Phytolacca americana L.-POKEWEED. Rare in disturbed prairies. PINACEAE Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg.-SAND PINE. Rare in scrub. Two trees and two seedlings remain as of Sept. 1988. Pinus elliottii Engelm.-SLASH PINE. Rare in dry prairies. Five native trees, in the 30-40 yr range, remain as of Sept. 1988; there are several seed- lings associated with some. Two trees and two seedlings from a planted source are in the southern portion. POACEAE Amphicarpum muhlenbergianum (Schult.) Hitchc.-BLUE MAIDEN- CANE. Occasional in prairies. Andropogon ternarius var. cabanisii (Hackel) Fern. & Grisc.-SPLIT- BEARD BLUESTEM. Frequent in wet prairies. 94 FLORIDA SCIENTIST [Vol. 53 Andropogon virginicus L._BROOMGRASS. Occasional in disturbed margins. Andropogon virginicus var. glaucus Hackel. (=A. capillipes Nash). CHALKY BLUESTEM. Infrequent in dry prairie. Aristida condensata Chapm.-BIG THREEAWN. Rare in scrub. Aristida spiciformis Ell.-WIRE GRASS. Infrequent in dry prairies. Aristida stricta Michx.-WIRE GRASS. Occasional in scrub. Cenchrus echinatus L.-SAND BUR. Rare in disturbed margins. Cenchrus incertus M.A. Curtis-SAND BUR. Abundant in disturbed mar- gins. Chloris petraea Sw. (=Eustachys petraea (Sw.) Desv.)-CROW’S FOOT GRASS. Occasional in disturbed margins. Dactyloctenium aegyptium (L.) Beauv.-EGYPTIAN GRASS. Rare in dis- turbed margins. Introduced. Digitaria longiflora (Retz.) Pers.-FINGER GRASS. Abundant in dis- turbed margins. Introduced; first noted in early 1980s. Elusine indica (L.) Gaertn.-GOOSEGRASS. Infrequent on disturbed margins. Introduced. Eragrostis ciliaris (L.) R. Br.-GOPHERTAIL LOVEGRASS. Abundant in disturbed margins and paths. Eragrostis refracta (Muhl.) Scribn.-COASTAL LOVEGRASS. Occa- sional in paths and wet prairies. Muhlenbergia capillaris (Lam.) Trin.-HAIRGRASS. Rare in wet prairies. Panicum maximum Jacq.-GUINEA GRASS. Occasional in disturbed prairies. Introduced. Panicum patentifolium Nash (= Dichanthelium sabulorum (Lam.) Gould & Clark)-PANIC GRASS. Occasional in scrub and dry prairies. Panicum purpurascens Raddi (=Brachiaria mutica (Forsk.) Stapf.)- PARA GRASS. Abundant in disturbed prairies. Introduced. Panicum repens L.-TORPEDO GRASS. Occasional in disturbed prairies. Introduced. Paspalum notatum Fluegge-BAHIA GRASS. Common on disturbed mar- gins. Introduced. Paspalum setaceum Michx.-CUTTHROAT GRASS. Occasional on dis- turbed margins. Rhynchelytrum repens (Willd.) C.E. Hubb.-NATAL GRASS. Abundant on disturbed margins. Introduced. Setaria geniculata (Lam.) Beauv.-LITTLE FOXTAIL. Occasional on disturbed margins. Sorghastrum secundum (Ell.) Nash-INDIAN GRASS. Infrequent in dry prairies. Spartina bakeri Merr.-CORDGRASS. Common in wet prairies. Sporobolus indicus (L.) R. Br.-SMUT GRASS. Abundant in disturbed margins. Introduced. Triplasis purpurea (Walt.) Chapman-PURPLE SANDGRASS. Infrequent in dry prairie beside paths. No. 1, 1990] AUSTIN—FAU VEGETATION 95 POLYGALACEAE Polygala grandiflora Walt.-MILKWORT. Occasional in disturbed margins. POLYGONACEAE Coccoloba uvifera (L.) L.-SEA GRAPE. One seedling found Sept. 1985. Polygonella gracilis (Nutt.) Meisn.-WIREWEED. Infrequent in scrub. Polygonella polygama (Vent.) Engelm. & Gray-JOINTWEED. Common in scrub. PORTULACACEAE Portulaca pilosa L.-PURSLANE. Occasional in disturbed margins. PTERIDACEAE Pteridium aquilinum (L.) Huhn-BRACKEN FERN. Occasional in dis- turbed sites throughout. RUBIACEAE Cephalanthus occidentalis L._BUTTON BUSH. Rare in wet prairie. Diodia teres Walt.-BUTTONWEED. Infrequent on disturbed margins. Galium hispidulum Michx.-BEDSTRAW. Rare in disturbed areas. Hedyotis nigricans (Lam.) Fosb.—Infrequent on disturbed margins. Hedyotis procumbens (J.F. Gmelin) Fosberg-INNOCENCE. Occasional in scrub and dry prairies. Richardia brasiliensis (Moq.) Gomez-RICHARDIA. Occasional in dis- turbed margins. Introduced. Richardia grandiflora (Cham. & Schlecht.) Steud.-RICHARDIA. Occa- sional in disturbed margins. Introduced; first appeared on campus in 1979. Richardia scabra L.—RICHARDIA. Occasional in disturbed margins. In- troduced. SCROPHULARIACEAE Buchnera americana L.-BLUEHEARTS. Occasional in dry prairies. Linaria canadensis (L.) Dum.-BLUE TOADFLAX. Occasional in scrub prairies. Scoparia dulcis L._SWEET BROOM. Occasional on disturbed margins. SELAGINELLACEAE Selaginella arenicola Underw.-SCRUB SPIKEMOSS. Rare in scrub. Considered threatened by the Florida Department of Agriculture. SMILACACEAE Smilax auriculata Walt.-CATBRIAR. Occasional throughout. SOLANACEAE Physalis walteri Nutt. (=P. viscosa sensu auct., non L.)-GROUND CHERRY. Occasional in scrub and prairies. Solanum americanum L.-~-NIGHTSHADE. Rare on disturbed margins. TURNERACEAE Piriqueta caroliniana (Walt.) Urban. Occasional on disturbed margins. URTICACEAE Pilea microphylla (L.) Liebm.-ARTILLERY PLANT. Rare on disturbed margins. Introduced. 96 FLORIDA SCIENTIST [Vol. 53 VERBENACEAE Callicarpa americana L.-BEAUTY BERRY. Infrequent in scrub; occa- sional in prairies. Lantana camara L.-LANTANA. Rare on disturbed margins. Lippia nodiflora (L.) Michx. (=Phyla nodiflora (L.) Greene)-CREEP- ING CHARLIE. Common on disturbed margins. VITACEAE Ampelopsis arborea (L.) Koehne-PEPPER VINE. Rare in dry prairies. Parthenocissus quinquefolia (L.) Planch.-VIRGINIA CREEPER. Infre- quent in dry prairies. Vitis rotundifolia Michx. (= Vitis munsoniana Simpson)-MUSCADINE GRAPE. Very abundant throughout. Vitis shuttleworthii House-CALUSA GRAPE. Abundant in prairies. ZYGOPHYLLACEAE Tribulus cistoides L._PUNCTURE VINE. Occasional on disturbed mar- gins. Introduced. ACKNOWLEDGMENTS— Thanks are due to students who studied various aspects of the site dur- ing the years, including Mary Brooks, J. N. Burch, E. V. Conroy, Jr., F. R. Posin, Donna Ritchey, and B. E. Tatje. I would also like to thank Mary Hoefs, Grace B. Iverson, Winifred Park, James W. Whitley, and my wife Sandra for reading early drafts of the manuscript and offering sugges- tions. T. T. Sturrock (Florida Atlantic University) provided information on the early use of build- ings on campus. Thomas Mann (Clemson Univ.) augmented my information on gopher tortoise populations, and gave helpful comments on the manuscript, Duncan M. Porter (Virginia Poly- technic Institute and State University) on Tribulus, and I. J. Stout (Univ. Central Florida) on feral cat effects on the Florida mouse. R. P. Wunderlin (University of South Florida) identified specimens of Eragrostis and made suggestions on the manuscript. LITERATURE CITED AUFFENBERG, W. AND R. FRANz. 1982. The status and distribution of the gopher tortoise (Gopherus polyphemus). Pp. 95-126 In: Bury, R. B. (ed.), North American Tortoises: Conservation and Ecology. Wildlife Research Report 12: U.S. Dept. Interior, Washington D.C. Austin, D. F. 1978. Exotic plants and their effects in southeastern Florida. Environm. Cons. 5(1):25-34. ., K. CoLEMAN-Marols AND D. R. RicHArpson. 1977(1978). Vegetation of southeast- ern Florida II-IV. Florida Scient. 48(4):331-361. ., F. R. Posin ano J. N. Burcu. 1987. Scrub species patterns on the Atlantic Coastal Ridge, Florida. Coastal Res. 3:491-498. Brooks, M. 1988. Species of a parcel of the FAU Preserve. Unpublished report. Florida Atlantic Univ., Boca Raton, FL. Brown, D. P. 1985. World War II in Boca Raton: The home front. Spanish River Papers 14(1):25 pages unnumbered. Burcu, J. N. anp G. A. Marsu. 1987. Survey of Natural Resources within the Limits of the Florida Atlantic University Campus. FAU/FIU Joint Center for Environmental and Ur- ban Problems. Convoy, E. V., Jr. 1970. Survey of vegetation on the FAU Campus. Unpublished report. Florida Atlantic Univ., Boca Raton, FL. Curt, D. W. 1976. Building Boca Raton’s airport. Spanish River Papers 5(1):15 pages unnum- bered. FrANz, R. AnD D. ToNNESSEN. 1983. The slow and steady decline of the Gopher Tortoise. Fla. State Mus. Notes 12(1):4. No. 1, 1990] AUSTIN—FAU VEGETATION 27 Gooprick, R. L. 1974. The wet prairies of the northern Everglades. Pp 47-52 In: GuEason, P. J. (ed.). Environments of South Florida: Present and Past. Mem. Miami Geol. Soc. 2. GopHER TorTOISE CouncIL. n.d. The Gopher Tortoise: A Species in Decline. leaflet. Jackson, D. R. 1985. Florida’s “Desert” tortoise. The Nature Conservancy News, Sept./Oct. p. 24-26. Mann, T. 1988. Clemson Univ., Clemson, $.C., Pers. Comm. Porter, D. M. 1970. Virginia Polytechnic Institute and State Univ., Blacksburg, Pers. Comm. RicHarpson, D. R., I. J. Srour, R. E. Roserts, D. F. Austin anp T. R. ALEXANDER. 1986. Design and Management Recommendations for a Sand Pine Scrub Preserve: The Yamato Scrub. 142 pp. RitcHey, D. M. 1988. Survey of a strip of the FAU “Triangle.” Unpublished report. Florida Atlantic Univ., Boca Raton, FL. Strout, I. J. 1986. Univ. Central Florida, Orlando, Pers. Comm. Sturrock, T. T. 1988. Florida Atlantic Univ., Boca Raton, Pers. Comm., Jan. TatyeE, B. E. 1979. A study of the burrowing owl population at Florida Atlantic University. Unpublished report. Florida Atlantic Univ., Boca Raton, FL. Tuomas, T. M. 1974. A detailed analysis of climatological and hydrological records of South Florida with reference to man’s influence upon ecosystem evolution. Pp 82-112. In: GLEa- son, P. J., (ed.) Environments of South Florida: Present and Past. Mem. Miami Geol. Surv. 2. U.S. Derr. or AcricuLTurRE. 1978. A Soil Survey of Palm Beach County Area, Florida. Washing- ton, D.C. Woop, D. A. 1985. Official lists of endangered and potentially endangered fauna and flora in Florida. Florida Game & Freshwater Fish Commission, Tallahassee, FL. WUuNDERLIN, R. P. 1982. Guide to the Vascular Plants of Central Florida. Univ. Presses of Florida, Tampa. , B. E. HANSEN AND D. W. Hatt. 1985. The vascular flora of Central Florida: taxo- nomic and nomenclatural changes, additional taxa, Sida 11(2):232-244. , B. E. HANSEN AND D. W. HA. . 1988. The vascular flora of Central Florida: taxo- nomic and nomenclatural changes, additional taxa, II. Sida 13(1):83-91. Florida Sci. 53(1):11-27. 1990. Accepted: January 30, 1989. Environmental Chemistry NONPOINT SOURCE PHOSPHORUS CONTROL BY A COMBINATION WET DETENTION/FILTRATION FACILITY IN KISSIMMEE, FLORIDA JEFFREY DEE HOLLER Smith and Gillespie Engineers, Inc., 7406 Manatee Street, Suite 1, Sarasota, Florida AssTRACT: Water quality investigations were conducted to assess the treatment potential (con- centration reduction) of a dual-component wet detention/filtration-berm stormwater manage- ment system, located in Kissimmee, Florida. Phosphorus concentrations are indicative of non- point source pollution in urban and commercial stormwater runoff. Therefore, orthophosphorus and total phosphorus concentrations were monitored at three different sampling stations within the system: 1) surface runoff influent channel; 2) wet detention basin standing pool; and 3) filtration-berm effluent collection box. Routine monthly data were collected to characterize prevalent ambient conditions. In addition, six distinct storm events were monitored with auto- matic samplers to characterize episodic phosphorus variations during the period November, 1985 to November, 1986. Statistical analyses (t-test) of routine monthly concentration data showed significant differences (p<0.05) between the stormwater influent and the wet detention basin standing pool samples for both orthophosphorus and total phosphorus. However, similar analyses between detention basin standing pool and filtration-berm effluent samples showed no signifi- cant differences. These results suggest positive treatment potential attained through wet deten- tion, but significant additional treatment was not realized through berm filtration. Storm event results reinforced these conclusions, indicating wet detention treatment potential far superior to filtration-berm treatment potential. The average storm event treatment potential realized by wet detention during six events for orthophosphorus and total phosphorus was 77%. The average treatment potentials realized by filtration for orthophosphorus and total phosphorus were -91 % and 16%, respectively. The average treatment potentials realized by the overall combined system for orthophosphorus and total phosphorus were 55 % and 85 % , respectively. In 1982, the South Florida Water Management District (SFWMD) initi- ated conceptual design of a pilot-scale stormwater management demonstra- tion project in the Lake Tohopekaliga watershed in the City of Kissimmee, Florida. A cooperative agreement was reached between SFWMD and the City of Kissimmee to construct and maintain a combination wet detention/ permeable filter-berm prototype stormwater management system for captur- ing and treating surface runoff from an urban/commercial watershed. This project became known as the Lake Tohopekaliga Demonstration (LTD) project. The purpose of the LTD project was to collect specific water quality (phosphorus), hydraulic, and other operational and maintenance data neces- sary to evaluate the effectiveness of a combination wet detention/filtration type facility to mitigate eutrophication problems resulting from nonpoint source stormwater runoff entering Lake Tohopekaliga. A major source of nutrient enrichment in surface waters is from nonpoint source stormwater runoff (Mattraw, et al., 1978). Researchers in central Florida found that the major source of phosphorus entering Lake Eola was from stormwater runoff (Wanielista et al., 1982). Accelerated eutrophication of Lake Tohopekaliga was suspected to be resulting from elevated phosphorus concentrations associ- No. 1, 1990] HOLLER— PHOSPHORUS CONTROL 29 KISSIMMEE, FL parkbox 4 | road parkin Fic. 1. LTD site plan ated with untreated stormwater runoff entering the lake. Therefore, the LTD facility was monitored to evaluate phosphorus treatment potential, which is defined here as the percent reduction in phosphorus concentration. This pa- per will examine routine and storm event investigations concerning facility performance to contain phosphorus associated with surface water runoff. 30 FLORIDA SCIENTIST [Vol. 53 Stupy AREA—The LTD project drainage basin comprises 75 acres of ur- ban/commercial land usage in Kissimmee, Florida, which is maintained by a conventional curb and gutter storm sewer collection system. The LTD project is generally located between Smith Street on the west and the East City Ditch on the east. The specific coordinates are the south one-half of Government Lot #2 (S22, T25S, R29E), Osceola County, Florida. The vicinity location and study site design schematic is indicated (Fig. 1). The treatment facility consists of a 200 x 400 ft rectangular pond, con- structed 100 ft east of the Park Street storm sewer outlet. The pond was constructed by building a levee (3 ft high) of native fill taken directly from the excavated depression. The pond sides were constructed at a ratio of 1:6. Wet detention storage was provided for the first one-half inch of runoff (1.46 ac-{t). The filtration berms were constructed in a radial finger pattern at the north end of the basin. Five individual finger-berms, each approximately 100 ft long, extended outward from one of two concrete drop boxes (total filter length = 1,000 ft). Each filtration-berm was constructed from six-inch diame- ter perforated PVC pipe, which was laid on the pond bottom and covered by one foot of filtration media. The filtration media consisted of native fill, limestone rock, and sand. The berm under-drains were laid at a slight slope (towards the drop boxes) of from 0.1-0.2 percent. A cross sectional design detail of the filtration berms is provided (Fig. 2). The combined wet detention/filtration-berm facility was designed to pro- vide a residence time of approximately two days. An emergency spillway was located on the southeast end of the pond to provide flood relief from the 25- year storm event (elevation=57.5 ft). Filtration-berm overflow discharge was provided by two screened inlet structures on top of each drop box, posi- tioned at 0.75 ft below the emergency spillway elevation (56.75 ft). MATERIALS AND MetHops—An original objective of this study was to monitor hydrology/ hydraulics characteristics to estimate water flow budgets for each storm event. A critical depth flume was installed at the basin inflow and a Parshall flume was installed at the basin outflow. The critical depth flume provided a well-defined channel section to facilitate the calculation inflow rates based on water depth. Similarly, the Parshall flume was intended to provide system surface water discharge flow data based on water depth differential (headwater-tailwater rela- tionships). Surface water stage recorders (Stevens Series 7000 digital punched-tape type) were installed on the upstream and downstream side of the flumes. Routine monthly grab samples were collected from three distinct sampling stations during the period March, 1984 through November, 1986. These stations included the basin inflow flume (Parkin); the wet detention basin standing pool adjacent to the filter-berms (Parkberm); and the westernmost filter-berm drop box structure (Parkbox). Sampling station locales are shown (Fig. DY. Storm events were defined as occurring at the beginning of stormwater runoff generated inflow entering the detention/filtration facility and ending at the cessation of outflow from the basin. Events were sampled on a wet season/dry season basis, with the goal of monitoring four distinct wet season and two distinct dry season events. The wet season was defined as running from 01 June through 31 October. Each distinct event monitored was separated from the previous one by a minimum of at least a four-day antecedent dry period of no significant rainfall in the subject watershed (less than 0.25”). No. 1, 1990] HOLLER— PHOSPHORUS CONTROL 31 —— ELEV. 56.0 2 FILTER MATERIAL F.D.0.T. NO.57 VE 1/3 SAND, 1/3 CLAY, ; : GRAVEL 1/3 TOPSOIL oh 4— PERMEABLE LINER 6" PVC (PERF.) RETENTION BTM. ELEV. 54.5 Fic. 2. LTD filter-berm section Due to geographical and logistical constraints, all storm event sampling duties were con- tracted to a private consultant (Aqua Chem Analyses, Inc. of Melbourne, Florida). Aqua Chem installed and maintained three automatic samplers (Sigmamotor 6200 series) at the above-men- tioned sampling stations. These samplers were activated by a single water level actuator, result- ing from a rise in water level within the basin. All samples were collected on a discrete, time- proportional basis. Sampling was terminated when 24 discrete samples were collected or the basin water level receded below the critical actuator elevation. A succession of discrete 500 ml samples were collected at each station according to the following time intervals: Parkin: 10 minutes, except first sample bottle, which was filled two minutes from when liquid level actuator was tripped; Parkberm: 2 hours, except first sample bottle which was filled one hour from when liquid level actuator was tripped. Parkbox: Same as Parkberm. 32 FLORIDA SCIENTIST [Vol. 53 All laboratory methods employed in sample analyses were performed according to “Standard Methods” or accepted EPA methodologies. A separate aliquot from each sample was frozen within 24 hours from the end of the sampling period for orthophosphorus (o-PO,-P) analysis, which was determined by molybdenum blue colorimetry. Total phosphorus (T-PO,-P) analyses were determined by acid digestion followed by molybdenum blue colorimetry (APHA, 1985). RESULTS AND Discusslon—Surface water stage data proved inconclusive. Stage data recorded during the study period indicated minimal head differ- ences across the Parshall flume (max. 0.1 ft), making it impossible to compute surface discharge rates. These minimal head differences resulted from severe submerged conditions persisting for extended periods. Submergence occurred because the water level in the basin receded at a much slower rate than originally intended. Typical head losses of 0.11 ft/day were common. Field studies were conducted in September, 1985 for the purpose of mea- suring filtration-berm percolation rates. Using a two-inch pump, a five-liter bucket, and a stop watch, one filtration-berm drop box was pumped out and measured under submerged conditions. These measurements represented the maximum filtration rate possible under free flow conditions. The actual flow rate for the entire system was much less due to submergence, and could not be determined due to a leak in one of the drop box pipe connections and incomplete installation of the Parshall flume. The percolation or filtration rates measured ranged from 0.016-0.083 cfs, and averaged 0.045 cfs (7.8 cu ft/ft/day). The under-drains were designed to provide a hydraulic surface loading rate of 100-150 cu ft/ft/day, with a maximum head over the filters of 1-3 ft. The actual reduced filtration-berm surface loading may be explained as resulting from sediment fines blocking the interstitial pore spaces of the media. In addition, massive growths of vegetation quickly inhabited the berm tops and surrounding pond area, serving to reduce available filtration capacity. Due to these unforeseen complications concerning the hydrology compo- nents of the system, it was not possible to estimate meaningful water budgets. Therefore, the system performance was evaluated based upon water quality considerations only. Phosphorus concentration data were used to determine treatment potential (percent concentration reduction) of both the wet deten- tion and filtration-berm components. Routine orthophosphorus (o-PO,-P) concentration data have been sum- marized in Table 1. The mean orthophosphorus concentration at the Parkin TABLE 1. LTD routine monthly phosphorus: summary statistics (1984-1987) Station Parkin Parkberm Parkbox o-PO,-P T-PO,-P o-PO,-P T-PO,-P o-PO,-P T-PO,-P n on i! 30 30 28 28 x 0.062 0.215 0.015 0.092 0.026 0.073 S 0.057 0.192 0.015 0.067 0.016 0.037 CV 92 89 100 73 38 51 aA]l values mg/L Except CV (%) No. 1, 1990] HOLLER— PHOSPHORUS CONTROL ao station was 0.062 mg/L. The mean orthophosphorus concentration within the water column at the Parkberm station (0.015 mg/L) was roughly four times less than the Parkin value. This observation is indicative of excellent soluble phosphorus removal. However, no additional treatment could be at- tributed to filtration, as the mean routine Parkbox concentration was 0.026 mg/L. The mean routine total phosphorus concentration at the Parkin station was 0.215 mg/L, approximately three times greater than the mean orthophosphorus value. The mean routine total phosphorus concentration at the Parkberm station was 0.092 mg/L, or approximately one-half the Parkin concentration. The Parkbox concentration (0.073 mg/L) was slightly less than the Parkberm value. These concentration data were subjected to the parametric t-test analysis to discern if any statistically significant differences existed between station mean values. Statistical significant differences between Parkin and Parkberm stations would indicate positive treatment attained through wet detention. Similarly, a comparison between stations Parkberm and Parkbox would re- veal possible significant treatment affected by filtration. A comparison be- tween stations Parkin and Parkbox would provide an indication of total over- all system performance. The results of these comparisons appear in Table 2. TABLE 2. LTD routine monthly data t-test results Station Comparison ( Treatment ) Parkin vs. Parkberm Parkberm vs. Parkbox Parkin vs. Parkbox Parameter (Detention) (Filtration) (Overall) o-PO,-P Sa NS S T-PO-,-P S NS S a §=Significant difference between means NS =No significant difference between means (p <0.05) Table 2 results indicate statistically significant (p <0.05) treatment attrib- uted to wet detention for both orthophosphorus and total phosphorus (Parkin vs. Parkberm). There was also significant positive treatment of both parame- ters shown for the overall system (Parkin vs. Parkbox). However, when the wet detention pond sample results (Parkberm) were compared with the filtra- tion-berm results (Parkbox), no statistically significant differences were shown between mean values for either parameter, indicating no additional treatment attributable to filtration. Six storm events were monitored from November, 1985 to November, 1986. These events ranged from 0.50-2.10 inches total precipitation. Ante- cedent dry periods ranged from 4 to 17 days. Table 3 shows meterological data associated with each event monitored. Storm event orthophosphorus and total phosphorus concentration data were summarized in Table 4. Surface water inflow (Parkin) event mean orthophosphorus concentrations ranged from 0.052-0.109 mg/L during the 34 FLORIDA SCIENTIST [Vol. 53 TaBLE 3. LTD storm event meterological data Event No. Date Time Rainfall (in Antecedent Dry Days I 20 NOV 85 14:00 0.05 5 2 14 MAR 86 11:00 2.10 10 3 12 AUG 86 18:00 1.75 t 4 13 SEP 86 17:30 1.00 4 5 07 OCT 86 04:00 0.60 17 6 24 NOV 86 16:30 0.70 9 six events monitored. These data were highly variable, as coefficients of vari- ation ranged from 15-95 percent. The mean orthophosphorus concentrations present in pond samples (Parkberm) during these events ranged from 0.012- 0.026 mg/L, and showed low variability (CV range = 21-58 percent). Filtra- tion-berm effluent sample (Parkbox) orthophosphorus event mean concentra- tions ranged from 0.009-0.059 mg/L, yielding a degree of variability consistent with pond samples (CV range = 15-59 percent). Inflow (Parkin) total phosphorus event mean concentrations ranged from 0.55-1.21 mg/L, with a degree of variability ranging from 33-107 percent. Wet detention pond (Parkberm) event mean total phosphorus concentrations ranged from 0.07-0.43 mg/L, and showed low variability (CV range = 26-56 percent). Filtration-berm (Parkbox) sample event mean concentrations ranged from 0.08-0.17 mg/L. These data showed a relatively high degree of variation (CV range = 18-100 percent). To gain an understanding of how these data (Table 5) relate to perform- ance of the detention, filtration, and overall system components, treatment potential was computed from event mean values. Orthophosphorus concen- trations were reduced 69-81 percent through wet detention. The mean treat- ment potential for orthophosphorus over the course of the six events was 77 percent, with an extremely low standard deviation of only 4 percent. Filtra- tion performance treatment potentials ranged from -321-50 percent, yielding a mean value of —91 percent. Filtration-berm treatment potential was ex- tremely variable (s= 147 percent). The filtration-berm performance vastly improved during events numbers 4-6, as compared to events numbers 1-3, possibly indicating stabilization of the filtration media as the system aged. Overall system performance (detention plus filtration) was intermediate be- tween detention and filtration component performance. Orthophosphorus treatment potentials ranged from 19-91 percent, with a mean value of 60 (s = 28 percent) percent over the span of all six events. No. 1, 1990] HOLLER— PHOSPHORUS CONTROL 35 TasBLe 4. LTD storm event phosphorus concentration summary statistics (Nov., 1985- Nov., 1986) eee aii ee ee Dern een er atkbox 77) Event No. o-PO,-P T-PO,-P o-PO,-P T-PO,-P o-PO,-P T-PO,-P 1 n 24 24 24 24 24 24 z 0.084 0.76 0.014 0.07 0.059 0.13 5 0.036 0.33 0.003 0.02 0.009 0.13 CV 43 43 21 28 15 100 2 n 24 94 Q4 24 94 24 Xx OF052 0.61 0.012 0.12 0.042 0.11 S 0.008 0.20 0.003 0.06 0.013 0.02 CV 15 oo 29 50 31 18 3 n 94 24 24 24 24 24 x 0.109 0.90 0.026 0.43 0.050 0.17 5 0.104 0.46 0.012 0.24 0.009 0.03 CV 95 5] 45 56 18 18 4 n 24 24 24 24 24 24 xX 0.087 eA 0.019 0.19 0.014 0.12 5 0.029 0.61 0.006 0.05 0.009 0.05 CV 33 50 33 26 59 42 5 n 94 24 24 94 24 24 x 0.095 0.73 0.018 0.13 0.009 0.08 5 0.069 0.78 0.010 0.05 0.003 0.02 CV 73 107 58 38 32 25 6 n 24 24 94 94 94 94 x 0.077 0.55 0.024 0.16 0.014 0.10 5 0.019 0.42 0.005 0.05 0.008 0.04 CV 24 76 21 31 59 40 All values mg/L Except CV (%) TABLE 5. LTD storm event phosphorus treatment potentialé Treatment Event No. Wet Detention Filtration Overall 0-EO)-P T-PO,-P o-PO,-P T-PO,-P o-PO,-P T-PO,-P 1 83 91 — 321 — 86 29 83 2, U7 80 — 250 8 19 82 3 76 52 — 92 60 54 81 4 78 84 26 $h7/ 84 90 5 81 a8 50 38 91 89 6 69 ral 42 38 82 82 K+s 77+4 77+13 —91+147 16+48 60 + 28 85+4 aAll Values Percent 36 FLORIDA SCIENTIST [Vol. 53 Total phosphorus treatment potentials are provided in Table 5. Wet deten- tion treatment potentials ranged from 52-91 percent (mean=77+s= 13 per- cent). Filtration provided total phosphorus treatment potentials ranging from —86-60 percent (mean = 16+s=48 percent). The total phosphorus treat- ment potential was enhanced by filtration, as values ranged from 81-90 per- cent (mean = 85+s=4 percent). Based upon phosphorus treatment potentials stated herein, an estimate of phosphorus loads may be obtained. If hypothetical flow values are utilized, pollutant mass can be determined, which enables the calculation of treat- ment efficiency. It has been previously shown that wet detention systems may typically detain 50% of surface inflow volume (Holler, 1984). If it is assumed that a chosen storm event produced 1.5 inches of precipi- tation and yielded approximate runoff inflows of 32 1/s, it follows that 16 1/s of surface outflow would result. Furthermore, phosphorus concentration data obtained during LTD storm event No. 2 may be used to compute storm event-based P-loads, and consequently treatment efficiency estimations. These assumptions yielded P-load-based treatment efficiencies of 88% and 90% for orthophosphorus and total phosphorus, respectively. Previously- stated treatment potentials of 77% for both parameters compare conserva- tively well with these treatment efficiency estimates. SUMMARY AND CoONCLuSIONS— The effectiveness of the LTD facility was evaluated based upon limited hydrological and extensive water quality data. Based on the design hydraulic loading rate (100-150 cu ft/ft/day) and the filter configuration, this system should have yielded a hydraulic conductivity in the range of 5.6-8.3 in/hr. This range would be typical of fine aggregate under-drains. However, given the high percentage of fine materials present in the LTD filter media, these values are probably optimistic. The LTD system filtration component performed poorly on a hydrologi- cal basis. There apparently was not enough difference between headwater and tailwater to effectively drive the system. Also, the filter-berms most likely became clogged with sediment fines shortly after construction was com- pleted. Phosphorus concentration data supported the poor hydrological perform- ance of the filter-berms. Routine phosphorus data showed significant reduc- tions in both orthophosphorus and total phosphorus concentrations attained through wet detention. However, no significant additional treatment was attained for either parameter through filtration. The overall system showed significant reductions of both parameters. Storm event data yielded the greatest treatment potential for orthophos- phorus through the process of wet detention (77 percent). Filtration actually served to increase orthophosphorus concentrations (mean treatment poten- tial=-91 percent). The overall system orthophosphorus treatment potential was 60 percent. Total phosphorus treatment potentials for detention, filtration, and over- all system components were 77, 16, and 85 percent, respectively. These No. 1, 1990] HOLLER— PHOSPHORUS CONTROL 37 results clearly indicate that the wet detention component was far superior to the filtration component for phosphorus control in the LTD stormwater man- agement system. Treatment efficiency estimates (based on assumed flow) yielded o-PO,-P and T-PO,-P values of 88% and 90%, respectively. Both indicators (treatment potential and treatment efficiency) proved compatible and comparable for the purpose of assessing phosphorus dynamics within the LTD system. ACKNOWLEDGMENTS— The author wishes to recognize the technical and financial support of the South Florida Water Management District and the analytical and logistical services of Aqua Chem Analyses, Inc., Melbourne, Florida. LITERATURE CITED AMERICAN PuBLic HEALTH ASSOCIATION. 1985. Standard Methods for the Examination of Water and Wastewater. 16th ed., APHA-AWWA-WPCF, New York. Ho.ter, J. D. 1989. “Stormwater detention basin nutrient removal efficiency.’ J. Water Res. Plan. Mgt., 115, (1):52-63. Mattraw, Jr., H. C., J. HARDEE, AND R. A. MILLER. 1978. “Urban stormwater runoff data for a residential area, Broward County, Fla.” Report 78-324, U.S. Geolog. Survey, Tallahassee, EL WanieLista, M. P., Y. A. YousEF, AND J. S. TayLor. 1982. “Stormwater management to improve lake water quality.” USEPA Grant 600/52-82-048. Univ. Central Florida, Orlando, FL. Florida Sci. 53(1): 28-37. 1990. Accepted: February 6, 1989. Biological Sciences SURVIVAL OF FLORIDA BAY FISH TAGGED WITH INTERNALLY ANCHORED SPAGHETTI TAGS GERALD M. Lupwic', JORGAN E. SKJEVELAND’, AND NICHOLAS A. FUNICELLI® 1U.S. Fish and Wildlife Service, Fish Farming Experimental Laboratory, P.O. Box 860, Stuttgart, AR 72160 2U.S. Fish and Wildlife Service, Chesapeake Bay Fisheries Coordination, 1825 Virginia St., Annapolis, MD 21401 3U.S. Fish and Wildlife Service, National Fishery Research Center— Gainesville, 7920 N.W. 71st St., Gainesville, FL 32606 Asstract: The percent survival of Florida Bay white mullet, striped mullet, spotted seatrout, and gray snapper marked with internally anchored spaghetti tags was 22, 67, 75, and 100, respectively. Unmarked fish of the same species had survival percentages of 27, 77, 75, and 100, respectively. The differences between the tagged and untagged fish were not significant. Trans- porting the fish up to 16 km from the point of capture did not significantly decrease survival in tagged or untagged fish either. THE use of tags to identify fish in mark-and-recapture studies to deter- mine movement or estimate population size is a common practice in fishery biology. To make accurate population estimates, one must know the mortality that results from the tagging operation, in addition to that from other causes (Lagler, 1956; Ricker, 1975). Studies on the initial mortality caused by tag- ging have been conducted on Atlantic herring, Clupea harengus harengus (Jensen, 1955; Winters, 1977), North Sea sole, Solea solea (Kotthaus, 1963), and Atlantic salmon, Salmo salar (Hiatt, 1963), among others. In some of these studies, tagging mortality was substantial. The U.S. Fish and Wildlife Service conducted experiments from January 1984 to September 1985 to determine movement and estimate population sizes of certain commercial and sport fishes in the marine waters of Ever- glades National Park. The area studied included most of Florida Bay and the Ten Thousand Islands section of the park. Experiments to determine the ini- tial mortality caused by tagging and the amount of tag shedding were con- ducted as a part of this research. MatTerIALs AND MerHops— Tagging mortality experiments were conducted at National Park ranger stations at Flamingo and Key Largo, and at several secluded sites in Florida Bay. Fish species marked were white mullet (Mugil curema), striped mullet (M. cephalus), spotted sea trout (Cynoscion nebulosus), and gray snapper (Lutjanus griseus). Fish were collected with gill or trammel nets, or by angling. The fish were then either trans- ported to holding tanks by boat in 150-liter, aerated plastic containers or placed into a frame cage covered with semirigid plastic mesh and located in the bay near the point of capture. The trans- ported fish were then held in a high-sided fiberglass tank supplied with continuously running sea water. Volume of the holding containers was about 800 liters. Water temperatures during the tests averaged 24.6°C and ranged from 13-31°C. For each experiment, half of the fish in each lot were measured and tagged with internally anchored tags and then placed in the cage or holding tank, the rest were placed directly into the No. 1, 1990] LUDWIG, ET AL.—TAGGED FISH SURVIVAL 39 holding container and served as a control. The number of fish in each test varied from three to 15 per treatment (Table 1), depending upon how many fish we were able to obtain during a single day. Containers were checked daily for 10 days (weather permitting, for fish held in the field) and fish survival and temperature were recorded. Dead fish were removed immediately. At the end of each experiment, ali fish were also examined to determine if they had shed tags. The mean, minimum and maximum total lengths in millimeters of the fish used for these experiments were: M. curema, 286, 181 and 335; M. cephalus, 358, 266 and 441; C. nebulosus, 372, 280 and 424; L. griseus, 252, 191 and 295. The tags used in this experiment were anchored internally. An incision about 10 mm long was made through the abdominal wall just anterior and slightly lateral to the vent. The plastic tag anchor, 25.4 x 6.4 x 0.5 mm, was inserted through the incision into the abdominal cavity. A yellow tubular plastic streamer, 50 mm long and 0.5 mm in diameter that was attached to the tag anchor extended externally through the abdominal wall. The percent survival of the tagged fish was compared with that of the untagged fish by analysis of variance with the SAS statistical software package (SAS Institute, Inc., 1985). The percent survival in each experiment was subjected to an arcsin transformation prior to the ANOVA (Sokal and Rohlf, 1981). Comparison of mortality between fish that had been trans- ported to a holding pen and those left in a pen near the point of capture was done with an odds ratio test (Fleiss, 1973). RESULTS AND DiscussioN—Survival was not significantly different (p =0.05) between tagged and untagged fish for white mullet, striped mullet, spotted sea trout or gray snapper (Table 1). Percent survival ranged from 0 to 100 for white mullet and striped mullet, 50 to 100 for spotted seatrout and TABLE |. Percent survival of tagged and untagged white mullet, striped mullet, spotted sea- trout and gray snapper captured and held for ten days at Key Largo or Flamingo, Florida, during 1984 and 1985. Tagged Fish Untagged Fish Species Location Date Number Survival (%) Number Survival(%) White Mullet Key Largo Mar. 2, 1984 8 100 10 100 Key Largo May 2, 1984 8 0 8 13 Key Largo Jan. 16, 1985 10 40 10 50 Key Largo Feb. 4, 1985 10 0 10 0 Key Largo Feb. 19, 1985 15 0 15 0 Key Largo Mar. 1, 1985 10 30 10 30 Key Largo’ Mar. 11, 1985 9 33 8 25 Key Largo’ Mar. 11, 1985 10 0 10 0 Key Largo’ Mar. 11, 1985 3 0 3 0 Flamingo Sep. 26, 1984 5 20 5 60 Total 88 22.3 89 27.0 Striped Mullet Key Largo Apr. 13, 1984 12 67 13 100 Flamingo Sep. 10, 1984 10 100 10 100 Flamingo Oct. 10, 1984 10 100 10 100 Flamingo Jan.7, 1985 10 0 10 0 Total 42 66.7 43 76.7 Spotted Seatrout Key Largo Mar. 16, 1984 10 50 10 60 Key Largo May 16, 1985 10 100 10 90 Total 20 75 20 75 Gray Snapper Flamingo Sep. 20, 1984 6 100 by 100 Flamingo Oct. 10, 1984 5 100 3 100 Total 11 100 8 100 40 FLORIDA SCIENTIST [Vol. 53 100% for gray snappers. The high degree of variability in the results of the experiment and the small sample size for three of the four species tested prob- ably reduced the chance of finding significant differences. However, because the percent survival of the tagged and untagged fish of each species within each experiment was similar, although highly variable between experiments, we concluded provisionally, that the difference in mortality between tagged and untagged fish probably was insignificant, and that the mortality ob- served was probably a result of capturing and handling. In an effort to determine if low survival of both tagged and untagged fish was the result of transporting the fish between the point of capture and the holding tanks, we conducted experiments in cages situated close to the area of capture and compared survival there with survival in the tank. Percent sur- vival in the cages was slightly higher than in the tank (Table 2) but it was not significantly so (odds ratio test) for white mullet, striped mullet, or spotted seatrout. The lack of significance was probably again related to the small sample size for the last two species. Of 45 striped mullet tagged in Everglades City, five (11%) lost their tags. Fish held at other sites lost no tags while in captivity. Table 2. Percent survival of tagged and untagged white mullet, striped mullet, spotted sea- trout and gray snapper held in a fiberglass holding tank at Key Largo or in net cages at the point of capture in Florida Bay, Florida, during 1984 and 1985. Species and Tagged Fish Untagged Fish location Number Survival (% ) Number Survival(% ) White Mullet Tank 40 28 40 32 Cage 43 16 43 JIS) Striped Mullet Tank 12 67 13 100 Cage 30 67 30 67 Spotted Seatrout Tank 10 50 10 60 Cage 10 100 10 90 Gray Snapper Cage 11 100 8 100 Total Tank 62 39 63 51 Cage 94 51 91 50 Conc.usions— This study suggests that marking white and striped mul- let, spotted seatrout and gray snapper with internally anchored tags does not increase mortality significantly over that of unmarked fish. In addition, al- though we attempted to handle all fish in a similar manner, the percent survival of tagged and untagged fish was extremely variable and, for white mullet, very low. We suspected that the highly variable but low survival of white mullet may have been related to the distances (sometimes greater than 16 km) that the fish were transported between the point of capture and the holding tanks. A comparison of survival of tagged and untagged fish that had No. 1, 1990] LUDWIG, ET AL.—TAGGED FISH SURVIVAL 4] been transported long distances and those that were not, indicated a consist- ently lower, but not significantly so, survival of the transported fish (Table 2). Although there have been extensive tagging studies in the coastal waters of Florida, few have quantified initial tagging mortality. The Schlitz tagging program of 1961-1965 included 18,000 fish tagged throughout Florida’s ma- rine waters (Beaumariage and Wittach, 1966; Beaumariage, 1969). That program was principally concerned with migration and growth and did not include experiments on survival of tagged fish. Broadhead and Mefford (1956) determined migration and exploitation of striped mullet in Florida during 1949-1953 on the basis of tag returns from 12,647 fish. As the tagging program progressed, it “became evident early in the tagging program that the tagged fish were not being subjected to the same mortality forces as were the untagged fish.” Although the authors did not conduct experiments on mortal- ity, they graphically analyzed the mortality of tagged fish on the basis of returns of tagged fish over time (30-day periods). An instantaneous mortality rate of 5.4 was determined by this method. Spotted seatrout have been the subject of two experiments on tagging mortality. Iverson and Moffett (1962) reported a tagging mortality of 4% in 94 fish (caught with gill and trammel nets and by angling) held for 3 weeks in submerged cages (half the fish were tagged with internally anchored tags and the rest were untagged). Moffett (1961), who held 15 spotted seatrout—5 tagged with internally anchored tags, 5 with body cavity tags, and 5 un- tagged—in a plastic cage for 66 days reported no mortality occurred; how- ever, one tag of each type was lost. Vogelbein and Overstreet (1987) studied long term (4 hr to 4 mo) pathological tissue changes caused by internal an- chor tags. They reported minor complications with spotted seatrout but rec- ommended continued use of the internal anchor tag for this species. ACKNOWLEDGMENTS— We thank Alex Gonzales, Patrick Mangan, Perry Oldenburg, James Rockowski, Donald Meineke, Michael Dewey, Leslie Mengel and Horace Bryant for helping us capture and transport the fish and construct the holding cages and Jane Dayton for typing the manuscript. LITERATURE CITED BEauManrliAGE, D. S. 1969. Returns from the 1965 Schlitz tagging program including a cumulative analysis of previous results. Technical Series No. 59. Florida Department of Natural Re- sources. Tallahassee. 38 pp. AND A. C. Wirticu. 1966. Return from the 1964 “Schlitz Tagging Program.” Technical Series No. 47. Florida State Board of Conservation. Marine Resources Laboratory. St. Petersburg. 51 pp. BROADHEAD, G. C. AND H. P. Merrorp. 1956. The migration and exploitation of the black mullet, Mugil cephalus L., in Florida as determined from tagging during 1949-1953. Technical Series No. 7. Florida State Board of Conservation. Marine Resources Laboratory. St. Petersburg. 33 pp. Fieiss, J. L. 1973. Statistical Methods for Rates and Proportions. John Wiley and Sons. New York, New York. 223 pp. Hairrt, S. C. 1963. Problems in tagging salmon at sea. Special Publication 4:144-155. Interna- tional Commission on Northwestern Atlantic Fisheries. Dartmouth, NS, Canada. Iverson, E. S. anp A. W. Morrerr. 1962. Estimates of abundance and mortality of a spotted seatrout population. Trans. Amer. Fish. Soc. 91(4):39. 42 FLORIDA SCIENTIST [Vol. 53 JENSEN, A. J. C. 1955. Danish herring tagging experiments inside the scaw. Rapp. P. V. Reun., Cons. Int. Explor. Mer 140(2):30-32. Kotrtnaus, A. 1963. Tagging experiments in the North Sea sole (Solea solea) in 1959 and 1960. Special Publication 4:123-129. International Commission on Northwest Atlantic Fish- eries. Dartmouth, NS, Canada. Lac.ter, K. F. 1956. Freshwater Fishery Biology. Wm. C. Brown Company. Dubuque, Iowa. 421 pp. MorrettT, A. W. 1961. Movements and growth of spotted seatrout, Cynoscion nebulosus (Cuvier) in West Florida. Technical Series No. 36. Florida State Board of Conservation, Marine Research Laboratory, St. Petersburg. 33 pp. SAS InstiruTE, Inc. 1985. SAS/STAT Guide for Personal Computers, Version 6 Edition. SAS Institute, Inc., Cary, North Carolina. 378 pp. SoKAL, R. R. AND F. J. RoHir. 1981. Biometry. W. H. Freeman and Company. San Francisco, California. 859 pp. Ricker, W. E. 1975. Computation and Interpretation of Biological Statistics of Fish populations. Bull. Fish. Res. Bd. Can. Bull. 191. 382 pp. VoGELBEIN, W. K. AND R. M. OverstreEET. 1987. Histopathology of the internal anchor tag in spot and spotted seatrout. Trans. Amer. Fish. Soc. 116:745-756. Winters, G. H. 1977. Estimates of tag extrusion and initial tagging mortality in Atlantic herring (Clupea harengus harengus) released with abdominally inserted magnetic tags. J. Fish. Res. Bd. Can. 34(3):354-359. Florida Sci. 53(1):38-42. 1990. Accepted: March 29, 1989. Biological Sciences POSTURES ASSOCIATED WITH IMMOBILE WOODLAND SALAMANDERS, GENUS PLETHODON C. KENNETH Dopp, Jr. National Ecology Research Center, U.S. Fish and Wildlife Service 412N.E. 16th Avenue, Room 250, Gainesville, Florida 32601 Asstract: The postures of immobile woodland salamanders (15 species of the genus Pletho- don) were observed in the field throughout the southeastern United States. There were a few significant differences among species in the frequency of three generalized posture categories, but these differences are not interpretable in terms of morphological, phylogentic, or ecological dif- ferences between species. Linear configurations were most commonly observed. Small-bodied species coiled more than large-bodied species. The duration of immobility varied significantly among posture categories, but salamander length was only marginally different among catego- ries. There probably are no specific immobility postures in Plethodon that serve solely antipreda- tor functions. Individuals remain immobile in the position in which they are encountered. IMmopi.ity (death-feigning, thanatosis) is a frequent behavior in pletho- dontid salamanders, both in temperate and tropical habitats (Brodie et al., 1974; Dodd and Brodie, 1976; Dodd, 1989). Presumably the behavior has selective value because of the frequency with which it is observed under field conditions. While the frequency and duration of occurrence of immobility have been quantified, there have been no detailed accounts of the postures that accompany the behavior. Coiling has been described in a variety of spe- cies (Brodie et al., 1974), and postures in which the body is kept immobile while the tail is undulated to attract predators to the part of the body which contains the greatest concentration of noxious secretions have been described and illustrated in tropical bolitoglossines (Dodd and Brodie, 1976). During the last 10 years, I observed salamanders of the genus Plethodon in the field to quantify various aspects of immobility (Dodd, 1989). No com- plex antipredator postures such as those of neotropical species have been seen, although some larger bodied Plethodon exhibit tail displays in proximity to predators under laboratory conditions (Brodie, 1977; Brodie et al., 1979). Here, I address whether salamanders assume specific postures during immo- bility which might enhance the effectiveness of the behavior as a defense against predators. MerHops—Salamanders were observed in the field by lightly raking litter and surface debris and by turning logs, rocks, or other large objects. A salamander was considered immobile if it did not move when touched, closely approached, or when nearby microhabitat was disturbed (Bro- die et al., 1974; Dodd and Brodie, 1976). Immobility terminated when the salamander first walked or moved, or when it righted itself from its back or side. I sketched the body configuration of the immobile salamander, and recorded a variety of measurements on salamander size and environmental conditions. Observations were made at locations throughout the southeastern United States from 1975 to 1985 (Dodd, 1989). 44 FLORIDA SCIENTIST [Vol. 53 Oo a Li SS ee ee Fic. 1. Schematic representation of 13 postural categories observed in immobile salamanders (Plethodon spp.) in the southeastern United States. Based on sketches of immobile salamanders, I initially recognized 13 postures (Fig. 1), as follows: (1) body in complete coil with head over or under tail; (2) body in partial coil with head near but not overlapping tail; (3) trunk of body straight but head and tail curved toward each other; (4) trunk of body broadly U-shaped with head and tail facing outward; (5) entire body broadly U-shaped; (6) main part of body straight but head and tail facing outward; (7) body in tight S shape; (8) body in loose S-shape with head and tail straight; (9) body slightly bowed, tail slightly flexed, head straight; (10) head and body in slight curve but tail coiled toward body; (11) body in a broad curve; (12) head and body straight but tail slightly curved; and (13) head, body, and tail completely straight. Salamander postures cannot be measured on an ordinal scale because there is not a continuous measurable progression in postures from | to 13, and there is a degree of subjectivity involved in placing them in each category. In addition X2 analysis of these categorical data, using a 13 x 15 contingency table, would provide many cells with expected values <5.0 thus providing a poor estimation of X2. For statistical analysis, the 13 posture categories were combined into three general categories to reflect a progression from coiled to straight (1-2, body coiled or semi-coiled; 3-8, various degrees of body contortions; 9-13, head and body more or less straight). Differences between species in posture frequencies were tested using a 3 x 15 X? contingency table. Differ- ences between posture categories in terms of duration of immobility and snout-vent length were tested using a 2-way analysis of variance. Statistical procedures were carried out using the SAS program for microcomputers (SAS Institute, 1985); the level of significance was set at a=0.05. RresuLts—Posture data were recorded for 868 individuals of 15 species. Immobile salamanders were likely to be found in any of the 13 posture cate- gories, although the linear body form was most common (Table 1). There were significant differences among species in the number of animals found in the three major immobility categories (x?= 105.1, p. <0.01, 28 df). However, there were no clear differences in postures between large (glutinosus, jordani, 45 DODD—SALAMANDER POSTURES No. 1, 1990] 6°06 69 OL c'6 © Or 0°9 ve £6 9°6 15S SS oS 69 % GLI LS €9 8L c8 6P 86 61 GL LY cY eV ES TROL [21040 1039}€') Aq [8101qNS aassojyouoh g lapyom snqpjound J 1upp.ol g snsouruns J satoeds as1e'T (% 8¥) 79% (% PE) E8T % (81) 96 03978) Aq [B102qNS ENP EMUD al 1ajsqan J yoopunuays J SNIDLIAS J ipuoWwYIU J isulqjau q oe Ua runuiffoy a SYDSLOP J I snalauio J satoadg ]]BwsS wt OwUWn SG I~ - WON ~-ontron OtaONns a4 4 organo ANA oO omnMmoo OONA a4 Oto NtANO aN OO oooc;eCc Il YHOO MOdOONAN OS NTO tOT On AS SOMNOWMWNTMNO CO maoowtrwaomonead Sr OoNnnNwWnnA ATTN TWH ONWRNTON rae TONnWmooon oonoowtono- OwWMMDODOVOMNINANANN © OA ONAN TR TNO TFNANWMANOCOhR- ATO ft maa OMNMDOAON © COI-NHTNDOOTM ©O G G G CG 1 él IT Ol 6 8 L 9 SI P t YysIeI}S po}10}U07) ee) A1089}') 91N}sog satoads N lm "$911039}¥9 UTBUI 9914} JY} JopuN siaquInu Aq pazeorpur sainysod jo SUOI}CIIOSap 10F 4X9} 99g ‘S9}e}S po}HU, UIO}seayyNOs ay} Ul (‘dds uopoyjajg) s1apueUWL]es o[IqouIUT Aq pournsse sainjsod JUarosfIp Jo Aouonbaiy *[ ATAV], 46 FLORIDA SCIENTIST [Vol. 53 punctatus, wehrlei, yonahlossee) and small species, although large-bodied species (sensu Highton and Larson, 1979) were more likely to occur in linear configurations than small-bodied species, such as P. cinereus or P. welleri. Whereas small species tended to display more body curvature while immobile than large species, they were not found in any one posture more frequently than any other posture. Only small-bodied species were found fully coiled (Table 1). When the expected values were compared with observed values within the contingency table, P. punctatus was found coiled less often, and P. hu- brichti more often, than expected. P. glutinosus was either contorted or straight, but rarely coiled. P. jordani was generally straight, and less con- torted and coiled than expected, whereas P. richmondi was often coiled or contorted, but less often straight. P. welleri was generally coiled, and less contorted or straight than expected. None of these differences reflect morpho- logical, phylogenetic, or ecological differences between species. The length of the salamander varied among the three posture categories, but the relationship was only marginally significant (F = 2.88, p=0.057, 781 df). The smallest individuals were in Category 1 (N=100, x=38.8 mm), the largest in Category 2 (N= 269, x=45.1 mm), and intermediate sizes in Cate- gory 3 (N=455, x=43.0 mm). The duration of immobility varied signifi- cantly among the posture categories (F = 7.38, p=0.0007, 781 df). Salaman- ders with postures in category 1 remained immobile longest (X=84.6 s) whereas those in category 3 were immobile for the shortest time (X=60.9 s). Salamanders in category 2 were intermediate (x=79.3 s). In no cases were there significant differences among species (length: F=0.92, p=0.5764; tine: F—1..36. p—O0n1090): No complex antipredator postures were observed in Plethodon as have been reported for neotropical species (Dodd and Brodie, 1976). Only infre- quently did salamanders flip and then become immobile, regurgitate the stomach contents, void fluid from the vent, or wrap the tail around the ob- server's hands during handling (Table 2). All species exuded a viscous secre- tion from the body and tail when handled. Immobile salamanders sometimes clasped limbs to the body when dis- turbed, but more frequently the limbs were extended away from the body (Table 2). Salamanders accidentally knocked onto their backs or sides usually remained immobile in that position rather than immediately righting them- selves. This was especially true of smaller species and small individuals of larger species. Discusston—Among Plethodon, the duration of immobility is inversely correlated with substrate temperature but not with the snout-vent length (SVL) of the salamander, although r’ values are low (Dodd, 1989). I postu- lated that variation in the intensity of disturbance, though unquantified in my study, may explain variation in the time salamanders remain immobile rather than inherent species-specific, size-related, or temperature-related in- fluences (Dodd, 1989). No. 1, 1990] DODD—SALAMANDER POSTURES 47 TaBLeE 2. Behavior and body configuration associated with immobile salamanders (Plethodon spp.) observed in the field (N = 868). Terminology follows Dodd and Brodie (1976). Behavior N Species (Numbers of individuals) Flipping 3 glutinosus (1), shenandoah (2) Legs Clasped 4 cinereus (1), glutinosus (1), punctatus (1), wehrlei (1) Legs Extended 30 cinereus (16), dorsalis (2), hubrichti (1), nettingi (2), punctatus (5), richmondi (1), serratus (1), websteri (1), welleri (1) On Back 12 cinereus (5), dorsalis (4), punctatus (1), websteri (2) On Side 10 cinereus (7), jordani (1), punctatus (1), welleri (1) Regurgitation 2 welleri (2) Void Fluid 2 punctatus (2) Wrap Tail 2 hubrichti (1), welleri (1) As an additional hypothesis, I suggest that the prior activity of the sala- mander may influence the length of time it remains immobile in response to a predator. Animals in a coiled or strongly curved body position presumably had been inactive prior to disturbance and thus remained immobile longer than animals in a linear posture. The smallest salamanders were coiled more often, but these were species such as P. cinereus and P. richmondi with slen- der, elongate trunks and relatively small limbs. Thus, they mechanically are better able to coil or contort their bodies than more robust, larger-limbed species, such as P. glutinosus or P. jordani. Presumably, salamanders are confronted by predators during routine daily activities such as feeding or reproductive behavior. To maximize feeding efficiency, the animal may assume a straight or minimally-curved body posi- tion that allows effective use of the lingual musculature and hyobranchial apparatus to secure prey (Lombard and Wake, 1976, 1977) and to maximize the chance of encountering prey. A salamander in a tight coil would be at an obvious disadvantage when feeding. In a straight or minimally-curved pos- ture, an immobile salamander is concealed from both its prey and predators. Conversely, a salamander in a tight coil may minimize water loss (Ray, 1958) and at the same time be smaller and less conspicuous to predators. Immobil- ity again may have a dual function, one antipredator and the other physio- logical. Small-bodied salamanders in both the temperate zone and the neotropics are found coiled more often than large-bodied species (Brodie et al., 1974; Dodd and Brodie, 1976; this study). A flip from a coiled or curved position followed by a resumption of immobility could startle and confuse a predator, and the immobile animal would then be difficult to find. The flip-immobility behavior may offset the lack of large granular glands, which contain noxious secretions, in small-bodied species. However, flipping is rare in small-sized 48 FLORIDA SCIENTIST [Vol. 53 temperate Plethodon, and some large-bodied species also flip in response to predators (Brodie et al., 1979; Arnold, 1982). Thus, defense against preda- tion is not a satisfactory explanation for the greater numbers of small sala- manders found in coiled positions. Brodie (1977) categorized salamander antipredator postures as active (i.e., postures which orient parts of the body in such a manner that glands containing noxious secretions are directed at a predator) and passive. Passive postures such as immobility enhance survival by not drawing a predator’s attention to the animal, or by minimizing the intensity of the attack (Ratner, 1967; Edmunds, 1974). No body orientation is involved, and the “posture” is actually the maintenance of a non-moving body position when a predator is present. As such, no unique body positions are associated with immobility, and one may question whether it is appropriate to speak of “immobility pos- tures.” Morphological constraints, the type of activity when disturbed, and physiological adaptations are more likely to explain the body position of im- mobile salamanders than antipredator explanations. Postures of immobile Plethodon are indeed passive (sensu Brodie, 1977), although they may be preadaptive for the active postural displays of other groups of salamanders (Brodie et al., 1974; Dodd and Brodie, 1976). ACKNOWLEDGMENTS—I thank Howard I. Kochman and John Oldemeyer for providing advice and guidance on statistical procedures. Ronn Altig, Edmund D. Brodie, Jr., R. Bruce Bury, William Cooper, Robert Jaeger and John Oldemeyer read early drafts of the manuscript and gave valuable criticism. I thank the superintendents of the Blue Ridge Parkway, Great Smoky Moun- tains National Park, and Shenandoah National Park for allowing observations within their juris- diction. Hugh Morton, owner of Grandfather Mountain, Linville, North Carolina, graciously allowed access without charge. The South Carolina Wildlife Department granted permission to conduct research on P. websteri in the Stevens Creek Natural Area. This research was funded, in part, by grants from the Highlands Biological Station, Highlands, North Carolina. LITERATURE CITED ARNOLD, S. J. 1982. A quantitative approach to antipredator performance: salamander defense against snake attack. Copeia 1982:247-253. BroniE, E. D., Jr. 1977. Salamander antipredator postures. Copeia 1977:523-535. , J. A. JOHNSON, AND C. K. Dopp, Jr. 1974. Immobility as a defensive behavior in salamanders. Herpetologica 30:79-85. , R. T. Nowak, AND W. R. Harvey. 1979. The effectiveness of antipredator secretions and behavior of selected salamanders against shrews. Copeia 1979:270-274. Dopp, C. K., Jr. 1989. Duration of immobility in salamanders, genus Plethodon (Caudata: Plethodontidae). Herpetologica 45:467-473. AND E. D. Broptg, Jr. 1976. Defensive mechanisms of neotropical salamanders with an experimental analysis of immobility and the effect of temperature on immobility. Her- petologica 32:269-290. Epmunps, M. 1974. Defence in Animals. A Survey of Anti-predator Defences. Longman Group Ltd., Essex, Great Britain. HicuTon, R. anp A. Larson. 1979. The genetic relationships of the salamanders of the genus Plethodon. Syst. Zool. 28:579-599. Lomsarb, R. E. anp D. B. Wake. 1976. Tongue evolution in the lungless salamanders, family Plethodontidae. I. Introduction, theory and a general model of dynamics. J. Morphol. 148:265-286. No. 1, 1990] ACKNOWLEDGMENT OF REVIEWERS 4Y . 1977. Tongue evolution in the lungless salamanders, family Plethodontidae. II. Func- tion and evolutionary diversity. J. Morphol. 153:39-80. Ratner, S. C. 1967. Comparative aspects of hypnosis. Pp. 550-583. In: Gorpon, J. E. (ed.), Handbook of Clinical and Experimental Hypnosis, Macmillan, New York. Ray, C. 1958. Vital limits and rates of desiccation in salamanders. Ecology 39:75-83. SAS InstiTuTE, INc. 1985. SAS Introductory Guide for Personal Computers, Version 6 Edition. SAS Institute, Inc., Cary, North Carolina. Florida Sci. 53(1):43-49. 1990. Accepted: April 4, 1989. ACKNOWLEDGMENT OF REVIEWERS It is a pleasure to acknowledge the service, dedication, and cooperation of our reviewers, who include the following persons. They gave generously of their time and expertise in reviewing manuscripts for Volume 52 of the Fiorida Scientist. Some reviewed more than one manuscript. Robert S. Braman Kendall L. Carder Walter J. Conley Bruce C. Cowell Patricia M. Dooris Jack W. Frankel Richard Franz Patrick J. Gleason Joseph C. Joyce John M. Lawrence James N. Layne Paul E. Moler George M. Padilla Louis A. Penner Peter C. H. Pritchard Harold L. Schramn, Jr. William S. Seiler Joseph L. Simon Jack I. Stout William H. Taft Daniel B. Ward Robert J. Wimmert Richard P. Wunderlin 50 FLORIDA SCIENTIST [Vol. 53 NOTE ON THE FEEDING BEHAVIOR OF THE COMMON ATLAN- TIC MARGINELLA PRUNUM APICINUM’ (GASTROPODA, MARGINELLIDAE)—Thomas M. Baugh, Department of Biological Sciences, Jacksonville University, Jacksonville, FL, USA! ABstTRACT—A report of Prunum apicinum Menke (1828) feeding on the calcareous algae Jania pumila. On March 29, 1989 a mass of the coralline algae Jania pumila Lamoufoux, attached to the seagrass Syringodium filiforme Kuntzing, was collected in shallow water adjacent to the NASA Causeway, Florida (State Route 405). The algal mass measured about 8 cm tall and 4 cm wide, at its widest. In the laboratory, the seagrass and the attached algae were placed in a 38 liter aquarium already containing a number of different species includ- ing one 10 mm long common Atlantic marginella (Prunum apicinum). On March 30, the Prunum apicinum was observed feeding on the algal mass. Feeding began at the top and continued, apparently uninterrupted, until the animal reached the bottom of the mass on April 8. By April 8, the algae had been reduced to a small amount of calcareous detritus, composed of uneaten cylindrical segments, at the base of the Syringodium filiforme. There is little published information on the feeding ecology of Prunum apicinum (Coovert, 1988). Perry and Schwengel (1956) refer to this species as a carnivore while D’asaro (1970) adds that it is a nonselective carnivore and a scavenger. In aquaria, Raeihle (1965) fed Prunum apicinum the mussel Myti- lus edulis, while Winner (1983) fed cut shrimp. This note, however, appears to be the first report of this species feeding on a coralline algae. ACKNOWLEDGMENTS—I thank Conrad White and Joel Snodgrass, Brevard County, Florida, Office of Natural Resources Management for helping in the identification of Jania pumila, the Merritt Island National Wildlife Refuge for their support of this and related work, and Gary Coovert, Dayton Museum of Natural History, for information on the Marginellidae. LITERATURE CITED Coovert, G. A. 1988. A bibliography of the recent marginellidae. Marginella Marginalia. Day- ton Museum of Natural History 5(1-5). Dayton, Ohio. D’asaro, C. N. 1970. Egg capsules of prosobranch mollusks from South Florida and the Bahamas and notes on spawning in the laboratory. Bull Mar. Sci. 20(2):414-440. MENKE, K. T. 1828. Synopsis methodica molluscorum generum omnium et specierum eorum, quae in Museo Menbeana Adservantur; cum synoymia critica et noverum specirum diagnosibus, i-Xii, 1-91. G. Uslar, Pyrimonti. Perry, L. M. AND J. S. SCHWENGLE. 1956. Marine Shells of the Western Coast of Florida. Palentol. Res. Inst. RAEIHLE, D. 1965. Studies of captive Prunum apicinum Menke. Am. Malacol. Union 32:21-22. Winner, B. E. 1983. The direct development of Marginella (Prunum) apicinum. Of Sea and Shore. 13(1):37-38. Florida Sci. 53(1):50. 1990. Accepted: June 20, 1989. 1Address correspondence to the author at 2115 Nathan Drive West, Jacksonville, Florida 32216 No. 1, 1990] DAWES— BOOK REVIEW ill REVIEW Diane S. Littler, Mark M. Littler, Katina E. Bucher, James N. Norris. Marine Plants of the Caribbean. A Field Guide from Florida to Brazil. Smithsonian Institution Press, Washington D.C. 1989. Pp. vii + 263. Price $14.95, paperback. The small (7.5 x 5”) field guide contains attractive color photographs of 204 algae and 5 seagrasses with a brief (15 pages) introductory chapter and a final chapter (8 pages) on underwater photography. There is also a small glossary to cover the limited terminology used in the species descriptions. The layout of the guide includes a color photograph and a single descriptive para- graph for each plant. The species descriptions are limited to the gross morphology of the plants as are the macrophotographs. Thus the text, although pleasing from a photo- graphic standpoint, is not useful in species identification because of the lack of more critical morphological features (e.g. Bryopsis plumosa, Colpomenia sinuosa). The species selected for the guide are, for the most part, the more common ones, although Sargassum filipendula, a common large brown sea- weed, was not included while two unknown algae were included. The species descriptions are written for the layperson (“slimy, gooey, pink- ish”) but are helpful. The underwater photographs are quite good, and thus the guide should be useful for the casual observer who would like to match some of the Caribbean algae found with names. The suggested price is quite reasonable considering the color photographs—Clinton J. Dawes, Depart- ment of Biology, University of South Florida, Tampa, FL 33620. Biological Sciences NOTES ON PLANTS ENDEMIC TO FLORIDA SCRUB STEVEN P. CHRISTMAN"”) AND WALTER S. JUDD” 1Department of Natural Sciences, Florida Museum of Natural History, Gainesville, FL 32611 2Department of Botany, University of Florida, Gainesville, FL 32611 AsstractT: A three-year field survey of Florida interior sand pine scrubs, conducted by the first author and funded by the Florida Nongame Wildlife Program, has resulted in a better understanding of the distributions of many Central Florida scrub endemic plants. The status and distributions of 21 species (or varieties) of plants that are endemic (or nearly endemic) to the Lake Wales, Lake Henry, and Winter Haven ridges in central peninsular Florida are reported. Habitat notes, range extensions, and corrections of erroneous literature records are documented. Several species of scrub plants seem threatened with imminent extinction in the wild, and several others are rapidly approaching that condition. FLoripa’s unique scrub habitat develops on wind-swept sand dunes be- hind active coastal beaches, in the interior of the peninsula in natural fire shadows adjacent to aquatic or wetland systems, and in disturbed sandy up- lands such as overgrazed flatwoods or sandhills in which the frequency of fire has been much reduced (Myers, 1985; Christman, 1988a). Florida scrub can occur on yellow (Astatula, Paola soil series) or on white (St. Lucie, Archbold, Satellite soil series) sands. Along the hydrologic gradient, scrubs vary from low scrubs, dominated by Serenoa repens (Bartr.) Small, Lyonia lucida (Lam.) K. Koch, and Ilex glabra (L.) A. Gray, to high rosemary balds, domi- nated by Ceratiola ericoides Michx. Between these extremes, and depending on fire frequency, scrubs can be dominated by Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg., P elliottii Engelm., or by one or more of four species of sclerophyllous oaks (Quercus inopina Ashe, Q. geminata Small, Q. chap- manii Sarg., and Q. myrtifolia Willd.). Turkey oak (i.e., Quercus laevis Walt.) barrens are a natural plant com- munity intermediate between high pine and scrub and characterized by elements of both, as well as several distinctive plant species (Myers and Boett- cher, 1987; Christman, 1988a, 1988b). Scrubs on the Lake Wales, Winter Haven, and Lake Henry Ridges (see White, 1970) of central peninsular Florida occupy remnants of beach and sand dune systems that were associated with Miocene, Pliocene or Early Pleistocene shorelines (White, 1970; Winker and Howard, 1977). The antiq- uity of Florida scrub is suggested by the large number of plant and animal species that are endemic to this habitat. Scrubs on the Lake Wales, Lake Henry, and Winter Haven Ridges (hereafter called the Central Ridges, Fig. 1) are home to ca. 40 species (or varieties) of plants that occur nowhere but scrubs in Florida. About 17 of these species occur only on the Central Ridges. The scrubs on the Central Ridges that support populations of these endemic species may be referred to as ancient scrubs. About 200 ancient scrubs have No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 53 been identified in Highlands, Polk, Orange, and Osceola Counties, all above 25 m MSL and most on private lands (Fig. 2). AA _ESPANOLANL\ — 3 4 P oe " 2 Leal \ = n x 2 z q re \7 q Ow 9 LAKELANO ORDONVILLE RIOGE f} z S RIDGE Ye fe} s BOMBING re ° 2 i N « Deter \ py APPROX. SCALE —_— te oe \ ~ hey Fic. 1. The Lake Wales, Lake Henry, and Winter Haven Ridges (from White, 1970) are considered to be biogeographically related, and are treated together as the Central Ridges. Among the animals, the Florida scrub jay (Aphelocoma c. coerulescens), the scrub lizard (Sceloporus woodi), the sand skink (Neoseps reynoldsi), and the blue-tailed mole skink (Eumeces egregius lividus) are restricted to Florida scrubs, and the last two occur only in scrubs on the central Florida peninsula. Mark Deyrup (1987) estimates that as many as 40 species of arthropods are endemic to central Florida scrubs. Central Florida is undergoing rapid and intense development. Many of the plant species restricted to Florida scrub are listed (Table 1) as endangered or threatened species by the U.S. Fish and Wildlife Service (USFWS) or the state of Florida, and many others are under review for listing (Martin, 1987). About 80% of the upland habitats on the southern Lake Wales Ridge have been converted to citrus groves or residential developments (Peroni and Abra- 54 FLORIDA SCIENTIST [Vol. 53 Fic. 2. Distribution of Scrub Islands on the Lake Wales, Lake Henry, and Winter Haven Ridges. The route of US 27 follows the Lake Wales Ridge, and the population centers of Haines City, Lake Wales, Avon Park, Sebring, Lake Placid, and Venus are indicated with two-letter abbreviations that are each 3 km tall, by scale, on this and all succeeding plots. hamson, 1985a & b), and about two-thirds of the ancient scrubs on the Cen- tral Ridges have been lost (Christman, 1988a). MetTHops— Scrubs in Central Florida were identified on black and white aerial photographs (Hurd, Mark, Inc., 1973), county soil survey maps and USGS topographic maps. Over 400 scrub “islands” were visited and searched for scrub endemic plants. Voucher specimens were deposited in the herbarium of the Florida Museum of Natural History (FLAS). Areas of scrub islands were calculated electronically from outlines drawn on USGS topographic maps, and represent the areas of entire scrubs. Rarely, of course, does a plant species occur throughout an entire scrub. Most scrub plants have specific microhabitat requirements that are not met everywhere in a particular scrub. Therefore, the area estimates presented below should be considered as maxi- mum potential habitat. RESULTS AND Discussion—Table 1 provides the current listed status and the number of protected populations of 40 species of vascular plants that are No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 55 basically restricted to Florida scrub, including 17 that are restricted to scrub or natural turkey oak barrens on the Central Ridges. USGS topographic maps with the ancient scrubs delineated on them are on file at the Tallahassee office of the Florida Natural Areas Inventory (FNAI), as are the precise localities known for the rare scrub plants. TABLE 1. Species of vascular plants that are basically restricted to peninsular Florida scrub. Those marked with an asterisk are restricted to scrubs on the Lake Wales, Lake Henry, and Winter Haven ridges in Central Florida. The counties of occurrence are provided for species restricted to three or fewer. Scientific name Listed status! Protected Common name (family) FNAI FLA USA populations *Ziziphus celata G1/Sl — PE none Florida jujube (Rhamnaceae; Polk & Highlands Cos.) *Lupinus westianus var. aridorum Gl/Sl -E E none Scrub lupine (Fabaceae; Orange & Polk Cos.) *Dicerandra christmanii G1/Sl_ E E none Yellow scrub balm (Lamiaceae; Highlands Co.) Chrysopsis floridana G1/Sl]_ E E none Florida golden aster (Asteraceae; Hillsborough Co.) *Crotalaria sp. nov. —- — = none Avon Park Crotalaria (Fabaceae; Highlands Co.) Dicerandra cornutissima Cl/Sil E none Long-spurred scrub palm (Lamiaceae; Marion & Sumter Cos.) Dicerandra immaculata G1/S1l] E E none Lakela’s mint (Lamiaceae; St. Lucie & Indian River Cos.) *Dicerandra frutescens G1/Sl E E one Scrub balm (Lamiaceae; Highlands Co.) *Eryngium cuneifolium GIS! E E one Wedge-leaved button-snakeroot (Apiaceae; Highlands Co.) Polygala lewtonii G1?/Sl E Cl one Lewton’s polygala (Polygalaceae; scrub/high pine ecotone) *Conradina brevifolia G2/S2. — C2 two Short-leaved rosemary (Lamiaceae; Polk & Highlands Cos.) Warea carteri Gl/Sl E E two Carter’s warea (Brassicaceae; scrub/high pine ecotone) Eriogonum longifolium var. gnaphalifolium G3Q/S3_ T C2 two Scrub buckwheat (Polygonaceae; scrub/high pine ecotone) Calamintha ashei G3/S3. TT Cl two Ashe’s savory (Lamiaceae) *Chionanthus pygmaeus C7526 E two Pygmy fringe-tree (Oleaceae; also in xeric hammocks) *Asimina tetramera GI/Sl] E E three Four-petaled pawpaw (Annonaceae; Martin & Palm Beach Cos.) *Hypericum cumulicola G2/S2 E E three Highlands scrub hypericum (Clusiaceae; Polk & Highlands Cos.) *Bonamia grandiflora G3/S3 E alt four Scrub morning glory (Convolvulaceae) *Liatris ohlingerae C3iS3n aE PE four Scrub blazing-star (Asteraceae; Polk & Highlands Cos.) *Polygonella myriophylla G2/S2. — Cl four Sand-lace (Polygonaceae) *Polygonella basiramia G3/S3 iE E four Hairy jointweed (Polygonaceae; Highlands and Polk Cos., also on Bombing Range Ridge) “Paronychia chartacea G2/S2. — T five Papery whitlow-wort (Caryophyllaceae) 56 FLORIDA SCIENTIST [Vol. 53 TABLE |. Continued Scientific name Listed status! Protected Common name (family) FNAI FLA USA populations Conradina grandiflora G3/S3, 0 — C2 <10 Large-flowered rosemary (Lamiaceae) *Schizachyrium niveum G1/S1?_ E C2 <10 Riparian autumngrass (Poaceae) *Prunus geniculata G2/S2 E E <10 Scrub plum (Rosaceae) Lechea cernua G3/S3. — C2 <10 Nodding pinweed (Cistaceae) *Nolina brittoniana G2/S2. — C2 <10 Scrub beargrass (Nolinaceae; also in high pine) *Clitoria fragrans G3/S3. iE Cl <10 Pigeon-wing (Fabaceae; Highlands & Polk Cos., scrub/high pine ecotone) Persea humilus? G3/S3. — C3C >10 Silk bay (Lauraceae) Pinus clausa _ — >10 Sand pine (Pinaceae; also Florida panhandle, SW Alabama) Carya floridana — — >10 Scrub hickory (Juglandaceae) Ceratiola ericoides — — — >10 Florida rosemary (Empetraceae) Asclepias curtissii G3/S3.T — >10 Scrub milkweed (Asclepiadaceae) Garberia heterophylla — aly — >10 Garberia (Asteraceae) Sabal etonia = T _ >10 Scrub palmetto (Arecaceae) Ilex opaca var. arenicola G3/S3. — C3C >10 Scrub holly (Aquifoliaceae) Osmanthus megacarpus — — — Ie Scrub wild-olive (Oleaceae) Quercus inopina _ o “= ip Scrub oak (Fagaceae) Sisyrinchium xerophyllum — — — ? Scrub blue-eyed grass (Iridaceae) *Bumelia tenax “lacuum entity” _ — — I? Scrub buckthorn (Sapotaceae; at most only a distinctive ecotype/geographic race of B. tenax; considered specifically distinct by Lakela (1963) but conspecific by Godfrey (1988) and Whetstone (1983)). Status key —FNAI: Gx/Sx=Global and State endangerment rankings for Special Element species of the Florida Natural Areas Inventory, from 1 (critically imperiled) to 5 (demonstrably secure). T = Rank for taxo- nomic subgroup; Q = Validity of taxon questioned. FLA: E, T, = Listed by the State of Florida as Endangered or Threatened. USA: E, T=Protected by federal law as Endangered or Threatened. PE, PT =Proposed for listing as Endangered or Threatened. Cl, C2, C3C =Under review or formerly under review for federal listing, with—(1) Substantial evidence indicating vulnerability and threat(s) supporting the appropriateness of proposing to list the species; (2) not enough information on hand to support a listing proposal at the present time; (3C) evidence that the taxon is more abundant or widespread than formerly b Here and/or not subject to any identifiable threat. 2Taxon that sometimes is treated as a variety of Persea borbonia (L.) Spreng. (e.g., Little, 1979), but here is considered to be specifically distinct because of significant differences from P. borbonia in density and length of appressed ferrugineous hairs on its abaxial leaf surfaces and in flavonoid complement (see Wofford, 1973). No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 57 Several natural area preserves on the Central Ridges provide protection for ancient scrubs and some of the scrub endemics. The Archbold Biological Station in southern Highlands County protects about 315 ha of scrub and 13 Central Ridge endemic species. The Nature Conservancy’s (TNC’s) Saddle Blanket Lakes Preserve covers about 30 ha in Polk County and offers protec- tion for 11 Central Ridge endemics. The proposed Lake Arbuckle State Park will include about 300 ha of scrub and populations of 10 endemic scrub plants. TNC’s Tiger Creek Preserve does not contain any true scrub, but does have natural turkey oak barrens and populations of five species of Central Ridge endemics. At Catfish Creek in Polk County, TNC has recently acquired about 120 ha, including scrub and securing protection for about five scrub endemics. Two central Ridge scrub endemics are protected at Highlands Hammock State Park. Two or three are protected at Turkey Lake and Lakes Cain and Marsha Parks, small parks in Orange County. Four species of Cen- tral Ridge endemics are not protected anywhere, and 12 have fewer than five protected populations. Species for which new distributional information has recently become available are discussed below. Ziziphus celata Judd and Hall Thought to be extinct (Judd and Hall, 1984), this unusual shrub was recently rediscovered in two small scrub sites on the Lake Wales Ridge in Polk and Highlands Counties (Wunderlin, 1985, 1989). The status of these small popu- lations, both on privately owned land, is said to be precarious. Lupinus westianus Small var. aridorum (McFarlin ex Beckner) Isley This Federally endangered scrub endemic occurs only rarely and sporadically along road shoulders and on private building lots within two small regions: about 230 hectares around Winter Haven in Polk County, and about 1000 hectares around Vineland and Windermere in Orange County, on the devel- oping outskirts of Orlando (Fig. 3). About 15 populations are known. The Winter Haven localities are on the Winter Haven Ridge, and the Orange County localities are on the northern end of the Lake Wales Ridge and the adjacent southern tip of the Mount Dora Ridge (White, 1970). Populations are known only from disturbed white sand scrubs. An undocumented record from southwest of Lake Arbuckle (USFWS, 1987b), some 50 km from the nearest known population, is probably based on a misidentification of L. diffusus Nutt., which is common there. The scrub lupine is not known to occur on any protected lands, nor is it likely that any preserves can be established for it. Lupinus westianus var. aridorum will not likely make it into the 21st century, at least not in the wild. Dicerandra christmanii Huck and Judd In 1948 an amateur botanist named Ray Garrett collected specimens of an unusual woody mint from just east of Sebring on the Lake Wales Ridge in Highlands County. He believed the material represented a new genus and species related to Dicerandra, a genus then known to contain but three her- baceous species in Georgia and northern Florida. Apparently Garrett’s speci- 58 FLORIDA SCIENTIST [Vol. 53 a N 0 5 10 KM LW Fic. 3. Distribution of Lupinus aridorum. mens were not seen by Shinners (1962b) when he described D. frutescens from “20 miles south of Sebring,’ or by Kral (1982) in his revision of the genus. Subsequent authorities (Wunderlin, 1984; Huck, 1987) referred Gar- rett’s specimens, now at FLAS, to D. frutescens Shinners. Although reported as extirpated (Wunderlin, 1984; USFWS, 1985), the Sebring population of Dicerandra is still present. In fact, some 40 ha of high sandy uplands supporting a thriving population of Dicerandra and other Lake Wales Ridge endemics are present at what is undoubtedly Garrett’s original locality. However these mints cannot be referred to D. frutescens. They represent another, hitherto unrecognized, species that was rediscovered by the first author during the course of the present study, and recently was named D. christmanii (see Huck et al. 1989). No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 59 Fic. 4. Distribution of Dicerandra christmanii. This new mint has since been found at four additional small yellow sand scrubs covering about 16 ha scattered along 5 km of the eastern flank of the Lake Wales Ridge (Fig. 4; Huck et al., 1989). Unless the first locality can be secured, there is likely no possibility of preventing the extinction in the wild of this species within the next few years. The largest site is part of a tract that is planned to be converted to citrus, and the other four sites are either very small (one has fewer than 20 individuals) or part of a planned subdivision. Dicerandra frutescens Shinners Scrub balm occurs only in the vicinity of Lake Placid in Highlands County, where it is restricted to yellow sand scrubs on Astatula and Paola soils. It has not been observed in sandhills or in white sand scrub, except along ecotones with the distinctive yellow sand scrub (Christman, 1988a). Characterizations of D. frutescens habitat as sandhills (Wunderlin, 1984; USFWS, 1985) and the soils as St. Lucie fine sand (Huck, 1987) are misleading. Depending on 60 FLORIDA SCIENTIST [Vol. 53 how they are counted, there are 9-15 known sites for D. frutescens covering about 60 ha and distributed over about 30 km of the Lake Wales Ridge (Fig. 5; Huck et al., 1989). Protected populations of this Federally endangered species occur only at Archbold Biological Station, at the southern terminus of the plant’s range. All other populations are on private lands awaiting devel- opment. The prospects for securing and/or protecting another natural popu- lation of D. frutescens are very slim. The survival of this species in the wild will probably depend entirely on the plants growing at Archbold. i Se Fic. 5. Distribution of Dicerandra frutescens. A specimen (Fulton 237, 1978) of Dicerandra frutescens in the Herbarium of the University of South Florida (USF) bears the locality data: “Lake Placid turn-off from Highway 70 E, ca. 1 mi. in.” This has been interpreted as “one mile north of SR 70 on Placid View Drive” by FNAI (Hardin, 1986) and the U.S. Fish and Wildlife Service (Martin, 1987). As understood, this locality represents the westernmost reported for the species and the only one in low white sand scrub. Intensive searching by the first author, Wunderlin, and others has failed to find Dicerandra at this locality. Until confirmed, this locality should be considered doubtful. No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 61 N | ++ , | ' 0 5 10 KM VIN Fic. 6. Distribution of Eryngium cuneifolium. Eryngium cuneifolium Small This Federally endangered perennial occurs only in high and dry open white sand scrubs (rosemary balds) on the southern tip of the Lake Wales Ridge in Highlands County (Fig. 6). Within this limited area, E. cuneifolium is re- stricted to blowouts and other highly disturbed soil surfaces. There are about 20 known localities (but some of these nearly adjacent) on private lands and road shoulders, totaling less than 2000 hectares, from Lake Placid to Venus, a distance of about 28 km. Within these scrubs, E. cuneifolium occupies less than 10% of the area. Another remnant population (about 3 ha) occurs about 10 km farther north, just south of Lake Jackson. This disjunct population, on vacant building lots, will soon be extirpated. Records from Collier and Put- nam Counties (Johnson, 1981; USFWS, 1987a) were based on misidentifica- tions (Hardin, 1986). Kral (1983) plotted the species in Polk County, but no records exist. Eryngium cuneifolium is protected only in rosemary balds that total about 20 ha at the Archbold Biological Station. The Nature Conserv- ancy is investigating the possibility of protecting a second population at a privately owned ranch near Venus. 62 FLORIDA SCIENTIST [Vol. 53 Conradina brevifolia Shinners Another scrub-endemic woody mint, the mis-named short-leaved rosemary occurs sporadically in white sand scrubs from Sun Ray to Sebring, a distance of just 30 km, in southern Polk and northern Highlands Counties (Fig. 7). Currently under review for Federal listing (Martin, 1987), C. brevifolia is considered by one authority (Wunderlin, 1982) to be conspecific with C. canescens (Torrey & A. Gray) A. Gray of the Florida panhandle. In his syn- opsis of the genus, Shinners (1962a) recognized the taxon; it was also recog- nized by Kral (1983). Whatever its taxonomic status, the Lake Wales Ridge Conradina is distinctive and highly endangered. Conradina brevifolia is pro- tected only at TNC’s small (30 ha) Saddle Blanket Lakes Scrub Preserve in Polk County. Populations also occur on Florida’s newly-acquired Lake Ar- buckle State Forest, where they may or may not be protected, depending on timber management priorities. \ LW Fic. 7. Distribution of Conradina brevifolia. No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 63 A record in FNAI from Lake Placid, about 35 km south of known locali- ties, was based on a misidentified specimen (Judd 2888, 1980) of Calamintha ashei. Hypericum cumulicola (Small) P. Adams One of the most bizarre of the scrub endemic plants, this little perennial St. John’s wort is completely restricted to the high and dry white sand scrubs and rosemary balds of the Central Ridges, where it occupies highly disturbed bare sand surfaces. Over a distance of about 90 km between Lizzie Lake, south- west of Lake Wales in Polk County, and Venus in Highlands County (Fig. 8), H. cumulicola populations have been found on about 70 scrubs whose total areas cover about 6000 ha. Several populations of this species are known to have disappeared within the past few years, and some as yet unknown vul- nerability may cause H. cumulicola to be one of the first of the scrub endem- ics to go extinct. Protected populations of this Federally endangered species occur only at Arbuckle State Park, Saddle Blanket Lakes Scrub Preserve, and Archbold Biological Station within white sand scrubs that total about 300 ha. LW Fic. 8. Distribution of Hypericum cumulicola. VN 64 FLORIDA SCIENTIST [Vol. 53 Fic. 9. Distribution of Liatris ohlingerae. VN Liatris ohlingerae (Blake) B. L. Robinson The endangered scrub blazing star occurs nowhere but scrubs (white and yellow sand) on the Central Ridges in Polk and Highlands Counties from Lake Wales to Venus, a distance of about 90 km (Fig. 9). Ninety-five scrubs that harbor L. ohlingerae populations cover about 8000 ha. Protected popu- lations occur at Arbuckle State Park, Saddle Blanket Lakes Scrub Preserve, Highlands Hammock State Park, and the Archbold Biological Station, in scrubs that total less than 200 ha. Calamintha ashei (Weatherby) Shinners The woody mint, Ashe’s savory, is currently under Federal review for listing (Martin, 1987) and is listed by the State of Florida as a Threatened species. Populations of C. ashei have been found at 64 white sand scrubs on the Lake Wales Ridge, from Haines City to Venus (about 115 km), totaling less than 7000 ha. A disjunct population occurs in openings and road shoulders within sand pine scrub in the southwestern corner of the Ocala National Forest in Marion County. The plants on the Ocala occur within an area of about 7000 No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 65 ha. Records from Georgia (Wharton, 1978; Kral, 1983) need further study (Wunderlin et al., 1980a), but the species undoubtedly occurs in Tattnall Co., Georgia. Arbuckle State Park and Archbold Biological Station have the only protected populations of C. ashei. Although the plant may continue to persist along road shoulders within the intensively-managed sand pine plantations of the Ocala National Forest, it will probably be extirpated within the planta- tions as the Forest Service increases its use of chemical herbicides as detailed in the recent National Forest vegetation management plan (USDA, 1988). The first author has been unable to locate this species at a reported local- ity near Citra in Marion County (D’Arcy s.n., 1968: cited in Wunderlin et al., 1980c), and the locality may be in error because no scrub could be found at that location. Also, many of the older herbarium records have locality data that should not be taken literally. For example, the type specimen from “near Ocala,” and an early collection from “near Astor Park” probably both refer to the known population half way between those two cities. Godfrey (Godfrey 78279, 1980) reported C. ashei near Juniper Run on SR 19, some 13 km east of known localities. Efforts to relocate the species in this area have been unsuccessful. The species was also reported by J. Stout (field survey, 1981, FNAI data) from Surveyor’s Scrub on the Winter Haven Ridge in Polk County, but A. Laessle (field survey, 1951), G. Schultz (field survey, 1983), and Christman (1988a) did not see the species there. Polygonella myriophylla (Small) Horton Sand-lace occupies portions of about 119 scrubs, which cover about 10,000 ha on the Central Ridges from Vineland in the southwestern corner of Orange County to Lake Placid in Highlands County (Fig. 10). Formerly under re- view for federal listing, P myriophylla was withdrawn from consideration because it was believed that it was distributed widely throughout peninsular Florida (Wunderlin et al., 1980b; Martin, 1987). This was an unfortunate error, because P. myriophylla is actually much rarer, and more habitat-re- stricted, than several Federally listed scrub species, such as Paronychia char- tacea Fernald, Chionanthus pygmaeus Small, Polygonella basiramia (Small) Nesom & Bates, and Prunus geniculata Harper. A report of Polygonella myriophylla from Lake County (Hansen & Ri- chardson 5270, 1979; cited in Wunderlin et al., 1980b) was based on a mis- identification (Hansen, 1986). Records from Orange and Osceola Counties are correct, but these localities are still on the Lake Wales Ridge, and are all less than 15 km from the Polk County line. Records plotted from DeSoto County in Kral (1983) are based on specimens collected in Highlands County before it was split off from DeSoto County. Populations of P. myriophylla are protected only at Lake Arbuckle State Park, Saddle Blanket Lakes Scrub Preserve, and Archbold Biological Station, in scrubs whose total area is less than 300 ha. Curiously, the species does not range all the way to the southern end of the Lake Wales Ridge, and the population at Archbold consists of widely scattered individuals, just barely persisting in the shade of large sand pines, a result of long-term fire exclusion. 66 FLORIDA SCIENTIST [Vol. 53 VN Fic. 10. Distribution of Polygonella myriophylla. Polygonella basiramia (Small) Nesom & Bates Occurring in scrubs on the Central Ridges and nearby Bombing Range Ridge in Polk and Highlands Counties (Fig. 11), Polygonella basiramia disperses and colonizes new scrubs better than the other endemics treated here. Poly- gonella basiramia ranges from the southern tip of the Lake Wales Ridge to Venus to the vicinity of Lake Pierce in Polk County. This Federally endangered species is protected at Highlands Hammock and Lake Arbuckle State Parks, Archbold Biological Station, and Saddle Blanket Lakes Scrub Preserve. Not withstanding accounts to the contrary (Wunderlin et al., 1982; USFWS, 1987a), Polygonella basiramia is common in the scrubs on the Avon Park Bombing Range, the only Central Florida scrub endemic that occurs on the Bombing Range Ridge. Paronychia chartacea Fernald Papery whitlow-wort (Federally threatened) occurs in scrub and open dandy habitats on the Central Ridges from Windermere in Orange County to Venus No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 67 in Highlands County. The Windermere localities are actually on the southern tip of the Mount Dora Ridge as delineated by White (1970), less than 10 km north of the line that separates it from the Lake Wales Ridge. This annual colonizes more readily than most of the other scrub endemics, and has been found in newly created habitats such as road shoulders and cleared sandhills. Paronychia chartacea is restricted to disturbed sand surfaces. A record from longleaf pine—wiregrass sandhills just west of Lake Apopka in Lake County (Ward 8334, 1972; cited in Wunderlin et al., 1981) is based on a misidentified P. americana (Nutt.) Fenzl ex Walpers. However, the species is known from a single Lake County locality in a scrub on the north- west edge of Lake Louisa (Christman 2223, 1989). Nolina brittoniana Nash Scrub beargrass is not restricted to scrub, but occurs in sandhills, hammocks, turkey oak barrens and scrubs scattered throughout the Central Ridges. This species is not quite as restricted geographically, either, and is known from Lake Orange, Osceola, Polk, Hernando, and Highlands Counties in Central Florida. A record (West s.n., 1928; cited in Wunderlin et al., 1980f) from 68 FLORIDA SCIENTIST [Vol. 53 “low ground” at Velleview in Marion County is questioned (and may be N. atopocarpa Bartl.). Prunus geniculata Harper The Federally endangered scrub plum is not restricted to scrub as indicated by Ward (1979b), but occurs in sandhills, scrub, and intermediate habitats on the Central Ridges in Highlands, Polk, Orange, Osceola, and Lake Counties. The species is protected at several places including Tiger Creek, Saddle Blan- ket Lakes, Arbuckle State Park, Bok Tower Gardens, and Babson Park Audu- bon Nature Park. Fewer than 50 individuals still persist in scrubs planned for development in Orange and Osceola Counties. Chionanthus pygmaeus Small The Federally endangered pygmy fringe tree is not restricted to scrub as indicated by Ward and Godfrey (1979a) and Wunderlin et al. (1980c), but occurs in xeric hammocks, sandhills, scrub, and intermediate habitats on the Central Ridges in Highlands, Polk, Osceola, and Lake Counties. Records from Hillsborough and Citrus Counties (FNAI data base) need verification. This species does not occur at Archbold Biological Station and is known to be protected only at Tiger Creek and Saddle Blanket Lakes Preserves. About 42 populations are known on private lands. Bonamia grandiflora (A. Gray) Heller Scrub morning glory (Federally threatened) occurs in about 50 scrubs on the Lake Wales Ridge that total about 3000 ha. A disjunct population in the vicinity of Bradenton in Sarasota and Manatee Counties has been reported extirpated (USFWS, 1987c). Although it formerly occurred there, scrub morning glory has not been seen at Archbold Biological Station in decades (Austin, 1979). Another disjunct population occupies portions of sand pine scrub in the southwestern corner of the Ocala National Forest in Marion County, but is threatened by commercial pulp wood activities including site preparation and potential use of herbicides. Although it has a wider geographic range than some of the other scrub endemics treated here, Bonamia grandiflora is very spotty in its occurrence (Fig. 12) and individual populations are apparently quite vulnerable to ex- tinction. Scrub morning glory is protected only at Turkey Lake and Lakes Cain and Marsha Parks near Orlando, Lake Arbuckle State Park, and Tiger Creek Nature Preserve where no more than a dozen individuals have been found. Protected populations occur in scrubs that total less than 200 ha. Undocumented reports from Hardee County (Johnson, 1981) and Char- lotte County (Darden, 1987) need verification. Warea carteri Small Carter’s mustard is listed as endangered by USFWS, and is considered to be “restricted to sand pine scrub vegetation” (USFWS, 1987a). However, this species actually occurs in natural turkey oak barrens and the ecotone between scrub and high pinelands. Warea carteri is rarely found in pure scrub. This species is an annual that probably maintains a seed bank in the soil, and in some years, populations do not appear above ground (Bissett, 1986; pers. No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 69 Fic. 12. Distribution of Bonamia grandiflora on the Central Ridges. The species occurs also in a disjunct population in Marion County, and formerly occurred in Sarasota and Bradenton counties. observ.). Warea carteri is not a strict Lake Wales Ridge endemic, and has been found in oak scrub in Brevard County (USFWS, 1988) and formerly occurred in Dade County, but has not been seen there in 50 years and is now considered extirpated (Nauman, 1980). A record from Liberty County (Nauman, 1980) is undoubtedly based on a misidentification of the similar W. cuneifolia (Muhl. ex Nutt.) Nutt. The species is known to be protected only at Archbold Biological Station and Tiger Creek Nature Preserve, where it is abundant. The related, and also very rare, W. amplexifolia (Nutt.) Small (see Judd, 1980) is found mainly in high pinelands or turkey oak barrens. Clitoria fragrans Small Butterfly pea occurs in scrub and in habitats intermediate between scrub and sandhills (turkey oak barrens) in Polk and Highlands Counties (Wunderlin et 70 FLORIDA SCIENTIST [Vol. 53 al., 1980e). The species is known to be protected only at Archbold Biological Station and Tiger Creek Preserve. Thirty-three populations are known on private lands. Eriogonum longifolium Nutt. var. gnaphalifolium Gand. Scrub buckwheat occurs in habitats intermediate between scrub and sand- hills (turkey oak barrens) from Marion County to Highlands County. About 20 populations covering less than 1500 ha are known. A population in the southwestern corner of the Ocala National Forest is threatened by forestry operations. Scrub buckwheat is known to be protected only at Archbold Bio- logical Station and Tiger Creek Preserve. Polygala lewtonii Small This species is not restricted to scrub, and, in fact, is most commonly encoun- tered in turkey oak barrens and disturbed high pine. It is known from about 15 localities in Marion, Lake, Orange, Osceola, Polk, and Highlands Coun- ties. Recent records from Orange County (Christman 2199, 1987, Christman 2214, 1988) show that the report of its extirpation there (Wunderlin et al., 1980d) was premature. Lewton’s polygala is known to be protected only at Tiger Creek Nature Preserve, but the plant is difficult to find when not in bloom, and additional localities may exist. Schizachirium niveum (Swallen) Gould This distinctive scrub endemic grass has been found at only 31 sites in white sand scrubs that total about 3700 ha from Lake Wales to Venus in Highlands and Polk Counties. It has not been found in turkey oak or longleaf pine sandhill habitats, and appears to be one of the rarer of the Central Ridge scrub endemics. It is known to be protected only at Archbold Biological Sta- tion where it occurs within scrubs that total about 100 ha. Conc.usions— There are 40 species of vascular plants that are restricted to scrubs, natural turkey oak barrens, and scrub/high pine ecotones in central peninsular Florida. Seventeen of these species are restricted to the Lake Wales, Lake Henry, and Winter Haven ridges in Orange, Osceola, Polk, and Highlands Counties. These three Central Florida ridges encompass about 280,000 ha from Lake Apopka to Venus, a distance of about 60 km. Within these ridges, scrub habitats occur as disjunct and isolated “islands” within a matrix of high pinelands, pine flatwoods, prairie, wetlands, and developed lands. The Central Florida ridges are over 25 m above MSL, and have had a longer history of emergence above sea level than other areas of peninsular Florida. Because of this and because of the high degree of endemism among animals and plants, the scrubs on these ridges are referred to as ancient scrubs in order to distinguish them from coastal scrubs and scrubs that have devel- oped following human disturbance, which do not harbor the rare endemics discussed here. There are about 200 ancient scrubs, and most are smaller than 80 ha. None of the scrub plants endemic to the Central Ridges occur within all of No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 1p! the ancient scrubs, and no single scrub contains populations of each of the endemic plant species. Thus, only a system of multiple scrub preserves throughout the Central Ridges will prevent the extinction of Florida’s unique scrub biota. A private biological research station, two Nature Conservancy preserves, two small urban parks, and a proposed state park contain the only protected ancient scrubs. Three Central Ridge scrub endemics are protected nowhere, two have only one protected population, and 12 have five or fewer protected populations. Because of human activities, the ancient scrubs of Central Florida have declined in total area from an estimated 32,000 ha prior to settlement to about 11,000 ha today (Christman, 1988a). The conversion of scrubs to citrus groves on the southern Lake Wales Ridge has accelerated since the devastat- ing freezes in 1983 and 1985 (Layne, 1987). The development of residential subdivisions and tourism facilities will soon eliminate all remaining ancient scrubs not already protected. Many times during the course of field work the first author found himself conducting plant surview in “hard hat areas,” dodging bulldozers, and searching undeveloped lots within subdivisions. About a quarter of the scrubs surveyed between 1985 and 1987 no longer support native vegetation. Efforts to cultivate some of the scrub endemics have begun at Bok Tower Gardens (Wallace and McMahan, 1988). The Nature Conservancy and Florida’s Conservation and Recreational Lands Program are attempting to acquire ancient scrubs for protection. Both organizations are seriously under- funded. The discovery of an unrecognized species of scrub mint in 1987 (i.e., Dicerandra christmanii; see Huck et al., 1989), the rediscovery in 1986 of a scrub endemic believed to be extinct (i.e., Ziziphus celata), and the recent discovery of yet another scrub endemic (i.e., Crotalaria sp. nov.; Wunderlin, 1989) while this manuscript was in review, along with the numerous taxo- nomic, biogeographic, and ecological questions still largely uninvestigated should stimulate researchers to work fast. Scrub is considered prime residen- tial and citrus land and low agricultural tax rates encourage its destruction. Florida’s state-wide policy of maximum sustained economic development leaves little room for preservation of biodiversity. ACKNOWLEDGMENTS—Funding was provided by a grant from Florida’s Non-game Wildlife Program (GFC-84-101) to the first author. Publication costs were provided by the Department of Botany, University of Florida. We thank David Martin and Richard Wunderlin for their helpful comments on an earlier draft of this paper. LITERATURE CITED Austin, D. F. 1979. Florida Bonamia. Pp. 71-72. In: Warp, D. B., ed. Rare and Endangered Biota of Florida. Vol. 5, Plants. Univ. Presses of Florida, Gainesville, FL. 175 pp. Bissett, N. 1986. Haines City, FL. Pers. Comm. V2 FLORIDA SCIENTIST [Vol. 53 CurIsTMAN, S. P. 1988a. Endemism and Florida’s Interior Scrub. Final Report to Florida Game and Fresh Water Fish Commission, Tallahassee, Contract No. GFC-84-101. 247 pp. + maps, tables, appendices. . 1988b. Tiger Creek Preserve monitoring studies, initial element occurrences: rare vascular plants and vetebrate animals of the uplands of Tiger Creek Nature Preserve. Unpublished report to the Nature Conservancy, Winter Park, FL. Darben, D. 1987. Port Charlotte, FL. Pers. comm. Deyrup, M. 1987. Archbold Biological Station, Lake Placid, FL. Pers. comm. Goprrey, R. K. 1988. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and Alabama. The Univ. of Georgia Press, Athens, GA. 734 pp. Hansen, B. 1986. Univ. of South Florida, Tampa, FL. Pers. comm. Haron, D. 1986. Florida Natural Areas Inventory, Tallahassee, FL. Pers. comm. Huck, R. 1987. Systematics and Evolution of Dicerandra (Labiatae). J. Cramer, Berlin. 343 pp. , W. S. Jupp, W. M. Wuirten, J. D. SKEAN, Jr., R. P. WuNDERLIN, AND K. R. DELANEY. 1989. A new Dicerandra (Labiatae) from the Lake Wales Ridge of Florida, with a cladis- tic analysis and discussion of endemism. Syst. Bot. 14:197-213. Hurp, Mark, INc. 1973. Aerial photographs. Minneapolis, MN. Jounson, A. F. 1981. Scrub endemics of the Central Ridge, Florida. Unpublished Report pre- pared for the U.S. Fish and Wildlife Service, Jacksonville, FL. Jupp, W. S. 1980. Warea amplexifolia (Nutt.) Small. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, At- lanta, GA. 28 pp. AND D. W. HALL. 1984. A new species of Ziziphus (Rhamnaceae) from Florida. Rho- dora 86:381-387. KRAL, R. 1982. Some notes on Dicerandra (Lamiaceae). Sida 9:238-262. . 1983. A Report on some Rare, Threatened, or Endangered Forest-related Vascular Plants of the South. Tech. Publ. R8-TP 2, U.S.D.A. Forest Service, Atlanta, GA. 2 vols., 1305 pp. LakeELA, O. 1963. On the identity of Bumelia lacuum Small. Rhodora 65:280-282. Layneg, J. R. 1987. Archbold Biological Station, Lake Placid, FL. Pers. comm. Lirtte, E. L., Jr. 1979. Checklist of United States Trees. Agriculture Handbook No. 541. Forest Service, U.S. Dept. of Agriculture, Washington, D.C. 375 pp. Martin, D. 1987. U.S. Fish and Wildlife Svc., Jacksonville, FL. Pers. comm. Myers, R. L. 1985. Fire and the dynamic relationship between Florida sandhill and sand pine scrub vegetation. Bull. Torrey Bot. Club 112:241-252. AND S. E. BorertcuHer. 1987. Provisional fire management plan for a portion of Tiger Creek Preserve, Florida. Report to The Nature Conservancy, Winter Park, FL. Nauman, C. E. 1980. Warea carteri Small. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 24 pp. Peroni, P. A. AND W. G. ABRAHAMSON. 1985a. Vegetation loss on the southern Lake Wales Ridge. Palmetto 5:6-7. . 1985b. A rapid method for determining losses of native vegetation. Natural Areas J. 5:20-24. SHINNERS, L. H. 1962a. Synopsis of Conradina (Labiatae). Sida 1:84-88. . 1962b. Synopsis of Dicerandra (Labiatae). Sida 1:89-91. USDA. 1988. Draft Environmental Impact Statement: vegetation management in the Coastal Plain/Piedmont, Vol. 1, USDA Forest Service, Southern Region, Atlanta, GA. USFWS. no date. Endangered and Threatened Species of Region IV. Regional and state lists with periodic revisions. Atlanta, GA. . 1985. Endangered and Threatened Wildlife and Plants: Determination of Endan- gered Status for two Florida Mints. Fed. Reg. 50(212):45621-45624. . 1987a. Endangered and Threatened Wildlife and Plants: Determination of Endan- gered or Threatened Status for Seven Florida Scrub Plants. Fed. Reg. 52(13):2227-2234. . 1987b. Endangered and Threatened Wildlife and Plants: Endangered Status for Lu- pinus uaridorum (Scrun Lupine). Fed. Reg. 52(66)11172-11175. . 1987c. Endangered and Threatened Wildlife and Plants: Determination of Threat- ened Status for Bonamia grandiflora (Florida Bonamia). Fed. Reg. 52(211):42068-42071. . 1987d. Endangered and Threatened Wildlife and Plants: Endangered Status for Wa- rea amplexifolia (Wide-leaf Warea). Fed. Reg. 52(82):15501-15505. No. 1, 1990] CHRISTMAN AND JUDD—ENDEMIC SCRUB PLANTS 73 . 1988. Regional News, Region 4. Endangered Species Technical Bulletin 13(1):1-4. Wa..ace, S. R. AND L. R. McManan. 1988. A place in the sun for the plants. Garden, Jan./Feb. 1988:20-23. Warp, D. B., ed. 1979a. Rare and Endangered Biota of Florida. Vol. 5, Plants, Univ. Presses of Florida, Gainesville, FL. 175 pp. . 1979b. Scrub plum. Pp. 53, 54. In: Warp, D. B., ed., Rare and Endangered Biota of Florida. Vol. 5, Plants, Univ. Presses of Florida, Gainesville, FL. 175 pp. AND R. K. Goprrey. 1979. Pygmy fringe-tree. P. 20. In: Warp, D. B., ed. Rare and Endangered Biota of Florida. Vol. 5, Plants. Univ. Presses of Florida, Gainesville, FL. 175 pp. Wuarton, C. H. 1978. The Natural Environments of Georgia. Georgia Dept. Natur. Res., Atlanta, GA. 227 pp. WHETSTONE, D. 1983. Jacksonville State Univ., Jacksonville, AL. Pers. comm. Wuire, W. A. 1970. The geomorphology of the Florida peninsula. Geol. Bull. 51, Bureau of Geology, Florida Dept. Natural Resources, Tallahassee, FL. 164 pp. Winker, C. C. ann J. D. Howarp. 1977. Correlation of tectonically deformed shorelines on the southern Atlantic coastal plain. Geology 5:124-127. Worrorp, B. 1973. A biosystematic study of the genus Persea (Lauraceae) in the southeastern United States. Ph.D. dissert., Univ. of Tennessee, Knoxville, TN. 160 pp. WUuNDERLIN, R. P. 1982. Guide to the Vascular Plants of Central Florida. Univ. Presses of Florida, Gainesville, FL. 472 pp. . 1984. Dicerandra frutescens Shinners. Endangered and threatened plant status sur- veys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 29 pp. . 1989. Univ. of South Florida, Tampa, FL. Pers. comm. , B. E. HANSEN, AND D. W. HA tt. 1985. The vascular flora of central Florida: taxo- nomic and nomenclatural changes, additional taxa. Sida 11:232-244. , D. RICHARDSON, AND B. Hansen. 1980a. Calamintha ashei (Weatherby) Shinners. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Serv- ice, Office of Endangered Species, Atlanta, GA. 48 pp. . 1980b. Polygonella myriophylla (Small) Horton. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 54 pp. . 1980c. Chionanthus pygmaea Small. Endangered and threatened plant status sur- veys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 28 pp. . 1980d. Polygala lewtonii Small. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 42 Pp. . 1980e. Clitoria fragrans Small. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 31 . 1980f. Nolina brittoniana Nash. Endangered and threatened plant status surveys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 47 . 1981. Paronychia chartacea Fernald. Endangered and threatened plant status sur- veys, Region IV, U.S. Fish and Wildlife Service, Office of Endangered Species, Atlanta, GA. 39 pp. Florida Sci 53(1):52-73. 1990. Accepted: June 23, 1989. Engineering Sciences LANDFILLS—A THING OF THE PAST? RICHARD C. JOHNSON, SR. Camp Dresser & McKee Inc., 1305 U.S. Highway 19 South, Suite 400 Clearwater, Florida 34624 Asstract: Generally, ground water contamination is a reality associated with landfills con- structed prior to about 1980-1984. In some cases this contamination has reached public and private drinking water supplies. State laws adopted in the mid-1980s requiring ground water monitoring at all landfills have accelerated discovery of ground water contamination problems associated with landfills. In addition, federal investigations under Superfund have identified sources of contamination and in some cases provided funds for remedial action at these sites. Remedial action is also being conducted by the state of Florida, municipalities and the private sector. For the past 4-5 years, landfills have been constructed in a manner that greatly reduces the potential for ground water contamination through the use of liner systems, slurry walls, leachate collection and detection systems, and mandatory ground water monitoring. However, there is a trend in Florida to reduce our reliance on the use of landfills by increasing our use of recycling and energy recovery systems. Are landfills a thing of the past? This question has been asked repeatedly in recent years, although with different emphasis depending on the point of view. To many environmentalists, the question is more of a statement reflect- ing what they hope is an end to the practice of placing solid waste in or on sources of drinking water supply (aquifers). To many Florida residents living near existing or proposed landfills, the question is usually asked with incredu- lity that continually developing technology has not produced more appealing solutions to waste disposal. Finally, to many public and private officials re- sponsible for community and industrial waste disposal, the question reflects the increasing difficulty and dwindling options associated with landfill and other methods of waste control. From 1974 to 1988, the number of active landfills in the state has de- creased from 508 to 164, representing a 60 percent reduction in active land- fills (Reese, 1988). In addition, most of these inactive sites have never been properly “closed” and are continuing sources of potential drinking water con- tamination. Nine of these landfills are on the Environmental Protection Agency’s (EPA) national priority list (NPL) of the nation’s hazardous waste sites eligible for cleanup under Superfund (U.S. EPA, 1988). Landfills consti- tute 25 percent of the total number of Florida sites on the NPL. Approxi- mately 54 additional landfills are under consent order with the state to ad- dress potential contamination problems. The waste disposal issues are complicated by an influx of about 800 new citizens a day to the state of Florida. Most of the disposal crunch has occurred in the urban coastal areas, where the population is growing rapidly and land- fill capacity is disappearing fast. While about 500 acres of new landfill space is sited per year in the state, the new facilities lie mostly outside of the rapidly growing urban areas. Additional landfill development is desperately needed No. 1, 1990] JOHNSON—LANDFILLS TS in these areas, even though recent costs of development have soared to $400,000 an acre, depending on design characteristics. The cost increases are primarily a result of increased regulatory requirements for landfill design and permitting. Regulatory requirements have increased dramatically as a result of past ground water contamination by landfills. REGULATORY ContTrots—Numerous regulatory controls are available to state environmental control officials to regulate landfill operation and clo- sure. However, there are four laws that typically govern in landfill environ- mental review and permitting. First and foremost, proposed landfills must comply with the Florida Resource Recovery and Management Act (Florida Statutes, 1988). FDER’ is the responsible agency for administering this act, which authorizes regulation of the construction and operation of solid waste handling, volume reduction, and/or energy recovery facilities. Specifically, the act mandates that a permit is required to construct and operate a solid waste or resource recovery and management facility. The details of the per- mitting requirements are specified in the FDER Rules and Regulations. Additional state acts of regulation that typically address aspects of landfill operation include the Florida State Lands Act, the Florida Pollution Control Act, and the Florida Water Resources Act (CDM, 1988). The State Lands Act governs dredging or filling in “waters of the state,’ which includes navigable waterways, estuaries, bays, bayous, rivers, streams and natural tributaries. Dredge and/or fill permits must be obtained from the FDER if the named actions are involved in landfill development. The Air and Water Pollution Control Act is administered both by the FDER and the Florida Water Man- agement Districts and governs the discharge of stormwater runoff from im- pervious surfaces. Stormwater discharge permits are required to ensure stormwater runoff is collected in adequately sized detention facilities. Fi- nally, the Water Resources Act requires issuance of a Consumptive Use Permit by the Water Management Districts for any use of water which reduces the supply from which it is withdrawn or diverted. Many other laws could potentially apply to landfill development and op- eration (CDM, 1988). As many as 13 federal laws and/or agencies could po- tentially be involved, including the National Environmental Policy Act, Clean Air Act, Clean Water Act, Corps of Engineers, the Federal Emergency Management Authority, Housing and Urban Development, and the Occupa- tional Safety and Health Act. An additional 12 state laws could potentially be involved, including the National Historic Preservation Act administered by the National Park Service, the Beach and Shore Preservation Act adminis- tered by the Department of Natural Resources, the Florida Coastal Manage- ment Act administered jointly by FDER and DNR, and others. At least four regional/local acts of regulation may also require compliance and are usually administered by local planning departments and regional planning councils. In some circumstances, even local code and ordinance enforcement permits must be obtained for landfill construction and operation. The effect of regu- 1Florida Department of Environmental Regulation. 76 FLORIDA SCIENTIST [Vol. 53 lation by these numerous acts and regulatory agencies is to make landfill permitting a long and uncertain activity. Permits can take anywhere from three to twelve months to obtain, but typically at least six to eight months are required. Additional solid waste regulations pertaining to landfill development and operation are a certainty. At the time of this paper (1988), three solid waste management planning bills were before the State Legislature. In addition, the federal government is beginning to get heavily involved in state solid waste management issues. In a recent draft report to Congress, EPA con- cluded that existing federal criteria for state solid waste regulation are inade- quate for municipal solid waste landfills because they lack essential require- ments for ground water monitoring, corrective action, and post-closure care. The draft report says that gaps exist in information on solid waste facilities and suggests that legislation is required which would expand EPA’s role over regulating the facilities to close the gaps. EPA’s office of solid waste an- nounced the formation of a municipal solid waste task force to develop a national strategy on management of municipal solid waste. The task force is scheduled to issue a final strategy document to address: 1) The role of source reduction, recycling, waste treatment and separa- tion, and landfills; 2) The health and environmental impacts; 3) The role of government procurement of recycled materials; 4) The need for federal in- centives for source reduction and recycling markets; 5) The need and role of product and raw materials substitution to increase waste recyclability or re- duce the toxicity or volume of waste. Finally, the issue of municipal incinerator ash disposal has caused consid- erable federal discussion which will likely result in state development of new EPA regulations for ash disposal. CriTIcAL Issues—Existing and proposed environmental regulations are focused on five critical issues surrounding landfill development and opera- tion. Two of these issues are associated with the problem of ground water contamination: the use of liners and leachate control systems. The remaining three critical issues are co-disposal landfills, landfill closure, and financing. Each of these issues is briefly discussed in this section. Proposed landfills in the state must demonstrate that they will not con- taminate ground water aquifers underlying the landfill area. All landfills in the last four to five years include liner systems with a minimum of 30 millime- ter (mil) thick liner, usually some form of polyethylene material. Most land- fills, however, have liners to 46 mil thicknesses and some to 80 mil thicknesses in an effort to ensure ground water protection and allay community fears. Natural material such as clay can be used as a liner in which case liners can attain thicknesses of three to four feet. The great increase in cost in develop- ing landfills is largely due to the cost of development and installation of liner systems. However, the liner is the primary means of preventing contaminated water or leachate from entering an aquifer system (Giroud, 1985). In addi- tion, landfill design is moving toward composite liners and multiple liner systems. No. 1, 1990] JOHNSON—LANDFILLS Th The second critical issue is the application of adequate leachate control measures to complement successful liner installation. Leachate is formed when water (usually rainwater) comes into contact with materials disposed of in a landfill. This typically happens when rainwater infiltrates the landfill and moves downward through the landfill materials. The liner is designed to keep this leachate from reaching the ground water system, but the leachate must be collected and handled properly for the liner system to work success- fully. What to do with the leachate once collected is one of the big issues currently being considered by regulatory agencies. Many active landfills take the collected leachate and recirculate it through the landfill. This procedure has several benefits. It tends to help decompose and stabilize the landfill quickly (through anaerobic treatment), while promoting leachate removal through evapotranspiration. However, both EPA and the state are moving toward banning recirculation since it is improperly done in many landfills and results in leachate ponding and breakout of leachate through the sides of the landfill. Other methods for handling collected leachate are much more expensive. One alternative disposal method is to put the leachate in a wastewater treat- ment plant (following a negative test for hazardous constituents). This is generally difficult due to the remote location of landfills and the associated non-availability of wastewater treatment plants. The solution to a remote location is to haul leachate away in tankers. However, this procedure now must be preceded by leachate pretreatment typically required to lower sus- pended solids, adjust pH, and remove heavy metals. Pretreatment systems and the use of tankers greatly increase landfill operating costs. On-site treat- ment and discharge of leachate is a last resort since it is very expensive. How- ever, on-site treatment of leachate and spray discharge is currently being done in Alachua County. Orange County is mixing leachate with the stormwater for dilution for eventual disposal in wetlands treatment systems. The disposal methods identified above have economic and technical ad- vantages and disadvantages which are all improved through leachate mini- mization (Tchobanoglous, et al., 1977). The best way to minimize leachate development is to operate a good landfill, which essentially has the effect of minimizing the amount of landfill material that can come into contact with rainwater. A good operation will minimize the working face, or the area of current landfill disposal. This can be accomplished by good compaction, en- suring that the working face is covered daily, and by rapid capping and seal- ing of landfill cells as they are completed. Overall, landfill operation should be designed to promote runoff and to minimize contact and contact times between landfill material and water. Co-disposal generally refers to the practice of mixing two distinct sources of material in a single landfill. The critical co-disposal issue of present con- cern is mixing standard garbage with incinerator ash, usually from municipal incinerators. In general, the trend is to require separate landfills or separate areas for the disposal of different materials. The concern is that ash or other 78 FLORIDA SCIENTIST [Vol. 53 materials may react with liquids and other household material and municipal solid waste to create a greater potential for leaching. There are currently draft bills before Congress which would regulate incinerator ash as a “special waste,” eventually requiring disposal in multiple-lined monofills (landfills ac- cepting only ash waste) with monitoring wells and leachate controls. The remaining critical issues of landfill closure and financing must be discussed concurrently. The problem that must be addressed is that proper and successful landfill closure is expensive. For example, a smaller county in north central Florida discovered that closure of the county landfill in con- formance with state rules exceeded the annual budget for the county. The lack of available funds is a serious obstacle to successful landfill closure in many areas of the state. One way this problem is being overcome is that the FDER currently requires communities to set up closure funds for new landfill sites. Part of the new permit application is a statement certifying that the landfill owner will pay to close, and requiring estimates of eventual closure cost. This requirement is designed to eliminate the excuse of ignorance and lack of money once closure is required. These new policies are working to some extent, but it is difficult to force public entities to set up closure funds at the start of landfill operations. Ultimately, cost of closure will have to be passed on to landfill users in the form of user fees. Future TRENpS—Future trends for landfills will focus on stricter regula- tions and stronger siting requirements, both required by new laws and by public concern (U.S. EPA, 1988). Examples of stricter regulations include a probable ban on co-disposal landfills. Another example is probable require- ment for liners for Class 3 (trash sites) landfill sites which currently do not require lining. Stronger siting requirements are evidenced by an increasing decision by landfill developers for composite (clay and synthetic) or double liner systems even though single liner systems are the minimum required by law. This is an additional example of how public concern and demand are affecting landfill development. As a result of these stricter controls and the cost of landfill closure dis- cussed above, the trend will continue toward fewer but bigger landfills. Smaller communities that cannot afford landfills individually will band to- gether to pool resources and create regional systems. In the northern part of the state, for instance, 13 counties banded together last year to conduct a study of solid waste disposal in the region. These counties will now divide into three groups and form regional disposal compacts, each with its own landfill. A regional approach is also being considered among counties in the state’s panhandle. The concern over potential ground water contamination and the cost of landfill development and operation is resulting in increasing attention on waste minimization. There is a strong trend in the state to minimize what goes into landfills. Waste minimization will primarily take two forms, entry controls and recycling. Entry controls will include closer inspection of indi- vidual landfill users, including more frequent inspection of dump trucks and No. 1, 1990] JOHNSON— LANDFILLS 10 stiffer penalties or banning from the landfill haulers who try to dispose of unapproved material. In addition, there will be a concentrated effort to elim- inate disposal of infectious waste (hospitals are supposed to have pathological incinerators, but not enough currently exist). The second form of waste minimization generally includes multiple types of recycling to minimize waste disposal in landfills. All solid waste bills before the Florida House and Senate at the time of this paper included recycling. The House bill specifies that 25-35% recycling must be achieved by the coun- ties. Recycling is generally approached at three levels with generally increas- ing difficulty and effectiveness. The first level requires the average citizen to haul garbage to recycling centers where newspaper, glass, metals, plastics, etc. can be separated from the basic garbage that proceeds on to a landfill. A second, more difficult level to enforce, is separation of garbage at the home with different pickups scheduled for different types of materials. The third, most expensive level, which essentially requires no commitment from the average citizen, is that all garbage collected is taken to separate collection facilities for separation. Several pilot programs are currently in place in Flor- ida including Palm Beach County, the City of Tallahassee and the City of Gainesville. Most programs are achieving no more than 10% effective recy- cling. The 25-35% specified in the House version of the bills will likely re- quire strong enforcement actions. Other types of waste minimization can add to the overall effectiveness of the total waste utilization program. Examples of these forms include the FDER amnesty days when citizens can collect household chemicals or other hazardous waste and take them to collection areas for proper disposal, and home composting of yard waste to keep it out of municipal landfills. Energy recovery in the form of resource recovery plants has been and will continue to be an important trend in solid waste disposal. This represents the highest degree of control in terms of potential ground water contamination, concern over co-disposal, etc. A number of resource recovery plants have been constructed in the past several years, although there has been a recent slowdown in construction due to changes in financial attractiveness (i.e., decrease in oil prices). Prior problems of resource recovery incinerator emis- sion and incinerator ash disposal problems are currently being resolved. The future trend is toward additional resource recovery or energy recovery facilities. ConcLusions— Overall, the state will have fewer, bigger, and better op- erated landfills in the future. Resource recovery will continue to be popular with the public and will be coupled with waste minimization and mandatory recycling to complete a comprehensive solid waste management picture in the state of Florida. The expense associated with these bigger and better facilities will cause regionalization of facilities as well as pooling of resources. Landfills are definitely not a thing of the past in Florida, but landfill facilities will be basically non-polluting and more sophisticated than ever before. 80 FLORIDA SCIENTIST [Vol. 53 LITERATURE CITED Camp Dresser & McKee, INc. 1988. Permitting and Regulatory Review. Technical Memoran- dum #7. Unpublished Report for Lee County, Florida. Tampa, Florida. FLoripa STaTuTEs. 1988. Chapter 88-130. Florida Resource Recovery Management Act. Giroup, J. P. 1985. Geotextiles and Geomembranes—Definitions, Properties and Design. GeoServices Inc. Consulting Engineers. Boca Raton, Florida. Reesr, J. A. 1988. Bureau of Waste Planning and Regulation. Florida Department of Waste Planning and Regulation. Florida Department of Environmental Regulation. Personal Com- munication. TCHOBANOGLOUS, G., H. THEISEN AND R. EiassENn. 1977. Solid Wastes— Engineering Principles and Management Issues. McGraw-Hill, New York. U.S. ENVIRONMENTAL PROTECTION AGENCY. 1988. Solid Waste Disposal: Faulty Criteria; EPA proposed rule. Federal Register. 53:#168. 33314. Florida Sci. 53(1):74-80. 1990. Accepted: May 12, 1989. INSTRUCTIONS TO AUTHORS Individuals who publish in the Florida Scientist must be active members in the Florida Academy of Sciences. 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Ad- dress all orders to: The Florida Academy of Sciences, Inc. c/o The Orlando Science Center 810 East Rollins Street Orlando, Florida 32803 Phone: (305) 896-7151 ISSN: 0098-4590 Florida Scientist Volume 53 Spring, 1990 ooo T Number 2 Q Pee aN { j ; } ‘Nf Oe J ‘ | CONTENTS \ N26 1999) X y, FG F63nmental Influences on the Distribution of BRP ics 7 NH Rats (Oryzomys palustris) in Coastal Marshes ........ a ———— James L. Wolfe 81 Recent Changes in the Distribution of Calulerpa prolifera mine tnesan River Lagoon, Florida ........ 0.0.00. .60. 2000.0 6: Conrad White and Joel W. Snodgrass 85 Vascular Plants of Fakahatchee Strand State Preserve ............. Daniel F. Austin, Julie L. Jones, and Bradley C. Bennett 89 rene Absorption in Florida Springs .. 2.0.66. 5 6c eee a es Carlos M. Duarte and Daniel E. Canfield, Jr. 118 Food Selection by Early Life Stages of Blue Tilapia, Oreochromis aureus, in Lake George, Florida: Overlap marmoymipatric Shad Warvae ee ee icles bv wee Alexander V. Zale and Richard W. Gregory 123 Development and Application of a Soil-Air Radon Analysis SSS Ti Tae EM GaN RT SRO) oe Ur a Robert S. Braman and Robert L. Sutton 130 A Note on the Fire Responses of Species in Rosemary Scrubs on the pemeactm ake Wales TIGDGE Fe ee ee eye al kn GCs Sale Has vv es Ann F. Johnson and Warren G. Abrahamson 138 LULZ SG OS eee A ee eo Richard P. Wunderlin 144 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1990 Editor: Dr. DEAN F. MarTIN Co-Editor: Mrs. BARBARA B. MARTIN Institute for Environmental Studies Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoripA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. 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Martin, Editor Volume 53 BarsBara B. MArTINs.Co-Editor ZG Spring 1990 Biological Sciences x ENVIRONMENTAL INFLUENCES ON THE Ue DISTRIBUTION OF RICE RATS (ORYZOMYS PALUSTRIS) IN COASTAL MARSHES JAMES L. WOLFE Department of Biological Sciences, Emporia State University, Emporia, KS 66801 Asstract: This study was designed to describe the distribution patterns of rice rats in coastal marshes and identify environmental variables that influence these patterns. Rice rat abundance increases from the forest-marsh ecotone toward the shoreline. Tidal flooding also increases along this gradient, but appears not to present a problem. Plant community composition, height, or cover density are not significantly correlated with distribution. These findings lead to hypotheses on the relation of predator and food resource densities to rice-rat distribution. CoasTAL marshes present unique environmental problems to terrestrial animals inhabiting them. Daily tidal fluctuations, storm surges that can in- undate entire marshes, and varying degrees of salinity are major problems to be faced. Generally these factors increase in intensity from the marsh-forest ecotone toward the shoreline. Cover and refuge sites for small mammals are most abundant in the ecotone and decrease in the open marsh. Maintaining a homesite in the ecotone and timing foraging in the marsh to avoid flooding and other hazards would seem to be a prudent course for such species to take. Most small and medium-sized mammals that forage in marshes seem to fol- low this strategy. Exceptions are the truly semi-aquatic forms such as the muskrat, Ondatra zibethicus and coypu, Myocastor coypus (see Perry, 1982 and Willner, 1982). The marsh rice rat, Oryzomys palustris, also lives in open marshlands (Wolfe, 1982a). Although it has few apparent physical adaptations, its behav- ior is that of a semi-aquatic form (Esher, et al., 1978). Individuals can survive large storm surges and return to their former home ranges (Wolfe, 1982b). They appropriate nests of marsh-nesting birds (Kale, 1965; Post, 1981; NeSmith and Cox, 1985) or construct their own above the high-water line. Otherwise little is known of the adaptations that afford this species almost exclusive access to the small mammal niche in exposed coastal marshlands within its geographic range. Documenting details of the distribution of indi- viduals and environmental influences on this distribution is needed to under- stand the factors that facilitate survival and resource use in this habitat. 82 FLORIDA SCIENTIST [Vol. 53 MetTHops— Data collected during a four-year population study on the Gulf coast of Missis- sippi (Wolfe, 1985) suggested that individuals were neither randomly nor uniformly distributed on trapping grids. Apparent trends on the grids were for rice rat captures to increase with dis- tance from the marsh-forest ecotone. Abundance did not seem to be influenced by water depth or plant community. As a result, the present study was designed to test the hypotheses that in large tracts of marsh bounded on one edge by a forest ecotone and on the other edge by an open shoreline, (1) rice rats increase in number from the ecotone toward the shoreline, (2) cover and characteristics of the plant community are not important determinants of distribution, and (3) water depth during normal tidal fluctuations is not limiting to distribution. The primary study area was on the northern shore of St. Louis Bay, Harrison County, Missis- sippi. A detailed description of the 400 ha. area is provided in Wolfe (1985). Twelve sampling stations were distributed over the entire marsh by a stratified random de- sign. Four stations were placed within 5 m of the ecotone, four within 5 m of the shoreline, and four in the marsh interior. Within each zone, sites were randomly selected. A 1-m? floating platform was placed at each location and tied to a concrete block. During sampling, an identical combination of Sherman live-traps and Museum Special snap traps, usually two of each, was used on each platform. Captured rats were removed from the study area. Platforms were trapped simultaneously a total of 21 days between January 1981 and October 1984. The months of Janu- ary, April, June, July, September, and October were included in the sampling effort. Environmental variables measured at each station included water depth at high tide, distance to ecotone, distance to shoreline, cover density, plant species richness, and height of vegetation. Sites were also classified by dominant plant species. The marsh was characterized by distinct zones, each dominated by a single species (Eleuterius and Eleuterius, 1979). Juncus romericanus, Scirpus olneyi, Spartina alterniflora and “mixed” zones, in order of relative abundance, were sampled by our trapping stations. The mixed zone was one of high plant diversity near the ecotone. Cladium jamiacensis, Panicum virgatum, and Spartina patens were interspersed with the shrubs Baccharis hamilifolia, Myrica cerifera, and Ilex vomitoria in this region. High tide was measured at three occasions during the study and the median value used as an index of relative depth. Plant species richness was measured by counting the number of species along a 5-m tran- sect adjacent to the west side of each station. Cover density was estimated at 100 cm above ground level on each transect, using a profile board (Nudds, 1977). Height above ground at which cover density decreased to 10% was recorded as a measure of vegetation height. In order to evaluate the results obtained at the Mississippi study site, two additional sites were selected for sampling. One was in southern Hernando County on the west coast of Florida and the other in Volusia County on the east coast of Florida. These sites were selected to have well- defined forest ecotones, and shorelines on open water (Gulf of Mexico on the west coast and the Halifax River estuary on the east coast) with at least 500 m of open marsh separating the two. Traps were placed along the ecotone, within 5 m of the shoreline, and approximately half way between. Sampling of the three areas was always done simultaneously with identical arrays of live and snap traps. TABLE 1. Rice rat captures in relation to location in coastal marshes. Zone Mississipi Gulf Florida Gulf Florida Atlantic Ecotone 18 4 0 Central Marsh 53 25 29 Shoreline marsh 89 35 29 Kruskal-Wallis H 6.8 9.6 10.6 P <0.04 <0.01 <0.01 RESULTS AND Discussilon— Captures in relation to location in the marshes are shown in Table 1. A significant site effect is evident on all three study areas (Kruskal-Wallis one-way analysis of variance, Conover, 1980). Table 2 shows a correlation matrix (Spearman rank correlation test, Conover, 1980) illustrating the relationship between environmental variables measured and rice rats captured. The strongest correlation is between captures and distance from the ecotone, followed closely by a strong negative correlation with the No. 2, 1990] WOLFE— DISTRIBUTION OF RICE RATS 83 distance from the shoreline. These relationships are not precisely inverse be- cause the shoreline, and to a lesser extent the ecotone, were irregular in shape. There is also a significant positive correlation between captures and water depth at high tide, because depth increases with distance from the ecotone. None of the variables related to measurements of the cover and vegetation are significantly correlated with captures. Several predictable cor- relations exist between the environmental variables (Table 2). TaBLe 2. Correlation matrix (Spearman’s rho) of environmental variables and rice rat cap- tures. Ricerat Distance Distanceto Water Plant species Percent Vegetation Variable captures toecotone shoreline depth richness cover height Distance to ecotone OLS Distance to shoreline -0.78** -0.64* Water depth 0.68 * OGns -0.41 Plant species richness 0.44 0.33 -0.36 -0.02 Percent cover -0.40 -0.39 -0.01 -0.58* -0.06 Vegetation height 0.13 0.13 -0.43 -0.28 0.44 44 ooh). 0)1 *P<0.05 Vegetation zones had no significant effect on rice rat abundance (Kruskal- Wallis H=4.1, P>0.05, Conover, 1980). These were not included in the cor- relation matrix because they were classified by type, and there was no basis for assigning them a numerical value. I conclude that rice rat abundance generally increases from the forest- marsh ecotone toward the shoreline in coastal marshes. Water depth, which generally increases with distance from the ecotone, appears not to present a problem. Increased numbers along the shoreline may be related to the fact that most shoreline areas sampled had a slightly higher elevation than the marsh interior. Mats of vegetation and other debris that are deposited along the shoreline provide refuge. During the course of the study, I found more than a dozen nests in these deposits. This distribution pattern could be influenced by a number of factors, all difficult to test. Near the ecotone, rice rats co-exist with several other species of small mammals (Wolfe, 1985), which are potential competitors. Fewer terrestrial (and possibly avian, but see Jemison and Chabreck, 1962) preda- tors venture far into the marsh than patrol the ecotonal areas. Easier to con- firm by additional study is the possibility that preferred food items have a similar distribution pattern to that of rice rats. ACKNOWLEDGMENTS— Financial support for the study was provided by DuPont’s DeLisle, Mississippi Pigment Plant, Mississippi State University, and the Archbold Biological Station. Ro- bert J. Esher and Dwight Bradshaw assisted with the field work. I thank Ronald L. Mumme for his comments on the manuscript, and Mr. and Mrs. Willis Butts for granting access to their property near Aripeka, Florida. LITERATURE CITED Conover, W. J. 1980. Practical Nonparametric Statistics. John Wiley & Sons, New York, 462 pp. 84 FLORIDA SCIENTIST [Vol. 53 ELeuTerius, L. N. AND C. K. ELeutertus. 1979. Tide levels and salt marsh zonation. Bull. Ma- rine Sci. 29:394-400. EsHer, R. J., J. L. Woure ano J. N. Layne. 1978. Swimming behavior of rice rats and cotton rats. J. Mamm. 59:551-558. Jemison, E. S. anp R. H. Cuasreck. 1962. Winter barn owl foods in a Louisiana coastal marsh. Wilson Bull. 74:95-96. Kate, H. W. 1965. Ecology and bioenergetics of the long-billed marsh wren Telmatodytes palus- tris griseus (Brewster) in Georgia salt marshes. Publ. Nuttall Ornith. Club 5:1-142. NeSmitu, C. C. anv J. Cox. 1985. Red-winged blackbird nest usurpation by rice rats in Florida and Mexico. Florida Field Nat. 13:35-36. Nupps, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildl. Soc. Bull. a1 1S-117, Perry, H. R., Jr. 1982. Muskrats, Pp. 282-325. In: CHAPMAN, J. A. AND G.A. FELDHAMER (eds.), Wild Mammals of North America. Johns Hopkins Univ. Press, Baltimore, 1147 pp. Post, W. 1981. The influence of rice rats Oryzomys palustris on the habitat use of the seaside sparrow Ammospiza maritima. Behav. Ecol. Sociobiol. 9:35-40. WILLNER, G. R. 1982. Nutria. Pp. 1059-1076. In: CHAPMAN, J. A. AND G. A. FELDHAMER, (eds.). Wild Mammals of North America. Johns Hopkins Univ. Press, Baltimore, 1147 pp. Wo tre, J. L. 1982a. Oryzomys palustris. Mamm. Species 176:1-5. . 1982b. Storm effects on rice rats inhabiting coastal marshes. Gulf Res. Rep. 7:169- 170: . 1985. Population ecology of the rice rat (Oryzomys palustris) in a coastal marsh. J. Zool. (Lond.) 206:235-244. Florida Sci. 53(2):81-84. 1990. Accepted: August 22, 1989. RECENT CHANGES IN THE DISTRIBUTION OF CAULERPA PROLIFERA IN THE INDIAN RIVER LAGOON, FLORIDA— Conrad White and Joel W. Snodgrass, Brevard County Office of Natural Resources Management, 2575 North Courtenay Parkway, Merritt Island, FL 32952. Asstract: The changes in distribution of the macroalga Caulerpa prolifera was determined between 1986 and 1989 in the northern and central portion of the Indian River lagoon, Florida. Aerial photography coupled with groundtruthing was used to map submerged aquatic vegetation. DurING a recent macrophyte mapping project in the Indian River lagoon system seven seagrass species and a number of attached macroalgae were identified (Virnstein and Cairns, 1986; White, 1986). The dominant sub- merged aquatic vegetation in large areas of the northern third of the lagoon was Caulerpa prolifera (Forsskal) Lamouroux. In the Banana River, White (1986) stated C. prolifera was the principal benthic vegetation. Previous mapping projects conducted in the lagoon during the mid 1970’s did not report C. prolifera (Thompson, 1976; Down, 1978, 1983). Jensen and Clark (1983), however, mention that C. prolifera and C. ashmeadii Harvey were found in the northern Indian River near Titusville. During the winter of 1986-87 C. prolifera disappeared from large areas of the northern Indian River. By the end of June 1987 C. prolifera had disap- peared from the Indian River north of Cocoa. The purpose of this paper is to report the changes in C. prolifera distribution over a relatively short period. MATERIALS AND METHops— Aerial photography was coupled with intensive groundtruthing to produce 1:24,000 scale maps of the submerged aquatic vegetation within the Indian River lagoon during 1986. Groundtruthing was accomplished by towing a diver behind a small boat at slow speed or by wading. Significant changes in species or densities were relayed to an observer on the boat. Density estimates (% coverage) were made utilizing a method used for terrestrial vegeta- tion crown densities (Daubeumire, 1968). Latitudinal transects (no aerial photography) and spot checks were used to determine the distribution and densities of C. prolifera from 1987 to 1989. ResuLts—In 1986 C. prolifera had a sparse (<5%) distribution in the Indian River from Turnbull Creek to State Road 402 (Fig. 1). The remainder of the Indian River north of Pineda Causeway displayed patchy distribution (<10%) in water depths greater than 1.0m and dense (10 to 40%) stands in water depths of 0.5 to 1.0m. Occasional large areas of 40 to 70% or 70 to 100% densities were observed. Densities in the Banana River and Newfound Harbor varied from 40 to 70% coverage over much of the bottom with the exception of the channel and nearshore area. The alga was not found south of Pineda Causeway during 1986 and 1987. By June of 1987 the majority of the alga had disappeared north of State Road 528 in the Indian River. During 1988 and 1989 spot checks showed further losses of C. prolifera had occurred south of SR 528 in the Indian River. Drastic reductions in C. prolifera coverage were observed in the Ba- nana River during late 1987 and 1988. In June 1988 a small patch (< 1m diameter) of the alga was found in the Turnbull area. Other small patches were found north of State Road 528 in the Indian River during January 1989 (Fig. 2). Discusston—The reasons for the loss of C. prolifera from the Indian 86 FLORIDA SCIENTIST [Vol. 53 Haulove Brevard County 4 mi. = Titusvilleg 8 Port Canaveral [FJ cauterpa. prolifera 1986 Caulerpa prolitera 1987 Fic. 1. Distribution of Caulerpa prolifera during 1986 and 1987. No. 2, 1990] WHITE AND SNODGRASS— DISTRIBUTION OF CAULERPA PROLIFERA ¢ Haulove Brevard County 4mi. Titusville @ Merritt island O O O Port Canaveral O O OO — ws O @ Caulerpa prolitera O Found O Caulerpa prolifera Not Found Fic. 2. Location of spot checks for Caulerpa prolifera in January, 1989. 88 FLORIDA SCIENTIST [Vol. 53 River lagoon have not been pinpointed. An ascoglossan, Elysia n.sp. (Jensen and Clark, 1983), was observed grazing on C. prolifera in large numbers (>100/m2) during the period of alga loss. A migration of Elysia southward from the Turnbull creek area mimicked the loss of C. prolifera. Although speculative at this point in time, Elysia may have accounted for the rapid disappearance of C. prolifera from both the Indian and Banana Rivers. It may be just as likely that changes in some physical factor not monitored during this period, coupled with overgrazing by Elysia led to the loss of C. prolifera. C. prolifera has, until recently, received only casual attention as a compo- nent of the lagoon, which makes assessing the effects of algal loss difficult. Traits normally attributed to seagrass areas- stabilization of the substrate, reduced turbidity, nutrient uptake, and biological habitat—may also be at- tributed to the blade and rhizoid structures of C. prolifera. Preliminary evi- dence indicates areas vegetated by C. prolifera have excellent macroinverte- brate productivity, although fish abundance in the alga beds was less than nearby Halodule wrightii beds (White and Snodgrass, 1988). The loss of one of the major habitats from a large estuarine-type system offers an excellent opportunity to study the structural and functional components of the Indian River lagoon system. ACKNOWLEDGMENTS— This project was financed in part by grants from the Florida Depart- ment of Environmental Regulation, Office of Coastal Zone Management under provisions of Contracts CM-121 and CM-177. The authors would like to thank Larry Bauer, Terry Peterman and Steve Cottrell for their help in the field during 1986 and 1987. LITERATURE CITED Down, C. 1978. Vegetation and other parameters in the Brevard County bar-built estuaries. Project Report 06-73. Brevard County Health Department, Environmental Engineering. . 1983. Use of aerial imagery in determining submerged features in three east-coast Florida lagoons. Florida Scientist 46(3/4) :355-362. DavuBEuMIRE, R. 1968. Plant communities, a textbook of plant synecology. Harper and Row, New York, NY. JENSEN, K. AND K. B. Crark. 1983. Annotated checklist of Florida ascoglossan Opisthobranchia. The Nautilus. 97(1):1-13. THompson, M. J. 1976. Photomapping and species composition of seagrass beds in Florida’s Indian River estuary. Harbor Branch Foundation Technical Report No. 10, 34 pp. VIRNSTEIN, R. W. anp K. D. Cairns. 1986. Seagrass maps of the Indian River Lagoon. Final report to Florida Department of Environmental Regulations, Office of Coastal Zone Management. | Wulite, C. 1986. Seagrass of the Indian and Banana Rivers. Final report to Florida Department of Environmental Regulations, Office of Coastal Zone Management. AND J. SNopcrass. 1988. Caulerpa prolifera versus seagrasses in the Indian River lagoon: A comparison of relative habitat values. Preliminary report to the Florida Department of Environmental Regulations, Office of Coastal Zone Management. Talla- hassee, FL. Florida Sci. 53(2):85-88. 1990. Accepted: May 5, 1989. Biological Sciences VASCULAR PLANTS OF FAKAHATCHEE STRAND STATE PRESERVE DanlieL F. Austin, JULIE L. JONES”, AND BRADLEY C. BENNETT”? () Department of Biological Sciences, Florida Atlantic University, Boca Raton, FL 33431; ()Florida Game and Freshwater Fish Commission, Immokalee, FL 33943; (3)New York Botanical Garden, Bronx, N.Y. 10458 Asstract: Study of the plants in the Fakahatchee Strand State Preserve during the period from 1978 to 1987 has resulted in a list of 477 species in 119 families. This species richness is unusual because it includes, within a matrix of temperate plant communities, a variety of tropi- cal species found nowhere else in the coterminous United States. Plant communities are discussed and mapped, and a list of species is given. FAKAHATCHEE Strand State Preserve is located in southwestern Florida in Collier County, and extends south from Alligator Alley (St. Rd. 84) to the coast (Fig. 1). The Preserve is about seven miles wide and about eighteen miles long, or about 126 square miles in total (32,640 hectares or 80,640 acres). The Fakahatchee Strand State Preserve has never been completely surveyed, only the Township lines having been established. In large part due to its wildness, the tract remains a home to Florida’s endangered panthers and minks, the only remaining healthy stand of the royal palms (Roystonea elata) in the state, and a large number of plant species. Through the almost two decades that the first author has been studying the Fakahatchee Strand, it has remained a fresh source of botanical surprises. Consideration of its recent history may account for part of the unusual flora, but surely older events have also been involved. The Seminole Indians gave it the current name (Fakahatchee = muddy creek), a subdivision of the larger region they called either Okholwake (bad water) or Archenaho-taphe (big cypress) (Ives, 1856; Simpson, 1956). Big Cypress subsequently became the English name for the entire southwestern region which includes the Fakahat- chee. Abundant Indian mounds scattered in and around the Strand are evi- dence that the Glades Indians also made use of the habitat. It will never be possible to determine all of the plants people introduced into the Strand. Even for the plants that were not introduced, known human usage and crea- tion of high land through mound building probably account for part of the older vegetation patterns. Although people have added a few species to the flora in the past 500 years or more, non-human events resulted in the estab- lishment of the majority of the species. When this study began in 1978, comparatively little was known about the Preserve region (Beard et al., 1948; Finn, 1966; Klein, et al., 1970; McElroy & Alvarez, 1975; Prestridge, 1947; Simpson, 1902; Small, 1937). Our original goal was to provide more data for resource management planning and future development of the Preserve. That goal has been partly achieved by us and 90 FLORIDA SCIENTIST [Vol. 53 ii Yj BONITA SPRINGS STATE ROAD 29 STATE ROAD 84 a Zz < sw Be oO w ” ew ale Ee = 1.0 if each species uses less-common resources more intensively than others and the preferences of the two species tend to coincide (Hurlbert, 1978). Dietary similarity of tide- water silversides and independent blue tilapia collected 12 May was moder- ate, as Bosmina and immature copepods were the two most important items in the diets of both (Table 2); however, the high relative abundance of imma- ture copepods in the lake decreased the intensity of overlap. Similarity of diets of shad and independent blue tilapia was relatively high as both species coincided in the consumption of a variety of less-abundant items. A high degree of similarity for brooded and independent blue tilapia resulted en- tirely from the intense use of Bosmina by both. Overlap in the diet of brooded blue tilapia with those of shad and tidewater silversides was negligible. Die- tary similarity of shad and tidewater silversides was moderate, as overlap was notable only for the relatively abundant immature copepods. TABLE 2. Diet overlap matrix for fishes captured 12 May 1983, Lake George, Florida. Independent Tidewater blue tilapia silversides Shad Tidewater silversides 1.733 Shad 2.596 1.679 Brooded blue tilapia 2.592 0.492 0.282 FLORIDA SCIENTIST [Vol. 53 126 = 0 5 0 = 0 = 0 10> spoor.nsQ (000>) —- 8% (l00';0>) - 9% (l00';0>) + G08 (l00'0>) - BZ T'S9 SIOJIOY (Z00'0) + 9'PG (100';0>) + 8'€6 (l00';0>) - Os (ZS0'0) Hw = -e8e SIE spodedoo ‘wul] = 0 = 0 (100 0>) - 80 = 0 10> sploonoedie yy (180'0) ieee - 0 si 0 (10°0) ues 80 r0 sprodopoAD (161 °O) ul BLK (10002) ~— — 10> (l00';0>) - ¥0 (L>0'0) i: ete rT sprouryeD (1Z0'0) ee SiG =e 20 . 0 (T00;0>) + 98 £0 puosounydoiq = 0 (T0002) = 102 = 0 (2S0°0) ie eed 10 pyuydoq (100'0>) - 60 (6£0°0) eS (STO'0) ey Pel (T00;0>) + 0'S2 ST puywusog AVANDIA YIP AWAIT y9Ip AWA y9Ip AWAIT y9Ip yeqqey 1ay,[ue[d00Z ul % Ul % ul % ul % ul % (VG = N) (0 = N) (63 = N) (0€ = N) peys SOpISsIOAIS eide[y on{q eidey[h oniq Joye MOpl pepoolg yuspusdapu] ‘(jSa} JULI PoUIS Ss UOXOO]I AA) UOTOVT9S WOPULI 9}BOIPUI 0} petapIsuod aIIM GQ'(< senyea ‘sasayquored ul UaAIs oe sonyea Ayyiqeqorg ‘AJeAIedser1 ‘uOTaTes aATVeB0U puke ‘WOpURI ‘aATyIsod Juaseide1 — pue “Y ‘+ sfoquids YUL ‘VPplopy ‘aB10945) aye] “Ege ABW ZI ‘SAYSIF JO SaNIATOV]a SuIpsajy pue ‘siapjUe[dooz peuinsuoo puke a[qe{leAe jo ssourpunge saAnepy ‘[ ATAVL, No. 2, 1990] ZALE AND GREGORY—FOOD OF BLUE TILAPIA | gy Discuss1on—Juvenile tilapia feed on zooplankton, especially microcrus- taceans (LeRoux, 1956; Drenner et al., 1982), and it has been assumed that early life stages also do so (Bowen, 1982). Fryer (1961) however, noted that first-feeding O. variabilis ate large quantities of planktonic algae. Planktonic algae and detrital particles were important components of the gut contents of young blue tilapia in Lake George, particularly in brooded fish. The relative importance of these items declined rapidly as size increased, however. Algae and detritus probably did not contribute significantly to the nutrition of these fish; the digestive physiology necessary to extract nutrients from these materi- als, although present in adults (Bowen, 1982), is almost certainly lacking during early stages of development (Fryer, 1961). We saw no evidence of algal digestion. The brooded fish probably were feeding indiscriminantly on all available particles small enough to ingest; with experience, they shifted to more nutritionally valuable forage. When provided microcrustaceans in the laboratory, young blue tilapia ate these organisms readily and grew rapidly (Zale, 1987). In Lake George, mi- crocrustaceans made up a large portion of the diet of blue tilapia, but rotifers were also a major forage, especially in brooded fish. Consumption of rotifers declined with fish size. As in laboratory trials (Zale, 1987), independent blue tilapia in Lake George positively selected for cladocerans and calanoid cope- pods. Immature copepods made up a larger fraction of the diets of Lake George fish than of laboratory-reared blue tilapia, but their relative abun- dance in the lake was also greater. Dietary overlap of independent blue tilapia and tidewater silversides was moderate and approximated that between shad and tidewater silversides. However, similarity of diets of shad and blue tilapia was relatively high. Declines in shad abundances that have followed introductions of blue tilapia (Germany, 1977; Hendricks and Noble, 1979; Taylor et al., 1984) have been postulated to be a result of the trophic similarity of adults of these species (Hendricks and Noble, 1979; Taylor et al., 1984). Perhaps the declines in shad abundances are also influenced by overlap during the early life period, when mortality is highest and definition of year-class strengths is most pronounced (Hjort, 1914, 1926; LeCren, 1962; Braum, 1978; Hunter, 1980). Zooplankton abundance typically declines during late spring and early summer (Cowell et al., 1975; Lemly and Dimmick, 1982), possibly as a result of predation by larval and juvenile fish (Shireman and Martin, 1978; Keast, 1980), suggesting that the zooplankton available to early life stages is limited. Larvae of differ- ent fishes occur sequentially (Amundrud et al., 1974; Keast, 1980; Conrow, 1984), possibly to avoid competition by temporally partitioning this resource (Keast, 1980). Introduction of an additional zooplanktivorous early life his- tory stage, particularly one that is relatively large and effective (Zale, 1987), may reduce abundance of zooplankton sufficiently to directly affect survival of larvae of native fishes. Indirectly, decreased food supply may retard the growth of native species, thereby protracting time spent at vulnerable sizes, decreasing age-specific fecundities, and delaying age at first reproduction. 128 FLORIDA SCIENTIST [Vol. 53 Overlap-mediated effects could occur only if pre-juvenile blue tilapia ate sufficient zooplankton to significantly reduce the food available to other spe- cies. This was unlikely in Lake George during summer 1983; young blue tilapia composed only a small fraction of the larval assemblage. Our data are limited and should be viewed cautiously; additional and intensive study in natural systems is needed to better evaluate the effects of early life history stages of blue tilapia on larvae of native species. Controlled, replicated com- petition experiments in research ponds also may be helpful. ACKNOWLEDGMENTS—M. W. Collopy, C. R. Gilbert, J. A. McCann, and P. H. Eschmeyer critically reviewed an earlier draft of the manuscipt. S. G. Merrifield assisted in its preparation. R. Conrow graciously permitted the use of her townet. Funding was provided by the National Fishery Research Laboratory of Gainesville (Contract No. 14-16-0009-78-912) and the Florida Cooperative Fish and Wildlife Research Unit. The Florida Unit is jointly supported by the U.S. Fish and Wildlife Service, the Florida Game and Fresh Water Fish Commission, the University of Florida, and the Wildlife Management Institute. LITERATURE CITED AMUNDRUD, J. R., D. J. FABER, AND A. Keast. 1974. Seasonal succession of free-swimming perci- form larvae in Lake Opinicon, Ontario. J. Fish. Res. Board Can. 31:1661-1665. Bowen, S. H. 1982. Feeding, digestion and growth—qualitative considerations. Pp. 141-156. In: Puuuin, R. S. V. AND R. H. LowE-McConneE zt, (eds.), The Biology and Culture of Tila- pias. ICLARM Conf. Proc. 7, Manila. BrauM, E. 1978. Ecological aspects of the survival of fish eggs, embryos and larvae. Pp. 102-131. In: Gerkinc, S. D. (ed.), Ecology of Freshwater Fish Production. John Wiley and Sons, New York. Conrow, R. 1984. Habitat preferences and seasonal succession of early life stages of fishes in Orange Lake, Florida, with an evaluation of sampling methods. M.S. thesis. Univ. Flor- ida, Gainesville. CoweE LL, B. C., C. W. Dye, AND R. C. Apams. 1975. A synoptic study of the limnology of Lake Thonotosassa, Florida. Part I. Effects of primary treated sewage and citrus wastes. Hy- drobiologia 46:301-345. DreNNER, R. W., G. L. Vinyarp, M. GopHEN, AND S. R. McComas. 1982. Feeding behavior of the cichlid, Sarotherodon galilaeum: selective predation on Lake Kinneret zooplankton. Hydrobiologia 87:17-20. Fryer, G. 1961. Observations on the biology of the cichlid fish Tilapia variabilis Boulenger in the northern waters of Lake Victoria (East Africa). Rev. Zool. Bot. Afr. 64:1-33. GERMANY, R. D. 1977. Population dynamics of blue tilapia and its effects on the fish populations of Trinidad Lake, Texas. Ph.D. dissert. Texas A & M Univ., College Station. Henopricks, M. K. anv R. L. Noste. 1979. Feeding interactions of three planktivorous fishes in Trinidad Lake, Texas. Proc. Ann. Conf. Southeast. Assoc. Fish Wildl. Agencies 33:324- 330. Hjort, J. 1914. Fluctuations in the great fisheries of northern Europe viewed in the light of biological research. Rapp. P.-v. Reun. Cons. Perm. Int. Explor. Mer 20:1-228. . 1926. Fluctuations in the year classes of important food fishes. J. Cons. Perm. Int. Explor. Mer 1:5-38. Houpr, E. D. ano R. C. ScHEKTER. 1980. Feeding by marine fish larvae: developmental and functional responses. Environ. Biol. Fishes 5:315-334. Hunter, J. R. 1980. The feeding behavior and ecology of marine fish larvae. Pp. 287-330. In: Barpacu, J. E., J. J. MacNuson, R. C. May, AND J. M. Remnuarr, (eds.), Fish Behavior and Its Use in the Capture and Culture of Fishes. ICLARM, Manila. Hurwsert, S. H. 1978. The measurement of niche overlap and some relatives, Ecology 59:67-77. Keast, A. 1980. Food and feeding relationships of young fish in the first weeks after the begin- ning of exogenous feeding in Lake Opinicon, Ontario. Environ. Biol. Fishes 5:305-314. Kou.er, C. C. anp J. J. Ney. 1982. A comparison of methods for quantitative analysis of feeding selection of fishes. Environ. Biol. Fishes 7:363-368. No. 2, 1990] ZALE AND GREGORY — FOOD OF BLUE TILAPIA 129 LeCren, E. E. 1962. The efficiency of reproduction and recruitment in freshwater fish. Pp. 283- 302. In: LECrEN, E. D. AnD M. W. Ho pearte, (eds.), The Exploitation of Natural Animal Populations. Blackwell Press, Oxford. LeMLy, A. D. ann J. F. Dimmick. 1982. Growth of young-of-the-year and yearling centrarchids in relation to zooplankton in the littoral zone of lakes. Copeia 1982:305-321. LeRoux, P. J. 1956. Feeding habits of the young of four species of tilapia. S. Afr. J. Sci. 53:33-37. Li, H. W., P. B. MoyLe, AND R. L. Garretr. 1976. Effect of the introduction of the Mississippi silverside (Menidia audens) on the growth of black crappie (Pomoxis nigromaculatus) and white crappie (P. annularis) in Clear Lake, California. Trans. Amer. Fish. Soc. 105:404- 408. SHIREMAN, J. V. AND R. G. Martin. 1978. Seasonal and diurnal zooplankton investigations of a south-central Florida lake. Florida Scient. 41:193-201. TayLor, J. N., W. R. Courtenay, JR., AND J. A. McCann. 1984. Known impacts of exotic fishes in the continental United States. Pp. 322-373. In: CourTENAy, W. R., JR. AND J. R. STAvF- FER, JR., (eds.), Distribution, Biology, and Management of Exotic Fishes. Johns Hopkins Univ. Press, Baltimore. VON GELDERN, C. E., JR. AND D. F. MircHELL. 1975. Largemouth bass and threadfin shad in California. Pp. 436-449. In: Stroup, R. H. ANp H. CLepper, (eds.), Black Bass Biology and Management. Sport Fishing Institute, Washington. Wyoposk1, R. S. anp D. H. BENNETT. 1981. Forage species in lakes and reservoirs of the western United States. Trans. Amer. Fish. Soc. 110:764-771. ZALE, A. V. 1984. Applied aspects of the thermal biology, ecology, and life history of the blue tilapia, Tilapia aurea (Pisces: Cichlidae). Ph.D. dissert. Univ. Florida, Gainesville. . 1987. Growth, survival, and foraging abilities of early life history stages of blue tilapia, Oreochromis aureus, and largemouth bass, Micropterus salmoides. Environ. Biol. Fishes 20:113-128. Florida Sci. 53(2):123-129. 1990 Accepted: September 26, 1989. Environmental Chemistry DEVELOPMENT AND APPLICATION OF A SOIL-AIR RADON ANALYSIS TECHNIQUE IN FLORIDA RoBERT S. BRAMAN AND Robert L. SUTTON Department of Chemistry, University of South Florida, Tampa, Florida 33620 Asstract: A probe method for measuring radon in soil-air has been developed and applied to studies of radon in Hillsborough County and Polk County, Florida. Application has been in a general survey of Hillsborough County and to selected homesites. In low radon locations, from 4 to 100 pCi/L alpha activity was found. In phosphate mineralized or reclaimed land, alpha activ- ity ranged from 200 to 8000 pCi/L. THE environmental hazard posed by radon, a consequence of the presence of uranium and radium, has been extensively investigated and reported. The National Council on Radiation Protection and Measurements (1984) has re- ported on occupational and environmental exposures. Ryan and co-workers (1983) have reviewed radon & radon daughters in dwellings as have Walsh and Lowder (1983). Radon is considered to be part of the indoor air pollution problem, entering homes from soil through cracks or openings in foundations and through use of radon-containing water (Hileman, 1983; Partridge et al., 1979; and Wilkening, 1985). Wilkening and Schery (1986) have studied physical processes in radon transport indoors. Seasonal variations in a west- ern states location have been studied by Wilkening and Wicke (1986). The work reported here was initiated in order to develop a reasonably rapid survey technique to assess the potential hazard represented by soil-air radon and as a preliminary to measurement of radon transport out of the ground. Measurement of radon in soil air has been reported on by several groups. The method of Vohra and co-workers (1964) involved drilling a bore hole into the ground, capping it, and inserting a suction probe. A balloon was filled with soil air, and the balloon air was analyzed for radon. Ghosh and Bhalla (1981) also used an auger and probe technique and found some seasonal and weather effects on radon concentrations in soil-air. They also measured thoron in soil-air and drew contour maps for thoron. Horiuchi and Mura- kame (1983) measured soil-air radon by placing a vial of liquid scintillation fluid in a covered bore hole in the soil. After several days, the radon concen- tration in the fluid became constant. In our approach, a metal probe was driven into the soil. Soil-air amounts sufficient to flush out the probe were pulled through the probe. Evacuated Lucas cells, see Lucas (1974) and the U.S. EPA (1976), were attached to the probe, and the soil-air was thereby sampled. The counting rate of the Lucas cells was determined after a 2.5-3.0 hour wait for equilibrium with the radon daughters and for thorons to completely decay. The method was used to sur- vey soil-air radon at various locations in Hillsborough County, Florida and to survey several homesites in Polk County, Florida. No. 2, 1990] BRAMAN AND SUTTON—SOIL-AIR RADON ANALYSIS 131 Driver Plug =— Sampling Port 2.2 cm ID 8 - 10 in. below ground Fic. 1. Soil-air probe design 132 FLORIDA SCIENTIST [Vol. 53 MATERIALS AND MErHops—Sampling Probe and Procedure. A sampling probe was con- structed of the design shown in Figure 1. The steel pipe was fitted with a brass driving point, sampling holes and a steel driving plug. The internal volume was approximately 315 mL. This pipe probe worked well in most soils encountered. After driving the probe into the ground ap- proximately 10 inches, the driving head was removed and replaced by a rubber stopper. A hand sampling pump having a stroke volume of 250 mL was then attached to the port at the top of the probe. Soil air was then pulled through the probe and into the sampling pump. Three strokes (750 mL) of soil-air were pulled through the probe prior to sampling with the Lucas cells. Evacuated Lucas cells of 100 mL volume were then attached to the side of the sampling port, opened for sampling and then closed. After 2.5-3.0 hours, the Lucas cells were placed in the counting chamber of a Ludlum Measurements, Inc. Model 2600 Spectrometer (Sweetwater, TX) and the alpha activity counted. Counting times were generally 10 to 15 minute periods. The counting rate is for both radon and its alpha emitting daughter products in equilibrium. After counting, the Lucas cells were evacuated, flushed with air two or three times and allowed to stand until a low background counting rate was obtained. This required overnight sampling in some cases. After sampling, the soil-air probe was pulled out of the ground and dirt inside removed. The probe was also, on occasion, washed out with water and dried. Lucas cell counting rates for ambient air pulled through a recently used probe were only slightly above background. The effect of the number of hand pump evacuations on radon in soil-air data was determined by experiment. Air samples were taken from a probe after 0, 1, 2 and more strokes. Results are shown in Table 1. After one stroke, the counting rate is within random counting error the same until 12 strokes. This experiment and several duplicate sample analyses from the same spot indi- cated good sampling reproducibility was obtained. The 10-inch depth is well into the equilibrium soil-air for radon as shown in the work of Vorha and co-workers (1964), who found that the top 5-10 cm contained essentially all of the concentration gradient from air-to-soil air. A Ludlum Measurements, Inc. model 12S Micro R meter (Sweetwater, TX) was used to measure ground level radiation at some locations to determine if it was related to soil-air radon. TABLE 1. Test of Probe Use With Pump. No. Pump Strokes Count/No. Minutes 0 1/10 7/60 ] 68/10 68/10 DY, 64/10 95/15 4 77/10 77/10 8 71/10 106/15 112 61/10 91/15 RESULTS AND Discusston—Soil-Air Survey—The soil-air radon method was used in two types of studies: a general survey of Hillsborough County, Florida and a comparison study of soil-air around homes versus radon inside homes by the Track-Etch method (Terradex Corp., Walnut Grove, CA). Table 2 gives the results of the general survey. Samples were taken at homes, parks, church yards and open fields. Duplicate samples were shown as two separate numbers. In nearly every case, the soil-air radon was substan- tially above the 4 pCi/L set as an indoor-air standard. Data for Hillsborough County are shown plotted on a map (Fig. 2). No. 2, 1990] BRAMAN AND SUTTON—SOIL-AIR RADON ANALYSIS 133 Of particular interest are the high counting rates on Davis Island, Hook- er’s Point, the eastern half of Hillsborough County and some locations near Lakeland, Florida. Davis Island and Hooker’s Point are low lying areas close to Tampa Bay, very likely filled many years ago with phosphate rock dredged out of the bay. Davis Island is a residential area. Lucas cell analyses of air inside one home tested gave 1-2 pCi/L radon. Soil-air at this location was approximately 800 pCi/L. Hooker’s Point is an industrial area, again on land filled with phosphate rock from the Bay, which likely accounts for the high soil-air radon found. The eastern Hillsborough County area radon levels were generally higher than found in the western part of the county. This likely reflects more phos- phate mineralization with its attendant radon on that side of the county. Radon at a Phosphate Processing Plant—Table 2 also gives data on soil-air radon obtained at a typical phosphate processing plant location. Soil-air ra- don concentrations were reasonably low outside of an office building. The rock pile consisted of pulverized phosphate ore ready for processing. This pile had been standing for several months to one year at the locations sampled for radon. Note that the radon air samples at the rock pile were lower in concen- tration than some other sites surveyed in Hillsborough and Polk counties. The gypsum pile was also moderate in radon concentration. Radon in Homes vs. Radon in Soil Air—Five houses on reclaimed phos- phate land in Polk County with persons in residence were available for analy- sis. Track-Etch interior-air radon data and soil-air radon data were obtained on four of the houses. Ground level radiation data were also obtained at some of the locations. Table 3 presents the data obtained. Track-Etch data were not as high as expected for interior radon at locations having a high soil-air radon. The highest reading recorded, at house “A”, was in a low spot on the side lawn; apparently the radiation comes from leaching off of a nearby old phosphate pile (hill). A similar observation was noted at house “B”. The soil-air radon variation was measured over two small plots of open ground. A substantial variation was noted for the five homesites on re- claimed phosphate land with a high radon area near the center of the land sites. Less variation was noted on the sandy soil location near the University of South Florida campus. Both plots of land were reasonably level with no low areas which might collect drainage water and give pockets of high radon concentrations such as were noted at houses “A” and “B”. A closer spacing of sampling could likely result in development of soil-air radon maps with con- tours if desired. This would require a larger number of Lucas cells so as to provide for flushing out of radon and decay of radon daughter products after each use. Radon vs. Ground-Level Radiation—Ground-level radiation values, mi- crorads/hour, were obtained with a Ludlum Model 12S meter. These data, shown in Table 2, were obtained from a variety of sampling locations and so several soil types are involved. As is indicated, a general rise in soil-air radon accompanies the ground level radiation. Ground level radiation data could 134 FLORIDA SCIENTIST [Vol. 53 Fic 2. Radon in soil-air, Hillsborough County, Florida serve as a crude indicator of radon with any levels above 5 microrads indica- ting a probable high radon and requiring soil air sampling. Conc.Lusions—The soil-air probe technique combined with Lucas cell sampling proved to be a comparatively easy method for determining soil-air radon. With multiple Lucas cells, area surveys can be performed in one day. Replicate sampling from one probe location gave good reproducibility. Nev- No. 2, 1990] BRAMAN AND SUTTON—SOIL-AIR RADON ANALYSIS 135 TABLE 2. Soil-air radon data, general survey. Location Soil-Air Radon, pCi/L Residence-Carrollwood Dr. Tampa-front yd. 29 rear yd. 62 Residence-124th Ave. Tampa 73 Residence, Knight’s Rd. 53 Park at US 41 & Alafia River 50 Simon-Bower Park 12 Ballast Point Park 2 eeeoll Hyde Park (Albany & Swan Ave.) 1), (il) Horizon Park (Dale Mabry Hwy.) ope v4 USF (Near Engineering Building) 32 USF (Near Physics Building) 40 USF (Near Administration Building) 42, 33 Davis Island (tennis courts) 700, 717 Davis Island (S.E. end) 2633, 2536 Davis Island Park (South end) 589, 555 Davis Island Residence #1 823, 858 Davis Island Residence #2 770, 788 Davis Island (near airport) 845, 900 Hooker’s Point (20th Street) AAs PANE Hooker’s Point (end of Maritime Street) 2175, 1949 DeSota Park (near Hooker’s Point) 245, 253 Temple Terrace (Gillette Ave.) Me}, B83 Temple Terrace (Florida College) 49, 39 Temple Terrace Annex Park 31 Residence (Oaklawaha Ave.) 23, 18 Robles Park (Tampa) 57 Thonotasassa (Main Street Park) 574, 600 Valrico (Dover Road) 240, 182 Knights (Park, SR 582 & SR 39) 443, 407 Old Hopewell Road (SR 39 & SR 60) 973 Intersection, SR 39 and SR 640 15 Oee lor Rowlett Park (at Hillsborough River) 61 Lettuce Lake Park (Fletcher Ave.) Ao Sheldon Park (Sheldon Road) 25, 44 Buffalo Estates (Danny Brian Road) 592, 545 Citrus Park 29 Lake Park (N. Hillsborough County) 82, 93 US 41 North of Ruskin 180, 134 SR 674, Wimauma AV Parl Fort Lonesome SR 674 321, 311 Pinellas County—Fort Desoto Park (at Gulf) 23 Lakeland, Mobile Home Park, East Side— 23 West Side— 5098, 4617 Polk County—at Phosphate Ore Processing Plant: “Rock Pile” — West Side 74) “Rock Pile” —East Side 942 Gypsum Pile— West Side 555 Gypsum Pile—FEast Side 120 Admin. Building—West Side 116 Admin. Building—East Side 33 ertheless, sampling over small plots of land (Table 3) and the homesite studies indicated that one can observe substantial variations in soil-air radon in a small area. Consequently, several samples should be taken when studying homesites. 136 FLORIDA SCIENTIST [Vol. 53 TaBLeE 3. Homesites and fields, alpha activity, pCi/L and background radiation, microrads/hr Location Polk County— Homesites House A— Indoor Air—6.03 pCi/L Soil Air— Backyard L. Rear Rt. Side (in a depression) Rt. Side (hill) Rt. Front Front House B—Indoor Air—1.71 pCi/L House C—Soil Air— Back L. Rear Rt. Rear L. Front Rt. Front Front Back Rt. Front Close Front House D—Indoor Air—1.69 pCi/L Rt. Rear Rear Left Side Close Left Side Far Right Side House E—Indoor Air—1.65, 4.77 pCi/L Rt. Rear Rear Right Left Close Left Side (Across Ditch) Front Homesite (Field) (100 ft apart) 26-30 urad/hr— (all sites) Open Field (Sandy) (50 ft. apart) (3 urad/hr— (all sites) #1 (left) #2 #3 #4 #5 (right) Front L. Rear Rear Center R. Rear L. Front Front Center R. Rear Radon Background ov i gor Reclaimed phosphate land or filled land, as in Davis Island and on Hook- ers Point, were sharply higher in soil-air radon, and this could have an im- pact on indoor radon, especially if surface water runoff collects radioactive materials near a homesite. Although indoor-air radon should increase with soil-air radon, more comparative studies are needed to better determine this. Obviously, home construction characteristics also have an influence. The No. 2, 1990] BRAMAN AND SUTTON—SOIL-AIR RADON ANALYSIS 137 finding of some locations having 1000 times more soil-air radon than the 4 pCi/L recommended action levels being considered lends impetus to this type of study. Also of interest will be the impact of micrometeorology on air radon at the high soil-air radon locations. Although Wilkening (1985) has done some of this type of work, the soil conditions, vegetation and weather in Florida contrast sharply with the locations studied by Wilkening. The nighttime loss of wind at a tree canopied location should result in a very substantial increase in air concentration of radon even at window heights above the ground as we found to be the case with mercury diffusing out of the ground (Johnson & Braman, 1974). ACKNOWLEDGMENTS— The authors wish to thank International Minerals and Chemical Cor- poration for financial aid. The authors also wish to acknowledge helpful discussions of their work with H. Ralph Brooker and Dale Spurgin of the Department of Physics, University of South Florida. LITERATURE CITED Guosu, P. C. anDN. S. BHALLA. 1981. The behavior of thoron (Rn-220) in soil. Indian J. Earth Sci. 8:1-9. Hiteman, B. 1983. Indoor air pollution. Environ. Sci. Technol. 17:469A-472A. Horivcui, K. anD A. Murakami. 1983. A new method for the determination of radon in soil air by the open-phial (method) and integral counting with a liquid scintillation counter. J. Radioanal. Chem. 80:153-163. Jounson, D. L. ann R. S. Braman. 1974. Distribution of atmospheric mercury species near the ground. Environ. Sci. Technol. 8:1003-1009. Lucas, H. F., Jr. 1957. Improved low-level alpha-scintillation counter for radon. Rev. Sci. Instr. 28:680-683. NATIONAL CoUNCIL ON RADIATION PROTECTION AND MEASUREMENTS. 1984. Evaluation of Occu- pational and Environmental Exposures to Radon and Radon Daughters in the United States. Washington, D.C. ParTRIDGE, J. E., T. R. Horron, AND E. L. SENsINTAFFER. 1979. A Study of Radon-222 Released from Water During Typical Household Activities. Eastern Environmental Radiation Fa- cility, U.S. Environmental Protection Agency, Washington, D.C. Ryan, M. T., W. A. Gotpsmitn, J. W. Poston, F. F. Haywoop, anp J. P. WiTHERsPooN. 1983. Rapon Dosimetry: A REVIEW OF RADON AND RADON DAUGHTER EXPOSURE CONDITIONS IN DWELLINGS AND OTHER STRUCTURES. ORNL/TM-5286, Oak RipcE NATIONAL LABORA- TORY, Oak RipGE, TENNESSEE. U.S. ENVIRONMENTAL PROTECTION AGENCY. 1976. Interim Radiochemical Methodology for Drinking Water. Washington, D.C. Voura, K. G., M. C. SuBBARAMU, AND A. M. Mouan Rao. 1964. Measurement of radon in soil gas. Nature 201:37-39. Watsu, P. J. AnD W. M. Lownper. 1983. Assessing the Risk from Exposure to Radon in Dwellings. ORNL/TM 8824, Oak Ridge National Laboratory, Oak Ridge, Tennessee. WILKENING, M. 1985. Radon transport in soil and its relation to indoor radioactivity. Sci. Total Environ. 45:219-226. AND S. D. Scurry. 1986. Physical Processes Affecting Levels of Radon, Thoron, and Their Decay Products in an Indoor Environment. Report 1986, DOE/ER-60216. Wash- ington, D.C. AND A. WicxkeE. 1986. Seasonal variation of indoor radon at a location in the South- western United States. Health Physics 51:427-436. Florida Sci. 53(2):130-137. 1990. Accepted: October 3, 1989. Biological Sciences A NOTE ON THE FIRE RESPONSES OF SPECIES IN ROSEMARY SCRUBS ON THE SOUTHERN LAKE WALES RIDGE ANN F. JOHNSON” AND WARREN G. ABRAHAMSON” ()Florida Natural Areas Inventory, Tallahassee, FL 32303; 2) Department of Biology, Bucknell University, Lewisburg, PA 17837. Asstract— Three herbaceous species (Lechea deckertii, L. cernua, and Paronychia chartacea) appeared in postburn samples of plots in rosemary scrubs, that were rare or absent in preburn samples of the same plots and were also rare in samples of plots in unburned scrubs. Changes in cover levels with stand age suggest that these species are displaced by the increasing cover of rose- mary and reindeer lichens within 9 to 12 years, their populations then being rejuvenated by periodic fires. Two major types of shrubby vegetation are found on the gently rolling topography of nutrient-poor acid sands comprising the southern Lake Wales Ridge (27°11’N Lat., 81°21’W long.): rosemary scrubs (Ceratiola ericoides) which tend to occupy the knolls, and scrubby flatwoods (Quercus inopina), on the surrounding slopes (Abrahamson et al., 1984). Studies have shown the latter type to be resilient to fire which is common in this lightning-prone re- gion—scrub oak (Quercus inopina) resprouts rapidly postfire, as do most of its associated species, and stands of scrubby flatwoods show little or no change in species composition or abundance after fire (Abrahamson, 1984a,b). It was not known, however, whether rosemary scrubs would show a similar response, particularly since rosemary, in contrast to scrub oak, is killed by fire and regen- erates from seed (Johnson, 1982)—leaving a period of time as the seedlings are maturing when space is available for colonization by other species. The ques- tion of the fire response of rosemary scrubs is of timely interest for scrub man- agement due to a high concentration of species endemic to this vegetation type that are federally listed as threatened or endangered (Christman, 1988). An opportunity to begin to answer this question arose when a series of three inadvertent fires at Archbold Biological Station burned rosemary stands in which permanent belt transects had been sampled 6 years previously. The belt transects, as well as belt transects in unburned stands, were resampled to docu- ment postfire changes in burned rosemary stands in comparison with changes in unburned stands over the same period. Supplementary data on the fire re- sponses of three species were taken from samples of permanent line transects that were burned in prescribed and accidential fires. MeErHops—Belt transects: Thirteen rosemary stands, scattered over the 1300 hectares com- prising the western section of Archbold Biological Station, were sampled in November-December 1979. These were initially chosen as part of a demographic study to represent a range of stand ages from 7 to more than 30 years (Johnson, 1982). After 30 years, the shrubs begin to layer, precluding further aging of the stands. Percent cover of all species was visually estimated in every 2X2 m2 quadrat along a transect 2 m wide and from 10 to 20 m long, in each stand. The transects No. 2, 1990] JOHNSON AND ABRAHAMSON—FIRE RESPONSES 139 were located in the densest portion of each rosemary stand and were oriented from the crest downslope into the adjacent vegetation type (i.e., scrubby flatwoods). Initially each transect was marked with two wooden end stakes. After the fires, these were replaced with permanent metal stakes. The two palmettos, Serenoa repens and Sabal etonia, were not distinguished in the initial sample and thus cover values for the two were combined. Seven of the 13 stands sampled were burned in three fires—in September 1984 a prescribed burn escaped (stand nos. 5, 11-1, 11-2); in January 1985 a prescribed burn escaped (stand nos. 18, 19, 20), and in June 1986 a lightning strike started a fire (stand no. 39). Burned stands were resampled 2 years postfire; the six unburned stands were resampled 7-9 years later, in 1987 (stand nos. 26, 27B) and 1989 (stand nos. 1, 16, 25, 29). Line transects: Permanent line transects, set up as part of a larger study to determine vegeta- tion responses to fire (Abrahamson, 1984a,b), were sampled using two 100-m parallel lines in the scrubby flatwoods sites and 4 parallel lines (60,60,60, and 20m) in the rosemary scrub site. The length of the line intercepted by each shoot, whether over, under, or touching the line, was recorded. Percent cover was calculated as: (total length of line intercepted per species/total length of line sampled) X 100. None of these transects had been burned for at least 25 years prior to sampling. Three of the four scrubby flatwoods transects were burned twice (1977 and 1985) during the sampling period, while the remaining scrubby flatwoods transect (WS P2) and the rosemary scrub were burned once (1985). ResuLts—Belt transects: A total of 37 species of vascular plants and 7 species of reindeer lichens were found in the belt transects. Numbers of spe- cies per transect were evenly divided across a range of from 8 to 19 per transect. Of the 37 species of vascular plants, casual observations suggest that 11 species are “seeders”, i.e. are destroyed by fire and recolonize solely from seeds or spores (Balduina angustifolia, Calamintha ashei, Ceratiola ericoides, Helianthemum nashii, Hypericum cumulicola, Lechea cernua, L. deckertii, Paronychia chartacea, Pinus clausa, Selaginella arenicola, Stipulicida seta- cea); 20 are “sprouters” (Andropogon brachystachyus, Bumelia tenax, Cnido- scolus stimulosus, Cyperus sp, Gaylussacia dumosa, Licania michauxii, Lyonia lucida, L. fruticosa, Opuntia compressa, Persea humilis, Polygonella polygama, Quercus chapmanii, Q. geminata, Q. inopina, Rhynchospora me- galocarpa, Sabal etonia, Serenoa repens, Smilax auriculata, Vaccinium myr- sinites, and Ximenia americana) and the reactions of the remaining 6 are uncertain (Aristida gyrans, Bulbostylis ciliatifolia, Palafoxia feayi, Poly- gonella robusta, Sisyrinchium sp., and Tradescantia sp.). The 7 reindeer li- chens—4 common species (Cladina subtenuis, C. evansii, Cladonia pro- strata, and C. leporina) and 3 uncommon (Cladonia pachycladodes, C. perforata, and C. subsetacea)—are all destroyed by fire, recolonizing by spores or fragments, and thus behave as “seeders.” The responses to fire of the 6 species that make up over eighty per cent of the cover in the stands are shown (Fig. 1). In the preburn and unburned samples, seeder species (rosemary and the lichens) make up the bulk of the cover (20-80 percent), while cover of sprouter species (primarily scrub oak and the palmettos) ranges from 2 to 25 percent (average=11 percent) of the total. In the two-year postfire samples the seeder species show little recovery, while the sprouter species equal or surpass preburn levels (Fig. 1). Seedling density of rosemary (0.6 to 15.7/m?), although adequate to re- place the stands at the normal range of shrub densities (0.3-2.1/m’; Johnson, 1982), was not related to previously measured seed production per m? of FLORIDA SCIENTIST [Vol. 53 140 YA S6< GCHNYNANN YA S6< CHNYUNANN YA S¢< GCHNYNANN YASG< GCHNYNANN AY AUVANVI = 80°0 = O10 = i LEO BO = a 10°0 = SN SN SN SN SN SN AYIA AUVANVI SN SN SN SN SN SN 410 ~=10'0 = SN SN SN Id PT IT 93 SMA = = £0°0 = oe'0 = SN SN SN AYA AUVANVI AYA AUVANVI SN SN SN = = 10°0 = oro ~=3— €0'0 WA SZ< GANYNANN a = TEE = SFO =r TT'0 = £0'0 = = = 80°0 = 160 SEO = = = = 60°0 10°0 GOW = O81.) =87 0 = 100 §8©10'°0> = 90'°0 — C00 -GL0— ZS0 = r0'0 ~=—- 600 = = a 200 3680s ST‘ = 100 §0°0 SN SN SN SN SN SN SN SN SN SN SN SN SN SN SN AYA HAAN aALdAS aAYa AYVOANVI AY AUVANVI = = = SN SN SN SN SN SN 30°0 = = SN SN SN SN SN SN = €0°0 = G00 = CFO 60T 980 €8'°0 = 300 = SN SN SN SN SN SN SN SN SN Id PT IT Id PT IT Id PT IT £3 SM Go SM od SM SpOOM}P [HY Aqqniosg q-ue[ 71 re ., /LLOI e-uel SN SN SN /LLOI Aqn{ = 80°;0 =r 0 /LLOI ars aa a 0) SL6I jars = “es 6L61 = a 600 O86I SN SN SN I86I —% —7 se 6861 SN SN SN C861 SN SN SN V86I AW AUVANVI C86I SN SN SN 9861 SN SN SN L861 SN SN SN 8861 680 SEG 607 6861 Id PT IT Ivak LG SM qniog Arewasoy ‘I1lf a1OJaq = q ‘a1lj Joye =e “IBAA YEU) po[dures jou sem o}IS JY} SOPBOIPUI GN ATI ‘JOosuBI] OUT] BY} UO potd}UNODUI }OU SBA\ SaIdads ay} }eY} 9JVOIpPUT SoUT] poyse(] ‘Says spoomyep Aqqnios Mo}; pue dPS Gn1ds auo 7 SyoasueI} BUT] JUOURUIEd LW (IZ UO (2g) DaID}ADYI DIYDhuOLDg pur ‘(p'T) Wj4ayIap DaYIaT ‘(I'T) DNULII DIYIAT JO 19A09 YUIDIOg *[ ATAV], No. 2, 1990] JOHNSON AND ABRAHAMSON—FIRE RESPONSES 141 BURNED UNBURNED 50- 40- Per cent ae cover 20- 10- i no no Ceratiola ericoides ae (Zo GS data standno. 39 11(2)18 19 20 5 11(1) stand age (yrs) 7 15 20 20 20 30 ia 30- -= Cladonia/Cladina spp. , 90 —0 Ho fo Go | a 20- ae : NS Quercus inopina , ANY 0 oo YN oo oo oo NW — a Sabal/Serenoa por eee RY a. NY BN wo a 10- Lechea cernua o- 0O 00 O41 Oo. OF. on OW oo 00 oO of 00 00 10- Lechea deckertii o- J of OQ O73 OO 0... of OO OO Oo. oO 00 00 10- Paronychia chartacea 4. ny 0. o4 OF O» O. Oo Oo. Oo. OO oO OO OO Zo eecier N sprouters Fic. 1. Per cent cover of selected species in belt transect plots in burned and unburned rose- mary stands ordered by age of stand. First bar of each pair shows cover in the initial sample (1979); second bar shows cover when resampled in 1987-1989, i.e. 2 yr postfire in burned stands; 7-9 yr later in unburned stands. La = layered stands (age greater than 30 yr). shrubs in the same transects (Johnson, 1982). An exception was the extreme case of the youngest stand (no. 39), which was non-reproductive (7 yr) when first sampled in 1979 and would have just barely reached reproductive matu- rity (14 yr) when it was burned in 1986. Only one seedling was found in the postfire sample of this transect. Such a density (0.03/m7’) is an order of magni- tude too low to replace the stand at normal shrub densities and serves to support the prediction (Johnson, 1982) that burning rosemary scrub at inter- 142 FLORIDA SCIENTIST [Vol. 53 vals shorter than 10 years (or in this case 14 years) can cause local extinction of the stands. Three species, Lechea deckerii, L. cernua, and Paronychia chartacea, not present (or present at low cover levels—0.3 per cent or less) in preburn and unburned samples, appear at slightly higher cover levels in the postburn sam- ples (Fig. 1). The time course of recovery after fire can be followed in two ways (Fig. 1): by comparing changes in the same stand sampled at different times, or by comparing stands of different ages, since all these even-aged stands can be assumed to date from a fire event (Johnson, 1982). In comparing stands of different ages, it appears that cover of rosemary shrubs can reach 20 per cent by age 7 years—which is 9 years postfire, since rosemary seeds do not germi- nate until the second growing season postfire (Johnson, 1982). After 11 years postfire, cover of rosemary shows no increase with increasing age, ranging between 30 and 50 per cent in stands from 9 to 30* years old. This lack of increase in rosemary cover is also seen in comparing samples from the 6 un- burned stands taken 8-10 years apart. Although cover remains relatively con- stant, structural changes continue as the rosemary shrubs grow and thin out (Johnson, 1982). The lichens, when compared over stands of different ages (Fig. 1), are seen to regain their preburn cover more slowly than rosemary, reaching maximum levels (25 to 35 percent) only in stands over 30 years in age. Although lichens are thought to be slow-growing, cover in three un- burned stands (Fig. 1) increased by about 10 per cent over the 9 year period between samples. Most of this increase was contributed by Cladonia leporina in the two younger, unshaded stands (nos. 26, 27B) and by Cladina evansii in the older (30 yr), more shaded stand (no 1). Cover of sprouter species shows little change with stand age and no trends over the 7-9 year time interval in unburned stands. Line transects: Cover responses of Lechea cernua, L. deckertii, and Paronychia chartacea from a line transect in rosemary scrub (Table 1) show the same postfire increases as seen in the belt transects. In this case Lechea cernua reaches higher cover levels than the other two species. In contrast to their consistent behavior in scrub samples, the three species showed postfire cover increases in only one (WS25) of the four stands of scrubby flatwoods sampled (Table 1), and these increases were substantially less than those observed in the scrub sample. This scrubby flatwoods stand (WS25) was adjacent to and contained elements of rosemary scrub. Discuss1oN—In contrast to the pattern in stands dominated by sprouter species which recover their preburn aspect in 2-4 years postfire (Abrahamson, 1984a,b), the major contributors to cover in rosemary scrubs (Ceratiola and Cladonia/Cladina spp.) are destroyed by fire and take 10 to 12 years to re- cover to preburn levels, leaving a window of time when other species may colonize the area. This opportunity is taken by an endemic annual cushion No. 2, 1990] JOHNSON AND ABRAHAMSON—FIRE RESPONSES 143 plant (Paronychia chartacea) and two suffrutescent forbs (Lechea deckertii and L. cernua) and possibly others. Gradually rosemary and the lichens re- gain their preburn cover, displacing the fire followers which are seen to per- sist in sand roads and in the larger openings in the stands. Postfire cover increases of these three species in scrubby flatwoods are damped or absent presumably because the rapid resprouting of the dominant shrubs preempts the available space. Other vegetation samples in scrubby flatwoods have shown these species to be less frequent in this vegetation type than the line transect samples shown here might suggest. None of the three were found in samples (four 2X10 m plots) from 9 stands of scrubby flatwoods from a biomass study (Johnson et al., 1986), and only L. deckertii was fre- quently encountered (5/12) in the belt transect extensions into scrubby flat- woods. The results from this limited sample raise a number of questions that could be addressed experimentally or by more extensive sampling, such as: 1) do other species come in after fire?, 2) how do rosemary and lichens displace the fire followers (allelopathy, shading)?, 3) do the fire followers persist as a seed bank between fires or must they seed in from refugia?, the answers to which would be helpful in managing these unique communities containing many federally endangered plant (and animal) species. ACKNOWLEDGMENTS— This work was supported in part by grants from Archbold Biological Station and Bucknell University. LITERATURE CITED ABRAHAMSON, W. G. 1984a. Post-fire recovery of Lake Wales Ridge vegetation in south-central Florida: a five-year study. Amer. J. Bot. 71:9-21. 1984b. Species responses to fire on the Florida Lake Wales Ridge: a five-year study. Amer. J. Bot. 71:35-43. , A. F. Jounson, J. N. Layne, anv P. A. Peroni. 1984. Vegetation of the Archbold Biological Station, Florida: an example of the southern Lake Wales Ridge. Florida Scient. 47:209-250. CuHRIsTMAN, S. P. 1988. Endemism and Florida’s interior sand pine scrub. Final report, project no. GFC-84-101, Florida Game and Freshwater Fish Commission, Tallahassee. Jounson, A. F. 1982. Some demographic characteristics of the Florida rosemary. Amer. Mid. Nat. 108:170-174. , W. G. ABRAHAMSON, AND K. D. McCrea. 1986. Comparison of biomass recovery after fire of a seeder (Ceratiola ericoides) and a sprouter (Quercus inopina) species from south-central Florida. Amer. Midl. Nat. 116:423-428. Florida Sci. 53(2):138-143. 1990. Accepted: October 6, 1989. 144 FLORIDA SCIENTIST [Vol. 53 REVIEW Robert K. Godfrey. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and Alabama. The University of Georgia Press, Athens and London, 1988. Pp. ix + 734. $50.00. Tus is a technical manual intended to assist in the identification of the native and naturalized trees, shrubs, and woody vines of northern Florida, southern Georgia, and southern Alabama. The distinction between a shrub and a tree and between a woody and non-woody perennial herb is often not evident. Some, such as Lupinus westianus, seem hardly woody. However, this does not distract from the book. The species are divided into three groups: gymnosperms, monocotyledons, and dicotyledons. Within each group, the families are arranged alphabetically, and within each family, the genera are also alphabetical. However, the species are sometimes alphabetical, some- times not, depending upon the author’s inclination. It is difficult to locate species in large genera that are not alphabetically arranged. For example, it is particularly inconvenient having to locate oak species (28 species distrib- uted over 30 pages of text) by having to refer to the key for the species number or to the index. Emphasis is at the species level with each described in detail. Family descriptions and descriptions of genera in which a single species oc- curs in the range of the manual are not provided. The species and generic descriptions are usually in a three paragraph format, one each for the habit, leaf, or floral/fruit characters. A few (e.g. Lonicera) are in a single paragraph format, thus making presentation inconsistent. Following the species descrip- tions is a generalized habitat statement along with the distribution of the species for the range of the manual, then the distribution over the species’ full range. The inclusion of the National Champion Big Tree is of limited interest use, as more often than not, the individual tree is outside of the manual range. Many names used in the more common manuals, such as Small’s 1933 Manual of the Southeastern Flora, are listed as synonyms so that cross refer- ence of names can be made. Unfortunately, this information is incomplete as many names found in Small or even some (e.g. Carya ovalis) that are found in Clewell’s 1985 Guide to the Vascular Plants of the Florida Panhandle are unexplained absences. The work is often frustrating to use because of the incomplete synonymy. No species are treated in the genus Pyracantha al- though a good description of the genus is given. Many species are illustrated which greatly facilitates identification. The illustrations, line drawings mostly prepared by Melanie Darst, are of excellent quality. A few that were prepared by Priscilla Fawcett did not reproduce well and look over-inked. Although regional in scope, the book is useful for some distance outside its prescribed range because of the wide distribution of many species. The ample descriptions, excellent illustrations, and simple format make it easy to use. I highly recommend it.—Richard P. Wunderlin, Department of Biology, Uni- versity of South Florida, Tampa, FL 33620. 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Florida's Estuaries — Management or Mismanagement? — Academy Symposium FLoripa SCIENTIST 37(4) — $5.00 Land Spreading of Secondary Effluent —Academy Symposium FLoripa SCIENTIST 38(4) — $5.00 Solar Energy — Academy Symposium FLORIDA SCIENTIST 39(3) — $5.00 (includes do-it-yourself instructions) Anthropology — Academy Symposium FLORIDA SCIENTIST 43(3) — $7.50 Shark Biology — Academy Symposium FLoripaA SCIENTIST 45(1) — $8.00 Future of the Indian River System — Academy Symposium FLoripa SCIENTIST 46(3/4) — $15.00 Individual orders should be sent with payment. A statement will be sent in re- sponse to a bona fide purchase order over $10.00 from a recognized institution. Ad- dress all orders to: The Florida Academy of Sciences, Inc. c/o The Orlando Science Center 810 East Rollins Street Orlando, Florida 32803 Phone: (305) 896-7151 Florida ISSN: 0098-4590 a in / as (ers 1 5 1990 \ Volume 53 Summer, 1990 Number 3 THE SECOND INDIAN RIVER RESEARCH SYMPOSIUM CONTENTS 2 OES TCG a eee Ned P. Smith and Richard L. Turner Numerical Modeling of Tidal Hydrodynamics and Salinity ®ransport in the Indian River Lagoon ..............0..20.005: Y. Peter Sheng, S. Peene, and Y. M. Liu Groundwater Seepage into the Indian River Lagoon at son MPT CL Pero che) fan aN A Need 48 ae ale Ashok Pandit and Clovis Clovis El-Khazen Water Quality Changes Associated with Rotary Ditched and Breached Mosquito Control Impoundments in MMSMRCOMEAC OOM 2 foes oiled ce ease Se a a a J. C. Gamble, J. P. Stewart, and W. R. Ehrhardt Precipitation Chemistry: Atmospheric Loadings to the Surface Waters of the Indian River Lagoon Basin by 2171112) Thomas W. Dreschel, Brooks C. Madsen, Lee A. Maull, C. Ross Hinkle, and William M. Knott III Salt Marsh Mitigation: An Example of the Process of Balancing Mosquito Control, Natural Resource, and Development EEE: ah a a eee er om ee Cee Peter D. O’Bryan, Douglas B. Carlson, and R. Grant Gilmore Holocene Evolution of Indian River Lagoon in Central ERRCe ECU OUTIL FP LOTIGA ie og sich s bs a shale ey deeie(erel's pute 6 ide oles Sharon F. Bader and Randall W. Parkinson An Introduction to the Tides of Florida’s Indian River Lagoon. II. “LENS CB i, OOD ROUT ROH AEE ae aaa Ned P. Smith Ichthyofauna Associated with Spoil Islands in Indian River AMO FIONA 6 10 o. Bee sig ae os Nancy Brown-Peterson and Ross W. Eames Adaptive Specializations of the Cyprinodont Fish EMMOIEES TIVOTIIOTOUUS irc ie 6 ibs ak gle tyelas's bv a's D. Scott Taylor The Large Spatial and Temporal Biological Variability of MADE ERIVED CACOON. 2... he) 6 i ws boa ees Robert W. Virnstein 145 147 169 180 184 189 204 216 226 239 249 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1990 Editor: Dr. DEAN F. MARTIN Co-Editor: Mrs. BARBARA B. MARTIN Institute for Environmental Studies Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoripA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. Applica- tions may be obtained from the Executive Secretary. Both individual and institutional members receive a subscription to the FLoripa ScIENTIST. Direct subscription is avail- able at $20.00 per calendar year. Original articles containing new knowledge, or new interpretation of knowledge, are welcomed in any field of Science as represented by the sections of the Academy, viz., Biological Sciences, Conservation, Earth and Planetary Sciences, Medical Sci- ences, Physical Sciences, Science Teaching, and Social Sciences. Also, contributions will be considered which present new applications of scientific knowledge to practical problems within fields of interest to the Academy. Articles must not duplicate in any substantial way material that is published elsewhere. Contributions are accepted only from members of the Academy and so papers submitted by non-members will be accepted only after the authors join the Academy. Instructions for preparation of manuscripts are inside the back cover. Officers for 1990-91 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. FREDERICK BUONI Treasurer: Dr. ANTHONY F. WALSH Operations Research Program 5636 Satel Drive Computer Science Department Orlando, Florida 32810 Florida Institute of Technology Melbourne, FL 32901 Executive Secretary: Dr. ALEXANDER DICKISON Department of Physical Sciences President-Elect: Dr. GEorcE M. Doorts Seminole Community College Division of Science and Mathematics Sanford, Florida 32771 St. Leo College St. Leo, FL 33574 Program Chair: Dr. DEL DELUMYEA Millar Wilson Laboratory for Chemical Research Secretary: Dr. PATRICK J. GLEASON Jacksonville University 1131 North Parkway Jacksonville, FL 32211 Lake Worth, Florida 33460 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Dean F. Martin, Editor BARBARA B. Martin, Co-Editor Volume 53 Summer 1990 Number 3 THE SECOND INDIAN RIVER RESEARCH SYMPOSIUM FOREWORD ALTHOUGH research in Indian River lagoon has been conducted on a wide range of topics for many decades, studies have mostly been done indepen- dently. Results are scattered through many scientific journals, theses, disser- tations, and reports. Within the past several years, however, there have been efforts to collect, integrate, and summarize studies of the lagoon. The Marine Resources Council of East Florida (MRC) has undertaken and completed a task as onerous as it is vital by compiling a list of 2007 citations of Indian River research (Morris, 1989). Currently, MRC is spearheading the produc- tion of a multivolume monograph that summarizes the state of knowledge of the biology, chemistry, geology, and physics of the lagoon. The reporting of research findings has progressed on yet another front in the form of symposium proceedings. Just as the symposium is necessary to assemble scientists and managers to discuss work related to the management of the lagoon, symposium proceedings provide a direct route to the scientific literature for recently completed studies. The first volume of proceedings was published as results of the Future of the Indian River System Symposium (FIRST; Montgomery and Smith, 1983). The present volume contains 10 of the 29 oral presentations and 7 posters given at the SECOND Indian River Research Symposium, held on the Florida Institute of Technology (FIT) cam- pus on 12-13 September 1988. The purpose of these proceedings is to inject new findings into the scientific literature. While this volume does not indi- cate either the magnitude or scope of current research activity in Indian River lagoon, it takes its place in the growing body of literature needed to support proper management and, thus, to sustain this valuable natural resource. The symposium was sponsored by MRC, FIT, St. Johns River Water Man- agement District (SJRWMD), and South Florida Water Management District (SFWMD), for whose support we are grateful. We are especially appreciative of the help of Diane D. Barile and her staff at MRC in organizing and han- dling local arrangements for the symposium. Preparation of this symposium volume would have been difficult without the guidance, encouragement, wisdom, and prodding given by Dean and Barbara Martin. Publication of this issue was aided by funds provided by SJRWMD and SFWMD under the Surface Water Improvement and Management (SWIM) Program. 146 FLORIDA SCIENTIST [Vol. 53 LITERATURE CITED Montcomery, J. R., AND N. P. SmitH (eds.). 1983. Future of the Indian River System. Fla. Scient. 46:129-431. Morris, F. W., IV. 1989. Bibliography of the Indian River Lagoon Scientific Information System. St. Johns River Water Management District and South Florida Water Management Dis- trict, Special Publ. SJ 89-SP2. Ned P. Smith” Richard L. Turner” Associate Editors “Harbor Branch Oceanographic Institution, 5600 Old Dixie Highway, Fort Pierce, FL 34946; Department of Biological Sciences, Florida Institute of Technology, Melbourne, FL 32901 4 May 1990 Oceanographic Sciences NUMERICAL MODELING OF TIDAL HYDRODYNAMICS AND SALINITY TRANSPORT IN THE INDIAN RIVER LAGOON Y. PETER SHENG, S. PEENE, AND Y. M. Liu’ Coastal and Oceanographic Engineering Department, University of Florida, Gainesville, Florida 32611 Asstract: A one-dimensional model of circulation and salinity transport within the Indian River lagoon system is presented. The model is based on the cross-sectionally averaged continuity equation, momentum equation, mass conservation equation for salt, and equation of state. A fractional step method and time-implicit scheme are used in the finite-difference model. The model is used to examine the role of pure barotropic tidal hydrodynamics and to explore the time scales of salinity transport in the lagoon. Model simulation of barotropic tidal circulation in Indian River lagoon successfully reproduced the water level measured during a 3-day episode in September 1984. Model simulation of tide-induced salinity transport in Indian River lagoon indicates a very long response time on the order of 100 tidal cycles. The presence of a uniform wind stress of 1 dyne/cm? from the South enhances the salinity transport and reduces the response time to 10 to 20 tidal cycles. Results indicate that long-term flow and salinity data are needed for understanding and modeling the circulation and salinity transport within the Indian River la- goon. GROwING human activities have led to the deterioration in water quality and biological resources in estuaries, including portions of Indian River la- goon (Fig. 1). In order to formulate a sound management plan for Indian River lagoon (Barile, 1984), it is essential to have a quantitative understand- ing of its hydrodynamic, chemical, and biological processes. The distribution of water quality and biological parameters of the lagoon are predominantly affected by the hydrodynamics. Hydrodynamic processes of Indian River lagoon are primarily driven by tide, wind, fresh water, and evaporation. While much work has been done previously and a literature survey was presented by Dombrowski, Morris and Reichard (1987), there have been few comprehensive hydrodynamic studies of the entire lagoon system. At present, there is a lack of quantitative under- standing of the hydrodynamic processes, and there exists no comprehensive hydrodynamic model which has been sufficiently validated for the entire lagoon. While several modeling studies (e.g., Morris, 1985 and 1987a; Smith, 1982) have investigated various parts of Indian River lagoon, only one model (Williams, 1985) attempted to include the entire lagoon. Williams (1985) utilized a link-node type model developed by the South Florida Water Man- agement District, DYNTRAN, to study the effect of high fresh-water dis- charge into Indian River lagoon from the St. Lucie estuary. The model was calibrated for the period September 13-16, 1984 and further tested for the period March 10-14, 1983. On the basis of limited available data, the model 1Graduate Student, Ocean Engineering Department, Massachusetts Institute of Technology, Cambridge, MA 02139 148 FLORIDA SCIENTIST [Vol. 53 80°30" Mosquito Lagoon Haulover Canal Canaveral Barge Canal Banana River 28°00' St. Lucie X= River jem St. Lucie ‘\ Canal — “St. Lucie Inlet 27°00" Fic. 1. Location of area under investigation. No. 3, 1990] SHENG ET AL.— NUMERICAL MODELING 149 was shown to reasonably reproduce the measured water level when proper forcing conditions are applied at the three tidal inlets (Sebastian, Ft. Pierce, and St. Lucie). Despite its simplicity, however, this type of link-node model cannot be readily expanded into a multi-dimensional model. This paper presents a one-dimensional numerical model of estuarine cir- culation and salinity transport, which can be readily expanded into a multi- dimensional model. In the following, the formulation of a one-dimensional hydrodynamic/salinity model is described first. The model is then used to examine the pure barotropic tidal hydrodynamics in Indian River lagoon. In particular, model simulation of measured water level in September 13-16, 1984 is presented. The model is then used to examine the time scales of salin- ity transport in the lagoon due to pure tidal forcing and combined tidal and wind forcing. MopEL DEVELOPMENT AND FORMULATION—The starting point of the present hydrodynamic model is the three-dimensional time-dependent Na- vier-Stokes equations for incompressible fluid with hydrostatic pressure dis- tribution and Boussinesq approximation. Although three-dimensional models have been successfully developed to describe circulations driven by wind, tide, and density gradients in estuaries, lakes, and marine waters (e.g., Sheng et. al., 1978; Sheng, 1986 and 1987), it is not necessary to use three-dimen- sional models at all times. For example, for certain simpler geometry and bathymetry, it is possible to use a two-dimensional laterally-averaged model and even a one-dimensional laterally-and vertically-averaged model. In the absence of strong stratification, it may also be possible to use a two-dimen- sional vertically-averaged model. For these simpler models, equations and numerical solutions become significantly simpler than those for a three-di- mensional model. Due to its relatively simple geometry, the majority of Indian River lagoon can be modeled with a one-dimensional laterally- and vertically-averaged hydrodynamic model. The use of a one-dimensional model is also consistent with the fact that little data exists to allow the calibration and validation of a multi-dimensional model for Indian River lagoon. A One-Dimensional Model—The equations of motion for the free surface elevation, (n), vertically-and laterally-averaged velocity (u), salinity (S), and density (p) are: On | 10(Hub) _ bi Eh een (1) OHu 10(Huub) _ On gu|u| O*Hu gH’ dp pumurmmyree = 9 gee! grt AN 3 5, a, (2) dHS | 10HuSb_, DHS ; Aube we” ase * ahs (3) p = p(S) (4) 150 FLORIDA SCIENTIST [Vol. 53 where q is the inflow through lateral estuarine boundaries, H = n +h repre- sents the total depth while h is the mean depth, b is the mean width of the lagoon at a particular longitudinal location x, 1,,, is the wind stress along the longitudinal axis, A,, and D,, are the longitudinal turbulent diffusion coeffi- cients, p is the density, S, is the salinity at the lateral inflow boundaries, and c is the Chezy coefficient defined as: C= — HME (5) where n is the Manning’s n. Numerical Scheme—The above system of equation is dynamically coup- led, i.e., the flow field can affect the salinity distribution which can in turn affect the flow field. To solve the above equations, a fractional-step finite- difference method (Yanenko, 1971) is used. Basically, the first fractional step treats the momentum equation with only the advection term in a time-im- plicit manner (advection step), the second fractional step treats the momen- tum equation with only the diffusion term in a time-implicit manner (diffu- sion step), and the last fractional step treats both the continuity equation and the momentum equation with all the terms (propagation step): Advection Step: oti ue tus =0 (6) urts = urtsh” (7) Diffusion Step: Propagation Step: — lay = =0 (9) ynti _ pnt? - yO 4 gh Oty Meats , fH oo (10) At ot Ox p 2p, Or No. 3, 1990] SHENG ET AL.— NUMERICAL MODELING 151 where uw is the vertically averaged velocity, U is the vertically-integrated ve- locity, and the term uct , arising from the nonlinear inertial term, is gener- ally negligible compared to the other terms in Equation (10). During the propagation step, all terms associated with the gravity wave propagation are treated implicitly such that sufficiently large time steps can be used. After the continuity equation and the momentum equation are solved, the finite-differenced salinity equation is solved using the latest veloc- ity field in a time-implicit manner. Because of the use of implicit schemes and the absence of any time-step constraint, the numerical integration of the sys- tem of equations is very efficient. Details of the numerical schemes can be found elsewhere (Liu and Sheng, 1988) and are not presented here. SIMULATION OF BAROTROPIC TIDAL HypRODYNAMIcS—AS a first step, the one-dimensional model is used to examine the purely barotropic tidal hydro- dynamics in Indian River lagoon. In particular, a preliminary model simula- tion was performed to check out the accuracy and efficiency of the 1-D model for tidal circulation. For simplicity, a scenario during September 13-16, 1984 simulated by Williams (1985) was used. Instead of the channel-junction con- figurations setup by Williams (1985) which contained very abrupt changes in bathymetry and width, however, we used the more recent and complete ge- ometry/bathymetry data provided by Fred Morris to produce the width (b), depth (h), and grid size (Ax) needed for our study. SOBRSTIAN INLET ST. LUCIE INLET BOTTOM PROFILE HORIZONTAL SCALE: ‘’—2HUES__, VERTICAL SCALE: aloo MEAN WATER LEVEL Fic. 2a. Actual bathymetry and geometry of the Indian River lagoon to the south of Banana River. 152 FLORIDA SCIENTIST [Vol. 53 HAN_DVER CRNAL ——— N BOTTOM PROFILE HORIZONTAL SCALE: ‘—I02-BUES _, VERTICAL SCALE: StH _; MEAN WSTER LEVEL Fic. 28. Actual bathymetry and geometry of the Indian River lagoon to the north of Banana River. With a total of 77 grid points, the raw data as shown in Fig. 2(a) and 2(b) contain small scale oscillations which could deteriorate the numerical solu- tion of the finite differenced equations. The grid spacing Ax varies from ap- proximately 150 m to 6100 m and the depth ranges from 0.15 m to 0.24 m. The grid indices of the three tidal inlets are J=1 (St. Lucie), J=19 (Ft. Pierce), and I[=41 (Sebastian). The spatially smoothed data used for the ac- tual model simulation are shown in Fig. 3(a) and 3(b). No. 3, 1990] SHENG ET AL.—NUMERICAL MODELING Vays LONCITUOINAL SCALE: Pen GMAT ReGen WIOTH OISTORTEO 8Y FACTOR OF Y U(Ie*l) [J ax (1-1) ——}— a «11 —— Fic. 3a. Smoothed grid of the Indian River lagoon to the south of Banana River. LONCITUOINAL SCALE: p—_lOHiLfS WIOTH OLSTORTED Br FACTOR OF | U(I+l) \ = \ BU (I *1) S (1-1) S (t+) }— axu(t) § = —}—— aku t+ 1 Fic. 38. Smoothed grid of the Indian River lagoon to the north of Banana River. 154 FLORIDA SCIENTIST [Vol. 53 FORCING TIDE St. Lucie Inlet Ft. Plerce Inlet — —-— Sebastian Inlet TIDE HEIGHT (Ft. ABOVE MLW) 0.0 10.0 20.0 30.0 40.0 50.0 60.0 70.0 80.0 90.0 100.0 TIME (Hour) Fic. 4. Water level at three tidal inlets in Indian River lagoon starting September, 1984. As shown in Fig. 3, a staggered grid system is used with the water level and salinity computed at the cell centers and the velocity computed at the cell sides. For simplicity, freshwater inflows and the St. Lucie estuary are not considered in this preliminary study. Flow in the tidal inlets is resolved in this study by solving the same set of one-dimensional equations (1) through (5) subject to a tidal forcing at the ocean entrance (Figure 4). The 1-D lagoon- wide model and the 1-D inlet model are dynamically coupled to allow propa- gation of information between the lagoon and the 3 inlets throughout the simulation period. The results of our model simulation are compared with the data at 3 selected locations in Figures 5, 6, and 7 (West St. Lucie Inlet, Jensen Beach, and North Ft. Pierce Inlet, respectively). The simulated water elevations (light solid lines) agree quite well with data (dashed lines) and Williams’ model results (heavy solid lines). For simplicity, no density effect was in- cluded in this simulation. Figure 8 shows the tidal range along the longitudi- nal axis of the lagoon based on the model simulation on September 13-16, 1984. The results clearly indicate that, to the north of Sebastian Inlet, tidal mixing is negligible. No. 3, 1990] SHENG ET AL.—NUMERICAL MODELING 155 WEST ST. LUCIE INLET = 1.0 = ——Sjimulated (This Study, 1988) 2 —— Simulated (Willlams, 1985) O a < 5 - a6 © LW Te Ww a == 0:0 goms0 "60°. 9:0" 120 -15.0 180) 21.0, 24.0 27.0 30.0 TIME (Hour) Sept. 13 Sept. 14 Fic. 5. Computed and measured tides at West St. Lucie Inlet. JENSEN BEACH = 1.0 = = Ww 0.8 > O a < 0.6 L te 0.4 Ss —— Simulated (This Study, 1988) ul 52 —— Simulated (Willlams, 1985) WwW S255 [BG fa) F100 24.0 27.0 30.0 33.0 36.0 39.0 42.0 45.0 48.0 51.0 54.0 TIME (Hour) Sept. 14 Sept. 15 Fic. 6. Computed and measured tides at Jensen Beach. 156 FLORIDA SCIENTIST [Vol. 53 S 0 NORTH FT. PIERCE INLET al = —— Simulated (This Study, 1988) = — Simulated (Williams, 1985) O ---- Data jaa) < 5 a6 © Lu a8 Lu Q = 6.0 48.0 51.0 54.0 57.0 60.0 63.0 66.0 69.0 72.0 75.0 78.0 TIME (Hour) Sept. 15 Sept. 16 Fic. 7. Computed and measured tides at North Ft. Pierce Inlet. m) TIDE RANGE ( 0.00 20.0 40.0 60.0 80.0 100.0 120.0 140.0 160.0 180.0 200.0 DISTANCE FROM ST. LUCIE INLET (km) Fic. 8. Computed tidal range along the entire Indian River lagoon. No. 3, 1990] SHENG ET AL.— NUMERICAL MODELING | Veal SIMULATION OF SALINITY TRANSPORT—Insufficient salinity data exist in Indian River lagoon to warrant a realistic simulation of salinity transport over the entire lagoon. Instead, we choose to study the dynamic response of the lagoon to ocean salinity, i.e., time scales of salinity transport due to tidal forcing alone and combined tidal and wind forcing. The lagoon is assumed to be quiescent and of zero salinity initially. At the ocean end of the three tidal inlets, a simple harmonic M, tide of amplitude 30 cm and a salinity of 30 ppt are imposed as boundary conditions. In the first model simulation, we con- sider the case of no wind. In the second simulation, we consider the case of a uniform wind stress of 1 dyne/cm? from the south. In both cases, we neglect fresh water inflow and evaporation, which are not expected to have major effect on the time scales of salinity transport. The values of A,, and D,, are chosen to be 10 m’/sec, which appears to be reasonable for a minimum grid size of 150 m and a mean tidal current of 0.1 m/sec. The open boundary condition for salinity at the ocean end of the tidal inlet depends on the flow direction. During inflow, the salinity is set to the ocean salinity value. During outflow, however, the salinity at the open boundary is calculated from the flow velocity and salinity inside the inlet via: Co ORE is ot Or The model with the above boundary condition is then run for more than 125 tidal cycles. Results indicate that the salinity field satisfies global mass conservation at all times, i.e., the total amount of salinity in the lagoon at any time is equal to the total amount of salinity at a previous time plus the total incremental salinity fluxes through tidal inlets and tributaries. How- ever, as shown in Figure 9, it takes over 100 tidal cycles for the salinity field to reach a quasi steady state, i.e., when salinity changes less than a few percent from one tidal cycle to the next. Due to the weak tidal currents north of Sebastian Inlet, results at grid point 60 on the northern end of the lagoon show little salinity transport. The quasi-steady results after one hundred tidal cycles are shown in Figs. 10, 11, 12, and 13 at four 6-hour intervals within one tidal cycle. Currents on the order of 20 cm/sec exist in the southern and middle portions of the lagoon, while currents in the northern part of the lagoon are less than 5 cm/sec with little variation in the water level. At 1200 hours, when the ocean water level is zero as shown in Figs. 10(a) and (b), water level is relatively flat and convergent currents on the order of 10 cm/sec are found near the 3 tidal inlets. At 1203 hours, when the ocean water level is at the peak as shown in Figs. 11(a) and (b), significant surface slope is found near Ft. Pierce Inlet which causes significant divergent currents up to 40 cm/sec. On the other hand, currents in the vicinity of Sebastian Inlet are less than, 10 cm/sec con- sistent with the relatively mild surface slope there. At 1206 hours, as shown in Figs. 12(a) and (b), currents in most parts of the lagoon are almost opposite of 158 SALINITY SALINITY SALINITY 2s es 2s se) 50 sO FLORIDA SCIENTIST POSITION: GRIO POINT 10 78 100 TIOAL CTCLE POSITION: GAIO POINT 30 7S 100 TIOAL CYCLE POSITION: GRIO POINT 60 7S 100 TIOAL CYCLE 125 125 125 [Vol. 53 1S0 1S0 130 Fic. 9. Temporal evolution of salinity at three points (see Fig. 11 for positions) in Indian River lagoon during the longterm salinity simulation with forcing by tide only. No. 3, 1990] SHENG ET AL.— NUMERICAL MODELING TIDAL AMPLITUDE SEBRSTIAN INLET l2te HAS FT. PIERCE INLET ST. LUCIE INLET SUAFACE PROFILE BOTTOM PROFILE SALINITY PPT HORIZONTAL SCALE: —I2-SUES _, VERTICAL SCALE: pA EEL __4 MEAN WATER LEVEL 159 Fic. 10a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1200 hours after the initiation of the longterm salinity simulation with forcing by tide only. U.Q A TIDAL AMPLITUDE -30.0 CH HR HA 1290 $s lei2 Ss HALLSVER CANAL —__» N SURFACE PROFILE SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE: VERTICAL SCALE: rl MEAN WATER LEVEL Fic. 108. Same as Figure 10a, but north of Banana River. 160 FLORIDA SCIENTIST [Vol. 53 TLOAL AMPLITUOE SEBASTIAN INLET FT. PIERCE INLET 20 CM/S ST. LUCIE INLET SUAFACE PROFILE BOTTOM PROFIL : SALINITY PPT HORIZONTAL SCALE: -—t02-BULES VERTICAL SCALE: eo MEAN RATER LEVEL Fic. 11a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1203 hours after the initiation of the longterm salinity simulation with forcing by tide only. TIOAL AMPLITUDE 1212 HRS HAULOVEAR CANAL ——— > N SURFACE PROFILE SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE: VERTICAL SCALE: pa EL _, MEAN WATER LEVEL Fic. 118. Same as Figure lla, but north of Banana River. SHENG ET AL.—NUMERICAL MODELING 161 No. 3, 1990] TIDAL AMPLITUDE SEBASTIAN INLET -30.0 CH 1200 HRS 1212 HRS FT. PIERCE INLET —- N ——+ 20 ns SURFACE PROFILE ST. LUCIE INLET 10 20 30 uo 50 z]s}s}=|3 [>] [> ]=]2 [2 [12 ]3 [2 ]s l= /= [2 lll eels |3 (212 [21s s{xls/a{alo (sls iva = -lel-lejelelelelelel[@ jell leo [e jolele jeluiciejeisieisioi~iw~iel le jyeye;eleyeiwielte SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE: -—2-SLULES__, RISES VERTICAL SCALE: MEAN WATER LEVEL Fic. 12a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1206 hours after the initiation of the longterm salinity simulation with forcing by tide only. TIDAL AMPLITUDE HAULSVEA CANAL —> N SURFACE PAOFILE BOTTOM PROFILE HORiZCNTAL SCALE: VERTICAL SCALE: p—_LLEEL _y MEAN WATER LEVEL Fic. 128. Same as Figure 12a, but north of Banana River. 162 FLORIDA SCIENTIST [Vol. 53 TIOAL AMPLITUDE SEBASTIAN INLET V2i2 HRS - PIERCE INLET 20 CM/S ST. LUCIE INLET SURFACE PROFILE OTTOM PROFILE SALINITY PPT HORIZONTAL SCALE: -—0-BULES 4 VERTICAL SCALE: pA ffl _y MEAN WATER LEVEL Fic. 13a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1209 hours after the initiation of the longterm salinity simulation with forcing by tide only. TIOAL AMPLITUDE Vel2 HAS HAULOVEA CANAL nt N SURFACE PROFILE SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE: VERTICAL SCALE: Fic. 138. Same as Figure 13a, but north of Banana River. SALINITY SALINITY SALINITY . 3, 1990] 2S SHENG ET AL.— NUMERICAL MODELING 163 50 POSITION: GRIO 8 75 100 125 150 MUGRE ENCE POSITION: GRIO 28 25 50 7$ 100 125 150 TIORL CYCLE POSITION: GRIO 60 25 30 ie) 100 125 150 TIOAL CYCLE Fic. 14. Temporal evolution of salinity at three points (see Figure 10 for positions) in Indian River lagoon during the longterm salinity simulation with forcing by tide and wind. those at 1200 hours. However, this is not exactly true everywhere, due to nonlinearity and differing inlet dynamics. At 1209 hours, when the ocean tide is lowest, results in Figs. 13(a) and (b) show strong convergent currents near the tidal inlets, consistent with the significant surface slopes. 164 FLORIDA SCIENTIST [Vol. 53 aan TIDAL AMPLITUDE SEBASTIAN INLET -30.0 Cm 1200 HRS 1212 HRS —») N —— 20 CH/s ST. LUCIE INLET SURFACE PROFILE BOTTOM PROFILE SALINITY PPT HORIZONTAL SCALE; -—02SU£S VERTICAL SCALE: pel _y MEAN WATER LEVEL Fic. 15a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1200 hours after the initiation of the longterm salinity simulation with forcing by tide and wind. In addition to the forcing of tide and salinity at the ocean boundaries, a wind stress of 1 dyne/cm? from the south is imposed on the 1-D model grid. The results as shown in Fig. 14 indicate a significant reduction of the time for reaching quasi steady state to less than 25 tidal cycles. The currents, as shown in Figs. 15(a), 15(b), 16(a), and 16(b), are somewhat stronger than those without the wind. There is apparently some northward transport of salinity as shown in Fig. 14. However, the wind causes a significant set-up in the northern portion of the lagoon which in turn causes a southward flow to reduce the net northerly current. Comparing the results at 1200 hours as shown in Figs. 15(a) and (b) with those in Figs. 10(a) and (b), it is apparent that salinity in the southern portion of the lagoon is much higher and more uniform when the wind is present. As shown in Fig. 15, the 10 ppt salinity value is formed 15 miles further north than in the no-wind case. The wind stress of 1 dyne/cm’ used in this model simulation is somewhat brisk for this area. A more representative value is probably 0.5 dyne/cm’. Thus, it is expected that the time scale of salinity transport should be some- where between 25 and 100 tidal cycles. No. 3, 1990] SHENG ET AL.— NUMERICAL MODELING 165 TIDAL AMFLITUDE -30.0 CM l2le HAS HAILOVER CANAL ——! N —— 20 CK/S SURFACE PROFILE SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE: -—0-SULES 4 VERTICAL SCALE: - A-LEfl _, MEAN WATER LEVEL Fic. 158. Same as Figure 15a, but north of Banana River. SUMMARY AND Discussion—A one-dimensional model for describing the lagoon-wide circulation and salinity transport in the Indian River Lagoon has been developed. Model simulation of barotropic tidal circulation success- fully reproduced measured water level during a 3 day episode in September 1984. However, existing data are insufficient for calibration and validation of more detailed and multi-dimensional model simulations. Time scales of salinity transport have been studied by means of the one- dimensional model, and were found to be on the order of 25 to 100 tydal cycles. Lagoon-wide data of water level, current and salinity need to be col- lected over extended time periods (from months to years) to allow elucidation and modeling of the hydrodynamics and salinity transport. Multi-Dimensional Model—Smith (1987) recently analyzed water level data collected over a 25-year period of time in the lagoon. He found that tidal harmonic constants calculated for the interior of the central and southern segments can be interpreted in terms of two interacting waves from adjacent inlets, rather than as a single, phase lagged tidal wave. Thus he suggested the use of a two-dimensional (x-y) numerical model for a more thorough investi- gation. In another study, Smith (1989) found that the model resolution of vertical flow structure significantly affected the predicted longterm wind- driven circulation in the lagoon, and thus implied the need for a multi-layer 166 FLORIDA SCIENTIST [Vol. 53 TIDAL RMPLITUDE SEBASTIAN INLET -30.0 CH 1200 HAS tei2 HAS FT. PIERCE INLET —~> N —— 26 CH/S ST. LUCIE INLET SURFACE PROFILE BOTTOM PROFILE SALINITY PPT HORIZONTAL SCALE: - 1D Hues _ VERTICAL SCALE: ——a soo | MEAN WATER LEVEL Fic. 16a. Simulated horizontal velocity, surface elevation and salinity distribution in Indian River lagoon (to the south of Banana River) at 1206 hours after the initiation of the longterm salinity simulation with forcing by tide and wind. model. Morris (1987b personal communication) also suggested the use of a multi-dimensional model for the St. Lucie estuary. Moreover, while the tidal inlets have significant effects on the hydrodynamics of the lagoon, the one- dimensional model cannot adequately resolve the detailed hydrodynamic processes in the vicinity of the inlets. The one-dimensional model presented here can be expanded into two- dimensional and even three-dimensional models. Long-term flow and salin- ity data should be collected for validating these multi-dimensional models. Watershed Approach—Recently, Barile (1987) suggested a watershed ap- proach to formulate a sound management plan of the lagoon, thus implying the need of a rather detailed fine resolution model for a particular watershed which is to be coupled to a coarser resolution model for the rest of the lagoon. In order to drive a fine-resolution model for a particular watershed, however, it is necessary to provide the needed boundary conditions over various ex- tended time periods. One option is to collect extensive field data for the vari- ous watershed studies, which will undoubtedly be rather costly. Instead of that, however, it is possible to use a well-calibrated lagoon-wide model to provide the necessary boundary conditions for various fine-resolution water- shed models. No. 3, 1990] SHENG ET AL.—NUMERICAL MODELING 167 TIDAL AMPLITUDE 1212 HRS HAULOVER CANAL —» iN —— 20 CH/S SURFACE PROFILE SALINITY PPT BOTTOM PROFILE HORIZONTAL SCALE; -—2BUES ___; VERTICAL SCALE: p—_-LEEL _, MEAN WATER LEVEL Fic. 168. Same as Figure 16a, but north of Banana River. The one-dimensional lagoon-wide model presented is a first step towards the watershed approach to understand and manage Indian River lagoon. ACKNOWLEDGMENT— We would like to thank Fred Morris for providing the bathymetry data, Ned Smith for providing the tidal data, and Diane Barile, Fred Morris, and Ned Smith for useful discussions. 168 FLORIDA SCIENTIST [Vol. 53 LITERATURE CITED Bari.e, D. D. 1984. The Indian River lagoon initiative. In: Proc. Indian River Resources Sympo- sium, Marine Resources Council of East Central Florida, Melbourne, Florida. . 1987. Managing cumulative effects in Florida wetlands. In: Proc. Indian River Re- sources Symposium, Marine Resources Council of East Central Florida, Melbourne, Flor- ida. DomsrowskI, M., F. Morris AND R. REICHARD. 1987. Hydrodynamics. Pp. 3-1 through 3-27, In: Indian River Lagoon Joint Reconnaissance Rept. (J. Steward and J. VanArmen, eds.). St. Johns River Water Management Dist. and South Florida Water Management Dist. Liv, Y. M. ANDY. P. SHENG. 1988. A two-dimensional finite-difference model for moving bound- ary hydrodynamic processes. UF/COE Rept., Coastal and Oceanogr. Eng. Dept., Univ. Florida. Morris, F. W. 1985. The St. Lucie Estuary model. In: Proc. Fifth St. Lucie Estuary Coordinating Conf., Florida Oceanogr. Society, Stuart, Florida. . 1987a. Modeling of hydrodynamics and salinity in the St. Lucie Estuary. Tech. Publ. 87-1, South Florida Water Management Dist., West Palm Beach, Florida. _________. 1987b. Personal communication. SHENG, Y. P. 1986. Finite-difference models for hydrodynamics of lakes and shallow seas. Pp. 146- 228, In: Physics-based modeling of lakes, reservoirs and impoundments. (W. Gray, ed.) American Soc. Civil Engr., New York. . 1987. On modeling three-dimensional estuarine and marine hydrodynamics. Pp. 35- 54, In: Three-dimensional Models of Marine and Estuarine Dynamics (N1HOUL, J. C. J., AND B. JAMAkT, eds.). Elsevier, New York. SHENG, Y. P. AND W. Lick. 1978. Numerical computation of three-dimensional circulation in Lake Erie: a comparison of a free-surface model and a rigid lid model. J. Phys. Oceanogr. 8:713-727. SmiTH, N. P. 1987. An introduction to the tides of Florida’s Indian River lagoon. 1. Water levels. Florida Scient. 50:49-61. . 1989. Computer simulation of wind-driven currents in a coastal lagoon. Pp. 113-131, In: Estuarine Circulation (NEILSoN, B. J., J. BRUBAKER AND A. Kuo, eds.). Humana Press, Clifton, N.J. WituraMs, J. L. 1985. Computer simulation of the hydrodynamics of the Indian River lagoon. M.S. Thesis, Florida Institute of Technology, Melbourne, 145 pp. YANENKO, N. N. 1971. The method of fractional steps. Springer-Verlag. New York. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3):147-168. 1990. Accepted: June 6, 1989. Engineering Sciences GROUNDWATER SEEPAGE INTO THE INDIAN RIVER LAGOON AT PORT ST. LUCIE ASHOK PANDIT AND CLovis CLovis EL-KHAZEN Department of Civil Engineering, Florida Institute of Technology, Melbourne, FL 32901 Asstract: A finite element program, GROSEEP, was developed to estimate the groundwater seepage from the surficial aquifer into Indian River lagoon along a selected cross-section in Port St. Lucie, Florida. The model is a general purpose model and can be used at other cross-sections. Model results using a single layer indicate that groundwater seepage could be an important freshwater source into the lagoon. Sensitivity analysis using a three-layer model predicted that among other parameters the presence of windows in the clay layers could significantly affect seepage rates into the lagoon. ESTUARIES are areas where sea water is measurably diluted with fresh- water from land drainage. One of the most important characteristics of an estuary is its effectiveness as a habitat for marine species. Since species toler- ances for salinity vary, it is possible for salinity to be too high or too low for specific species at certain times of the year. Gunter, Ballard, and Venkatara- maiah (1973) found that dilution of salt water by excess fresh water or vice versa can have a definite and sometimes drastic effect on the flora and fauna of the affected area. The Indian River lagoon, a semi-enclosed area where freshwater mixes with saltwater, can be defined as an estuary or as a lagoon. It extends 155 miles from Volusia to Palm Beach County and is approximately 227 square miles in area. The width and depth of the lagoon vary between 0.5 to five miles and three to ten feet, respectively. Circulation and flushing of Indian River lagoon are greatly influenced by freshwater inflows. Princi- pal freshwater sources are natural streams, rainfall, direct land runoff, a number of wastewater treatment plants and groundwater. Glatzel (1986) estimated annual values for inputs due to precipitation, controlled discharge (through canals), anthropogenic flows, and non-point runoff to be 32.0, 49.0, 26.0 and 13.0 billions of cubic feet, respectively. Glatzel also noted that groundwater input from the surficial aquifer could not be estimated because of lack of sufficient data. South Florida Water Man- agement District (SEWMD, 1987) concluded that only groundwater could account for a relatively constant, background flow of freshwater into the St. Lucie Estuary. Annual seepage was estimated to be between 60 and 600 ft’ per foot of shoreline. Gu, Iricanin and Trefry (1987), reported that in samples taken from Eau Gallie Harbor interstitial water chlorinity decreased linearly from 10700 ppm to 3500 ppm over a length of a core approximately 11.8 inches long. This decrease suggested that groundwater may be seeping through the sediments. The results of these studies, indicating that ground- water flow could be a major freshwater input into the lagoon, lead to the present research study. 170 FLORIDA SCIENTIST [Vol. 53 Indian River lagoon has two watersheds: a natural watershed formed by natural topographic highs on both sides of the lagoon, and a more extensive watershed drained by several canals that link the mainland and the barrier islands to the lagoon. The boundaries of the natural watershed are generally considered to be the Atlantic Coastal Ridge on the mainland and the high ridge in the barrier islands east of the lagoon. Since water table elevations usually conform to the natural topography, the groundwater divide in the surficial aquifer is considered to be along these ridge lines. The direction of regional groundwater flow in the surficial aquifer, within the confines of the groundwater divides on either side of the lagoon, is perpendicular to the lagoon (Toth, 1987). Thus, the surficial aquifer can provide a continuous flow of groundwater into the lagoon and act as a non-point source. METHops—Groundwater seepages can be estimated by solving partial differential equations that describe the conservation of fluid mass during flow through a porous medium. A detailed description of these equations is provided by Pinder and Gray (1977). The equations are solved by using numerical methods such as the finite difference or the finite element methods. Eyre (1985) simulated the flow of groundwater and the effects of future groundwater development by using a two-dimensional finite element model. Attia and co-workers (1986) used a two-dimensional finite element model to simulate groundwater flows in the aquifer underlying the Nile Valley of Egypt. A general purpose Galerkin two-dimensional finite element model developed by Pandit (1982) was also used to predict steady and unsteady groundwater flow rates below an impervious dam. The results from this model were quite close to analytical results. In this study, a finite element model GROSEEP (Groundwater Seepage) was developed to quantify the non-point groundwater flow into the lagoon. GROSEEFP is a modified version of the model developed by Pandit (1982). DESCRIPTION OF SELECTED SITE AND LirHoLoGy— The subsurface system in central coastal Florida consists of a surficial aquifer and the confined Flori- dan Aquifer, separated by the relatively impermeable Hawthorn Formation. Groundwater levels in the surficial aquifer, along the groundwater flow di- rection, are required to operate the finite element model, GROSEEP. A sec- tion on the southern end of St. Lucie County was selected for this study after assessment of all available data (Fig. 1). This section was selected mainly because of the existence of five wells, STL 173, STL 174, STL 175, STL 176, and STL 177, which were located along the direction of regional ground- water flow. Three additional two inch diameter PVC pipe shallow monitor- ing wells, IRL 1 (Indian River Lagoon 1) through IRL 3, were installed by the authors on the barrier island to determine the location of the water-table- divide on the east side of the lagoon. Well IRL 4 was installed on the main- land to pinpoint the location of the water table on the west side of the lagoon. An added advantage in using this cross-section was that groundwater eleva- tion in wells STL 175 and STL 176 had been measured by a water stage recorder from March 1975 to October 1978 by the USGS and these data were available for analysis. The average monthly groundwater elevations in STL 176 for 1975 and 1976 are shown in Table 1. aol PANDIT AND EL-KHAZEN—GROUNDWATER SEEPAGE No. 3, 1990] IUI[AIOYS UOOBP] JO 00} J9d5 dUT[ALOYs UOOSE] JO 00} 19d 1aAe] [[aYs pue pues oy} Jo APWANONpuOd orpneIpAY [VoVI0A puke [eyUOzTIOY ay} ae FAY ‘EHH Jake] ABO ayy Jo AWANONPUOO oT[NeIpAY [eOI7I9A pur [e}UOZIIOY Vy} ae ZA Y ‘SHY JaAR] pues ay} Jo AYATONPUOO oTNeIpAY [VONJIOA pue [eUOZIIOY 9y} Be TAY ‘TH Ajaanoedsai ‘s1aAe] [[PYS pue pues pure ‘Aeyo ‘pues ay} Jo Sassauyxory} oy} o1e Ep puke ‘Zp ‘Tpe 9 13'0 G00'0 00°0S GIP G00'0 00°01 00°ST GI EI 13'0 G0'0 00°0S SI? S000 00°01 00°ST tI 13 130 G0'0 00°0S GIP G00'0 00°0S 00°ST EI I 98°0 GO'0 O1'0 GI'F1 G0'0 00°01 00°6 eal 8 98°0 GO'0 O10 GI'P G0'0 00°0S 00°6 Il 6 83'0 G0'0 00°01 a ial G0'0 00°01 00°6 OI ZI 83'0 G0'0 00°01 co G0'0 00°0S 00'6 6 6 CHO G0'0 00'°0S co al GO'0 00°01 00°6 8 gI 13'0 G0'0 00°0S i GO'0 00°0S 00°6 i Ol 98°0 GO'0 01'0 GIF GO'0 00°01 00°ST 9 rI 98°0 G0'0 O10 GIF GO'0 00°0S 00°ST G EI 83'0 GO'0 00°01 GI'FI G0'0 00°01 00°ST 7 03 830 G0'0 00°01 GI G0'0 00°0S 00°ST g ET 13'0 G0'0 00°0S al G0'0 00°0S 00°ST 6 13 13'0 G0'0 00°0S co G0'0 00'°0S 00°ST I (0'93-Aep/g"y) (Aep/"34) (Aep/"3y) (Aep/"3) (Aep/"3) (Aep/"3y) (Aep/"3y) (4) "ON adedaag “CAM qAM alAM aSHX aH alH TY asery) 0=SS ‘44 0°03T =eEP ‘4 0°01 =20P “YH 0'0E =eIP “YH S'0=ZY 10J sonfea asedaag °Z ATAV]L, 9 ¢T 6 Cl 9CI Lot LOt UIT SIT 9 TT esl 6cI 6 él eel LL6I € 1 Sel vel IGT 611 Vol Col cl vil eit S00 8 TT 9L61 ‘09q ‘AON ‘po "ydas ‘sny Ajn{ oun{ Aeyw idy ‘ey xe Xs | ‘ue Ivak (ISW 2eA0qe) ("3J) SUOTBAVTS 19}B2MpUNOIS 9) T TLS Ul UOBAV]S I9}eMpuUNOIS A[YUOUT IBVIOAY *[ ATAV], 172 FLORIDA SCIENTIST [Vol. 53 N INDIAN RIVER LAGOON BANANA RIVER CAPE CANAVERAL ATLANTIC \N OCEAN s“ SEBASTIAN INLET BREVARD COUNTY INDIAN RIVER & COUNTY INDIAN RIVER LAGOON PG 4 PG 1 PG12 ~ OFT. PIERCE INLET ST. LUCIE STL 173,174,1735,176,177 COUNTY lib IRL 1,2,3 ANS SELECTED \ [CROSS-SECTION M 1031 .1030 MARTIN , af EN) ST. LUCIE INLET 1047,1058 yess = COUNTY WW 43-51320 M 1053,1052 Fic. 1. Location and nomenclature of shallow wells at selected cross-section and other shal- low wells along Indian River lagoon, Florida. No. 3, 1990] PANDIT AND EL-KHAZEN— GROUNDWATER SEEPAGE 173 N eee. | eee 2400 ft. 600 h.->| STL 176 40 |) 400 \160 STL 175 ft Ler tt. — IRL 4 ee ieee: IRL3 OCEAN STL 177 2 POPE EEE EE EEE A A A A ia a ee ‘ NONNN NN NNN NNN NNN NN SN SN NS ‘ fof tie ‘ ¢ See aR RRR RR SERRE SS 7 ON ¢ é Ss CP o Caer, ¢ ae ae YS SAND & SHELL a oN .ON é . . NUN NENIEN ba Tees bey Tee ¢ CCEA) CAMEL CALA x x x x x x ‘ x x x x x x, N x AY s x x ‘ ‘x x x x iS ~~ x an en Pee AA ee ¢ ¢ ¢ ¢ ¢ ¢ ¢ ¢ ¢ ¢ 7 4 ¢ ¢ ta ¢ ¢ a ¢ 7 ts ? 7 é ¢ é ? HAWTHORN e aS % Fic. 2. Description of idealized cross-section and well location. The natural watershed on the west side of the lagoon is very narrow in the Port St. Lucie area due to the presence of some local hills up to 40 feet high (Fig. 2). Groundwater elevation measurements indicated that although the topographic high exists at the location of well STL 177 the groundwater divide exists at the location of well STL 176. Based on the lithology at the location of STL 175, STL 176, STL 177, reported by Miller (1979) and the lithology at the new wells, IRL 1, IRL 2, IRL 3, and IRL 4 installed by the authors, the hydrogeologic cross-section was idealized for modeling purposes (Fig. 2). The surficial aquifer has a medium to fine grained sand layer, a clay layer, and a sand and shell layer. The medium to fine grained sand layer extends to a depth of approximately 20 to 30 feet from the MSL (mean sea level). It usually consists of some combination of fine-to-medium tan sand, fine-to-medium dark brown sand, medium-grained dark brown sand, gray silty sand, and yellowish or orange silty sand. The lithologic section of STL 175 reported by Miller (1979) shows the presence of a clay layer at a depth of 30 feet to 37 feet from the MSL. Discussion with local drillers indicated that the thickness of the clay layer is usually within two to ten feet and varies considerably within a short distance. The fact that the lithologic description of STL 177 (Miller 1979) does not report the presence of a clay layer supports this observation. It is possible that the clay layer was too thin to be located by split spoon samples at this location. The sand and shell layer has a thickness of about 100 feet to 120 feet and connects the clay layer to the Hawthorn For- mation. The percentage of shells in the sand and shell layer increases from about 75 percent to 90 percent with increasing depth. The distance between STL 175 and STL 176 is 13 feet. 174 FLORIDA SCIENTIST [Vol. 53 / N 2400 ft: 600 el WATER TABLE, h=f (x) Pfr eee M.S.L. ay CONSTANT HEAD BOUNDARY h=160 ft. 160 ft. 1400 ft: WATER TABLE, h =f, (x) 7 > WATER DIVIDE WATER DIVIDE IMPERMEABLE BOUNDARY Fic. 3. Region size and boundary conditions assuming water-table-divide in the barrier is- land. RESULTS AND Discussilon— The partial differential equation that describes unsteady groundwater flow in a homogeneous but anisotropic medium is: K,,(07h/dx’) + K_,(0°h/0z?) = S. dh/odt (1) in which K,,, K,, (L/T) are the hydraulic conductivities of the aquifer in the x and z directions, respectively; h(L) is the freshwater hydraulic head; S,(L") is the specific storage of the aquifer; t is time; and x, z are the spatial coordi- nates. There are two ways to idealize the cross-section shown in Fig. 2. One way would be to bound the cross-section between the water table divides at the mainland and the barrier island. Groundwater measurements indicated that the water-table divide in the barrier island is approximately at the location of well IRL 3. The boundary conditions for the idealized cross-section between the water table divides (Fig. 3) are based on the following assumptions: 1) The shallow aquifer extends up to the Hawthorn Formation which is 160 feet below MSL. 2) A water divide exists at the location of well STL 176 in the mainland and at the location of well IRL 3 in the barrier island. 3) The Hawthorn Formation is impermeable. 4) The water level in Indian River lagoon is at mean sea level. 5) The lagoon has an average uniform depth of five feet at the selected cross section. 6) The density of the lagoon water is the same as the density of the freshwater. The boundary conditions for this ideal- ized cross-section can be mathematically stated as follows: No. 3, 1990] PANDIT AND EL-KHAZEN— GROUNDWATER SEEPAGE MS dh/dx(0, z, t) =0 h(0,160,t) =h, + 160 h(x, 160, t) =f,(x) + 160 (O h(1400, z, t) = 160.0 (15 h(x, 155, t) = 160.0 (14 400 - 40 ppt), temperature (>35°C.) or turbidity (>29 NTU’s over background). VEGETATION. The twice yearly over-flights were conducted once during the flooded season and once during the dry, open period. While not required in the operating permit, ground vegetation observations have been a routine part of this work as well. a 1.2 ha (3 ac) stand of Batis/Salicornia was staked at the perimeter of its coverage on May 30, 1986. In addition, four transect lines with five points per line were marked and photographs were taken at each point in May 1987, December 1987, and May 1988. Prior to impounding, high marshes were dominated by Batis/Salicornia beds with mixed black and white mangrove (Laguncularia racemosa). These transects have shown a net increase in vegetative cover, in particular by Batis, Salicornia, and black mangrove. When the stand was visited in May 1987, 200 FLORIDA SCIENTIST [Vol. 53 ae DISSOLVED OXYGEN (ppm) a IMP FLOOR ar PER DITCH © RIVER Fic. 4. Dissolved oxygen quarterly means for Indian River lagoon, perimeter ditch and im- poundment floor stations. one year after the initial staking, it had already spread up to 30 meters in some directions. The semi-annual transect pictures show there was die back in December after the flooding season but regrowth by May after the dry- down period. Adjacent to this staked stand, several hundred black mangroves established themselves and are now approximately one meter high. The installation of the permanent pump at Impoundment #6 has allowed IRMCD to maintain a consistently lower flooding elevation, thereby encour- aging high marsh vegetation growth. Mosquito Production: Rainfall (2.5 in.) during the week of May 11 re- sulted in a brood of larvae across the marsh surface that was treated by aerial larviciding with Altosand (an Altosid/sand mixture). No larval production was observed during the pump-up and therefore no additional larviciding was required. No aerial larviciding of the impoundment was required during the closed, management period. Ground ULV adulticiding was performed near the impoundment 11 times during the first year due to mosquito produc- tion from nearby, un-impounded marshes. Current Status: Based on results from the first year of water quality moni- toring and management, IRMCD requested and received two permit modifi- cations from the DER. One request was to change the fall culvert openings from a fixed date to a variable date based on water level equilibrium between the lagoon and impoundment. This opening procedure should help eliminate fish kills, as experienced in the first year due to unequal water levels. The second modification allowed a reduction in sampling stations, eliminating those that showed similar trends to nearby stations. No. 3, 1990] O BRYAN ET AL.—SALT MARSH MITIGATION 201 Discussion— After over two years of negotiating to obtain necessary per- mits, a management plan was devised that biological sampling indicates has been successful in improving wildlife usage of a formerly isolated wetland. Fish and macrocrustacean sampling, while limited, indicated increased spe- cies utilization of the marsh after re-connection to the estuary. No obvious negative effects on avian populations were observed during the dredging, filling and management activities. These results are in agreement with the exhaustive research on fish and avian usage of re-integrated impoundments conducted elsewhere in the region. This similarity in results should speed acceptance of future RIM plans as mitigation. Observations also showed that historical high marsh vegetation responded well to the carefully controlled flooding regime. The survival and enhance- ment of Batis/Salicornia beds and black mangroves, in lieu of red mangroves, should be a management goal in restoring impacted marshes back to a more natural state. The DER operating permit, while providing firm water management guidelines, is flexible enough to incorporate new scientific findings and man- agement experience. The water quality sampling required by the operating permit provides marsh managers with the data to monitor long term im- provement and/or deterioration. The willingness of DER to incorporate these findings into permit revisions is commendable and demonstrates how far the permitting process has progressed. Periodic fish kills due to hypoxic conditions and/or hydrogen sulfide build-up is still a problem during the closed and critical reopening periods. Management experience has led to modifications in the opening schedule to help alleviate this problem. In addition, current research is focusing on new techniques of aeration, over-pumping and bottom water releases. As more research and management experience becomes available, permit modifica- tions will be required to further fine tune the management of these re-inte- grated wetlands. Conc Lusions— This cooperative project proved to be beneficial to the private land owner, the local mosquito control district and natural resource interests. These benefits are: 1) Seacrest Estates obtained valuable single-family waterfront lots that otherwise would not have been developable. 2) The mosquito control district received better marsh management tools in the form of water control structures and a permanent pump station at no cost to local taxpayers. 3) Fish and macrocrustacean access to the impoundment was improved with the seasonal connection of this formerly isolated wetland to the estuary. Currently there is no incentive for a property owner to improve or en- hance impacted wetlands for altruistic reasons. Since most regional im- pounded wetlands are privately owned, mitigation projects such as this that can positively enhance salt marsh management need to be considered. While 202 FLORIDA SCIENTIST [Vol. 53 some agencies and individuals can justly argue there should be no additional filling of wetlands whatsoever, this project had a ratio of re-integrated marsh to filled wetlands of approximately 64 to 1. Though there have been some negative aspects of RIM, in most locations it is still considered to be the best current impoundment management option to open closed impoundments. RIM will no doubt be an important consideration of any diversified manage- ment plan for Indian River lagoon. ACKNOWLEDGMENTS— The authors wish to acknowledge that funding for the biological sam- pling was provided by Seacrest Estates, Inc. The assistance of Mr. J. Mason and Mr. T. Smoyer with photography and darkroom work is gratefully acknowledged. The authors appreciate the review comments of Mr. E. J. Beidler and Mr. G. Dodd, Mr. G. Dodd’s graphics work. This constitutes contribution number 763 to the Harbor Branch Oceanographic Institution, Inc. LITERATURE CITED Caruson, D. B. anno J. D. Carrot, Jr. 1985. Developing and implementing impoundment management methods benefitting mosquito control, fish and wildlife: a two year progress report about the Technical Subcommittee on Mosquito Impoundments. J. of Fl. Anti- Mosquito Ass. 56:24-32. Caruson, D. B., R. G. GitmMore, AND J. R. Rey. 1985. Salt marsh impoundment management on Florida’s central east coast: reintegrating isolated high marshes to the estuary. Pp. 47-63, In: Proc. 12th Annual Conf. on Wetlands Restoration and Creation. Carson, D. B., anv P. D. O’Bryan. 1988. Mosquito production in a rotationally managed impoundment compared to other management techniques. J. Am. Mosquito Control Ass. 4:146-151. CLEMENTS, B. W. AND A. J. Rocers. 1964. Studies of impounding for the control of salt-marsh mosquitoes in Florida, 1958-1963. Mosquito News 24:265-276. Gitmore, R. G. 1984. Fish and macrocrustacean population dynamics in a tidally influenced impounded subtropical salt marsh. Final Report: Florida Dep. Environmental Regula- tion—CZM 43:35 pp. . 1987. Fish, macrocrustacean and avian population dynamics and cohabitation in tidally influenced impounded subtropical wetlands. Pp 373-394. In: WHITMAN, W. R. AND W. H. MenrepiTH (eds.) Proceedings of a symposium on waterfowl and wetlands management in the coastal zone of the Atlantic flyway. Del. Dept. Nat. Res. and Envir. Control, Dover, Delaware. , D. W. Cooke AnD C. J. DoNoHoE. 1982. A comparison of the fish populations and habitat in open and closed salt-marsh impoundments in east-central Florida. Northeast Gulf Sci. 5:25-37. , D. J. Perens, J. L. Fyre anp P. D. O’Bryan, 1986. Fish, macrocrustacean and avian population dynamics in a tidally influenced impounded subtropical salt marsh. Final Rept. FDER—CZM 73,93. 25 pp, 17 Tbls., 29 Figs., Appendix. Haecer, J. S. 1960. Behavior preceding migration in the salt-marsh mosquito Aedes taeniorhy- nchus (Wiedemann). Mosquito News 20:136-147. HarrinctTon, R. W. AND E. S. HarrincTon. 1961. Food selection among fishes invading a high subtropical salt marsh: from onset of flooding through the progress of a mosquito brood. Ecology 42(4):646-666. . 1982. Effects on fishes and their forage organisms of impounding a Florida salt marsh to prevent breeding by salt marsh mosquitoes. Bull. Mar. Sci. 32(2):523-531. KusHLan, J. A. 1973. White Ibis nesting in the Florida Everglades. Wilson Bull., 85:230-231. Provost, M. W. 1967. Managing impounded salt marsh for mosquito control and estuarine re- source conservation. LSU Marsh and Estuary Symposium. pp. 163-171. . 1976. Tidal datum planes circumscribing salt marshes. Bull. Mar. Sci. 26:558-563. . 1977. Source reduction in salt-marsh mosquito control: Past and future. Mosquito News 37:689-698. No. 3, 1990] O BRYAN ET AL.—SALT MARSH MITIGATION 203 Rey, J. R., R. A. Crossman, T. R. Kain, ann D. S. Taytor. 1985. An overview of impounded mangrove forests along a subtropical lagoon in east-central Florida, U.S.A. The Man- groves: Proc. Nat. Symp. Biol. Util. Cons. Mangroves, Nov. 1985:341-350. Wane, R. A. 1962. The biology of the tarpon, Megalops atlanticus, and the ox-eye, Megalops cyprinoides, with emphasis on larval development. Bull. Mar. Sci. Gulf and Caribbean 12:545-622. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3): 189-203. 1990. Accepted: June 2, 1989. Atmospheric and Oceanographic Sciences HOLOCENE EVOLUTION OF INDIAN RIVER LAGOON IN CENTRAL BREVARD COUNTY, FLORIDA SHARON F. BADER AND RANDALL W. PARKINSON Department of Oceanography and Ocean Engineering, Florida Institute of Technology, Melbourne FL 32901 Asstract— This study examined the coastal response of central Brevard County to rising Holocene sea level as it has been recorded in sediments of Indian River lagoon. The typical Holocene sediment sequence, in ascending order, consists of: (1) depauperate marsh muds, (2) sandy muds and muddy sands that contain a restricted marine fauna, and (3) sands and muddy sands that contain an abundant, more normal marine fauna. This transgressive sequence devel- oped as rising Holocene sea level flooded a Pleistocene topographic depression (paleolagoon). It is often capped by (4) a coarser sand that exhibits a decrease in faunal abundance and diversity. The origin of this sediment type remains problematic. “Muck”, a fine-grained, organic-rich sedi- ment, occurs in bathymetric depressions of the lagoon and was not identified at any other strati- graphic position. This observation supports the hypothesis that “muck” is an anthropogenic sedi- ment type. Stratigraphic cross-sections along the axis of the lagoon reveal a concomitant increase in mud and decrease in sand from south to north. This sediment gradient is logically related to the morphology of the present barrier island system. In the northern area, the relatively broad width of the barrier island and its well-developed ridge-and-swale topography may have limited the flux of quartz sand into the lagoon during major storms and hurricanes. On the broadest scale, dramatic sea level fluctuations during the last 140,000 yr are primarily responsible for the vertical succession of sediments deposited in late Pleistocene and Holocene coastal environments (Kraft and Chrzastowski, 1985). These deposits and their associated morphology are currently being modified to varying degrees by the present Holocene sea-level rise. The nearly 250-km chain of barrier islands that rim the east coast of Florida was sculpted by these same processes. Although the large-scale dy- namics that produced Florida’s Atlantic barrier island system have been ex- amined in detail (Osmond et al., 1970; White, 1970; Kofoed, 1963; Brown et al., 1962), very little is known about the geologic history of its back-barrier lagoons (c.f., Almasi, 1983). This study is the first to document the Holocene evolution of Indian River lagoon in central Brevard County (Fig. 1). DESCRIPTION AND GEOLOGIC History oF Srupy AREA— The study area con- sists of a shallow, microtidal back-barrier lagoon (Indian River) that lies along central Florida’s Atlantic coast between the Eau Gallie River and Tur- key Creek (Fig. 1). Cuspate spits lie along the seaward margin of the lagoon in this area and define it as a unique geomorphologic unit. The well-defined symmetry of the spits suggests that the back-barrier lagoon system has re- mained relatively stable over time (Zenkovitch, 1958). The lagoon deepens from south to north, requiring the use of 12-ft (3.6-m) bathymetric contour lines north of the study area, but only spotty use of a 6-ft (1.8-m) contour south of the study area (USC&GS, 1980). Within the study area, the barrier island displays a well-developed, linear, beach-ridge-and-swale topography No. 3, 1990] BADER AND PARKINSON— HOLOCENE EVOLUTION 205 Atlantic Coastal Ridge (after Brown et al., 1962. FE] Cuspate Spits [KJ Overwash/Flood Tidal Delta ™ Radiocarbon Dates (Osmond et al., 1970) oN St. Johns River / Fic. 1. Regional map of Central Brevard County, Florida, illustrating the location of major geomorphic features and location of radiocarbon dates (modified from Osmond et al. 1970). Indian River lagoon study area lies between Eau Gallie River and Turkey Creek. Atlantic Ocean 28°00 that parallels the strike of the island (USC&GS, 1980). Just south of 28°N latitude (Fig. 1), the core of the barrier island narrows, and both the ridge- and-swale topography and cuspate spit morphology give way to a flood tidal delta/washover fan geomorphology. Many studies of barrier island and lagoonal sedimentation along the At- lantic seaboard emphasize that landforms created during Pleistocene high sea stands influenced the later deposition of Holocene coastal sediments (Davis, 206 FLORIDA SCIENTIST [Vol. 53 1985; Evans et al., 1985; Halsey, 1979; Moslow and Heron, 1979; Kraft et al., 1979). Similarly, relict Pleistocene landforms in central Brevard County have influenced the development of Holocene depositional sequences within In- dian River lagoon. Lying about 9.0 m above present sea level, the City of Melbourne strad- dles the highest elevations of the Atlantic Coastal Ridge (Fig. 1), a coast- parallel feature that developed about 140,000 to 120,000 yr B.P., when (Pamlico) sea level reached heights of 7.5 m to 9.0 m above present level (White, 1970). Merritt Island (Fig. 1) is closely age-related since its southern tip has a radiocarbon age of about 110,000 yr B.P. (Osmond et al., 1970). The next major transgression (Silver Bluff) occurred about 30,000 yr B.P. and brought sea level close to its present-day height, with a maximum not exceeding 2.5 m above present sea level (DuBar, 1974). Radiocarbon dates (Osmond et al., 1970) suggest that topographically lower portions of the At- lantic Coastal Ridge may have been inundated by the Silver Bluff highstand. Other late Pleistocene sediments, including the Anastasia Formation, were deposited to the east of the Atlantic Coastal Ridge and underlie the present- day barrier island (Mehta et al., 1976). Some of these deposits appear to have had sufficient elevation above sea level during Silver Bluff time to generate and protect a back-barrier lagoon, since paleolagoonal sediments, dated at 31,000 yr B.P., lie about 2 m below the sediment-water interface of the present-day lagoon at the southern end of Brevard County (Almasi, 1983). Following regression of the Silver Bluff sea to approximately 100 m below present level, the Holocene transgression began (Kraft and Chrzastowski, 1985). Sometime between 5,000 and 6,000 yr B.P., marine waters re-entered the paleolagoon (Almasi, 1983), and lagoonal sedimentation began anew. Contemporaneously, sand and shell material began to accumulate upon the Pleistocene paleotopographic high that now underlies the present barrier is- land (Mehta et al., 1976; Osmond et al., 1970). This study focused on the sediments and sedimentary sequences that accumulated within the back-bar- rier lagoon during the late Holocene sea-level rise. MATERIALS AND MErHops— Using equipment and procedures described by Lanesky and co- workers (1979), 16 vibracores were collected in the study area (Fig. 2). Six of these cores were part of Project “MUCK” (see Trefry et al. 1987). Ten other cores were taken within an 8.0-km long section of the lagoon between Eau Gallie River and Turkey Creek (Fig. 2). Vibracore posi- tions were located using triangulation of fixed reference points. Split cores were described with respect to color, sedimentary structures, mottling, and sedi- ment type. Sediment samples were taken at 33-cm intervals or from lithologic units that would otherwise have been unsampled. The samples were wet-sieved through 2.0-mm and 0.0625-mm screens to separate the gravel-, sand-, and fine-sized fractions. Gravel- and sand-sized fractions were oven dried and weighed. The sand fraction was dry sieved at 0.5-phi intervals to determine the modal sand size for each sample. The fine-grained (mud) portion of each sample was dispersed in a 4% solution of sodium hexametaphosphate in preparation for pipette analysis (Folk, 1974) to determine the relative proportions of silt and clay. The relative dry-weight percentages of gravel, sand, and mud were used to classify each sediment sample as one of four general sediment types (Fig. 3): (1) sand; (2) muddy sand; (3) sandy mud; (4)mud (Folk, 1974). Microscope slides of each mud sample were prepared and examined under a binocular microscope to check for framboidal pyrite, a common replacement mineral of rootlets in a reducing environment (Plint, 1983). In addition, the mud samples were subjected to HCl to determine if carbonate was present. If present, the samples No. 3, 1990] BADER AND PARKINSON—HOLOCENE EVOLUTION 207 Atlantic Ocean Vibrocore Site and Core Number Eau Gallie Causeway Core Borings (D.O.T., 1983 Melbourne Causeway Core Borings (D.O.T., 1978) Vibrocore Locations in Sixth Transect (Almasi, 1983) Merritt Island Eau Gallie River Crane Creek Turkey Creek (J Transect Identification Fic. 2. Map of the study area showing vibracore locations. Solid lines indicate location of stratigraphic transects, two of which are shown in Fig. 4. were analyzed chemically to determine the relative percent of carbonate and noncarbonate mate- rial (Flores, 1988). Sixty-four samples were taken from representative lithologic units for examination of mollus- can abundance and diversity. Each 10-cm sample was washed through a 2.0-mm screen and the fraction larger than 2.0 mm inspected. Each gastropod shell with an intact columella and each pelecypod valve with an undamaged hinge and/or a distinguishable pallial sinus were counted and identified to the species level. The combined faunal and lithologic data allowed sedimentary depositional units or facies to be described and upcore changes to be identified. Scaled stratigraphic cross-sections (c.f., Fig. 4A), drawn shore-parallel along the axis of the lagoon (Transect S) and shore-normal (Transect Y), provided a three-dimensional perspective of the lateral and vertical distribution of sediment types within the study area. 208 FLORIDA SCIENTIST [Vol. 53 KEY * Vibrocore Samples iso tee + Muck O Carbonate-rich Samples * Almasi's (1983) Samples PER CENT GRAVEL Trace (0.01%) Mud Sand (<0.0625 mm) (0.0625 - 2 mm) 0.1 1.0 9.0 SAND:MUD Fic. 3. Ternary diagram, adapted from Folk (1974), illustrating the distribution of the sedi- ment types collected during this study and Almasi’s (1983) Turkey Creek transect. G = gravel, g = gravelly, S = sand,s = sandy, M = mud, m = muddy. ResuLtts—The sediment samples contain four principal components: (1) quartz sand; (2) whole mollusc shells and angular shell hash; (3) mud with variable proportions of silt and clay; (4) carbonate nodules and mud chips. As expected, the gravel-sized fraction of the samples is dominated by skeletal material and, to a lesser degree, carbonate nodules and mud chips. The pri- mary component of the sand-sized fraction is quartz sand. Silt- and clay-sized particles, organics, finely disseminated calcium carbonate and framboidal pyrite comprise the fine-sized fraction. Based on the Folk (1974) classification scheme, 88% of the Indian River lagoon samples (excluding “muck”, an anthropogenic sediment type) con- sisted of sandy shell, sand, and muddy sand. Sandy muds comprise only 5% of the samples while the remaining 7 % are muds (Fig. 3). Although the sequence of sediment types varies from core to core, three general sediment trends are recognized. First, the sediment coarsens in the upcore direction, as evidenced by an increase in modal quartz sand size and the concomitant decrease in weight-percent mud (Fig. 5). Second, the Holo- cene sediment sequence appears to coarsen from north to south, along strati- graphically equivalent horizons (Fig. 4A). Third, sediments that have accu- mulated along the axis of the lagoon contain persistently higher concentrations of mud than those deposited along the lagoon margins (Fig. 4A). 209 BADER AND PARKINSON— HOLOCENE EVOLUTION No. 3, 1990] *SUOTBIO] 9.109 10J | “BIJ 99S *“JOAg] Bas UROUT st UINZVG “yUaTeAINbe jou are soTeOs [eJUOZIIOY 90N ‘oAIsso1ssues} st aoUINbas JUSUIIpes aUBD0[O}] SUNOIpU! (y) yo suonoas-ssor1o sAeIdI9qUy (gq) ‘eyep a1OOBIQIA BuIsN (7YBII) YOU 07 YINOs puL (391) Seo 0} SOM PaJONIYsSUOD sUOTaS-ssorO OTYdeIBHeNS (VY) “F “OY ’ ~~ KR ¢ S3dAL LNAWIGSS G31S8VINN OL A3y = = @® § 3 g 8 (9) 2 2 E g rad e 3 3 = : : PP n sy & (vy) Q g : : 3 5 210 FLORIDA SCIENTIST [Vol. 53 A total of 26 molluscan species were identified. However, two species accounted for 81% of the total number of individuals counted: Mulinia la- teralis (67% of the total number) and Anomalocardia auberiana (14%). These two species were nearly mutually exclusive, with A. auberiana occur- ring below M. lateralis (Fig. 5). In addition, molluscan species found within the A. auberiana zone were typically of low abundance and diversity. Most of these species were noted to be euryhaline and tolerant of brackish-water sa- linities. In contrast, the Mulinia lateralis zone contained a more abundant and diverse molluscan assemblage. While many of these species were euryhaline, several stenohaline species were identified in relatively large numbers. A dra- matic decrease in molluscan abundance and diversity was often noted to occur within the upper 0.5 to 1.0 m of the present sediment-water interface (Fig. 5). DEPOSITIONAL INTERPRETATION— The combined sediment and faunal data delineate four sedimentary facies: (1) subaerial Pleistocene sand, (2) coastal marsh mud, (3) restricted lagoonal muddy sand, (4) open lagoonal sand (Fig. 4B). This upcore facies transition from subaerial to open lagoonal is clearly transgressive and described briefly below (for additional details see Bader, 1988). Pleistocene sand: Two cores taken along the margins of the lagoon (2C and 4C; see Fig. 2) recovered clean, iron-stained, fine-grained quartz sands at their base (Fig. 4A). These sands are mottled, contain <2% skeletal mate- rial, and are locally cemented by carbonate. Core 4C, taken along the west- ern margin of the lagoon, encountered basal quartz sand identical to the Pleistocene quartz sands that form the Atlantic Coastal Ridge along the main- land shoreline (Fig. 1). Hence, they are interpreted to be Pleistocene strandline deposits (Fig. 4B) that were submerged during the initial stages of coastal submergence. Based on the presence of an iron-stain, the basal quartz sands encountered in Core 2C are also interpreted to be Pleistocene deposits (see Almasi, 1983). Marsh mud: The basal Holocene sediments along the axis of the lagoon (Cores 1 and 6; Fig. 2) contain between 63% and 93% mud. The muds are clay-rich, silt-poor, and lack visible layering. Very fine- to fine-grained quartz sand is present in trace amounts. Carbonate mud was identified at the base of Core 1 (Flores, 1988). Framboidal pyrite and wood fragments are common constituents at the base of Core 6. Although the muds are essentially depauperate with respect to molluscs, small numbers of the barnacle Balanus eburneus are sometimes present. The observed sediment characteristics, in conjunction with the strati- graphic position of this unit, are interpreted to reflect deposition in a coastal marsh environment (Fig. 4B). Because only two cores penetrated this sedi- ment type, it was not possible to determine what type of coastal marsh (i.e., freshwater, salt) produced this deposit. Restricted lagoon muddy sand: Generally overlying the marsh deposits No. 3, 1990] BADER AND PARKINSON— HOLOCENE EVOLUTION Za Modal Per Modal Per Sediment Sand Cent R.A. No. of No. of Sediment Sand Cent R.A. No. of No. of Type Size Mud (%) Species Shells Type Size Mud (%) Species Shells Vv FMO 15 0 1000 20 O 2000 Vv FMO 15 0 1000 20 0 2000 o oO 1 1 == me) ¢ M4 e% @ td os eo e = = £ £ S & é é be s 2 2 3 3 CORE 1B SEDIMENT TYPES INDEX SPECIES ~-N shel ae A M. lateralis 356 cm sent ef] ane ae = Neither Species Fic. 5. Downcore sediment and faunal trends of two representative vibracores (see Fig. 2 for core locations). Vibracore graphically illustrated using symbols shown in key. Modal sand size: V = very fine; F = fine; M = medium. R. A. (%) indicates the relative abundance of Mulinia lateralis and Anomalocardia auberiana using symbols shown in key. Shadow graphs demonstrate downcore changes in the abundance (number of shells) and diversity (number of species) of molluscs. 212 FLORIDA SCIENTIST [Vol. 53 are muddy sands (Fig. 4A) that contain mud balls, thin layers of shell, and small pockets of sand. In addition, hydrogen sulfide odors and/or blackened shells are present towards the base of this unit. Hydrobia sp., oyster debris, and calcareous worm tubes co-occur with a restricted marine fauna, domi- nated by Anomalocardia auberiana. The A. auberiana faunal zone can be traced to the north, where the muddy sands grade into sandy muds (Fig. 4A). As mentioned previously, the A. auberiana faunal assemblage contains eury- haline, brackish-water species. These sediments are interpreted as intertidal to shallow subtidal (re- stricted) lagoonal deposits (Fig. 4B), where seasonal patterns of precipitation and evaporation strongly influenced salinity values. Open lagoon sand: Fine-grained, gray sands usually overlie the restricted lagoon sediments (Fig. 4A). The sands contain localized accumulations of mud chips, mud balls, and shell deposits. Framboidal pyrite and plant frag- ments are locally abundant. Molluscan skeletal debris is significantly more diverse and abundant than in the underlying Anomalocardia auberiana zone. In contrast to the A. auberiana assemblage, which contained brackish-water molluscan species, Mulinia lateralis is the predominant species in the open lagoon facies, co-occurring with species that prefer more normal marine sa- linities. Toward the north, the fine gray sands grade into muddy sands also characterized by M. lateralis. These sediments are interpreted to have been deposited in a more open lagoon environment (Fig. 4B), with improved cir- culation and salinity values approaching normal marine. Recent sediments: A medium-grained, grayish-brown sand often caps the Holocene sediment sequence, except in localized topographic depressions and within the intracoastal waterway, where “muck” has accumulated (Fig. 4A). The sediment hosts a molluscan assemblage similar to the underlying gray sands (M. lateralis zone), although species diversity and abundance is gener- ally reduced. As these sands lie at the present sediment-water interface they are included as part of the open lagoon facies (Fig. 4B). Surficial deposits of “muck”, a fine-grained, organic-rich sediment (Tre- fry et al., 1987) with variable quartz sand content (Bader, 1988), reverses the upward coarsening trend discussed above. Faunal abundance and diversity are minimal. As “muck” sediment began accumulating within the lagoon about 30 yr ago and is attributed to anthropogenic processes, its mode of origin was not analyzed during this study. Discussion— This study was intended to be a preliminary investigation of the back-barrier Holocene sediment sequence of central Brevard County. It is a logical extension of a similar study conducted by Almasi (1983) further south. In general, a coarsening upward, transgressive sediment sequence has developed within Indian River lagoon during the late-Holocene. Although this study is only the second to have focused on the geological history of Indian River lagoon, the recognition of a transgressive facies sequence is not surprising. Hence, the discussion to follow will focus on specific lateral and vertical anomalies. No. 3, 1990] BADER AND PARKINSON— HOLOCENE EVOLUTION 213 A concomitant increase in mud and decrease in sand was noted in south- to-north transects along the axis of Indian River lagoon (Fig. 4A). Because fluvial systems entering the lagoon do not provide a significant volume of bedload material, we argue that barrier island morphology is probably re- sponsible for this trend. Within the study area, the width of the barrier island and its well-developed ridge-and-swale topography (Fig. 1) may have mini- mized the influx of sand into the lagoon during major storms (Davis, 1985). Tidal inlets, which often form during storm events, are absent in the north- ern area, and there are no morphologic features to suggest that they had ever been present. The presence of a well-developed cuspate spit morphology along the lagoonal margin of the barrier island (Figs. 1 and 2) suggests this area has been relatively stable (Zenkovitch, 1958), adding additional support to our hypothesis. In contrast to the persistently muddier sediment observed in northern limits of this study area, the relatively sandy sediments to the south conform more closely to the lagoonal sequences described in Almasi’s (1983) Turkey Creek transect. These sandier sediments are logically related to the overwash and flood tidal delta geomorphology of the barrier island in this area (Fig. 1). Both processes responsible for the construction of these features would have contributed large quantities of sand to the lagoon. The relatively rapid rates of sedimentation generally associated with these events may be partially re- sponsible for the shallower water depths observed in this section of the la- goon. In transects from the axis of the lagoon toward its margins, quartz sand content increases, and the concentration of mud diminishes. This sediment distribution pattern is typical of microtidal lagoons (Davies, 1964). Sands are deposited closer to the periphery of the lagoon because winds impart a rela- tively higher energy to the shallower water. Silts and clays are winnowed from the sands and remain in suspension until they can settle out in the deeper waters of the central lagoon. In addition, the presence of local quartz sand sources along both margins of the lagoon has undoubtedly contributed to the observed grain size trends. Conc.usions—(1) The Holocene sediment sequence within Indian River lagoon of central Brevard County, Florida, is transgressive, reflecting a tran- sition from subaerial to open lagoon environments. (2) This sequence developed as rising Holocene sea level flooded a topo- graphic depression (paleolagoon) between 5,000 and 6,000 yr B.P. (3) Cross-sections along the axis of the lagoon reveal a concomitant in- crease in mud and decrease in sand from south to north at all stratigraphic levels. This gradient is interpreted to reflect the minimal role of overwash and flood tidal delta sedimentation in the northern portion of the study area where beach ridge sets and island width may have prevented island breach- ing during storm surges. (4) Marginal deposits are persistently sandier, reflecting both proximity to local quartz sand sources (Atlantic Coastal Ridge and present barrier island) 214 FLORIDA SCIENTIST [Vol. 53 and the energy gradient predicted for microtidal back-barrier lagoons. (5) “Muck”, a fine-grained, organic-rich deposit, was noted to occur only in bathymetric depressions of the present lagoonal substrate and is not present at any other stratigraphic position. This observation supports the contention that “muck” is of anthropogenic origin (Trefry et al., 1987). ACKNOWLEDGMENTS— This paper is based on work done for a M.S. degree by the senior au- thor, under the supervision of Dr. Nell Tyner. Dr. Edward Kalajian of the Florida Institute of Technology and Mr. Wilson Timmons of the Office of Natural Resources Management, Brevard County, provided additional support and guidance. Richard Gurlek, Kim D’Arcy, Garry Hollem, Karen Monroe, and Karen Pitchford were an able crew on the vibracoring trips. This paper is contribution no. 142 of the Department of Oceanography and Ocean Engineering, Florida Insti- tute of Technology. LITERATURE CITED AtmasI, M. N. 1983. Holocene sediments and evolution of the Indian River (Atlantic coast of Florida). Ph.D. dissert. Univ. Miami, Coral Gables. Baper, S. F. 1988. Stratigraphy of the Indian River lagoon: Central Brevard County, Florida. Masters thesis. Florida Inst. Technology, Melbourne. Brown, D. W., W. E. KENNER, J. W. Crooks, AND J. B. Foster. 1962. Water resources of Brevard County, Florida. Florida Geological Survey Report of Investigation No. 28, Tallahassee. Davies, J. L. 1964. A morphogenetic approach to world shorelines. Z. Geomorph. 8:127-142. Davis, R. A., JR. 1985. Beach and nearshore zone. Pp. 379-444. In: Davis, R. A., Jr. (ed.), Coastal Sedimentary Environments (Rev. Ed.). Springer-Verlag, New York. DEPARTMENT OF TRANSPORTATION (FLORIDA). 1978. Brevard County, bridge on State Road 516 over Indian River (I.W.W.) at Melbourne. Report of core borings for structures. DEPARTMENT OF TRANSPORTATION (FLORIDA). 1983. Brevard County, State Road 518, Eau Gallie relief bridge and swing span replacement. Report of core borings for structures. DvuBar, J. R. 1974. Summary of the Neogene stratigraphy of southern Florida. Pp. 206-231. In: Oaks, R. Q., AND J. R. DuBar (eds.), Post-Miocene Stratigraphy, Central and Southern Coastal Plain. Utah State Univ. Press, Logan. Evans, M. W., A. C. Hing, D. F. BELKNaP, AND R. A. Davis, Jr. 1985. Bedrock controls on barrier island development: west-central Florida coast. Mar. Geol. 63:263-283. Fores, C. E. 1988. Adsorption-desorption studies of phenylmercuric acetate onto model and natural sediments. Masters thesis. Florida Inst. Technology, Melbourne. Foxk, R. L. 1974. Petrology of Sedimentary Rocks. Hemphill Publishing Co., Austin. Ha sey, S. D. 1979. Nexus: new model of barrier island development. Pp. 185-210. In: LEATHER- MAN, S. P. (ed.), Barrier Islands from the Gulf of St. Lawrence to the Gulf of Mexico. Academic Press, New York. KoFoep, J. W. 1963. Coastal development in Volusia and Brevard Counties, Florida. Bull. Mar. Sci. Gulf Caribb. 13:1-10. Kraft, J. C., AND M. J. Curzastowski. 1985. Coastal stratigraphic sequences. Pp. 625-664. In: Davis, R. A., Jr. (ed.), Coastal Sedimentary Environments (Rev. Ed.). Springer-Verlag, New York. , E. A. ALLEN, D. F. BELKNap, C. J. JoHN, AND E. M. MaurmMeyer. 1979. Processes and morphologic evolution of an estuarine and coastal barrier system. Pp. 149-184. In: LEATHERMAN, S. P. (ed.), Barrier Islands from the Gulf of St. Lawrence to the Gulf of Mexico. Academic Press, New York. Lanesky, D. E., B. W. Locan, R. G. Brown, AND A. C. Hine. 1979. A new approach to portable vibracoring underwater and on land. J. Sediment. Petrol. 49:654-657. Menta, A. J., W. D. Apams, AND C. P. Jones. 1976. Sebastian Inlet: glossary of inlets, report no. 3. Univ. Florida, Gainesville. Mostow, T. F., AND S. D. Heron, Jr. 1979. Quaternary evolution of Core Banks, North Caro- lina: Cape Lookout to New Drum Inlet. Pp. 211-236. In: LEATHERMAN, S. P. (ed.), Barrier Islands from the Gulf of St. Lawrence to the Gulf of Mexico. Academic Press, New York. No. 3, 1990] BADER AND PARKINSON— HOLOCENE EVOLUTION 215 Osmonp, J. K., J. P. May, AND W. F. Tanner. 1970. Age of the Cape Kennedy barrier-and-lagoon complex. J. Geophys. Res. 75:469-479. Punt, A. G. 1983. Facies, environments and sedimentary cycles in the Middle Eocene, Brack- lesham Formation of the Hampshire Basin: evidence for global sea-level changes? Sedi- mentology 30:625-653. TreFry, J. H., D. K. SrauBue, M. A. Sister, D. TiERNAN, R. P. TRocINE, S. Merz, C. J. GLAscock, AND S. F. Baper. 1987. Origin, composition and fate of organic-rich sediments in coastal estuaries: Project Muck. Florida Inst. Technology, Melbourne. U.S. Coast AND GEopETIC Survey. 1980. Melbourne east quadrangle, Florida, Brevard County, 7.5 minute series (topographic). U.S. Geological Survey, Reston. White, W. A. 1970. The geomorphology of the Florida peninsula. Geol. Bull. No. 51, Florida Bureau of Geology, Tallahassee. ZENKOVITCH, V. P. 1958. On the genesis of cuspate spits along lagoon shores. J. Geol. 67:269-277. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3):204-215. 1990. Accepted: May 8, 1989. Atmospheric and Oceanographic Sciences AN INTRODUCTION TO THE TIDES OF FLORIDAS INDIAN RIVER LAGOON II. CURRENTS NEp P. SMITH Harbor Branch Oceanographic Institution, 5600 Old Dixie Highway, Fort Pierce, FL 34946 Asstract: Hourly current measurements recorded along the Atlantic Intracoastal Waterway are used to quantify amplitudes and local phase angles of the principal tidal constituents in Florida’s Indian River lagoon. Twenty-seven study sites located between latitudes 27°10' and 28°17'N were occupied for periods of 1-2 months each during 1976-1988. Six principal tidal constituents (M2, Sy, No, K;, O,, and P,) have amplitudes greater than the precision of the mea- surements used to make the calculations. The semi-diurnal M, constituent dominates all other tidal constituents. M, amplitudes exceed 60 cm/s in the vicinity of Ft. Pierce Inlet; the amplitude just north of St. Lucie Inlet is approximately 45 cm/s. M, amplitudes are relatively small in the vicinity of Sebastian Inlet: Values of 11 and 19 cm/s are computed from data obtained at study sites approximately 3 km north and south of the inlet, respectively. M, amplitudes north of 28°11'N are less than 1 cm/s. A comparison of phase angles indicates that tidal wave forms propagate at speeds of 5-10 km/n. In the central parts of the central and southern segments of the lagoon, the time delay of any phase of the tide is 6.5 and 5h, respectively, relative to Ft. Pierce Inlet. THE ebb and flood of the tide play a central role in the circulation of most estuaries by performing several important functions. At inlets and passes, they generally provide the dominant estuarine-shelf exchange mechanism. Within an estuary, tidal motions effect longitudinal, lateral, and vertical mixing, which serve to disperse pollutants and anything else that is dissolved or suspended in the water column. Tidal motions are thus of major impor- tance in maintaining or restoring water quality. The relative importance of tidal motions within an estuary varies accord- ing to the type of estuary. Coastal lagoons, in particular, have characteristi- cally low tidal amplitudes because of the restrictive effect inlets and passes have on exchanges. Thus, even under unexceptional wind conditions, the ebb and flood of the tide in the interior of a lagoon may be dominated by the wind-driven component of the current. Nevertheless, tidal motions are im- portant because they are as dependable as they are periodic, and thus predict- able. Tidal processes can be enhanced greatly by nontidal forcing, but the baseline level of transport, mixing, and flushing they provide should be inves- tigated and quantified. Indian River lagoon lies along the Atlantic coast of central Florida. The lagoon is 195 km long, characteristically 2-4 km wide, and water depths are generally 1-3 m on both sides of the Atlantic Intracoastal Waterway (AIW). The lagoon is of the “restricted” type, according to the definitions suggested by Kjerfve (1986). The lagoon can be subdivided into three segments: The northern segment, north of Sebastian Inlet, is 113 km long; the central seg- ment, between Sebastian and Ft. Pierce Inlets, is 45 km long; the southern segment, between Ft. Pierce and St. Lucie Inlets, is 37 km long. The land- No. 3, 1990] SMITH— INDIAN RIVER TIDES YAW ward side of the lagoon receives fresh water from natural creeks and rivers, from groundwater discharge (Pandit and El-Khazen, 1990), and from a series of man-made drainage canals that have added to drainage from the natural watershed of the lagoon. This paper is the second of a two-part survey of the tides of Florida’s Indian River lagoon. The first paper (Smith, 1987), using water-level data, showed that the M, constituent—the principal semi-diurnal constituent—is dominant throughout the lagoon. M, amplitudes in the northern, central, and southern segments are 0-5 cm, 5-10 cm, and 10-15 cm, respectively. The constricting effect of inlets, combined with the frictional damping of the tidal wave form, restricts significantly the portion of the lagoon that is influenced by tides. Thus, tidal ranges associated with diurnal and semi-diurnal period constituents decrease rapidly with distance inside each of the three inlets. Ft. Pierce Inlet, dredged to a depth of about 6 m, carries more water between the lagoon and the inner shelf than does either Sebastian Inlet or St. Lucie Inlet. As a result, tidal amplitudes in the lagoon are greatest in the northern part of the southern segment and in the southern part of the central segment. Tidal-current amplitudes presented in this paper are also larger in the vicinity of inlets—especially Ft. Pierce Inlet. Spatial irregularities in phase angles are apparently attributable largely to differences in weather condi- tions and/or inlet cross-sections during the 13-yr period over which data were obtained. THE OBsERVATIONS— Current data used to quantify tidal harmonic constants (amplitudes and local phase angles, referenced to Eastern Standard Time) of the principal diurnal and semi- diurnal constituents were recorded using General Oceanics Type 2010 film-recording inclinome- ters. Both inclination and azimuth angles were read to the nearest degree. Although the conver- sion from inclination angle to current speed is nonlinear, a precision of +0.5° corresponds to about + 1.5 cm/s, within the inclination range commonly found in this study. The accuracy of the azimuth reading is + 5° for inclination angles greater than 10°, according to instrument specifi- cations. Data were collected in field studies conducted at irregular intervals from 1976 through 1988. Time series were typically 30-45 da long, but records from AIW Markers 57, 133, and 207 were 223, 175, and 102 d long, respectively. During the 1976 and 1977 field studies, current meters were attached to concrete blocks, and the midpoint of the current meter was approximately 1 m above the bottom at 0° inclination. In later studies, the mooring design was altered such that the current meter recorded the flow about 1.5 m above the bottom, or just below mid-depth. All current-meter records were obtained from moorings in the AIW. Currents in the relatively shallow seagrass flats on both sides of the waterway are spatially variable because of the presence of spoil islands and local irregularities in bottom topography. By conducting field measurements in the AIW, results are more directly comparable, and the greater depth (3.5 m) reduces the possibility of vandalism and minimizes the contaminating effect of wind-wave motions. The study focuses specifically upon semi-diurnal and diurnal tidal constituents. With the exception of data collected at AIW Marker 57 (27°52’N), current-meter records were not long enough to quantify the harmonic constants of fortnightly, monthly, and longer constituents. The Marker 57 data, however, suggest that amplitudes of the long-period tidal constituents are at or within the accuracy of the measurements and, thus, cannot be quantified with confidence. MeErHops—Because of the large length-to-width ratio of Indian River lagoon, and because currents in the AIW are constrained significantly by the channel, only the longitudinal, along- channel component of the current is used in the analysis. The along-channel direction was deter- mined initially from NOAA Nautical Charts 11472 and 11485. In many instances, however, the indicated along-channel orientation was confirmed with polar coordinate plots of the raw data. “Along-channel” was defined by the line passing through the greatest concentrations of points 218 FLORIDA SCIENTIST [Vol. 53 representing flood and ebb current velocities. Timing errors in the raw data were generally minimal. When the total number of observa- tions did not correspond with the time interval between the known starting and ending times, however, the time series was “stretched” or “compressed” by fitting a natural cubic spline func- tion through the data points and then interpolating or extrapolating along the curve to extract values at hourly intervals. Tidal harmonic constants were computed from along-channel current components using the 29-d harmonic analysis computer program described by Dennis and Long (1971). Time series were usually long enough to permit several 29-d analyses, with each successive computation starting several days farther into the time series. When several harmonic constant pairs were available, they were vector-averaged, as suggested by Haurwitz and Cowley (1975), to provide the single value used to represent the study site. Vector probable errors provided a measure of the scatter about the vector mean. When the vector probable error exceeded 50% of the vector- averaged amplitude, the constituent was judged to be statistically insignificant at that location, and it was dropped from further consideration. AIW Marker a M2 K, O; | ! | | | | | | | | | | | | | | | | | | | | ! | | | | | | | | | | | | { | | | | | 1 1 ee ed Le) ee) Co eee ma Oo woo 5 00 5 0 680 5 00 5 00 5 -27 BOF Amplitude, cm/sec Fic. 1. Amplitudes of principal semi-diurnal and diurnal tidal constituents, in cm/s, recorded in Florida’s Indian River lagoon between Atlantic Intracoastal Waterway (AIW) Markers 86 and 232. Note that the M, constituent amplitude has been plotted on a compressed axis. Major tick marks indicate labeled A[W marker numbers; minor tick marks represent intermediate markers listed in Table 1. ResuLts—Harmonic constants of tidal constituents within the lagoon are listed in tabular form in Table 1 and shown in analog form in Figs. 1 and 2. Figure 1 shows tidal constituent amplitudes, in cm/s, along the longitudinal axis of the lagoon. The M, constituent was isolated and is presented above a compressed axis to the right of the lower-amplitude constituents. All constituents show locally higher amplitudes in the immediate vicinity of St. Lucie and Ft. Pierce Inlets. Amplitudes computed from data collected just inside Sebastian Inlet are not appreciably higher than those from adjacent study sites further north and south. The lower-amplitude con- stituents show locally higher amplitudes in the interior of the lagoon where constrictions reduce the cross-sectional area. The Mg, constituent is best suited for determining where tidal motion and, therefore, tidal flushing are minimal. In the southern segment, between St. Lucie and Ft. Pierce inlets, lowest M, amplitudes of approximately 7 cm/s are computed from data collected at AIW Marker 218 (27°16’N). In the central segment, between Ft. Pierce and Sebastian inlets, minimal amplitudes No. 3, 1990] SMITH—INDIAN RIVER TIDES 219 28°20'T AIW Marker M, S, K,&P, O; 86 | \ lo 99 | 3 9 | 28°00 93 | 57 | 50) 70 oe = 112 P= ’ 5 407 143 is 166 30’ 181 20' 202 218 lo 232 Le SSS) —EEEEE— sy _eEE———E— (oy 360° O° 360° O° 360° O° 360° Oz 360° 27°00" Local Phase Angle Fic. 2. Same as Fig. 1, except showing local phase angles. Note that the K, and P, constituents are represented by the same line. occur in the vicinity of AIW Marker 154 (27°36’N), just south of Vero Beach. North of Sebastian Inlet, in the northern segment of the lagoon, amplitudes continue to decrease northward. M, amplitudes reach 1 cm/s at ATW Marker 99 (28°11’N). This appears to be a real value, even though it is at the precision of the measurements. Multiple, overlapping, harmonic analyses produce consistent phase angles. Still, the tidal excursion—the horizontal displacement associ- ated with either the flood or ebb portion of the tidal cycle—is less than 150 m (the M, constituent tidal excursion is found by multiplying the amplitude by the 12.421-h period and dividing by =). Measurements were not made farther north, because tidal excursions on the order of 100 m are not physically important, even if the amplitudes are statistically significant. Local phase angles are shown in Fig. 2. The horizontal scale has been extended beyond a range of 0-360°, making it possible to offset curves that would otherwise overlap. In general, one would expect phase angles to be lower near inlets, thereby indicating a phase lead of flood, ebb, or slack water conditions. The M, constituent shows this best, because it is most prominent throughout the lagoon. Other constituents show greater spatial variability, because it is difficult to pinpoint the exact crest of a very low-amplitude sine wave. M, constituent phase angles increase by approximately 140° between either St. Lucie or Ft. Pierce Inlet and the interior of the southern segment. This corresponds to a time delay of approxi- mately 5 h. The phase lag in the interior of the central segment is somewhat greater, partly because the central segment is 8 km longer than the southern segment, and certainly because spoil islands in the interior decrease the speed of propagation. The 190° M, phase lag between Ft. Pierce Inlet and the area between AIW markers 75 and 112 (27°46’ to 27°43’N) corresponds to a time lag of over 6.5 h. It is of interest to note that maximal phase lags occur at stations well north of the midpoint of the central segment and over 20 km north of where the lowest amplitudes occurred. It appears that northward moving tidal waves from Ft. Pierce Inlet are dominant through most of the central segment, while waves moving south from Sebastian Inlet play a dominant role only relatively close to the inlet. In the northern segment of the lagoon, a phase lag of 130° is calculated between AIW markers 57 and 86 (27°52’ to 28°17’'N). This corresponds to a time lag of about 4.5 h from Sebastian Inlet to just south of Cocoa. o2 6ST 61 6ST 61 = 91 oe irs 13 S ILI E83 ILI 3 = oT ST Gg 9% G1Z 69% SIZ SPE 10 can 0% per SSI 6&3 SSI 800 91 Gg 6'P Nes 023 63E 023 610 $0 Ls a v's 281 GG 281 8E0 el es g's 8'¢ 933 163 933 €£0 G0 £3 at 9°¢ 99% Z6G $9 0£0 3 eI 63 gE 8S 2 LSI ELT LST SES 5 oT ce ce Zed SI GST 961 GST 186 o IT ge g€ 03 - OFT LLI OFT 062 ce 60 ST VS al 3 ZIT SOI SII L8G = G0 rT QT Cal SOG 133 SSG 183 ‘0 60 UT v0 3S QF 3G 830 10 9°0 v0 10 880 L0 880 LYE €'0 OT Ol v0 ‘d 'O Ly oN yuan}I}suo7) 220 060 60 €90 I'T 0) G0 060 8% 060 oe 180 iG G60 9% 090 8% EG 60 280 OT L10 Cal G60 OT 8£0 C50 €F0 r'0 SEE 10 EG 61 OL Cee OTT ShE 86 GZ0 191 1¥0 6°91 40) 9ST ZrO GSI crO LSI 6SZ OTT £66 FOr G6G CL ZI 8° 68S OG 600 OT 820 lean oW SR SF Re SF RSER OR SC RER ERS R OR LS RS 2S SR gL nf LL AeW 08 ABW gy nf LL ACW gz inf LL Aew 08 AeW €8 dag €g un{ €g unf 98 PO 18 dag 18 09q 98 1dy 23eq © 66008 ‘9 8EoLG 18°66 008 18 6 oL6 10° €6 008 ‘TOV OLE 8° 6 008 LEP OLS 16 VGo08 1G VV oLG 16 G6008 8° SV oLG 18°SG6 008 © OVoLG 10’ L6 008 B'LVoOLG :T°66008 16 CGoLG 10 TE.008 © GS oLG 16 6008 iL 89 oL6 ‘9' VE 008 © £0086 V'9E 008 18°90 086 © 8E 008 8°01 086 ‘S' T¥008 © LT o86 M N u01}]e00'T] eri vel cel oll DIN "SALI9S OUT} JY} JO JUIOdpIUI ay} 0} SIoJaI 9}ep VY, “WI0}}0q BY} VAOe 19}9UI }UILINI VY} JO (WI) }YSIaY 9} SI H{ “Sdeidap UI (7%) sa[sue aseyd [eo] ‘s/w ur (L) sapnydury ‘uoose] Joa ULIPU] OJ syUsNz}sUOO [ep [euINIp pue [euinip-ruras pedrioutsd jo syueysuoo olUOULIeH *[ ATAV], 221 SMITH—INDIAN RIVER TIDES No. 3, 1990] TL TT08 ‘S 61.08 VET 008 8°ST 008 iL 9T 008 0’ 61 008 ‘9°61 008 iL 06008 T'T6008 iL 16008 16° T6008 16 66008 M VOToLG VT oLG DST oLG T'060L6 1 TG oLG iV L6oLG 19 86 oLG 16 CEoLG OVE LG VIEL iL GE oLG DLE oLG N uonRo0'T SS SS SS eS SS SS Ee a Sa ee ee a ee ee ee ee ee en G&G GGG raul PET ZIT 681 83S (avd x 6'T 8'¢ 8's 69 a 2 L'&r u Tg Aqnf GI £01 c60 €0I 002 V6 11Z x 80 GS SV GI VT 8'8 u 18 3das GT £60 OFT 760 00Z Eb ra x r'0 L'0 or GT 9°0 OL u 9g AB CT E13 SSI ELZ 09 1840) CLS x 10 9'T 0% 6% rT 81 ue Lg uef GT 6IT GLO 6IT L6G €S0 oSE x eT E'S 0'F 19 g'¢ Ee u 18 mf Cor 163 C63 163 coo 9S0 3£0 x iG 9°8 G6 (Saal v8 8°OL u g sny GT €3I CEI ESI 961 BES ) H o's 8°g 0'6 26 L’S Z'6S u gz nf OT 631 Cel 631 80Z 1S OFZ x 0% 9°¢ 09 19 6'E 6'8E u 6L ABW co) EST oS EST OFZ PLZ EGG x ST s'¢ a vy GG G'0E u gz [nf OT 8ZI €9I 8ZI £83 613 99% x v0 L'0 TT IT VT VL u Luef O'T 690 1¥Z 690 G03 CES 643 x Z'0 6'0 80 TT 20 9° u Luef OT LOT GZS LOT ELZ SOE 863 H IT eT v's 0% 80 v6 u LL AB- OT ‘d 'O 'y oN g¢ A a7eq H yU9N}I}SUOT) penunuoy ‘| ITaVvy, 222 FLORIDA SCIENTIST [Vol. 53 When time lags such as those noted above are combined with distances separating stations, one can estimate phase speeds for any given tidal constit- uent. Again using the M, constituent as an example, in the southern segment, the nearly 5 h required for wave forms to move approximately 20 km trans- lates into a phase speed of just over 4 km/h. In the central segment, wave forms moving north from Ft. Pierce Inlet to A[W Marker 70 in about 6.5 h indicate a phase speed of approximately 5 km/h. In the northern segment of the lagoon, the average phase speed of the M, constituent moving north from Sebastian Inlet to near Cocoa is just under 10 km/h. Discussilon—Both the ebb and flood of the tide along the AIW and the rise and fall of water level in the lagoon arise fundamentally from tidal wave forms moving north to south along Florida’s Atlantic shelf. As high and low tides pass by the inlets, water is alternately forced into and drawn out of the lagoon. But the transition from shelf tides to lagoon tides is quite different for currents than for water levels. Part I of this study (Smith, 1987) showed tidal amplitudes decreasing substantially in the immediate vicinity of each inlet. Frictional effects in the shallow water of the lagoon combine with the con- stricting effect of an inlet to decrease tidal-period variations in water level as tidal waves enter the lagoon. For example, M, amplitudes estimated from bottom pressure records obtained 1 km seaward of Ft. Pierce and Sebastian inlets are just over 50 cm. For comparison, the M, amplitude in the lagoon near both Ft. Pierce and St. Lucie Inlets was found to be 17 cm; it was 8 cm near Sebastian Inlet. Current data presented here in Part II of this study reveal a quite differ- ent pattern. Tidal amplitudes increase dramatically at each inlet. Under fric- tionless conditions, phase speed is proportional to the square root of the water depth. As tidal waves enter the lagoon, the shallow water slows the speed of propagation substantially. As a result of the reduced phase speed, the wave- length becomes significantly shorter, the wave form is compressed, and the slope of the wave form ahead of and behind the crest increases. The pressure gradient associated with the surface slope is the primary driving force for the ebb and flood of the tide; and, thus, significantly higher current speeds result. The shallow water initially slows the tidal wave form much more than it decreases its amplitude, explaining the substantial, if local, increase in current speed. Within the lagoon, the water depth and, therefore, wavelengths remain about the same, and wave forms are not compressed further. Frictional losses continue to decrease the tidal range, however. Thus, in the interior of the lagoon, the pressure gradient and resulting current speeds decrease apprecia- bly. Figure 1 shows that at the midpoint of both the southern and central segments, M, current speeds are about 10 cm/s. In the northern segment, current amplitudes are reduced to less than 1 cm/s north of 28°10’N. Farther north, tides are physically unimportant in both water-level and current- meter records. No. 3, 1990] SMITH— INDIAN RIVER TIDES 223 Both Figures 1 and 2 show some significant differences in phase and am- plitude at adjacent stations. Any of several different explanations may ac- count for this. First, mass continuity considerations require that the ampli- tude vary inversely with the local cross-sectional area of the lagoon. The depth and width of the AIW are essentially constant along the length of the lagoon, but the width of the lagoon itself, as well as the mean depth outside the dredged channel, can vary by a factor of 2-3. Second, in view of the probability that the bottom friction layer extends through the entire water column, the indicated spatial variation in amplitude must reflect the mooring design used at the time the data were recorded. Specifically, at any given location, the amplitude of the tidal constituent in question may be directly but nonlinearly related to the height of the current meter above the bottom. This presents a problem when time series from many individual studies are combined. No attempt was made to correct for the position of the current meter above the bottom, but recent modeling work (Smith, in press) suggests that amplitudes computed from measurements made 1.5 m above the bottom in water 3 m deep are approximately 15% greater than amplitudes computed from measurements made 1 m above the bottom. A third reason why neighboring amplitudes may differ significantly stems from the seasonal and certainly annual variation in the cross-sectional area of the nearest inlet. Ft. Pierce Inlet receives annual maintenance dredging; both Sebastian Inlet and St. Lucie Inlet are dredged less frequently. Because data collected for this study were collected over a 13-yr period, it is probable that tidal exchanges through Sebastian and St. Lucie inlets varied measurably and that this had a direct influence on tidal amplitudes in the lagoon. No long time series are available from outside the lagoon to determine whether these differences are a result of tidal exchanges through inlets or a result of changes in the constituents themselves during the study period. Station-to-station variations in local phase angle along the axis of the lagoon (Fig. 2) may be influenced by any of several processes which influence the propagation speed of tidal wave forms. Both high-frequency wind-wave motions and low-frequency meteorological forcing can influence tidal condi- tions and thereby contribute noise to a composite which combines measure- ments separated in time and space. Grant and Madsen (1979) have shown that turbulence arising from oscillatory wave motions increases the effective bottom roughness and thus retards tidal wave propagation. Enhanced bot- tom stress might be especially important in the shallow waters of a coastal lagoon. Spatial variations in fetch from one study site to the next, together with temporal variations in mean wind speed from one study period to the next, can distort spatial patterns in local phase angles when results are compiled. Over longer time scales, the propagation of tidal waves can be influenced directly by nontidal water-level variations. As noted above, the phase speed in shallow water is proportional to the square root of the water depth. In Indian River lagoon, where seasonal variations in water level have an aver- 224 FLORIDA SCIENTIST [Vol. 53 age amplitude of 10 cm (Smith, 1987) and where meteorological forcing can raise or lower water levels an additional 0.5 m relative to the seasonal norm (Smith, 1986), the phase speed in water 1-2 m deep can vary on the order of + 15-20%. While data assembled in this study provide a good overview of the ebb and flood of the tide along the longitudinal axis of the lagoon, results pre- sented (Figs. 1 and 2) cannot be extrapolated laterally into the shallow waters of both sides of the AIW. A logical follow-up to this introduction to tides would involve the application of a lagoon-scale, two-dimensional hydrody- namic model. Once calibrated, the model could provide the time series neces- sary to construct co-tidal and co-range lines throughout the lagoon. In situ data might still be required to verify tidal conditions in areas of specific interest, but results of the model would improve considerably the one-dimen- sional picture presented here. Current-meter data from within Indian River lagoon are put in perspec- tive by noting tidal conditions in adjacent shelf waters. Unpublished current data were collected 1 km from the coast at a study site 15 km north of Sebas- tian Inlet between mid-July and mid-November, 1984. Computed harmonic constants include an M, amplitude of 1.5 cm/s in the along-shelf direction and 0.5 cm/s in the cross-shelf direction. Both values are well within the precision of the current-meter measurements used in the calculations. The associated along-shelf tidal excursion is less than 0.25 km. Thus, tidal cur- rents over the inner shelf are physically unimportant for most practical pur- poses. With the available data base, Indian River lagoon emerges as an unexcep- tional “restricted” type coastal lagoon. Tidal co-oscillations provide signifi- cant transport near each inlet. While tidal motions may stand out distinctly in current-meter records, they decrease substantially in importance in the interior of the lagoon. There, as in many coastal lagoons, tides provide a baseline level of transport and flushing but play an ancillary role to the local wind-driven circulation. ACKNOWLEDGMENTS— Any study that involves reading, key-punching, proofreading, and ana- lyzing over 100,000 current speed and direction pairs will understandably require and benefit from the assistance of many individuals. Dr. Ortwin von Zweck directed the data collection and reduction during 1976 and 1977 under a consulting agreement with Harbor Branch Foundation while he was a member of the Florida Institute of Technology faculty. Dove W. Green painstak- ingly digitized many time series collected during that same time, including 11 used in this study. Lew Gilliland took over from 1979-80; Mark Sternberger continued in 1981-83, as the study expanded into the southern segment of the lagoon. Elizabeth Smith assisted in data collection and analysis during 1982-84. Patrick Pitts has been playing a central role in the data collection and reduction since mid-1984. He was in charge of field studies designed to fill gaps in the southern segment, and he worked with Paul Mikkelsen in 1986 to extend spatial coverage into the northern segment of the lagoon. In addition, Patrick Pitts worked with summer intern Rusty Tarver in 1987 to complete the important but onerous task of double-checking and editing time series during the final compilation of the 27 time series used in this paper. Harbor Branch Oceano- graphic Institution, Inc., Contribution Number 772. No. 3, 1990] SMITH— INDIAN RIVER TIDES Pipts) LITERATURE CITED Dennis, R. E. AND E. E. Lone. 1971. A user’s guide to a computer program for harmonic analysis of data at tidal frequencies. NOAA Tech. Rept. 41, U.S. Dept. Comm., Rock- ville, MD. Grant, W. D. anp O. S. Mapsen. 1979. Combined wave and current interaction with a rough bottom. J. Geophys. Res. 84:1797-1808. Haurwitz, B. anp A. Cow.ey. 1975. The barometric tides at Zurich and on the summit of Santis. Pure Appl. Geophys. 113:355-364. Kyerrve, B. J. 1986. Comparative oceanography of coastal lagoons. Pp. 63-81. In: WourE, D. A. (ed.), Estuarine Variability. Academic Press, New York. PanpiT, A. AND C. C. EL-Kuazen. 1990. Groundwater seepage into the Indian River lagoon at Port St. Lucie. Fla. Scient. 53(3a). SmiTH, N. P. 1986. The rise and fall of the estuarine intertidal zone. Estuaries 9:95-101. . 1987. An introduction to the tides of Florida’s Indian River lagoon. I. Water levels. Fla. Scient. 50:49-61. . (in press) Computer simulation of residual tidal transport in a coastal lagoon. J. Geophys. Res. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3):216-225. 1990. Accepted: May 13, 1989. Biological Sciences ICHTHYOFAUNA ASSOCIATED WITH SPOIL ISLANDS IN INDIAN RIVER LAGOON, FLORIDA NANcyY BROWN-PETERSON”) AND Ross W. EAMES”) Florida Department of Natural Resources, Bureau of Aquatic Preserves,(!) 4842 South U.S. 1, Ft. Pierce, FL 34982 and 13 East Melbourne Ave., Melbourne, FL 32901. Asstract: The ichthyofauna associated with 90 spoil islands along 190 km of Indian River lagoon from Haulover Canal to St. Lucie Inlet was surveyed from October 1987 through July 1988. A total of 25,801 individuals representing 60 species and 32 families were collected, with 25% of the species of tropical or subtropical origin and the remainder from warm-temperate fauna. While Syngnathus scovelli, Eucinostomus spp., Lagodon rhomboides, and Menidia penin- sulae were most frequently captured, the percent occurrence of species varied seasonally due to recruitment of juvenile fishes. A UPGMA cluster analysis separated spoil islands into five distinct groups broadly differentiated by influence from ocean inlet waters but greatly affected by season- ality. When spoil islands were separated into four groups based on tidal excursion values and the number of tidal cycles from an inlet, species richness was significantly (p<0.05) lower around islands not affected by inlet waters (> 60 tidal cycles) than around islands moderately affected by inlets (within 3-6 tidal cycles). Juveniles of species that are offshore spawners were most common around islands affected by ocean inlets (1-6 tidal cycles) and were rarely or never found in the northernmost portion of the lagoon (>60 tidal cycles). In contrast, lagoonal spawners were generally more common around islands moderately or slightly affected by tidal action (3-60 tidal cycles). In general, spoil island ichthyofauna is similar to the ichthyofauna from other littoral areas within Indian River lagoon. Seasonal recruitment of juvenile fishes indicates the impor- tance of spoil islands as nursery habitat. Finally, distance from ocean inlets appears to affect ichthyofauna distribution around spoil islands. INDIAN River lagoon, Florida, has a diverse fish fauna, with many repre- sentatives from both the warm-temperate Carolinian and tropical Caribbean faunas. While the ichythyofauna within various habitats of the lagoon has been documented (Gilmore, 1977; Gilmore et al., 1983; Snelson, 1983; Mul- ligan and Snelson, 1983), only Gilmore (1977) collected fishes from the entire system. The majority of Gilmore’s (1977) collections, however, were in the southern portion of the estuary. Although Snelson (1983) discussed differences in fish distribution between the northern and southern portions of the lagoon, his collections were only from the northern section. Furthermore, the only information available concerning ichthyofauna associated with spoil islands is from 16 islands in Indian River County (Indian River County, 1981) and 3 islands in northern Brevard County (Schooley, 1977). There are 137 spoil islands in Indian River lagoon. These islands were created between 1951 and 1961 when the Atlantic Intracoastal Waterway was dredged. They occur sporadically along 190 km of the lagoon and pro- vide shallow-water habitat for juvenile and small resident fishes. The spoil islands extend through a faunal transition zone, with its approximate center at Cape Canaveral, that is characterized by a Carolinian-influenced fauna to the north and a Caribbean-influenced fauna to the south (Briggs, 1974; 1Present address: Department of Wildlife and Fisheries, P.O. Drawer LW, Mississippi State University, Missis- sippi State, MS 39762-5917 No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA 227 BREVARD co — Ar UNTY GROUP | INDI AN 933 “RIDE SEBASTIAN INLET VER Cou! 034 NG GROUP V GROUP VI MERRITT ISLAND _INDIAN | 065 ‘RIVER COUNTY ST. LUCIE COUNTY 67° « GROUP VII PIERCE/ °83 FORT PIERCE INLET °84 ATLANTIC OCEAN GROUP III GROUP VIII ST. LUCIE INLET Fic. 1. Study sites in Indian River lagoon, Florida, indicating the 90 spoil islands sampled. Islands within each of the groups (I-VIII) were sampled once during all four seasons. 228 FLORIDA SCIENTIST [Vol. 53 Gilmore 1977). The proximity of some islands to ocean inlets provides an opportunity to examine fish assemblage differences between inlet and non- inlet areas within the same lagoonal system. Studies by Snelson and Williams (1981), Snelson (1983), Schmid and co-workers (1988) and Gilmore (1988) suggest that distribution, species richness and composition of the fish fauna in Indian River lagoon differ between inlet and non-inlet areas; but there has been no quantitative documentation of factors that may contribute to the observed ichthyofaunal differences. Furthermore, all the previous studies concentrated on either the northern or southern portions of the lagoon. Thus, there is little published information on the effect of inlets on fish distribution in the lagoon as a whole. The purposes of this study are threefold: 1) to document the juvenile and small resident fish species associated with spoil islands, 2) to investigate seasonal differences in species groups around spoil islands and 3) to determine if fish assemblages associated with spoil islands vary with the amount of tidal influence (distance) from ocean inlets. Stupy AREA AND METHops—This study encompassed 90 of the 137 spoil islands in Indian River lagoon from Haulover Canal in northern Brevard County (latitude 28°45’N) through St. Lucie Inlet in Martin County (latitude 27°10’N; Fig. 1). The study area includes three major inlets (Sebastian, Ft. Pierce, and St. Lucie) as well as areas that are not influenced by inlet water. Tidal amplitude and excursion values vary among the three inlets, with maximal values occur- ring at Ft. Pierce Inlet and minimal values at Sebastian Inlet (Smith, 1990). Each island was sampled once over a 10-mo, four-season period from October 1987 through July 1988. Since each island was sampled only once due to the large number of islands in the survey, the study area was divided into eight sections (Groups I-VIII, Fig. 1), and approximately the same number of islands from each section was sampled during each season. Four 20-m, non- overlapping seine hauls were taken parallel to the shore at each island with a 4-m x 2-m net with 3.2-mm mesh. Water depth ranged from 0.31 to 0.76 m, and non-vegetated and vegetated (Halo- dule wrightii, Syringodium filiforme, or Thalassia testudinum) bottom types were sampled around each island. Whenever possible, an equal number of seine hauls were made in the vege- tated and non-vegetated habitats. Specimens were counted and identified in the field, and voucher specimens were preserved in 10% formalin, later transferred to 60% isopropanol, and retained by the Department of Natural Resources. Species comprising at least 1% of the total number of individuals captured were assigned to two categories based on the location of spawn- ing: oceanic (nearshore) spawning species; lagoonal spawning species. These categories were determined from life-history information in Bohlke and Chaplin (1968), Hoese and Moore (1977), McClane (1978), Manooch (1984), Jennings (1985) and Robins and coworkers (1986). Fish associated with spoil islands were analyzed using Jaccard’s qualitative similarity index and UPGMA cluster analysis. Species that occurred on fewer than five islands were eliminated from the analysis. Tidal excursion values (the average distance a parcel of water travels during one tidal cycle) were calculated for 27 locations in the lagoon using the formula given in Smith (1990). The calculated tidal excursion values were used to assign spoil islands to four categories based on the length of time (number of tidal cycles) required for oceanic larvae to migrate through inlets to the islands (Table 1). Analysis of variance (ANOVA) and Student-Newman- Keul’s (SNK) multiple comparison test were performed using SPSS/PC + (SPSS, Inc. 1988). He- teroscadasticity was examined with the F,,,, test (Sokal and Rohlf, 1969). RESULTS AND Discussion—A total of 25,801 individuals representing 60 species and 32 families were collected around spoil islands during the 10-mo period of the study (Table 2). This represents a total spoil island ichthyofauna from a combination of vegetated and unvegetated habitats. The eight most common species, representing 82% of the fish captured, were mojarras (Eucinostomus spp.), pinfish (Lagodon rhomboides), striped mullet (Mugil No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA 229 TABLE 1. Classification of spoil islands based on the number of tidal cycles necessary for water from ocean inlets to reach the islands. The number of tidal cycles was calculated based on tidal excursion values from Smith (1990). Island numbers refer to islands in Fig. 1. Tidal Cycle Groups 1 2 3 4 Number of tidal cycles 1-2 3-6 7-14 >60 Islands included 33-46 24-32 15-23 1-14 67-84 47-48 49-59 86-90 60-66 85 Number of islands 37 19 20 14 cephalus), bay anchovy (Anchoa mitchilli), spot (Leiostomus xanthurus), tidewater silverside (Menidia peninsulae), ladyfish (Elops saurus), and gulf pipefish (Syngnathus scovelli). The most specious families were Cyprinodon- tidae, Sciaenidae, Gobiidae, Syngnathidae and Tetraodontidae (Table 2). We captured more species around spoil islands than either of the two previous studies (Indian River County, 1981; Schooley, 1977) that investigated fish associated with spoil islands. Both studies used similar methods, and both sampled vegetated and unvegetated areas around the islands. Indian River County (1981) found a total of 37 species around 16 spoil islands in Indian River County, 33 of which species were common to our list. Furthermore, the three most specious families found in the present study and in the Indian River County (1981) study were Cyprinodontidae, Sciaenidae, and Gobi- idae. Three of the four species not in common represented a single occurrence in the Indian River County study, while the fourth species, Harengula ja- guana, although abundant, occurred on only one island. In contrast, Schooley (1977) documented 31 species around three spoil islands near Titus- ville, 10 of which species were not collected in the present study. The most specious families in Schooley’s (1977) study were Sciaenidae, Cyprinodonti- dae and Atherinidae. Although 32% of the species Schooley found were not caught in the present study, only two of his species, Menidia beryllina and Harengula pensacolae, occurred more than once or twice. Since only 14 of the 90 islands examined in this study occurred in the northern portion of the lagoon, it is not unexpected that the species captured are less comparable to Schooley’s (1977) study than to the Indian River County (1981) work in the lower lagoon. In general, it appears that fish communities associated with spoil islands are not very different from fish communities within the lagoon as a whole. However, direct comparison of this study with several studies of grassbed fishes in the lagoon (i.e., Snyder, 1984; McNeese, 1986; Gilmore, 1988) is not completely applicable since the present study includes ichthyofauna from both vegetated and unvegetated areas. All of the species captured in the present study were considered “frequent,” “common” or “abundant” in the lagoon by Gilmore (1977) and Gilmore and co-workers (1983) with the excep- tion of Gymnura micrura, Cyprinodon variegatus, Lobotes surinamensis, Micropogonius undulatus, Chasmodes bosquianus, C. saburrae, Peprilus 230 FLORIDA SCIENTIST [Vol. 53 paru, Citharichthys spilopterus, Trinectes maculatus, Lactophyrus triqueter and Chilomycterus antillarum. Of these species, only M. undulatus was rela- tively common in the present study (Table 2). Snelson (1983), however, re- ported that M. undulatus was frequent in the northern lagoon. TABLE 2. List of fish species associated with spoil islands in the Indian River Lagoon. *indi- cates species of tropical or subtropical origin (R. G. Gilmore, pers. comm.). Species Dasyatidae Dasyatis sabina Gymnura micrura Elopidae Elops saurus Clupeidae Brevoortia spp. Engraulidae Anchoa mitchilli Batrachoididae Opsanus tau Exocoetidae Hyporhamphus unifasciatus* Belonidae Strongylura notata* Strongylura timucu* Cyprinodontidae Cyprinodon variegatus Floridichthys carpio Fundulus grandis Fundulus majalis Fundulus similis Lucania parva Poeciliidae Poecilia latipinna Atherinidae Menidia peninsulae Syngnathidae Hippocampus zosterae* Syngnathus louisianae Syngnathus scovelli Centropomidae Centropomus undecimalis* Carangidae Oligoplites saurus Trachinotus falcatus Lutjanidae Lutjanus griseus * Ocyurus chrysurus* Lobotidae Lobotes surinamensis Gerreidae Diapterus auratus * Eucinostomus spp. Haemulidae Orthopristis chrysoptera Sparidae Archosargus probatocephalus Diplodus argenteus* Lagodon rhomboides Atlantic stingray Smooth butterfly ray Ladyfish Menhaden Bay anchovy Oyster toadfish Halfbeak Redfin needlefish Timucu Sheepshead minnow Goldspotted killifish Gulf killifish Striped killifish Longnose killifish Rainwater killifish Sailfin molly Tidewater silverside Dwarf seahorse Chain pipefish Gulf pipefish Snook Leatherjacket Permit Grey snapper Yellowtail snapper Tripletail Irish pompano Mojarra Pigfish Sheepshead Silver porgy Pinfish Abundance No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA Table 2. Continued Species Sciaenidae Bairdiella chrysoura Cynoscion nebulosus Leiostomus xanthurus Micropogonius undulatus Sciaenops ocellatus Ephippidae Chaetodipterus faber Mugilidae Mugil cephalus Mugil curema Sphyraenidae Sphyraena barracuda* Sphyraena borealis Blenniidae Chasmodes bosquianus Chasmodes saburrae* Gobiidae Bathygobius soporator Gobionellus boleosoma Gobiosoma bosci Gobiosoma robustum Microgobius gulosus Stromateidae Peprilus paru Bothidae Citharichthys spilopterus Paralichthys lethostigma Soleidae Trinectes maculatus Cynoglossidae Symphurus plagiusa Balistidae Monacanthus hispidus Ostraciidae Lactophrys triqueter* Tetraodontidae Sphoeroides nephelus Sphoeroides spengleri* Sphoeroides testudineus* Diodontidae Chilomycterus antillarum * Silver perch Spotted seatrout Spot Atlantic croaker Red drum Spadefish Striped mullet White mullet Greater barracuda Northern sennet Striped blenny Florida blenny Frillfin goby Darter goby Naked goby Code goby Clown goby Harvestfish Bay whiff Southern flounder Hogchoker Blackcheek tonguefish Planehead filefish Smooth trunkfish Southern puffer Bandtail puffer Checkered puffer Web burrfish 231 Abundance 232 FLORIDA SCIENTIST [Vol. 53 Fifteen species (25%) are of tropical or subtropical origin (Table 2), al- though the majority of these species were rarely collected. Gilmore (1977), in a thorough study of the lagoon and adjacent waters, found that 28% of the fish fauna was tropical, again suggesting the spoil-island fish fauna is similar to the lagoonal fauna. Tropical species important in this study include Diplo- dus argenteus and Strongylura notata. While S. notata was found around spoil islands throughout the entire lagoon, D. argenteus was only captured in the southern portion of the lagoon. Gilmore (1977) found both of these spe- cies occasionally or frequently in the southern lagoon, but Mulligan and Snelson (1983) did not record either species in the northern Indian River lagoon. This may be due in part to differences in collection gear between our study and Mulligan and Snelson’s (1983) work, since Snelson (1983) found S. notata was occasional in his study although his data do not differentiate be- tween Indian River lagoon, Banana River, or Mosquito Lagoon. The ichthyofauna associated with spoil islands varied seasonally. Overall, Syngnathus scovelli, Eucinostomus spp., Lagodon rhomboides and Menidia peninsulae had the highest frequency of occurrence; and these species, along with Gobiosoma robustum, did not display a marked seasonality in capture frequency (Table 3). In contrast, Sciaenops ocellatus was most common in fall while Gobiosoma bosci and Mugil cephalus were frequent in the winter. Mu- gil cephalus, Leiostomus xanthurus, and Orthopristis chrysoptera were fre- TABLE 3. Seasonal percent occurrence of fishes around spoil islands. A species was included if it occurred around at least 50% of the islands in any one season. Species Total Fall Winter Spring Summer Syngnathus scovelli 89 % 67 % 86 % 92% 100% Eucinostomus spp. 83 81 81 80 83 Lagodon rhomboides 12 43 o7 96 87 Menidia peninsulae 68 57 52 76 83 Gobiosoma robustum 32 10 38 52 26 Sciaenops ocellatus 20 52 28 4 4 Gobiosoma bosci 46 43 67 36 39 Mugil cephalus 34 0 67 68 4 Leiostomus xanthurus 48 0 24 96 61 Orthopristis chrysoptera 4] 28 5 52 78 Archosargus probatocephalus 22 5 0 16 61 Strongylura notata 37 5 10 48 78 Number of islands 90 23 21 25 21 quent in spring; and L. xanthurus, O. chrysoptera, Strongylura notata, and Archosargus probatocephalus were commonly captured in summer. These differences in species composition reflect seasonal recruitment to the spoil islands and are similar to patterns observed by Schooley (1977) and Gilmore (1988). The bias of the sampling gear toward the capture of juvenile fishes further emphasizes the seasonal variability in the present study. The cluster analysis (34 species on 90 spoil islands) produced five island groupings differentiated by influence from ocean inlet waters and seasonality (Fig. 2). Group A represents islands (N = 9) associated with all three inlets No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA 233 ) 0.2 04 06 08 JACCARD’S SIMILARITY INDEX Fic. 2. Results of a UPGMA cluster analysis of 34 species from 90 spoil islands using Jaccard’s Similarity Index. Numbers 1-90 correspond to islands in Fig. 1. 234 FLORIDA SCIENTIST [Vol. 53 (1-2 tidal cycles from an inlet) that were sampled during the fall (October and November). Group B represents 22 islands sampled during winter and spring (January-April). Sixteen of these islands are directly or moderately influenced by ocean inlet waters (1-6 tidal cycles), while one sub-cluster within group B is represented by islands only slightly affected by inlets (7-14 tidal cycles away). Group C includes 27 islands sampled during spring and summer (March-July), 21 of which receive substantial influence from inlet waters (1-6 tidal cycles away). However, Group C also contains a sub-cluster of islands only slightly affected by inlet waters (7-14 tidal cycles away). Group D represents 13 islands sampled during the fall and winter (November to January), 10 of which are moderately or only slightly influenced by inlet waters (3-14 tidal cycles away from inlet). Group E (N = 14) includes the 13 northernmost islands in the study that are not associated with inlets (>60 tidal cycles away from ocean inlet waters) and were sampled from November to July. Two patterns emerge from the cluster analysis. First, while there are dis- tinctions between islands that are affected by waters from an inlet (Groups A, B and C, Fig. 2) and islands that are not (Groups D and E), the effects of seasonal recruitment of juvenile fishes appear to overshadow the inlet/non- inlet trend. For example, Sciaenops ocellatus was frequently captured only in Group A, the fall inlet cluster group (Table 4). The time of capture coincides with peak red drum recruitment (Manooch 1984) and, therefore, may mask any effects due to the proximity of the islands to inlets. Similarly, the fre- quency of capture of Elops saurus, Mugil cephalus, Leiostomus xanthurus, and Archosargus probatocephalus in groups B and C (Table 4) may also be a result of peak recruitment periods rather than strong association with inlets. Although the islands were divided into eight sampling groups (Fig. 1), and islands from each group were sampled seasonally to minimize this problem, it is evident that each island needs to be sampled over several seasons before the inlet/non-inlet trend can clearly emerge. The most obvious pattern emerging from the cluster analysis is that is- lands near Titusville and Cocoa (islands 1-14, Fig. 1) form a distinctly sepa- rate group from all the other spoil islands (Group E, Fig. 2); seasonal varia- tions are not evident. Snelson (1983) documented that the northern half of the lagoon has not only a different but also a more depauperate fish fauna than the southern half. Indeed, Floridichthys carpio and Lucania parva, two of the five most frequently captured species in Group E, were rarely found in the other four cluster groups while Eucinostomus spp., common in Groups A- D, were rarely captured on islands in Group E (Table 4). Only Syngnathus scovelli, the most ubiquitously captured species overall, occurred around greater than 50 % of islands in each of the five cluster groups. The average species richness of spoil islands in each tidal cycle group was calculated in order to examine more quantitatively the apparent effect of inlet waters on spoil island ichthyofauna (Table 5). Species richness increased significantly (p<0.05) from a low value in islands >60 tidal cycles from an No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA 235 TABLE 4. Percent occurrence of fishes around spoil islands in cluster groups as defined in Fig. 2. A species was included if it occurred around at least 50% of the islands in any one cluster. Cluster Group Species A B C D E Eucinostomus spp. 100 % 91% 100% 100% 14% Syngnathus scovelli 60 95 96 69 100 Lagodon rhomboides 90 100 100 8 21 Sciaenops ocellatus 80 32 0 38 0 Anchoa mitchilli 80 41 4] 92 ai Orthopristis chrysoptera 60 18 100 8 0 Micropogonius undulatus 10 68 0 fh 7 Elops saurus 0 73 oe 0 0 Gobiosoma robustum 0 50 37 23 36 Mugil cephalus 0 95 st 8 36 Archosargus probatocephalus 0 0 67 15 0 Leiostomus xanthurus 0 68 78 0 36 Strongylura notata 0 23 89 8 36 Gobiosoma bosci 0 4] 33 69 86 Floridichthys carpio 0 36 18 8 78 Menidia peninsulae 20 59 78 85 100 Lucania parva 0 9 15 31 100 Number of islands 10 22 27 13 14 inlet to a high value in islands located 3-6 tidal cycles from an inlet (Table 5). Species richness values of islands closest to inlets (1-2 tidal cycles) were not significantly different from richness values of islands 7-14 tidal cycles from inlets or from islands furthest from inlets (>60 tidal cycles). In general, there is a trend for increasing species richness with increasing proximity to inlet waters. This increase may likely be attributed to greater accessibility of spoil islands near inlets to oceanic species. The apparent anomaly of decreased species richness on the islands closest to inlets (1-2 tidal cycles) was also ob- served by Gilmore (1988) and Snyder (1984) in grassbed fishes in the lower Indian River lagoon, although the differences they observed were also not statistically significant. TABLE 5. Average species richness (x + 1 S.E.) of ichthyofauna associated with spoil islands in four tidal groups. Islands included in each tidal cycle group are defined in Table 1. Groups connected by solid bar are not significantly different from each other at p<0.05. Number of tidal cycles 1-2 3-6 7-14 >60 Species richness 10.1+0.42 11.0+0.68 8.9+0.82 8.2+0.56 236 FLORIDA SCIENTIST [Vol. 53 TABLE 6. Average density of fishes in four tidal cycle groups. Species are divided into two categories based on spawning area. Values represent number of fish caught per island (seined area = 320 m2). Number of Tidal Cycles from Inlet Species 1-2 3-6 7-14 >60 OFFSHORE SPAWNERS Brevoortia spp. 16.97 0.74 1.00 0.00 Diplodus argenteus 4.22 0.05 0.00 0.00 Elops saurus 22.46 6.11 1.20 0.14 Sciaenops ocellatus 14.35 0.79 0.70 0.00 Orthopristis chrysoptera 4.73 5.42 1.95 0.00 Lagodon rhomboides 4.73 64.74 41.40 0.00 Trachinotus falcatus 0.32 1.42 1.50 0.00 Archosargus probatocephalus 0.43 2.53 0.65 0.00 Leiostomus xanthurus 35.05 32.42 27.35 19.36 Mugil cephalus 37.38 36.42 24.50 30.07 Mugil curema 1.11 4.53 0.15 1.00 Micropogonius undulatus 2.05 4.32 12.70 6.07 LAGOONAL SPAWNERS Floridichthys carpio 0.22 1.89 0.05 5.00 Menidia peninsulae 3.95 32.84 31.10 93.21 Lucania parva 0.30 0.21 1.10 68.36 Gobiosoma bosci 1.05 2.53 5.25 7.00 Gobiosoma robustum 1.30 1.26 0.90 2.57 Strongylura notata 0.43 1.37 0.35 1.71 Eucinostomus spp.? 67.19 33.79 49.95 0.36 Syngnathus scovelli 5.81 22.47 11.90 19.07 Syngnathus louisianae 0.59 0.79 0.55 0.07 Anchoa mitchilli 22.78 25.79 71.50 229 Bairdiella chrysoura 0.08 9.42 1.60 0.07 Microgobius gulosus 0.16 0.00 3.75 0.07 aFucinostomus gula and E. harengula, the two most commonly collected species, are lagoonal spawners (R. E. Matheson, pers. comm.) To examine more critically the inlet phenomenon on fish distribution, the 24 most abundant species were assigned to two groups: offshore and lagoonal spawners. The average density of each species occurring around islands in each of the four tidal cycle groups was calculated (Table 6). In general, juve- niles of species that spawn offshore were found most commonly on spoil is- lands within six tidal cycles of the inlet, while lagoonal spawners were dis- tributed fairly evenly among all the islands regardless of tidal influence. The offshore spawners Brevoortia spp., Diplodus argenteus, Elops saurus, and Sciaenops ocellatus were found almost exclusively at islands within two tidal cycles of the inlets. Only three species of offshore spawners had a high aver- age density on islands greater than seven tidal cycles from inlet waters (Table 6), and all commonly occur in the upstream portions of estuaries as juveniles (Moore, 1974; Miller et al., 1984; Smith et al., 1984). In contrast, the highest densities of the lagoonal spawners Floridichthys carpio, Menidia peninsulae, Lucania parva, and Gobiosoma robustum were found on islands farthest from inlets (>60 tidal cycles). The average density of lagoonal spawners was never highest at islands one to two tidal cycles from inlets, with the exception of Eucinostomus spp. These results are not unexpected since recruitment into No. 3, 1990] BROWN-PETERSON AND EAMES—ICHTHYOFAUNA PG estuaries by larvae and juveniles spawned offshore is largely dependent upon tidal movement. Furthermore, recruitment to spoil islands by larvae and juveniles that do not actively migrate upstream is limited by tidal excursion. In contrast, recruitment of larvae and juveniles of lagoonal spawners to spoil islands is not dependent upon tidal excursion. The importance of inlets in relation to fish distribution in Indian River lagoon has been discussed by Snelson and Williams (1981), Snelson (1983), Snyder (1984), Schmid and co-workers (1988), and Gilmore (1988). All con- cur that inlets enhance ichthyofauna diversity by providing access to the la- goon for oceanic species. However, these studies neither documented statisti- cally significant differences in diversity or species richness between inlet and upstream sites nor provided a quantitative measure of the actual effect of oceanic waters entering the estuary through inlets. While there are many factors governing the influence that inlets have on fish distribution, this is the first study to relate one such quantitative factor (tidal excursion values) to observed distributional information. Additionally, this is the first study that has examined the lagoon from Haulover Canal to St. Lucie Inlet and at- tempted to quantify the effect of inlets on the fish fauna. It appears that the distribution of ichthyofauna along the spoil-island chain is affected by tidal excursion and, thus, proximity to inlets. Both species richness (Table 5) and species composition (Table 6) appear to depend upon tidal influence, support- ing the trend suggested by cluster analysis (Fig. 2). While this study provides preliminary evidence that distance from inlets is an important factor in ich- thyofauna distribution around spoil islands in Indian River lagoon, a more quantitative study that is less affected by seasonal recruitment is necessary to confirm the trends suggested here. ACKNOWLEDGMENTS— Kellie Pendoley, David Singewald, and Mary Dominic helped with the field collections. Paul Mikkelsen provided valuable assistance in computer programming, and Charissa Baker drew the figures. Ned Smith kindly shared his data on tidal excursion values. We are grateful to an anonymous reviewer for noticing a taxonomic error. Finally, Mark Peterson made many helpful suggestions throughout the course of the study and reviewed the manuscript. This study was supported by a grant from the Florida Inland Navigation District. LITERATURE CITED BouHLkE, J. E., AnD C. C. G. Cuapuin. 1968. Fishes of the Bahamas and Adjacent Tropical Waters. Livingston Publ. Co., Wynnewood, PA. Briccs, J. C. 1974. Marine Zoogeography. McGraw-Hill Book Co., New York. Gitmore, R. G., Jr. 1977. Fishes of the Indian River lagoon and adjacent waters, Florida. Bull. Fla. State Mus. Biol. Sci. 22:101-148. . 1988. Subtropical seagrass communities: population dynamics, species guilds and microhabitat associations in the Indian River Lagoon, Florida. Ph.D. dissert. Florida Inst. Technology, Melbourne. , P. A. Hastincs, anp D. J. Herrema. 1983. Ichthyofaunal additions to the Indian River lagoon and adjacent waters, east-central Florida. Fla. Sci. 46:22-30. Hogrse, H. D., anp R. H. Moore. 1977. Fishes of the Gulf of Mexico, Texas, Louisiana and Adjacent Waters. Texas A&M Univ. Press, College Station. INDIAN River County. 1981. Spoil Island Study. Privately printed, 86 pp. 238 FLORIDA SCIENTIST [Vol. 53 JENNINGS, C. A. 1985. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (Gulf of Mexico)—sheepshead. U.S. Fish. Wildl. Serv. Biol. Rep. 82(11.29). U.S. Army Corps of Engineers, TR EL-82-4. Manoocu, C. S. III. 1984. Fishes of the Southeastern United States. N.C. State Museum of Natural History, Raleigh. McCLaneg, A. J. 1978. Field Guide to Salt Water Fishes. Holt, Rinehart and Winston, Atlanta. McNEEsE, P. L. 1986. Spatial and temporal variability of fish communities inhabiting Halodule wrightii seagrass beds in the lower Indian River estuary. Masters thesis, Florida Inst. Technology, Melbourne. MILter, J. M., J. P. REED, anp L. J. Prerraresa. 1984. Patterns, mechanisms and approaches to the study of migrations of estuarine-dependent fish larvae and juveniles. Pp. 209-225. In: McCLieaVve, J. D., G. R. RANDALL, J. J. Dopson, AND W. H. NEILt (eds.), Mechanisms of oa in Fishes. Plenum Press, New York. Moore, R. H. 1974. General ecology, distribution and relative abundance of Mugil cephalus and ‘Mugil curema on the south Texas coast. Contrib. Mar. Sci. 18:241-255. MUuLLIGAN, T. J., AND F. F. SNELSON, Jr. 1983. Summer-season populations of epibenthic marine fishes in the Indian River lagoon system, Florida. Fla. Sci. 46:250-276. Rosins, C. R., G. C. Ray, anp J. Doucuass. 1986. A Field Guide to Atlantic Coast Fishes of North America. Houghton Mifflin Co., Boston. ScHOOLEY, J. K. 1977. Factors affecting the distribution of the nearshore fishes in the lagoonal waters of the Indian River, Florida. Masters thesis, Univ. Florida, Gainesville. ScHMip, T. H., L. M. EHRHART, AND F. F. SNELSON, JR. 1988. Notes on the occurrence of rays (Elasmobranchii, Batoidea) in the Indian River lagoon system, Florida. Fla. Scient. 51:121-128. SmiTH, N. 1990. An introduction to the tides of Florida’s Indian River lagoon. II. Currents. Fla. Scient. 53:72-81. SmitH, S. M., J. G. Horr, S. P. O’NemL, AND M. P. WEINSTEIN. 1984. Community and trophic organization of nekton utilizing shallow marsh habitats, York River, Virginia. U.S. Natl. Mar. Fish. Serv. Fish. Bull. 82:455-467. SNELSON, F. F., Jr. 1983. Ichthyofauna of the northern part of the Indian River lagoon system, Florida. Fla. Scient. 46:187-206. , AND S. E. WituiaMs. 1981. Notes on the occurrence, distribution, and biology of elasmobranch fishes in the Indian River lagoon system, Florida. Estuaries 4:110-120. Snyper, D. B. 1984. Species richness, abundance and occurrence of grassbed fishes from Jupiter Inlet, Florida. Masters thesis, Florida Atlantic Univ., Boca Raton. SoKAL, R. R., AND F. J. RoHiF. 1969. Biometry. Freeman and Co., San Francisco. SPSS, Inc. 1988. SPSS/PC + v 2.0 Base Manual. SPSS, Inc., Chicago, IL. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3):226-238. 1990. Accepted: March 1, 1990. Biological Sciences ADAPTIVE SPECIALIZATIONS OF THE CYPRINODONT FISH RIVULUS MARMORATUS D. Scott TayLor Indian River Mosquito Control, PO Box 670, Vero Beach, Florida 32961-0670. Asstract: The killifish Rivulus marmoratus (Poey) uses the active and abandoned burrows of the great land crab Cardisoma guanhumi as its primary micro-habitat in the salt marshes of Indian River County, Florida. Over 250 specimens have been captured from burrows, and a maximum of 26 individuals was found in a single burrow, thus representing one of the largest single-point collections recorded in Florida. At higher elevations in the marsh, the fish may be confined to burrows for up to 9 months, but a repertoire of behaviors, documented through laboratory and field observations, allows survival in this specialized environment. The impor- tance of land crab burrows in the life cycle of this fish reinforces the need for habitat protection of both species. THE mangrove rivulus (Davis, 1986), Rivulus marmoratus (Poey), was first discovered in Cuba by Poey in 1880 and redescribed from Florida by Harrington and Rivas (1958). The species designation of rivulus is still under review, but the common scientific name is used here (American Fisheries Society, 1980). This small, cryptic fish has since been recorded from various points in the Caribbean (Harrington and Rivas, 1958) and sporadically col- lected on both coasts of Florida in mangrove swamps and salt-marshes as far north as the latitude of Vero Beach (Huehner et al., 1985). Rivulus marmoratus has attracted considerable interest among fish biolo- gists because of its infrequent collection: fewer than 50 specimens on the east coast of Florida prior to the collections of this author (Taylor, 1988; W. P. Davis, pers. comm.). In addition, Harrington’s (1961) study reporting that rivulus is a simultaneous functional hermaphrodite capable of self-fertiliza- tion resulted in extensive laboratory work on the reproductive biology of the fish (Huehner et al., 1985). The ecological information published on rivulus has been limited by the fish’s relative rarity and the difficulty of making observations and collecting in the heavily vegetated habitats where it occurs. Huehner et al., (1985) summarized the collection locales in Florida and the Caribbean, and more recently Abel and co-workers (1987) and Ritchie and Davis (1986) collected rivulus from heavily vegetated temporary ponds in southwest Florida. Taylor (1988) reported the collection of 44 specimens, utilizing a novel collection technique, from burrows of the great land crab Cardisoma guanhumi in Indian River County, Florida. The ability of rivulus to tolerate water quality that may be fatal to other fish has been noted by Kristensen (1970) and Abel and co-workers (1987). Kristensen (1970) was the first to describe emersion for this species, in which the fish left pools with deteriorating water quality for a damp terrestrial Present address: Brevard Mosquito Control District, PO Box 728, Titusville, Florida 32781-0728 240 FLORIDA SCIENTIST [Vol. 53 location. Resting emersed fish are torpid and display no movement of the gills or mouth (Abel et al., 1987), but they may also travel for a considerable distance overland by flipping or slithering (Kristensen, 1970; Huehner et al., 1985). Elevated levels of hydrogen sulfide, a common toxic component of drying pools in mangrove swamps, induced rivulus to leave the water in laboratory experiments, but reduced levels of dissolved oxygen did not result in emersion (Abel et al., 1987). The fish is apparently capable of cutaneous respiration while emersed via an extensive network of capillaries in the skin and fins (Grizzle and Thiyagarajah, 1987). Emersed rivulus can survive up to 30 days in a moist environment (Abel et al., 1987). The great land crab, which figures importantly in the following study, is a large, terrestrial, tropical species, found north to central Florida on both coasts (Gifford, 1962). Scattered colonies have been observed as far north as Flagler County on Florida’s east coast (pers. obs.). Land crabs construct bur- rows up to 1.5 m deep in or adjacent to salt marshes and mangrove swamps and in some cases several kilometers inland along freshwater tributaries (Gif- ford, 1962). Burrows are dug to the depth of ground water, as the crabs must occasionally moisten their gills to respire. Water level in the burrow varies with groundwater level, rainfall, or tides in adjacent estuaries (Herreid and Gifford 1963). Land crabs occasionally seal their burrows with a mound of mud, presumably during molting. Burrows may remain sealed for months with the crab inside (D. Wolcott, pers. comm.). Laboratory research on cloning, genetics and immunochemistry have taken precedence over field studies of rivulus. This study, combining field and laboratory observations, demonstrates that rivulus can be easily collected in unprecedented numbers from the burrows of the great land crab, where the fish uses its unusual capabilities to survive in the high salt marshes of Indian River County, Florida. MATERIALS AND MErHops—Observations were conducted at land crab burrows in salt marshes of Indian River County, Florida and in a specially designed aquarium in the laboratory. Field methods: Specimens were collected and observed over a 1-yr period, September 1987- September 1988. Rivulus marmoratus were collected by a hook-and-line technique (Taylor, 1988) and a funnel trap inserted in the land crab burrows. The trap consisted of a 350-ml plastic drinking cup (73-mm diameter) fitted with a funnel (painted blue) reducing to an opening of 14- mm diameter. The cup was drilled with holes to provide water circulation. The funnel was held on with rubber bands attached to “S” hooks and clipped onto the bottom of the cup. The trap was inserted funnel down into the burrow mouth to the water level and left in the burrow overnight. In larger burrows, the trap was wrapped with a strip of foam rubber for a snug fit. Three categories of burrows were sampled: 1) active: currently containing a crab, as evi- denced by fecal pellets deposited outside the burrow and the presence of fresh tracks in the mud; 2) inactive: no fresh tracks or fecal pellets present; 3) sealed: occupied burrows which had been sealed with a mud cap. Sealed burrows were sampled by carefully removing a portion of the mud cap. Artificial burrows were dug in several salt-marsh locations known to contain rivulus to deter- mine if the fish will readily colonize new habitat. Nine burrows were dug with a common post- hole digger (13-cm diameter hole) and five with an 8-cm PVC coring device. Holes were placed a minimum of 3 m from adjacent land crab burrows. Burrows were monitored 6 mo later with dip nets or traps. Laboratory methods: Observations were made 5 d/wk (7:00 am to 4:00 pm) over the period February 1987 to August 1988 from a specially designed aquarium intended to simulate the No. 3, 1990] TAYLOR— ADAPTIVE SPECIALIZATIONS 24] burrow habitat of the land crab. This tank was constructed of glass with dimensions of 31K31X52 cm and filled with mud from a salt marsh that contained land crab burrows. The tank sub- stratum had a depth of 43 cm, with the surface of the mud sloping gradually from one corner. Two artificial burrows (10 cm diameter and 36 cm deep) were constructed along two sides of the tank in the lower corner, allowing a view of the interior of each burrow. The two burrow openings were 8 cm apart. Plantings of glasswort Salicornia virginica and young seedlings of black mangrove Avicennia germinans were distributed throughout the tank. A small amount of mangrove leaf litter was also placed on the substratum. The burrows were filled with Indian River lagoon water (salinity 25 ppt). The tank was covered with a Plexiglass sheet to maintain high humidity and illuminated with a Gro Lux fluorescent bulb on a 12 L-12 D cycle. Water temperature varied with ambient room temperature (24-28 C). Burrow water was monitored for dissolved oxygen twice weekly with a YSI Model 57 D.O. meter. A test for hydrogen sulfide using a colorimetric method (Van Handel, 1987) was performed monthly. One large (40-50 mm SL) rivulus was placed in each burrow and fed mosquito larvae throughout the observation period. Observations were recorded several times daily, and the tank was filmed from above for two 12- hr periods with a slow-motion video recorder. The following aspects of behavior were noted: 1) incidence of emersion and factors inducing it; 2) movement of the fish around artificial barriers during emersion; 3) response to varying water levels in the burrows; 4) feeding behavior; 5) reproductive behavior. An ancillary experiment was also performed to determine the ability of rivulus to survive in a burrow environment during a period without standing water. Five 4-1 plastic containers were filled with salt-marsh mud and 8-cm diameter burrows were dug. The burrows were filled with lagoon water, and two rivulus of known weight and length were placed in each burrow. Water was gradually removed over a 3-d period until only damp mud remained in the bottom of the burrows. Containers were then loosely sealed with black plastic and left undisturbed. Rainwater was added as needed but only in amounts sufficient to keep the mud damp and to allow no standing water. Two containers were reflooded with lagoon water after 44 d and the remaining three after 66 d. Fish were reweighed, and survival was determined for both intervals. ResuLts—Field observations: Extensive surveys were conducted of the salt marsh (impounded and unimpounded) in Indian River County to deter- mine the presence of land crab burrows. These observations revealed that land crabs have two distinct zones of habitation in salt marshes bordering Indian River lagoon: 1) upland areas above the high marsh, or dikes and spoil mounds—locations flooded only by extreme tides or extraordinary rainfall events; 2) high marsh zones dominated by the saltwort Batis maritima and the glassworts Salicornia virginicia and S. bigeloveii. Land crabs prefer ele- vations in the high marsh of 0.35-0.46 m above mean sea level. Burrows are rare in heavy mangrove growth and are also uncommon in impounded marshes, with the exception of some impounded marshes with breached dikes allowing normal tidal fluctuation. Burrows were examined for rivulus at the second of these two zones, the high marsh. It is not known if rivulus is found in burrows in the upland and dike zone, as these burrows are much deeper and were not sampled. Using the two collection methods described above, over 250 rivulus were collected from 16 different marsh areas in Indian River County. Fish ranged in size from 9 to 55 mm SL. No male rivulus, thought to be a result of abnormally low incubation temperatures (Harrington, 1967), were collected. The hook-and-line technique proved to be a valuable survey tool and was often used as a means of determining which burrows to use for trap place- ment. Hook and line was also effective in sampling burrows too small to accommodate traps. 242 FLORIDA SCIENTIST [Vol. 53 Rivulus marmoratus readily entered the described trap, apparently seek- ing contact with the air-water interface. Numbers of fish trapped varied from location to location. Sampling was, of course, biased to burrows of a size which would accommodate the traps used. In some locations, traps averaged 2 fish/trap with a maximum of 15 fish/trap. All three categories of burrows (active, inactive, sealed) contained rivulus, although trapping in active bur- rows was affected by the crab pushing the trap out of the burrow. Rivulus marmoratus were found in sealed burrows on several occasions, apparently in good health in spite of the potential for months of entombment. The largest single-burrow collection of 26 individuals was obtained from a sealed burrow. When sealed burrows containing rivulus were opened, the fish often milled about at the surface in a disoriented fashion, sharply con- trasting with their typically extremely secretive behavior in non-sealed bur- TOWS. Emersion of rivulus at the mouth or sides of a crab burrow was observed on several occasions. It appears that emersion may be induced both by poor water quality (water quality was not tested in burrows) or aggression be- tween fish. The adaptive value of emersion was very apparent in one instance when receding fall tidal flooding left 400-500 mosquitofish Gambusia affinis and sailfin mollies Poecilia latipinna stranded in a single burrow that had contained rivulus prior to the flooding. As the stranded fish began to die and decompose, several small (<10 mm SL) rivulus were observed resting emersed on the burrow wall above the floating corpses. Two weeks later, when all dead fish were gone, 12 rivulus, the only living fish inhabitants of the burrow, were captured. In two other burrows, disturbance of sediments from deep within the burrow released a strong hydrogen sulfide odor and resulted in the emersion of several fish. The chasing of a smaller rivulus by a larger individual resulted in emersion of the smaller fish on several occasions. However, no evidence of injury resulting from aggressive encounters (torn fins or missing scales) was observed in any of the specimens collected in the field. Rivulus marmoratus readily accepted the artificial burrows placed in the field. Both varieties (8-cm and 13-cm diameter) of artificial burrow were colonized by rivulus. Nine large (13-cm diameter) burrows in two different marshes were occupied by 13 fish 6 mo after being dug, with a maximum of 6 individuals in a single burrow. Five small (8-cm diameter) burrows in two different marshes contained three rivulus after 6 mo. Colonization took place both in artificially flooded mosquito control impoundments and in unim- pounded marshes that became minimally, if at all, flooded. These data sug- gest that rivulus will travel at least 3 m without significant standing water to seek new habitat. Laboratory results: In discussing observations made in the crab burrow aquarium, a distinction must be made between resting emersion, in which rivulus remains motionless while out of the water, and active locomotion of the fish across the damp surface by flipping or wiggling. Both these behaviors No. 3, 1990] TAYLOR— ADAPTIVE SPECIALIZATIONS 243 were frequently observed in the aquarium. The two rivulus placed in the tank each switched burrows within 72 hr. by wiggling rapidly across the 8-cm surface separating the two burrows. This behavior continued throughout the 1.5-yr observation period; one individual switched burrows up to a maximum of 6 times in 24 h. Some of this move- ment probably occurred at night, as fish were often in different burrows when observations commenced at 7:00 AM. Resting emersion, in which the fish generally left the water and remained on the side of the burrow, was also noted throughout the observation period. No direct correlation between emersion and the water quality parameters monitored could be made. Hydrogen sulfide was not detected in the burrow water with the method used (detection limit: 0.05-0.10 mg/l). Dissolved oxy- gen ranged from a minimum of 0.6 ppm when the tank was initially set up to a maximum of 3.8 ppm several weeks later. Neither extreme of dissolved oxygen seemed to affect the incidence of emersion, although both rivulus spent up to 2-3 h/d emersed upon first being placed in the tank, when dis- solved oxygen levels were noticeably lower. Later, only the smaller of the two fish was observed to rest emersed, exclusively as a result of aggressive encoun- ters when both fish were in the same burrow. Eventually, the smaller fish was removed from the tank after suffering continual harassment from the larger fish. With the single remaining rivulus in the tank, a Plexiglass barrier was placed between the two burrows to determine the ability of rivulus to move around barriers. Initially, a barrier was emplaced between the burrows with an opening between the end of the barrier and the tank wall. The 7-cm opening was located 23 cm from the occupied burrow. Within 24 h, the fish had found the opening and resumed switching burrows. When the opening was reduced to 2 cm and then 1 cm, frequency of burrow switching was not reduced. The opening was then closed and a 1-cm hole was drilled in the barrier at ground level at a location between the burrows. Within 5 d, rivulus had located the hole and resumed switching burrows. The hole in the barrier was subsequently sealed, and a slow-motion video camera was placed above the tank for a 12-h period on two consecutive days to record the attempts of rivulus to traverse the intact barrier. The fish left the water on 17 occasions during these two video-taping periods and oriented itself to the location of the sealed hole, sometimes repeatedly flinging itself at the side of the barrier. All movements were toward the inaccessible burrow. The barrier was com- pletely removed after denying the fish access to the other burrow for 1 mo. The frequency of burrow switching then reached a maximum of 10 times/d. Varying the water level in the burrows revealed that rivulus responds quickly to simulated cycles of flood and drought. While both rivulus were in the same burrow, water levels were gradually reduced over a 2-wk period until complete dry-down. Aggressive actions between the fish ceased once the burrow was dry, and both fish buried themselves close together in the damp substratum. Neither fish attempted to leave the burrow once water levels had 244 FLORIDA SCIENTIST [Vol. 53 receded to within 8 cm of the burrow opening. When burrows were suddenly reflooded after dry-down, allowing fish swimming access to either burrow and the flooded “marsh” surface, they immediately left the burrow, thereaf- ter retreating into the burrows only occasionally, especially when alarmed. Rivulus marmoratus was an active predator both in and out of the burrow over the flooded surface. When denied food for a 25-d period, rivulus gorged heavily; two fish consumed 150 mosquito larvae in a 12-h period. When the tank was flooded following burrow dry-down, rivulus foraged for mosquito larvae in water too shallow for effective swimming. Larvae were nevertheless successfully captured by rivulus, and the fish used the same wiggling motions seen when moving over the mud surface. Rivulus marmoratus was also capa- ble of jumping 11 cm into the air to obtain small insects or earthworms held in a pair of forceps. Observations on the reproductive behavior of rivulus were also made with this aquarium. Four months after initial set-up of the tank, while only one fish was present, rivulus eggs were found on the damp mud surface 23 cm away from the burrows. Up to eight eggs were laid in a 24-h period, although actual egg laying was never observed directly or recorded with the video camera. New eggs were invariably present only when the tank was first checked at 7:00 am. Ninety-five eggs were laid by this single fish over a 3-mo period, all but eight on the mud surface out of the burrow. Of these eight eggs, two were found 2.0—2.5 cm from the mud surface sticking to a strand of glasswort and the glass wall of the tank, and 6 were found inside the burrow but above the waterline on the wall of the burrow. A significant proportion (21/95) of the eggs were laid in the corner of the tank farthest from the burrows and at the highest elevation. Eggs were often found in clusters of 2— 4. No eggs were ever found in the burrow beneath the surface of the water. Fertile eggs developed and hatched on the damp mud surface. Fully de- veloped eggs would hatch either spontaneously or could be rapidly induced to hatch within 10 min with a light sprinkling of tap water. Hatchling rivulus could tolerate at least several days on the damp mud surface and would wiggle under detritus if the surface was slightly dampened with a sprinkling of water. One hatchling wiggled into a burrow 8 cm away where the parent fish ate the young. Rivulus marmoratus demonstrated an ability to aestivate that exceeds ob- servations of Abel and co-workers (1987). Three of four fish survived 44 d without water and five of six fish for 66 d. Mean weight loss over the 44-d interval was 17.7% and over 66-d, 31.4%. Discussion— Rivulus marmoratus, previously thought to be rare, was ob- served to be abundant in a specific micro-habitat, land crab burrows, in locales along Indian River lagoon. Harrington and Rivas (1958), the pioneer collector of this species in Indian River lagoon, had concluded rivulus was rare after extensive icthyological collecting with conventional techniques in mangrove swamps. Gilmore (pers. comm.) has since collected over 1.5 mil- lion fishes in mangrove swamps using a variety of nets and traps but captured No. 3, 1990] TAYLOR— ADAPTIVE SPECIALIZATIONS 245 only 3 rivulus. The crab burrow habitat explains the apparent scarcity of this fish. The crab burrow allows rivulus the opportunity to survive marsh dry- down and then be the first fish species to appear on the marsh when water levels rise during high-marsh flooding. This could provide feeding opportuni- ties to compete successfully with later arriving fish ascending from lower elevations in the marsh. The ability of rivulus to consume a considerable number of mosquito larvae demonstrates that this species could be a valuable mosquito control agent. Salt-marsh mosquito larvae, which hatch from eggs deposited in the high marsh, would certainly provide a ready food resource for starved rivulus emerging from crab burrows. The Atlantic salt-marsh snake Nerodia fasciata taeniata was observed in three land crab burrows that contained rivulus and may prey on the fish. Land crabs, which are primarily herbivores, have been recorded preying on their own young, on fiddler crabs (Uca spp.) and feeding on carrion such as dead fish (Wolcott and Wolcott, 1987). It seems doubtful, however, that land crabs could capture an elusive fish such as rivulus in the confines of the bur- row. Taylor (1988) suggests that the use of an isolating habitat such as a crab burrow may explain the evolution of self-fertilizing simultaneous hermaphro- ditism in rivulus, yet the population levels observed in some burrows (maxi- mally 26 individuals) would seem to contradict this. This contradiction may be explained by the fact that land crabs are much less abundant now along Indian River lagoon as they were in the recent past (E. J. Beidler and J. Salmela, pers. comm.). This implies loss of habitat for rivulus and the poten- tial for crowding in available habitat. Only further studies of the fish in locations where land crab populations have not been impacted could verify this. Land crabs have an obvious preference for unimpounded high marsh veg- etated with saltwort and glasswort, although some large populations were encountered in impounded marsh subject to normal tidal fluctuation through dike breaches. Crabs may avoid impounded marsh due to abnormal flooding regimes, heavy mangrove growth, or the unsuitability of the heavy peat accu- mulation for burrow construction. Indeed, test burrows dug in impounded marshes with heavy peat deposits were found to collapse and fill with sedi- ment after 2-3 mo. Given the yearly tidal cycle in this part of Florida (Provost, 1976) and the elevations at which most rivulus-containing burrows are found, it is apparent that the fish are confined to the burrows for the greater part of the year, conceivably the entire period from January to September. The aquarium ob- servations indicate rivulus is able to leave the burrow at any time and seek new habitat, either other burrows or standing water. The infrequent collec- tion of rivulus in permanent standing water clearly contrasts with the consist- ent capture of the fish in crab burrows a few meters away from standing water in mosquito control ditches. 246 FLORIDA SCIENTIST [Vol. 53 Rivulus marmoratus appears to be capable of determining topographical features in its terrestrial wandering, as the observations with the Plexiglass barrier would indicate. It is also of interest that the burrow-switching behav- ior reached a maximum after the fish was deprived of the opportunity to switch. The motive for this burrow switching is unknown. Food-seeking was not a factor, as rivulus were fed in both burrows. This behavior may indicate that the fish frequently leaves crab burrows in the wild and travels over the dry or minimally flooded marsh surface. The colonization of the artificial burrows within 6 mo tends to verify this. Emersion was seen as a response to aggressive interactions both in the laboratory and in the field. Huehner and co-workers (1985) also documented this response to aggression in an aquarium divided by an earthern berm. The cessation of aggression following dry-down of burrow water was similarly noted by Huehner co-workers (1985) in their aquarium when water levels were reduced. The survival rate of rivulus after 66 d without free water is impressive, and the real limits of this ability may greatly exceed this. Fish were immediately active following reflooding and were feeding upon mos- quito larvae with 10 min. Knowledge of the reproductive habits of rivulus is not couplers W. P. Davis (pers. comm.) reports that rivulus occasionally flip eggs out of the water in breeding bowls, and the eggs adhere to the glass covers and subse- quently develop. Egg deposition observed in the burrow tank does not follow this behavior, as no eggs, with the exception of one, were found sticking to the sides or top of the tank. The clustering of eggs in groups and heavy deposition in certain locations also indicate that rivulus is physically depositing them on the surface. Although cyprinodonts, particularly other members of the genus Rivulus, are not known for nocturnal spawning, the fact that newly depos- ited rivulus eggs were only found in the early morning hours suggests noctur- nal spawning. Rivulus marmoratus eggs have not previously been found in the wild (Ri- chie and Davis, 1986; Abel et al., 1987). Based on the aquarium observations, an extensive search was conducted in the field for rivulus eggs, but only one infertile egg was found, adhering to the wall of a land crab burrow 7 cm above the water surface. Since rivulus will eat its eggs in captivity (Abel et al., 1987) and is cannibalistic in the wild (Taylor, 1988), it seems logical that rivulus either leaves the burrow for oviposition or waits until the fall marsh flood period for major spawning. The fact that no fish smaller than 7 mm have been captured or observed in burrows lends support to both of these hypotheses. The fate of fish hatched from stranded eggs by short-term flood- ing of the marsh surface is not known, although hatchlings demonstrated an ability to survive without standing water. Perhaps these juveniles occupy an as yet unidentified micro-habitat. Much remains to be learned of this elusive fish, which apparently spends much of its life hidden from view underground. This study, in effect, raises more questions than it answers, among them the exact population status of No. 3, 1990] TAYLOR— ADAPTIVE SPECIALIZATIONS 247 the fish. The identification of the niche of rivulus in the subtropical/tropical marsh and mangrove swamp habitats along Indian River lagoon will hope- fully contribute to continued and even more stringent protection of these ecosystems. SUMMARY AND CONCLUSION—Mangrove rivulus, Rivulus marmoratus, uses the active and abandoned burrows of the great land crab Cardisoma guanhumi as its primary habitat in the salt marshes of Indian River County, Florida. Over 250 specimens have been collected from burrows with hook- and-line and traps. Rivulus marmoratus can be abundant in some burrows: 26 specimens have been collected from a single burrow. Rivulus marmoratus may be confined to some burrows for months, and burrows provide many more specimens than adjacent standing water. A variety of behavioral and physiological adaptations allows full utilization of the crab burrow habitat. The importance of land crab burrows in the life cycle of rivulus reinforces and emphasizes the need for protection of high salt-marsh and mangrove swamp habitat for both fish and crab. ACKNOWLEDGMENTS— The author wishes to thank E. J. Beidler for his interest and support of this study. Also thanks to W. P. Davis for critical review and stimulation, R. G. Gilmore for helpful input, and D. B. Carlson for editorial comments. LITERATURE CITED ABEL, D. C., C. C. KoEnic, anp W. P. Davis. 1987. Emersion in the mangrove forest fish Rivulus marmoratus: a unique response to hydrogen sulfide. Environ. Biol. Fishes 18:67-72. AMERICAN FIsHERIES SocieTy. 1980. Common scientific names of fishes. Amer. Fish. Soc. Spec. Publ. 12:1-174. Davis, W. P. 1986. The role of Rivulus marmoratus in research on aquatic pollutants. J. Amer. Killifish Assoc. 19:70-80. Girrorp, C. A. 1962. Some observations on the general biology of the land crab, Cardisoma guanhumi (Latreille), in south Florida. Biol. Bull. 123:207-223. GrizzLE, J. M., AND A. TurvacarajAH. 1987. Skin histology of Rivulus ocellatus marmoratus: apparent adaptation for aerial respiration. Copeia 1987:237-240. HarrInctTon, R. W., Jr. 1961. Oviparous hermaphroditic fish with internal self fertilization. Science 134: 1749-1750. . 1967. Environmentally controlled induction of primary male gonochorists from eggs of the self-fertilizing hermaphroditic fish, Rivulus marmoratus Poey. Biol. Bull. 132:174- too , AND L. R. Rivas. 1958. The discovery of the cyprinodont fish, Rivulus marmoratus, with a redescription and ecological notes. Copeia 1958:125-130. Herreip, C. F., II] anp C. A. Girrorp. 1963. The burrow habitat of the land crab, Cardisoma guanhumi (Latreille). Ecology 44:773-75. HuEHNER, M. K., M. E. SCHRAMM, AND M. D. Hens. 1985. Notes on the behavior and ecology of the killifish Rivulus marmoratus Poey 1880 (Cyprinodontidae). Fla. Scient. 48:1-7. KRISTENSEN, I. 1970. Competition in three cyprinodont fish species in the Netherlands Antilles. Stud. Fauna Curacao other Caribb. Isl. 32:82-101. Provost, M. W. 1976. Tidal datum planes circumscribing salt marshes. Bull. Mar. Sci. 26:558- 563. Ricuie, S. A., AND W. P. Davis. 1986. Evidence for embryonic diapause in Rivulus marmoratus: laboratory and field observations. J. Amer. Killifish Assoc. 19: 103-08. Tay.or, D. S. 1988. Observations on the ecology of the killifish Rivoulus marmoratus (Cyprino- dontidae) in an infrequently flooded mangrove swamp. N.E. Gulf Sci. 10:63-68. VAN HANDEL, E. 1987. A sulfide detection test for field use. Mosq. News 3:644. 248 FLORIDA SCIENTIST [Vol. 53 Wo tcott, D. L., AND T. G. Wotcotr. 1987. Nitrogen limitation in the herbivorous land crab Cardisoma guanhumi. Physiol. Zool. 60:262-268. From the SECOND Indian River Research Symposium, Marine Resources Council, 12-13 September 1988, at Florida Institute of Technology, Melbourne. Florida Sci. 53(3):239-248. 1990. Accepted: May 5, 1989. Biological Sciences THE LARGE SPATIAL AND TEMPORAL BIOLOGICAL VARIABILITY OF INDIAN RIVER LAGOON! ROBERT W. VIRNSTEIN St. Johns River Water Management District, P.O. Box 1429, Palatka, FL 32178 Asstract: The biological variability of the Indian River lagoon system is huge, whether examined over distances from 100 km to centimeters or times from centuries to hours. Biotic communities in the northern end of the lagoon are temperate, whereas those in the southern part of the lagoon are tropical—e.g., Spartina marshes dominate in the north, and mangrove forests and seagrass in the south. The fish fauna of the northern part of the lagoon system is strikingly depauperate when compared to that of the southern. Over distances of only a few kilometers or even meters, species composition can change substantially, and densities can vary by two to three orders of magnitude. The lagoon of a few centuries ago was quite different from the lagoon of today. Large changes in submerged vegetation have occurred within the last decade. Changes between years often far exceed and, thus, obscure “seasonal” patterns. In the same habitat, samples taken one day can be very different from samples taken the next day or night. Some of these differences are explainable; others are not. Temporal and spatial variability, instead of being a disadvantage, can be used to identify the processes that produce the variability. Because of the huge variability of the lagoon system, samples from one area or one time period necessarily give an incomplete and often incorrect picture of the lagoon. It is unjustified to assume that a limited study represents biotic conditions in the entire lagoon. I must conclude that a “typical” or “representative” site or time period does not exist. To be representative, a sampling program must reflect the diversity of the system. ONE aspect of the Indian River lagoon system continues to astound me— the huge variability of biotic systems in both space and time. This paper describes biological variability of this lagoonal system and the need to con- sider this variability in establishing sampling design and interpreting data. This paper is based on the literature and my 13 yr experience working on the lagoon. Some examples are merely observational or anecdotal but were cho- sen to illustrate the range of variation that might be expected at a certain scale of space or time. For both spatial and temporal variability, I present examples ranging from large scale to small. The scales I have chosen span a range of six to seven orders of magnitude—spatially from hundreds of kilometers to centimeters and temporally from centuries to hours. Examples are presented in that or- der. To indicate the range and magnitude of biological variability that may be encountered, I have often chosen extreme examples. Once we know that variability exists, we must ask what the implications are. In any effort to characterize the biology of a system, certain assumptions must be made. For example, one almost always makes the tacit assumption that samples can be taken that are “representative” of the system under study. Such an assumption usually is unwarranted. The appropriateness of the as- sumption depends on a knowledge of the variability of the system and the consideration of this variability in the design of a sampling program and 1$.E.A. contribution no. 10 of Seagrass Ecosystems Analysts, Inc., Vero Beach, Florida. 250 FLORIDA SCIENTIST [Vol. 53 interpretation of data. Unless such consideration is made, we should be wary of the interpretation of results from the sampling program. I hope this paper will demonstrate that consideration of variability is especially necessary for the Indian River lagoon system, where large differences in biotic diversity exist between neighboring latitudes (Virnstein et al., 1984). SPATIAL VARIABILITY—The huge spatial variability of the Indian River lagoon system is largely due to its geographic location. The lagoon system spans 260 km along the east coast of Florida and includes a major biogeo- graphic transition zone from warm-temperate to subtropical (Gilmore, 1977). Figure 1 illustrates that minimal ocean water temperatures within only a few kilometers of lagoon coastline differ as much as along hundreds of kilometers of coastline further north. Specifically, minimal temperature dif- fers 8°C along the coast of the lagoon but only 2°C from northern Florida to southern North Carolina (Fig. 1). Thus, the northern part of Indian River lagoon is primarily warm-tem- perate; the southern part is primarily subtropical. It is, therefore, not surpris- ing that the biological systems are quite different at the geographic extremes of the lagoon. One obvious example is the shoreline plants. Marshes in the northern part are dominated by temperate grasses (Spartina alterniflora in high-salinity areas), whereas marshes in the southern part are dominated by mangroves. Submerged plants also show large latitudinal differences. At the northern end of the lagoon system (north of New Smyrna Beach, Fig. 2), there is no seagrass; in the southern part, there are seven species of seagrass. Only two of these species are found north of the lagoon, and then not until North Carolina. Because macrophytes often define habitats, associated ani- mal communities likewise show huge latitudinal differences in composition. At a scale of 100-200 km, Snelson (1983) concluded that, in comparing fish communities, “it is clear that the northern part of the lagoon is strikingly depauparate when compared with the southern part.” Nearly 200 species reported from the southern part of the lagoon system (Gilmore, 1977; Gilmore et al., 1983) were not collected in Snelson’s 1,800 collections of fish from the northern lagoon system. The southern fauna includes more fresh- water species, more reef-associated species, and more species with oceanic and tropical affinities (Snelson, 1983). Salinities are more stable in the south- ern half of the lagoon. A latitudinal gradient also exists for seagrass-associated macrobenthos. Young et al. (1976) reported average densities of 14,200/m? just northwest of Haulover Canal, 6,600/m? just north of Link Port, and 4,000/m? just north of St. Lucie Inlet (Fig. 2). At a smaller scale of 10 km, Gilmore and co-workers (1981) found differ- ent dominant species of seagrass-associated fishes at Fort Pierce Inlet com- pared to those at Link Port (Fig. 2). Fouling communities also differ. Com- munities on hard substrata at Fort Pierce Inlet are dominated by stenotopic colonial forms, whereas eurytopic solitary forms dominate areas north of the inlet. Number of species of fouling organisms decreases from 32 at Fort Pierce Inlet to 21 at Link Port, 14 at Vero Beach, and 9 at Eau Gallie (Mook, 1983) (Fig. 2). No. 3, 1990] VIRNSTEIN —BIOLOGICAL VARIABILITY 725) | ee ee ect! oO @ 6. © e < a Canaveral 16 Fic. 1. Map of Florida and southeast US coast showing mean minimal monthly temperature (°C) along the coast, illustrating a steep gradient of ocean temperature in the vicinity of Cape Canaveral and throughout the coast of the Indian River lagoon system. Arrows indicate direction of ocean currents. At the kilometer scale, Virnstein and co-workers (1983) sampled inverte- brates of seagrass beds at Link Port and at a site 2 km due north (Fig. 2). In the same number of samples taken over the same time period by the same method at both sites, there were 3,258 Modulus modulus and 3,051 Cerith- ium muscarum at the Link Port site but only 1 and 0, respectively, of these gastropods at the other site. Conclusions about dominant gastropod species would differ vastly, depending on the site chosen for study. 2o2 FLORIDA SCIENTIST [Vol. 53 3% PONCE DE LEON INLET New Smyra'| ® Beach | ee: =e \-ST. LUCIE a A \ INLET JUPITER A INLET Fic. 2. Map of the Indian River lagoon system. Major features and place names referred to in the text are included. No. 3, 1990] VIRNSTEIN— BIOLOGICAL VARIABILITY 253 At the meter scale, seagrass beds at the site north of Link Port contained three times the density of macrobenthic invertebrates found in unvegetated sediments only a few meters away (Virnstein et al., 1983). Epifaunal abun- dance was 13 times greater in seagrass than in sand. This obvious influence of habitat is understandable, knowing that seagrass beds provide increased food and refuge. But, even within the same seagrass bed, dipnetting for pa- laemonid shrimp can yield few in one area, whereas it can yield several times as many shrimp with the same sampling effort only a few meters away (Virn- stein, personal observation). At a scale of centimeters, two core samples taken next to each other in an apparently homogeneous habitat (e.g., a mud bottom or a continuous and apparently uniform grass bed) often differ in density of macrobenthic inver- tebrates by a factor of two to three. For four replicate cores, the standard deviation of density often exceeds the mean (e.g., Nelson, et al. 1982, p. 125). For individual species, variations can be even greater. TEMPORAL VARIABILITY—Geologically, Indian River lagoon is young, hav- ing existed as an estuarine lagoon for only a few thousand years (Almasi 1983; Stauble, in prep.). Over this time period, a continually rising sea level and opening and closing of several inlets have undoubtedly changed the lagoon. Since discovery of the New World less than 500 yr ago, sea level has risen a meter. Because 1 m is about the average water depth of seagrass beds in the lagoon, most of the present seagrass beds are at most only a few centuries old (Virnstein, in prep.). Because the average depth of the lagoon is only 1.5 m, a 1-m change in depth must have had profound impacts on salinity, hydrody- namics, marshes, seagrasses, and all associated animal communities. The na- ture and magnitude of most changes are, however, unknown. Besides sea level changes, other abiotic factors may affect the lagoon ir- regularly every few decades. The lagoon area has not experienced a severe hurricane for several decades now. In January 1977 and December 1989, it snowed throughout the lagoon region, and water temperatures in the lagoon were as low as 5°C (Gilmore et al., 1978; Snelson and Bradley 1978; Virn- stein, personal observation). In contrast, most recent winters (1985-1988) were mild. During the severe winters, much of the seagrass defoliated, leav- ing sediments exposed except for the stubble of short shoots. During milder winters, seagrass density decreased but retained a canopy of leaves (Virn- stein, personal observation). In the 1950s and 60s, most (13,000 ha) of the marshes of the lagoon were impounded (separated from the lagoon by dikes) for mosquito control (Hoff- man and Haddad, in prep.; Rey and Kain, 1989). The isolation of these marshes from the lagoon ecosystem has undoubtedly had a huge impact on those animals that depend on these marshes, e.g., for spawning and nursery areas (Gilmore et al., 1982). Also on the scale of decades, there have been large changes in composition and density of submerged vegetation. The attached green alga Caulerpa pro- lifera, a seagrass look-alike, has apparently proliferated from scattered plants 254 FLORIDA SCIENTIST [Vol. 53 in the early 1980s to submerged meadows covering about 100 km’ of lagoon bottom in 1986 (White, 1986). In the past few years, three species of Ha- lophila seagrass have also proliferated. Halophila englemannii is common in the northern part of the lagoon (White, 1986). In the southern part of the lagoon in 1986 and 1989, H. johnsonii and H. decipiens covered several square kilometers of lagoon bottom (Virnstein and Cairns, 1986; Virnstein, personal observation). In addition, unattached “drift” algae were very abun- dant in the Link Port area in winter/spring at least during 1978-1980 (Virn- stein and Carbonara, 1985) but were nearly absent during 1983-1988 (Virn- stein, personal observation). Over decades, animal populations also change. In the mid-1970s, the si- punculan worm Phascolion cryptus, which lives in gastropod shells, occurred at densities of a few thousand/m? at Link Port (Young and Young, 1977; Rice et al. 1983). In the 1980s, densities were only a few hundred/m? or less (Virn- stein, unpublished data). Seasonal variability can be large, but it is often masked by huge changes between years. Seasonal maximal density of seagrass-associated amphipods was 68 times the minimum, based on the average of five sites (Nelson et al., 1982). The seasonal pattern within 1 yr was, however, rarely repeated in subsequent years, especially in the southern part of the lagoon (Nelson et al. 1982). Months of peak abundance within a given year were rarely the same at any of the five sites, even for the most closely adjacent sites. Sampling schedules rarely measure biological variability on time scales shorter than a month, but there are a few examples from studies of seagrass- associated invertebrates. Clumps of defaunated algae, seagrass, or artificial seagrass are colonized rapidly, with abundance and diversity of motile epi- fauna (primarily amphipods and gastropods) reaching equilibrium levels within 1 wk (Virnstein and Curran, 1986). Even within a few hours, there can be large biological changes. Tidal elevation, for example, is obviously important to an organism that forages within the intertidal zone. And for many species of motile epifauna in sea- grass beds, most individuals are found at the sediment surface during the day and within the leaf canopy at night (Howard, 1988). There are also large diel changes in the fish fauna within seagrass beds (Gilmore and Donohoe, 1982; Gilmore, unpublished data), yet few sampling programs include nighttime sampling. Within a 0.6-m? patch of seagrass, Howard (1985) found that >50% of the individuals of a caridean shrimp were replaced by other indi- viduals within 3 h at night. Turnover rates of gastropods were lower, but some species approached 50% turnover within 6 h. IMPLICATIONS AND APPLICATIONS— Based on this huge variability over all selected scales of distance and time, I must conclude that there is no such thing as a “typical” or “representative” site or time period. No site can repre- sent the entire lagoon. But what do we do? How do we handle such huge variability over all scales? How do we set up a monitoring program that represents the biological No. 3, 1990] VIRNSTEIN— BIOLOGICAL VARIABILITY 255 conditions in the lagoon system? The only answer is to match the diversity of the biological system with comparable diversity in the sampling program. That is, the sampling program should be as diverse as the system under study. To represent the system fully, all habitat types should be sampled. And if diurnal, seasonal, and long-term trends are to be described, sampling must be spread over a schedule that covers all these time scales. Uniformly spaced samples can only measure variability at the shortest distance or time interval. If this diversity of sampling is impractical (e.g., for logistic or economic reasons), then hard decisions must be made about what scales are appropriate for answering the questions posed. For example, is it more important to de- tect seasonal or annual patterns? If only long-term trends are wanted, then sampling effort should be randomly allocated within years rather than uni- formly scheduled. One strategy I recommend is that investigators take advantage of varia- bility to understand or identify processes rather than to consider such varia- bility a problem. For example, if communities in two areas differ, don’t ask, “Why?” Rather ask, “How?” That is, ask what processes could produce such a difference, and then design experiments to distinguish among the alterna- tives. Variation is an inherent part of all estuarine systems and must be incor- porated into sampling design and interpretation. For the Indian River lagoon system, with its huge variability over all distance and time scales, such con- siderations are absolutely necessary. Assuming that a set of samples is repre- sentative of the lagoon is both misleading and foolhardy. LITERATURE CITED Autmas!, M. N. 1983. Holocene sediments and evolution of the Indian River (Atlantic coast of Florida). Ph.D. dissert. Univ. Miami, Coral Gables. Gitmore, R. G. 1977. Fishes of the Indian River lagoon and adjacent waters, Florida. Bull. Fla. State Mus. Biol. Sci. 22:101-147. , L. H. Buttock, ANnp F. H. Berry. 1978. Hypothermal mortality in marine fishes of south-central Florida, January, 1977. Northeast Gulf Sci. 2:77-97. , D. W. Cooke, anp C. J. DoNoHOoE. 1982. A comparison of the fish populations and habitat in open and closed salt marsh impoundments in east central Florida. Northeast Gulf Sci. 5:25-37. ., AND C. J. DoNouoe. 1982. Diel variation in seagrass bed fish populations. Fla. Scient. 45(Suppl. 1):27. —_______, D. W. Cooke, anp D. J. Herrema. 1981. Fishes of the Indian River lagoon and adjacent waters, Florida. Tech. Rep. No. 41, Harbor Branch Foundation, Inc., Ft. Pierce. , PA. HastTINGs, AND D. J. HerRrREMA. 1983. Ichthyofaunal additions to the Indian River lagoon and adjacent waters, east-central Florida. Fla. Scient. 46:22-30. HorrMan, B. A., AND K. D. Happap. (In prep.). Land use changes in the Indian River Lagoon region. In: Indian River Lagoon Monograph. Howarp, R. K. 1985. Measurements of short-term turnover of epifauna within seagrass beds using an in situ staining method. Mar. Ecol. Prog. Ser. 22:163-168. . 1988. Diel variation in the abundance of epifauna associated with seagrasses in the Indian River lagoon, Florida. Mar. Biol. 99:37-43. Mook, D. H. 1983. Indian River fouling organisms: A review. Fla. Scient. 46: 162-167. Netson, W. G., K. D. Cairns, ANp R. W. VirnsTEIN. 1982. Seasonality and spatial patterns of seagrass-associated amphipods of the Indian River lagoon, Florida. Bull. Mar. Sci. 32:121-129. 256 FLORIDA SCIENTIST [Vol. 53 Rey, J. R., AND T. Kain. 1989. A guide to the salt marsh impoundments of Florida. Florida Medical Entomology Laboratory, Vero Beach. Rice, M. E., J. Prraino, AND H. F. Retcuarpr. 1983. Observations on the ecology and reproduc- tion of the sipunculan Phascolion cryptus in the Indian River lagoon. Fla. Scient. 46:382- 396. SNELSON, F. F., Jr. 1983. Ichthyofauna of the northern part of the Indian River lagoon system, Florida, Fla. Scient. 46: 187-206. , AND W. K. Brapb.ey, Jr. 1978. Mortality of fishes due to cold on the east coast of Florida, January, 1977. Fla. Scient. 41:1-12. STAUBLE, D. K. (In prep.). Geological history of the Indian River Lagoon. In: Indian River Lagoon Monograph. ViRNSTEIN, R. W. (In prep.). History of seagrasses in the Indian River Lagoon. In: Indian River Lagoon Monograph. , AND K. D. Cairns. 1986. Seagrass maps of the Indian River lagoon. Report to Florida Department of Environmental Regulation. Seagrass Ecosystems Analysts, Inc., Vero Beach. , AND P. A. CarBonara. 1985. Seasonal abundance and distribution of drift algae and seagrasses in the mid-Indian River lagoon, Florida. Aquat. Bot. 23:67-82. , AND M. C. Curran. 1986. Colonization of artificial seagrass versus time and distance from source. Mar. Ecol. Prog. Ser. 29:279-288. , P. S. MIKKELSEN, K. D. Cairns, AND M. A. Capone. 1983. Seagrass beds versus sand bottoms: The trophic importance of their associated benthic invertebrates. Fla. Scient. 46:363-381. , W. G. NEtson, F. G. Lewis, III, anp R. K. Howarp. 1984. Latitudinal patterns in seagrass epifauna: Do patterns exist, and can they be explained? Estuaries 7:310-330. Wulite, C. B. 1986. Seagrass maps of the Indian and Banana Rivers. Report to Florida Depart- ment of Environmental Regulation. Brevard County Office of Natural Resources Man- agement, Merritt Island. Youn, D. K., M. A. Buzas, AND M. W. Younc. 1976. Species densities of macrobenthos associ- ated with seagrass: A field experimental study of predation. J. Mar. Res. 34:577-592. , AND M. W. Younc. 1977. Community structure of the macrobenthos associated with seagrass of the Indian River estuary. Pp. 359-382. In: Cou.., B. C. (ed.), Ecology of Marine Benthos. Univ. South Carolina Press, Columbia. 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Isolation and Structure Elucidation of A*-9, 11-seco-Gorgostene-38,11 248- triol-9-one from the Gorgonian (Pseudopterogorgia americana Ne OE ee rca el Gon Rs ge ios Gd aw oa oo wa eal M. J. Musmar and Alfred J. Weinheimer 257 OT ELS Sg Sn ee Richard Franz 262 LD LEE a aa er Richard Wunderlin 262 Reemee etme Rey IDCer 2... ct eee neces ees W. D. Klimstra and Allan L. Dooley 264 Scale Development in Ovuliferous Cones of Pinus Elliottii Lil, SOLES Ge ee area rarer R. F. Mente and S. D. Brack-Hanes 274 Distribution of the Eastern Chipmunk (Tamias striatus) in SLT LEL sso. sey Oh ERR Ns ae ee een Jeffrey A. Gore 280 A Review of the Florida Crayfish Fauna, with Comments on Nomenclature, Distribution, and Conservation ................ Richard Franz and Shelley E. Franz 286 Meaningful Environmental Data: Not Just the Laboratory’s Responsibility .....................0. Donald C. Anné 297 Ichthyofaunal Evaluation of the Peace River, Florida ............. Thomas R. Champeau 302 Behaviorial Characteristics of Squirrel Monkeys at the Bartlett Estate, Ft. Lauderdale............... Ryan J. Wheeler 312 ana PMENORRNC Sie ee aie GL aia os aoa ve Sw ww BG 317 SS I IEE IEE ET TT TI IE EI TY IEEE TEE AIT IE IE LOE IE EEO EEE EEE ENE EEE SEE QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright© by the Florida Academy of Sciences, Inc. 1990 Editor: Dr. DEAN F. MARTIN Co-Editor: Mrs. BARBARA B. MARTIN Institute for Environmental Studies Department of Chemistry University of South Florida Tampa, Florida 33620 THE FLoripDA SCIENTIST is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to indi- viduals or institutions interested in supporting science in its broadest sense. Applica- tions may be obtained from the Executive Secretary. 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Officers for 1990-91 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. FREDERICK BUONI Treasurer: Dr. ANTHONY F. WALSH Operations Research Program 5636 Satel Drive Computer Science Department Orlando, Florida 32810 Florida Institute of Technology Melbourne, FL 32901 Executive Secretary: Dr. ALEXANDER DICKISON Department of Physical Sciences President-Elect: Dk. GEorcE M. Dooris Seminole Community College Division of Science and Mathematics Sanford, Florida 32771 St. Leo College St. Leo, FL 33574 Program Chair: Dr. DEL DELUMYEA Millar Wilson Laboratory for Chemical Research Secretary: Dr. Patrick J. GLEASON Jacksonville University 1131 North Parkway Jacksonville, FL 32211 Lake Worth, Florida 33460 Published by the Florida Academy of Sciences, Inc. 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida 32602 Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BarBaARA B. Martin, Co-Editor eee reer LK Volume 53 Autumn 1990 Number 4 A EEN CU ee ee heey aces oe I 5, Eo ee eee Chemical Sciences NOVEL MARINE STEROIDS FROM THE FLORIDA KEYS. I. ISOLATION AND STRUCTURE ELUCIDATION OF A’-9,11-SECO-GORGOSTENE- 36,11,246-TRIOL-9-ONE FROM THE GORGONIAN (PSEUDOPTEROGORGIA AMERICANA (GMELIN)) M. J. MusMar’, AND ALFRED J. WEINHEIMER Department of Medicinal Chemistry, University of Houston, Houston, TX 77004 USA Asstracr—A large number of sterols has been characterized from marine organisms, and rela- tively few polyoxygenated steroids have been characterized due to their complexity. The gorgonian Pseudopterogorgia americana, was shown to contain several novel steroids that will be published in this journal. Ample spectral evidence, including X-ray crystallography, has been collected to sup- port the proposed structure of the title steriod. THE gorgonian, P. americana is a rich source of novel steroids. The A°-9,11- secogorgostene -36,11-diol-9-one was characterized, along with the corres- ponding 5,6a-epoxy derivative (Spraggins, 1970). Subsequent work (Haertle, 1971) led to the discovery of the new gorgost-5-ene-36,9,11-triol, which was converted to the 9,11-seco steroid previously isolated by Spraggins (1970). Re- cently a group of Australian workers (Kazlauskas, et al. 1982) reported the isolation of 36,11-dihydroxy-24-methyl-9,11-seco-cholest-5-ene-9-one and 3, 11-dihydroxy-24-methly-9, 11-seco-cholest-5-ene-9-one from a soft Coral Sinularia sp. Triusubstituted 22,23-cyclopropyl moieties have been encountered in a number of marine sterols since the first report describing the occurence of this unique structural feature in gorgosterol (Hale, et al., 1970) and acanthasterol (Skeikh, et al., 1971). Among the interesting features associated with the 22, 23- cyclopropyl unit is the dramatic shielding of the H24 resonance which was first confirmed by an autocorrelated two-dimensional proton (COSY) spectrum ob- tained for 5a,6a-epoxy-3,11-dihydroxy-9,1l-seco-5 -gorgostan-9-one 3,11- diacetate (Musmar, et al., 1983). As a part of our continuing studies on the sterols of marine invertebrates, specifically from the gorgonian P. americana, we would now like to report the isolation and structure elucidation of A®-9,11-seco-gorgostene-36, 11,246-triol- 9-one (Fig. 1) which crystallizes with two different forms in the unit cell. Present address: Department of Chemistry, University of South Florida, Tampa, FL 33620. USA. 258 FLORIDA SCIENTIST [Vol. 53 HO Fic. 1. A5-9,11-Seco-Gorgostene-36,11,246-triol-9-one. MATERIALS AND METrHops—General Methods: Melting points were taken on a Thomas Hoover capillary melting point apparatus and were not corrected. IR spectra were taken on a Perkin-Elmer Infrared Spectrometer 283 by using KBr pellets. |'H-nmr spectra were recorded on different nmr spectrometers, 100, 400 MHz at our Laboratories and at cooperating laboratories. CDC13 was used as a solvent for all !H-nmr and C-nmr spectral data. The chemical shifts are expressed in 5-values (ppm) relative to internal TMS. Coupling constants are reported in Hz. The 13C-nmr spectra were obtained on a Varian Associates Model XL-100 Fourier Transform Spec- trometer operating at 25.2 MHz, equipped with a Nicolet 1180 computer through a model 293A pulse programmer. High resolution mass spectra were provided by C. E. Costello, Chemistry Department, Massachusetts Institute of Technology, Cambridge, Mass. Analytical TLC was per- formed on precoated silica gel plates. Collection and Extraction: The gorgonian was collected on the coral reefs off the Florida Keys. This gorgonian grows firmly attached to the bottom and was cut from the reef with prun- ing shears, snipped into small pieces. Three pails containing 23 kilograms (wet basis) of the intact gorgonian were used. The gorgonian was soaked a few days in sea water/methanol, which was filtered. Fresh methanol/isopropanol (1:1) was added, and then filtered on the second day. The gorgonia was macerated daily with fresh solvent over a period of one week. The combined filtrates were concentrated under reduced pressure, and the water was removed by lyophilliza- tion. The resulting freeze-dried sample weighed 1136 g. Isolation of A5-9,11-seco-gorgosten-36,11,246-triol-9-one: 291.3 g of the solid freeze-dried aqueous alcoholic extract was treated with a mixture of 1.5 L of ethyl acetate and 150 mL of isopropyl alcohol. After stirring for 30 minutes, the solution was filtered under vacuum. The residue, following decantation, was extracted with more fresh ethyl acetate (350 mL). The sol- vent was then removed under reduced pressure, and the residue, weighing 30 g, was dissolved in 1L methanol. 20 mL of water was added to the methanol solution which was extracted with hexane three times, 200 mL each. An additional 270 mL of water was added, and the methanol solution, now 22.5% water (v/v) was extracted with 4 x 400 mL portions of chloroform. The hexane soluble material weighed 1.87 g, and the chloroform soluble material weighed 25.7 g. The freeze-dried material, which did not dissolve in ethyl acetate/isopropy! alcohol mixture and weighed 261.3 g, was disolved in 500 mL 10% HCl, filtered, and immediately extracted with chloroform four times, 200 mL each. The residue, following the removal of the chloroform was 6.5 g of an oily, viscous, red-colored material. The combined chloroform soluble material weighed 32.2 g and was chromatographed on 300 g of silica gel. The column was eluted in step gradients using the following series of increasing solvent polarities, and collecting 500 mL frac- tions: chloroform (fractions 1-4), 1% methanol in chloroform (fractons 5-8), 2% methanol in chloroform (fractions 9-11) and 7% methanol in chloroform (fractions 12-15). Preliminary TLC work indicated that the fractions 5 through 14 contained the desired com- pound. The combined fractions weighed 20.5 g and were chromatographed a second time on 360 g of silica gel using the solvent mixture ethyl acetate/hexane (3:7). Some 48 fractions were col- lected (70 mL each), followed by 22 more fractions (100 mL each). Fractions 52 through 59, weighing 2.78 g, were combined and chromatographed on silica gel using 25% acetone in hex- ane, collecting fractions of 70 mL each. Fractions 12 through 16 weighing 370 mg contained the No. 4, 1990] MUSMAR AND WEINHEIMER— NOVEL MARINE STEROIDS 259 sought compound. Following crystallization and recrystallization from acetone, 70 mg of pure material was obtained. The low-resolution mass spectrum, determined at 70 eV, showed the M-H,O at m/z 456 (2.9), and the fragments, m/z 441 (4.5, M-H,O-CHs3), 431 (1.6, M-C,H;), 413 (5.9, M-C3H,-H,O), 385 (4.3, M-C;H,;-H,O), 345 (45.4, M-H,O-C,H);), 331 (41.3, M-H,O- C,H,;), 313 (11.1, M-2H,O-C,H,7), 302 (9.1, M-C,H;9-CH3-CgHj,), 287 (6.8, M-2H,O-C,,H)j,), and 179 (13.6 for rings A and B). Other ions appeared at m/z 161 (39.7), 147 (42.9), 135 (41.5), 120 (100), and 107 (88.8). The ir spectrum (KBr) showed peaks at 3400, 2980, 2970, 2890, 1715 and 1705 unresolved, 1640, 1470, 1450, 1380, 1370, 1055, 970, and 820 cm—'. The proton noise decoupled 3C-nmr spectrum showed absorptions at 6= 217.28, 140.37, 121.33, 74.42, 71.34, 59.09, 50.26, 48.21, 45.66, 43.03, 41.70, 40.62, 40.52, 34.70, 34.49, 32.51, 31.09, 30.75, 28.57, 97.70, 25.94, 24.10, 22.89, 20.48, 48.17, 48.17, 17.19, 17.00, and 6.13. The !H-nmr spectrum at 400 MHz showed absorptions at 6=5.46 (1,d,J =5.6 Hz), 3.85 (1,m), 3.71 (1,m), 3.48 (1,m), 3.03 (1,dt), 1.35 (3,s), 1.17 (3,s), 1.04 (4,s), 0.97 (3,s), 0.88 (6,d, J=6.8 Hz), 0.83 (1,dd, J=4.5, 9.5 Hz), 0.67 (4,s), and —0.24 (1,dd, J =4.6, 5.9 Hz). Weal 140 120 1d0 go Fic. 2. The 25 MHz proton noise decoupled 3C-nmr spectrum (CDCl;) of A®-9,11-seco- gorgostene-36,11,246-triol-9-one, V, showing the region 0-140 ppm. RESULTS AND Discusslon—Among the fascinating structural features in this group of steroids is the presence of 22,23-cyclopropyl ring, the quater- nary nature of C23, the dramatic shielding of H24, and the overall stereo- chemistry of 22R, 23R, 24R (Faulkner, et al., 1988). The A®-9,11-seco-gorgos- tene-38, 11,246 triol-9-one was isolated in analytically pure form and crystallized from ether. The structure was determined by collective and am- ple spectral evidence, including comprehensive x-ray study (Musmar, 1983), which showed that the compound exists in two forms in the crystal state with extensive inter- and intra-molecular hydrogen bonding. The molecular for- mula was determined by High Resolution Mass Spectrometry (HRMS) analy- sis using, Fast Atom Bombardment (FAB) that showed a weak peak at m/z 567 corresponded to (475 (m+1) + glycerol)*:. Based on this and other spec- tral evaluation, the formula (C,,H,,.0,) was thus adopted for the molecular ion. Fragment ions with molecular compositions of (C,H,,0;, C3H,0., and C,,H,,0) were also established on the basis of the high resolution mass spec- trum, these ions representing the successive loss of three molecules of water. 260 FLORIDA SCIENTIST [Vol. 53 a en nn i) Fic. 3. The 400 MHz 'H-nma spectrum (CDCl,) of A5-9,11-seco-gorgostene-36, 11,24-triol- 9-one, V. The proton noise decoupled “C-nmr spectrum (Fig. 2) showed a resonance at 6 = 217.28 (carboxy, C9), two SP’ carbons, resonating at 6140.37 and 121.34 (C5 and C6, respectively). Three SP? carbons C24, C3 and C11 attached \o oxygens) at 6=74.42, 71.35, and 59.10, respectively. The 'H-nmr spectrum (Fig. 3) showed H6 at 6=5.46 as a double with apparent J =5.6 Hz. The two multipletes (C11-methylene protons) observed at 6=3.85 and 3.71 were mag- netically inequivalent due to intramolecular hydrogen bonding to the oxy carbon C9 which appear as a split ketone absorption in the ir spectrum (KBr) occurring at 1700 cm~'. These observations may be considered as characteris- tic features of the C9-one-C11-01 steroids. Further, diagnostic ions in the mass spectrum having the elemental composition of (C,,H,,O, and C,,H,,O,) allowed the inference of the carbon skeleton of the side chain, and restricted the locations of three of the four oxygen atoms to the A,B, and C rings. One of the most interesting features of the 'H-nmr is the presence of five singlet methyl groups resonating at 6=1.35, 1.17, 1.04, 0.97, and 0.67. By analogy to known sterols in this series only the lowest and the highest field methyls could be safely assigned as the 19 Me and 18 Me resonances, respectively. Two coincident secondary methyl! doublets appeared at 6=0.88, J=6.8 Hz. Thus, of the seven methyl groups in the molecule, five must be a part of the side chain. The 2DJ resolved 'H-nmr spectrum confirmed these observations. Per- haps the most difficult aspect of the structure elucidation was the establish- ment of the molecular connectivity and stereochemistry of the side chain. Based upon precendent, the cyclopropyl] ring was located at positions 22,23. The most obvious proton of this moiety resonating at 6= —0.24 as a (dd), No. 4, 1990] MUSMAR AND WEINHEIMER— NOVEL MARINE STEROIDS 261 J=5.9 and 4.6 Hz. The second proton resonated at 6=0.83 as a (dd), J=9.5 and 4.5 Hz was partially obscured by the resonance of the two methyls at = 0.88. The third potential cyclopropane proton resonance was obscured by the 18 Me resonance at 6 = 0.67, the location confirmed by baseline irregulari- ties and the integration which clearly showed the presence of four rather than three protons. The structural relationship between these cyclopropyl protons was readily determined by conventional decoupling experiments, confirming that the molecule contained a 1,1,2-trisubstituted cyclopropyl system located at C22-C23. The deshielding of two of the three cyclopropane protons from the “normal” region inferred the proximity of an electronegative substituent. The fourth oxygen atom needed to balance the molecule is predicted to be a tertiary alcohol due to lack of geminal proton resonances in the 'H-nmr spec- trum. The presence of only one methyl group at 6=1.17 gave added support to this arrangement. In comparison with the spectra of similar compounds, location of the hydroxyl group at C24 was logical, and appeared to be more likely than at the other potential sites in the side chain. These evaluations were compatible with the proposed side chain structure and its 22R, 23R, 24R stereochemistry were unequivocally confirmed by x-ray crystallography. The crystal structure has a number of unique features and properties. ACKNOWLEDGMENT: We wish to thank Josephine Aaron for typing this manuscript. LITERATURE CITED F’ uLKNER, D. J., E. Fany, C. H. Rao, K. V. RAMANA, AND D. V. Rao. 1988. Metabolites of the Gorgonian Isis hippuris from India. J.Nat. Prod. 51:954-958. Haert_e, W. R. 1971. A Novel Marine Steroid, Gorgest-5-ene-36,9,11-triol from Pseudoptero- gorgia americana. Masters Thesis, Univ. of Oklahoma, Norman. Hate, R. L., J. LEcLERQ, B. Turscu, C. Dyerassi, R. A. Gross, A. J. WEINHEIMER, K.C. Gupta, AND P. J. ScHever. 1970. Demonstration of a biogenetically unprecedented side chain in the marine sterol, gorgosterol. J. Amer. Chem. Soc. 92:2179-2180. Kaz.auskas, R., P. T. Murpny, P. N. Raut, R. L. SANDERS, AND R. J. WELLS. 1982. Spermidine derivatives and 9,11-secosteroids from the soft coral (Sinularia sp.). Aust. J. Chem. 35:69- 75. Musmar, M. J., A. J. WEINHEIMER, G. E. Martin, AND R. E. Hurp. 1983. Assignment of the high-field resonances of a gorgosterol derivative through the use of autocorrelated two- dimensional !'H-NMR spectroscopy. J. Org. Chem. 48:3580-3581. . 1983. Part I. Assignment of the High-Field Resonances of 56,66-Epoxy-9, 11-Seco- Gorgostan-36, 11-Diol-9-one 3,11-Diacetate Through the Use of Auto-Correlated Two- Dimensional and Proton-Carbon Correlated NMR Spectroscopy. Part II. Three Novel Marine Steroids: A®-9,11-Dihydroxy-9,11-Seco-Dinostan-9-One, and 38, 96, 116-Trihy- droxy-Dinostane. From the Gorgonian (Pseudopterogorgia americana). Ph.D. dissert., Univ. of Houston, Houston. SCHEUER, P. J. (ed.). 1978. Marine Natural Products, Vol. 2, p. 75. Academic Press, Inc. SHEIKH, Y. M., C. Djerassi AND B. Turscu. 1971. Acansterol: A cyclopropane-containing marine sterol from Acanthaster planci. Chem. Commun. 217-218. SpraccIns, R. L. 1970. Part I. Two New Prostaglandins: 15-EPI-PGA, and its acetate, methyl diester from the Gorgonian Plexaura homomalia. Part II]. Two extremely novel marine sterols: A°-3,11-dihydroxy-9,11-Secogorgosterene-9-one and its 56, 66-epoxide from the Gorgonian Pseudopterogorgia americana. Ph.D. dissert., Univ. of Oklahoma, Norman. Florida Sci. 53(4): 257-261. 1990. Accepted: October 10, 1989. 262 FLORIDA SCIENTIST [Vol. 53 REVIEW C. W. Hart, Jr. and Janice Clark. An Interdisciplinary Bibliography of Freshwater Crayfishes. Smithsonian Institution Press, Washington, D.C. 1989. Pp. 000. Price: $35.00 THis attractively-produced book provides 12,481 references on freshwater crayfishes in the superfamilies Astacoidea and Parastacoidea, from Aristole through 1987. Each bibliographic citation includes author(s), date, title of the paper or book, unabbreviated name of the journal or publisher, pagina- tion, subject codes and a reference number. The authors have included 122 subject codes that describe everything from distribution, ecology and physiol- ogy to myths, postage stamps and recipes. These codes indicate the general categories of information contained in each citation. This bibliography was originally published in the Smithsonian Contribution to Zoology series (455:1-437) and included citations through 1985. The new volume includes an additional 2,574 references. The original bibliography and its index are reprinted as they appeared in the initial volume; the additions are presented in a separate section with their own index. An important tool for students of crayfish biology, this excellent reference may be ordered from the Smithso- nian Institution Press, Department 900, Blue Ridge Summit, PA 17294 at a cost of $35, plus $2.25 for postage and handling.—Richard Franz, Florida Museum of Natural History, University of Florida, Gainesville, FL 32611. REVIEW Robert K. Godfrey. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia and Alabama. The University of Georgia Press, Athens and London, 1988. Pp. ix + 734. $50.00. Tus is a technical manual intended to assist in the identification of the native and naturalized trees, shrubs, and woody vines of northern Florida, southern Georgia, and southern Alabama. The geographic area covered by the manual does not follow political boundaries. The distinction between a shrub and a tree and between a woody and non-woody perennial herb is often not evident. It is only through familiarity that one can make these distinctions. The author draws upon his many years of experience to make decisions in this regard for the reader. If he has erred, it is that perhaps he has been too inclusive. Some, such as Lupinus westianus, seem hardly woody. However, this does not distract from the book. The species are divided into three groups: gymnosperms, monocotyledons, and dicotyledons. Within each group, the families are arranged alphabetically, and within each family, the genera are also alphabetical. However, the species are sometimes alphabeti- cal, sometimes not, depending upon the author’s inclination. It is difficult to No. 4, 1990] WUNDERLIN— BOOK REVIEW 263 locate species in large genera that are not alphabetically arranged. For exam- ple, it is particularly inconvenient having to locate oak species (28 species distributed over 30 pages of text) by having to refer to the key for the species number or to the index. Emphasis is at the species level with each described in detail. Family descriptions and descriptions of genera in which a single spe- cies occurs in the range of the manual are not provided. The species and generic descriptions are usually in a three paragraph format, one each for the habit, leaf, or floral/fruit characters. This makes it visually easy to locate the description of the feature of interest. A few (e.g. Lonicera) are in a single paragraph format, thus making presentation inconsistent. Following the spe- cies descriptions is a generalized habitat statement along with the distribution of the species for the range of the manual, then the distribution over the species’ full range. The inclusion of the National Champion Big Tree is of limited interest use, as more often than not, the individual tree is outside of the manual range. Many names used in the more common manuals, such as Small’s 1933 Manual of the Southeastern Flora are listed as synonyms so that cross reference of names can be made. Unfortunately, this information is incomplete as many names found in Small or even some (e.g. Carya ovalis) that are found in Clewell’s 1985 Guide to the Vascular Plants of the Florida Panhandle are unexplained absences. The work is often frustrating to use because of the incomplete synonymy. No species are treated in the genus Pyracantha although a good description of the genus is given. Many species are illustrated which greatly facilitates identification. The illustrations, line drawings mostly prepared by Melanie Darst, are of excellent quality. A few that were prepared by Priscilla Fawcett did not reproduce well and look over-inked. Although regional in scope, the book is useful for some distance outside its prescribed range because of the wide distribution of many species. The ample descriptions, excellent illustrations, and simple format make it easy to use. I highly recommend it.—Richard P. Wunderlin, Department of Biology, Uni- versity of South Florida, Tampa, FL 33620. Biological Sciences FOODS OF THE KEY DEER OW, D. KLIMSTRA AND ® ALLAN L.. DOOLEY ()Cooperative Wildlife Research Laboratory Southern Illinois University, Carbondale, IL 62901 and °2)103 Oakwood Dr., Tuttle, OK 73089 ABSTRACT: A comprehensive study (1967-1973) of Key deer resulted in 129 rumen samples from mortalities. Analyses yielded 164 plant foods of which 28 comprised about 75 % of the total volume. In order of importance value, red mangrove, black mangrove, Indian mulberry, silver palm, brittle thatch palm, blackbead, grasses, pencil flower, acacia, and sapodilla were the 10 top ranking. Woody browse contributed 42% of total volume; woody plant fruits 27%; palm flowers, fruits and spathes 14%; forbs 13%; and miscellaneous 3%. There was seasonal change in use, reflecting individual plant phenology as related to weather. Clearly evidenced was diver- sity in the deer’s diet suggesting habitat management must address variety between and within plant communities. THE first ecological investigation of the Key deer (Odocoileus virginianus clavium; Barbour and Allen, 1922), conducted June 1951-September 1952 (Dickson, 1955) included food habit studies based on direct observation of deer feeding, browse evidence, and pellet and stomach analyses. Fifty-two plant species were identified as Key deer foods. Direct observations of deer feeding yielded 21 plants; 17 were recorded for one animal. Dickson (1955) stated sign of browsing as rare and found only in certain areas; 19 plants were recorded as food. A total of 293 pellet groups was examined; based on seeds and seed fragments and histological characteristics 27 taxa were identified. The single stomach examined contained only fruits of silver palm (Coccothri- nax argentata) and tallowwood (Ximenia americana). During January 1968-September 1973 rumen samples from 129 Key deer mortalities were collected and analyzed to determine plants consumed, parts of plants utilized, seasonal dietary aspects, and local differences in vegetation utilization (Dooley, 1975). With increased development of privately-owned lands, excellent habitat is and will be altered resulting in extensive changes in available foods. Purpose of this study was to contribute a better understand- ing of Key deer habitat needs so as to enhance development of short-and long- range management plans for public lands in the Lower Florida Keys. Srupy ArEA— The Florida Keys form a crescent of small islands extending south/southwest approximately 208 km from peninsular Florida. Big Pine Key is the largest island (circa 2400 ha, 9.6 km long and 3.2 km wide) of the Lower Keys complex and the principal home of the Key deer. Soils vary from thick marl depositions to bare rock of the oolitic formation (Dickson, 1955). Average rainfall at Key West, Florida, 48 km west of Big Pine Key is 101.5 cm annually. Its occurrence is normally greater than 10 cm per month June- October and least during December-March, when a mean of 4.5 cm per month occurs (Klimstra et al., 1974). Characteristically, October-March rep- resents a dry period. The lack of organic materials in many areas results in rapid runoff and pooling of rainfall in depressions. Thus, many plants are adapted to relatively xeric conditions. No. 4, 1990] KLIMSTRA AND DOOLEY— KEY DEER 265 Islands near sea level are surrounded by thick growths of red mangrove (Rhizophora mangle) established in shallow salt water tidal zones. With in- crease in elevation, red mangrove is replaced at approximately high tide level by black mangrove (Avicennia germinans), white mangrove (Laguncularia racemosa), and buttonwood (Conocarpus erecta). These maritime zones, de- pending on island elevations, usually grade into hardwood and pineland hab- itats that are intolerant of salt water. Notable acreages of slash pines (Pinus elliottii var. densa) occur on Big Pine, No Name, Sugarloaf, Cudjoe, and Little Pine keys within the Key deer range; lesser stands are found on Big Knockemdown, Little Torch, Middle Torch, and Howe keys. The relative size, distribution, and composition of natural plant associations are primarily the result of elevation gradients and salt water influence (Dickson, 1955). Disturbed area habitats show greatest species diversity followed by hard- wood, pineland, cultivated, strand, and marsh (Table 1). Of all plant species, 46.3% are forbs, 42.3% woody, and 11.3% grasses and sedges. TABLE 1. Number and percent of total for plant species occurring in various cover types on Big Pine Key, Florida (Dooley 1975). Number of Percent Cover Type Species of Total Disturbed areas (roadsides, subdivisions) 190 29.8 Hardwood 126 19.8 Pinelands 118 18.6 Cultivated 94 ae Strand (maritime zone) 90 14.2 Marsh (fresh or brackish) 18 Deb’ The total population of Key deer 1967-1973 was estimated at 350-400 having increased from 25-80 animals since 1950-51. Big Pine Key, which supports 60-65% of the population, provides a great variety and number of plant species (Dickson 1955). All the major habitat types found in the Lower Florida Keys occur on Big Pine (Klimstra et al., 1974). METHops—Stomach contents from 129 Key deer mortalities available for study represented 83 males, 44 females and 2 unknown from Big Pine (112), No Name (6), Little Torch (5), Ramrod (3), and Big Torch (1) keys; 2 lacked site identification. The majority represented roadkills on U.S. Highway 1, Key Deer Boulevard (State Road 940), and Wilder Road on Big Pine Key (Fig. 1). Approximately 0.95 1 of rumen was taken from each deer collected; samples were placed in plastic containers and frozen until examined. Because of limited mortalities in any one year, rumen samples available were grouped by quarters (January-March (37), April-June (29), July- September (25), and October-December (36)) to evaluate seasonal changes in plant utilization. Rumen samples were prepared for analyses according to Robel and Watt (1970) utilizing three sieves constructed from standard 5-, 10-, 20-mesh/cm hardware wire to separate rumen contents. Plant material was identified, recorded, and placed in a drying oven at 60°C for 4 hours prior to quantitative determination. After drying, a volumetric measurement (cc) was made of materials from 5-mesh/cm screen (Robel and Watt, 1970). Dried materials from the 10- and 20-mesh/cm screen were analyzed using the point method described by Chamrad and Box (1964). Percent occurrence determined for each plant species and unknowns was translated into volumetric (cc) values by taking the percent determined by point analysis (Chamrad and Box, 1964), dividing by 100, and multiplying this value by the total volume of material examined. 266 FLORIDA SCIENTIST [Vol. 53 Lions Club Port Pine Heights Parking Lot Blue Hole Koehn Subdivision Watson Hammock Pine Heights Eden Pines Refuge Headquarters Doctors Arm 11 Tropical Bay Estates 12 Palm Villa 13 Watson Field 14 Road Prison 15 Shopping Center *6 Pine Channel Estates 17 Sea Center 18 Sea Camp 19 Sands Subdivision 20 Chamber Commerce 21 = St. Peters Church 22 Outward Bound 23 B. P. Fishing Camp Cactus Hammock oP ONOanawn = CHANNELS ROADS HAM MOCK PINE WOODS DEVELOPED AREA MIXED HARDWOODS MANGROVE THICKET SCRUB BUTTONWOOD — MANGROVE o> | Sais B BIG PINE KEY UEOBOT! Var Big Munson | Fic. 1. Major plant communities, subdivisions, roads and miscellaneous sites, Big Pine Key. No. 4, 1990] KLIMSTRA AND DOOLEY—KEY DEER 267 The percent composition of individual food items was calculated by adding values (cc) in each of the samples and dividing by the total volume of materials examined, then these multiplied by 100. The percent frequency of plants was calculated from the number of times an item was identified, divided by the total number of samples examined. Plant importance rating was calcu- lated by multiplying percent frequency and percent volume of the food item which provided a relative index for rating a given food item. A plant reference collection, assembled from within the Key deer range, aided identification of plant foods. Common and scientific names were based on Long and Lakela (1971), Small (1933) and Watkins (1961); when there was difference, priority was given the most recent publication. RESULTS AND DiscussiIon—A total of 164 food plants was identified in rumen samples (Table 2); 28 accounted for approximately 75% by volume while 10 comprised 56.6%. Red mangrove (Rhizophora mangle) leaves and fruits and black mangrove (Avicennia germinans) fruits yielded nearly 24%. Indian mulberry (Morinda royoc) leaves and fruits; silver palm (Coccothri- TABLE 2. Foods with importance values greater than 2.0 as documented in rumen samples from 129 Key deer mortalities during December 1967-June 1973 (Dooley 1975). Percent Percent Importance Plant Frequency Volume Value 1. Rhizophora mangle 63 12.15 765.45 2. Avicennia germinans 34 11.39 387.26 3. Morinda royoc 65 5.05 328.25 4. Coccothrinax argentata 34 7.39 251.26 5. Thrinax microcarpa 34 5.06 172.04 6. Pithecellobium keyense 48 3.33 159.84 7. Grasses 68 2.03 138.04 8. Stylosanthes hamata 49 PR tl 135.73 9. Acacia spp. 43 3.00 129.00 10. Manilkara spp. 28 4.49 125.72 11. Chamaesyce spp. 76 1.02 17.52 12. Erithalis fruticosa 24 3.22 77.28 13. Serenoa repens 22 Lael 42.02 14. Bumelia celastrina 28 1.31 36.68 15. Galactia spp. 46 0.78 35.88 16. Randia aculeata 29 1.12 32.48 17. Jacquinia keyensis 24 1.16 27.84 18. Smilax havanensis 37 0.62 22.94 19. Pinus elliottii var. densa 66 0.34 22.44 20. Mushroom 20 lel 22.20 21. Ipomoea spp. 10 2.12 21.20 22. Sida spp. 29 .61 17.69 23. Physalis spp. 44 All 9.24 24. Chiococca spp. 1 .76 9.12 25. Byrsonima cuneata 10 83 8.30 26. Lantana involucrata 23 .30 6.90 27. Myrtus verrucosa 9 12 6.48 28. Melanthera spp. ll NG) 3.99 29. Laguncularia racemosa 9 42 3.78 30. Desmodium canum 10 .28 2.80 31. Agalinis spp. 9 29 2.61 32. Guettarda scabra 6 38 2.28 33. Tillandsia sp. 15 14 2.10 1The complete list of 164 plant foods identified is on file in the office of the National Key Deer Refuge, Big Pine ive) the Cooperative Wildlife Research Laboratory, Southern Illinois University at Carbondale (Dooley 268 FLORIDA SCIENTIST [Vol. 53 nax argentata) flowers and fruits; brittle thatch palm (Thrinax microcarpa) fruits; blackbead (Pithecellobium keyense) leaves; graminid leaves; pencil flower (Stylosanthes hamata) leaves; acacia (Acacia spp.) fruits and leaves; and sapodilla (Manilkara spp.) fruits were also important. Red mangrove, which exhibited the greatest importance value, confirmed Dickson’s (1955) finding that >63% of 293 pellet groups contained this species. By contrast, mainland Florida deer stomach contents indicated that, although a variety of food plants was utilized, 20 items contributed 84% of volume (Harlow, 1959) in one study; another yielded 120 plant foods, with 10 items making up 90% and 4 items 75% of volume (Strode, 1954). TABLE 3. Plant type categories of 164 Key deer food plants as recorded from 129 rumen samples (Dooley 1975). Number of Percent of Plant Type Species Total Forbs 69 42.1 Woody Plants 65 39.6 Grasses and Sedges 23 14.0 Domesticated fe 4.3 Forbs contributed the greatest number of species used by deer followed closely by wood plants (Table 3). The variety of food used was representative of plants found in disturbed areas, followed by those of pinelands, hardwood, strand, and fresh and brackish marshes (Table 4). These data suggest a great variety and number of plant species (Klimstra et al., 1974) are subjected to Key deer use. TABLE 4. Occurrence of 164 Key deer food plants within specific habitat types (Dooley 1975). Number and Plant Type Total Species Percent Habitat Division Woody Forb Graminid Occurrences of Total Disturbed areas 26 51 17 94 40 (roadsides, subdivisions) Pinelands 22, 36 4 62 26 Hardwood 38 4 3 45 19 Strand (maritime zone) 13 6 6 25 11 Marsh (fresh or brackish) 3 1 4 8 4 Percentages of forage types in the overall deer diet indicated woody plant leaves and new growth stems were most important (Fig. 2), followed by woody plant fruits, palm fruits and flowers, forbs, and miscellaneous items i.e., grasses, mushrooms, and pine needles. Although hardened twigs of wood species were not important, leaves and new woody growth of the most important browse species including red mangrove, Indian mulberry, black- bead, acacia, erithalis (Erithalis fruticosa), saffron plum (Bumelia celas- trina), white indigo berry (Randia aculeata), joewood (Jacquinia keyensis), catbriar (Smilax havanensis), snowberry (Chiococca spp.) lantana (Lantana No. 4, 1990] KLIMSTRA AND DOOLEY—KEY DEER 269 42.1% WOODY PLANTS (LVS: & STEMS) 27.0% WOODY PLANTS (FRUITS) 3.5% GRAMINIDS MUSHROOM PINE Fic. 2. Categories of plant foods utilized by Key deer based on 129 rumen samples. involucrata), rough velvet-seed (Guettarda scabra), and torchwood (Amyris elemifera) provided 42% of the overall diet. The variety of fruits and flowers important in the deer diet suggested regular and extensive use of red man- grove, black mangrove, brittle thatch palm, sapodilla, acacia, silver palm, Indian mulberry, ground cherry (Physalis angustifolia), locust berry (Byrson- ima cuneata), white stopper (Myrtus verrucosa), tallowwood (Ximenia amer- icana), white mangrove (Laguncularia racemosa), seven-year apple (Casasia clusiifolia), guava (Psidium guajava), and Barbados cherry (Malpighia gla- bra). Flowers, stalk, and spathes from saw palmetto (Serenoa repens) and silver palm also contributed importantly to the Key deer diet. The 42% fruit and flower use is similar to that documented for mainland deer (Lay, 1965; Harlow, 1959; Strode, 1954; Harlow and Jones, 1965; and Harlow and Hooper, 1971). Seasonal Food Utilization—The 37 samples for January-March showed important foods to be red mangrove leaves and fruit; blackbead leaves; saw palmetto flowers, stalks, and spathes; erithalis leaves and stems; saffron plum leaves; grass and sedge leaves; joewood leaves, mushroom stems and caps; and white indigo berry leaves. These accounted for approximately 55% of the total food volume; unidentified leaves and herbaceous stems yielded 20.7% reflecting relative increase in browse in rumen samples for this period. 270 FLORIDA SCIENTIST [Vol. 53 59.5% WOODY PLANTS 35.1% F2 (LVS. & STEMS) eo fans s 37.7% WOODY PLANTS (FRUITS) SN 13.2% Se WOODY ~~s PLANTS (FRUITS) / 1.4% 4.7% GRAMINIDS GRAMINIDS MCROGn MUSHROOM STS PINE JANUARY - MARCH APRIL - JUNE i] 1 i} i] ( 25.9% ' ; WOODY PLANTS | eaters J (LVS.& STEMS) 4 SAG, i (LVS. & STEMS) 1 WOODY PLANTS ; cae i FRUITS / 0% ' ( / WOODY PLANTS (FRUITS) 4.0% GRAMINIDS MUSHROOM ae PINE ice GRAMINIDS MUSHROOM PINE OCTOBER - DECEMBER JULY - SEPTEMBER Fic. 3. Categories of plant foods utilized by Key deer based on 37 rumen samples for January- March, 29 April-June, 25 July-September, and 36 October-December. Of 84 food plants, woody browse from red mangrove, blackbead, indigo berry, saffron plum, erithalis, Indian mulberry, acacia, joewood, and cat briar accounted for almost 60% of the total volume (Fig. 3). Fruits of sweet acacia (Acacia farnesiana), red mangrove, and sapodilla comprised 13.1%, the lowest for woody plant fruits during the four seasonal periods. The 29 samples for April-June showed silver palm flowers, fruits, and stalk; sapodilla fruits and leaves; red mangrove leaves and fruits; Indian mul- berry leaves and fruits; acacia leaves and fruit pods; erithalis leaves and fruits; and blackbead leaves to be of importance. These accounted for 55.9 % of the total food volume while unidentified material yielded 16.5% . Of 67 plants recorded, woody plants accounted for 72.8%; forbs 10.5%; palms 15.4%; and grasses, pine, and mushrooms 1.4% (Fig. 3). Fruits of woody No. 4, 1990] KLIMSTRA AND DOOLEY— KEY DEER Za plants i.e., sapodilla, red mangrove, Indian mulberry, and acacia were most important, comprising 37.7% of the aggregate volume while browse made up 35.1%. Palms yielded 15.4% , with silver palm flowers, immature fruits, and spathe most important. The 25 samples for July-September indicated important foods to be brittle thatch palm fruits and stalks; black mangrove fruits; Indian mulberry leaves and fruits; silver palm fruits and flowers; erithalis leaves and fruits; and grass and sedge leaves. These accounted for 61.5% of the total aggregate volume; unidentified yielded 11.9% . Of 72 plants identified, woody plants contrib- uted 62.0%, forbs 6.5%, palms 27.5%, and grasses, mushrooms, and pine needles 4.0% (Fig. 3). Fruits were most important, contributing 36.1% of the total volume. Flowers and fruits of silver and brittle thatch palms pro- vided 27.5% to the volume, the highest percent of diet attributed to fruits and flowers (63.5 % ) of the four periods. The 36 rumen samples for October-December showed black mangrove fruits the most important food item. Others of importance included red man- grove leaves and fruits; silver palm fruits; pencil flower leaves and stems; brittle thatch palm fruits; and morning glory (Ipomoea spp.) flowers, leaves, and stems. These accounted for 58.1% of the aggregate volume; unknown leaves and stems made up 15.9% . Of 91 food plants recorded, leaves, herba- ceous stems, and fruits of woody plants contributed 68.8% of the total vol- ume; forbs 16.9%; and palms 10.4% (Fig. 3). Fruits of woody plants com- prised 36.0% while their leaves and herbaceous stems contributed 32.8%. Aspects of deer diet suggested increased availability and/or preference by Key deer for certain plant components on a seasonal basis. Flowers and fruits contributed approximately 21, 53, 63, and 46%, while browse from forbs and woody plants contributed 73, 45, 32, and 49% of the diet, respectively, during the four periods examined (Fig. 3). The variety and tropical affinities of many plants which contributed fruits and flowers represented factors af- fecting seasonal diet composition. Fruits and flowers of several plant species were found in greatest quantity April-November, while browse was greatest during December-March. The rainfall during quarterly sample collections indicated average rainfall was lowest during January-March; this corre- sponded to when browse was most utilized by Key deer. The pattern of rain- fall in this subtropical climate is an important factor affecting phenology of the majority of plants within the Key deer range and appears to correspond with “seasonal” patterns of forage utilization witnessed in the deer diet. Although monthly fluctuations occurred, several plants contributed some browse throughout the year. These included red mangrove, Indian mulberry, pencil flower, acacia, grasses, spurges (Chamaesyce spp.), milk peas (Galac- tia parvifolia), saffron plum, catbriar, slash pine (Pinus elliottii var. densa), ground cherry (Physalis angustifolia), lantana, and five-petalled leaf flower (Phyllanthus pentaphyllus). Other food plants present in samples from every month but one included: blackbead, erithalis, white indigo berry, joewood, small-leaved melanthera (Melanthera parvifolia), Spanish needle (Bidens pi- 212 FLORIDA SCIENTIST [Vol. 53 losa), teaweed (Sida acuta), and bromeliad (Tillandsia sp.). These data sup- port Klimstra et al. (1974) observation that certain plants were subjected to regular and continuous browsing. Also indicated was that greatest quantities of browse from certain plants were recorded when fruits and flowers were least available. Local Differences in Food Habits—To relate major vegetation type with foods eaten, samples were grouped to reflect three principal sources within the Key deer range. One source included 39 samples representative of the northern portion of Big Pine Key; another represented 40 samples from along U. S. 1 on Big Pine Key (Fig. 1); and, another represented 15 samples from other keys including No Name (6), Little Torch (5), Big Torch (1), and Ram- rod (3). Food items in samples representing the two areas of Big Pine Key were not substantially different; but, there was variation reflecting the im- portant food plants present on both segments. There was apparent difference in food utilization by deer of other keys as Pigeon plum (Coccoloba diversifo- lia) and devil’s potato (Echites umbellata) appeared in samples from No Name, and erithalis, grasses, morning glory showed greater use than on Big Pine. Acacia did not occur in samples from other Keys. This study substantiates Klimstra et al. (1974) observation based on browsing that foods utilized varied from one area of a Key to another and particularly from one Key to another. Variety of habitats and plants available to deer on certain Keys was more limited than on Big Pine Key. In part, differences in rumen sample composition and number may have reflected availability of certain plants; however; other unknown factors, unrelated to plant availability, were clearly indicated. Utilization of red mangrove is an example; although relatively abundant on all islands, it comprised a smaller portion of deer diets on keys other than Big Pine. Management Implications—The 164 documented Key deer plant foods was strong evidence of extensive use of all major habitat types (Table 4). Although a large number and variety of plants were recorded, relatively few provided the bulk of its diet (Table 1). There was obvious seasonal fluctuation in use of certain plant species and their parts. The above findings suggest maintaining a variety of habitat types and food plants available to Key deer is essential for effective habitat management. The continued development of islands, such as Big Pine Key, has resulted in clearing and loss of pinelands, hardwoods, and maritime zones. This has eliminated priority food plants such as red mangrove, black mangrove, acacia, brittle thatch palm, silver palm, sapodilla, and blackbead. Continued loss to prioritized human inter- ests and needs of important habitats and food plants requires public-owned lands be effectively managed to insure survival of a healthy Key deer popula- tion. Although the National Key Deer Refuge and associated government- owned lands provide the foundation for maintenance of a viable Key deer population, increased development of private properties restricts the amount and availability of quality native habitats. Although use of roadsides and subdivisions is evident in the diets of deer, such man-controlled areas are not No. 4, 1990] KLIMSTRA AND DOOLEY—KEY DEER 273 in their best long-term needs and interests. First, few of the priority foods used occur there and; second, deer use of these areas greatly increases their vulnerability to human interaction, especially automobile encounters. It is believed the attraction to such land uses is the edge and openness they provide rather than quality of food resource. Proper management of Key deer numbers in relationship to available habitat requires prediction and evaluations of qualitative trends in habitat condition; thus, utilization and abundance of plants and/or plant parts con- sidered important to Key deer should be monitored. Controlled burning, clearing, and reseeding techniques to maintain a variety of native habitat types, have been identified as management for selected areas within the Key deer range (Klimstra et al., 1974). The impact of such activities on essential habitats and food plant species must be continuously evaluated to insure short-and long-term management practices accommodate Key deer needs. ACKNOWLEDGMENTS— Appreciation is extended to Dr. J. W. Hardin, Dr. N. J. Silvy, the late Mr. Jack Watson, Sr., and Mr. John Roseberry for assistance with various aspects of this study. Southern Illinois University through the cooperative Wildlife Research Laboratory, United States Fish and Wildlife Service, National Geographic Society, North American Wildlife Foundation, and National Wildlife Federation contributed financial support. Dr. Alan Woolf reviewed the manuscript. The manuscript originates from a thesis by Allen Dooley in completing requirements for the M.A. degree in Zoology, Southern Illinois University and is a contribution from the Laboratory’s Proj. No. 15: Big Game Investigations. LITERATURE CITED Bargpour, T. AND G. M. ALLEN. 1922. The white-tailed deer of eastern United States. J. Mammal. 3(2):65-78. CuHamraD, A. D. Anp T. W. Box. 1964. A point frame for sampling rumen contents. J. Wildl. Manage. 28(3):473-477. Dickson, J. D., III. 1955. An Ecological Study of the Key Deer. Tech. Bull. 3. Florida Game and Fresh Water Fish Comm. Tallahassee. 104 pp. Dootey, A. L. 1975. Foods of the Key deer (Odocoileus virginianus clavium). M.A. Thesis. Southern Illinois Univ., Carbondale. 80 pp. Hariow, R. R. 1959. An Evaluation of White-tailed Habitat in Florida. Tech. Bull. 5. Florida Game and Fresh Water Fish Comm. Tallahassee. 64 pp. AND R. G. Hooper. 1971. Forages eaten by deer in the southeast. Proc. Southeast Assoc. Game and Fish Comm. 25:18-46. AND F. K. Jones, Jr. 1965. The White-tailed Deer in Florida. Tech. Bull. 9. Florida Game and Fresh Water Fish Comm. Tallahassee. 240 pp. Kurmstra, W. D., J. W. Harpin, N. J. Sitvy, B. N. JAcosson, AND V. A. TERPENING. 1974. Key deer investigations final report. Period of study: December 1967-June 1973. Cooperative Wildl. Res. Lab., Southern Illinois Univ., Carbondale. Mimeo. 184 pp. Lay, D. W. 1965. Fruit utilization by deer in southern forests. J. Wildl. Manage. 29(2):370-375. Lone, R. W. anv O. Laxketa. 1971. A flora of tropical Florida. Univ. Miami Press, Coral Gables. 962 pp. RoBEL, R. J. AND P. G. Watt. 1970. Comparison of volumetric and point analysis procedures to describe deer food habits. J. Wild]. Manage. 34(1):210-213. SMALL, J. K. 1933. Manual of the Southeastern Flora. Univ. North Carolina Press, Chapel Hill. 1554 pp. SrrobE, D. D. 1954. The Ocala deer herd. Florida Game and Fresh Water Fish Comm. Game Publ. No. 1, 42 pp. Warkins, J. V. 1961. Your Guide to Florida Landscape Plants. Univ. Florida Press, Gainesville, 292 pp. Florida Sci. 53(4): 264-273. 1990. Accepted: August 29, 1989. Biological Sciences SCALE DEVELOPMENT IN OVULIFEROUS CONES OF PINUS ELLIOTTII ENGELM., PINACEAE R. F. MENTE AND S. D. Brack-HANEs Collegium of Natural Sciences, Eckerd College, St. Petersburg 33733 Asstract— There has been no ontogenetic study of conifer ovuliferous scale development. This information is important for interpreting and understanding the fossil record of conifers and their ancestry. Mature and juvenile stages of seed cones and their fragments are found isolated in ancient sediments. Therefore, this ontogenetic study of cone scales in Pinus elliottii Engelm. (Diploxylon Section, Pinaceae) was initiated with special emphasis on the development of two prominent scale features; the apophysis and umbo. In this species, the two structures do not develop until the cone’s second year of growth, and during all subsequent stages, they are broader than they are thick. Furthermore, the dorsal umbo develops and matures before the apophysis. StupIEs of extant plants provide insight into developmental stages of related extinct plants (Florin, 1944; Karrfalt and Eggert, 1977). Florin (1944) contrib- uted ideas about the origin of ovulate conifer cones and their scales from his work with fossil and living forms. Karrfalt and Eggert (1977) compared basal meristem development of the living plant, Isoetes, with the underground axis Stigmaria, a fossil lycopod. A similar approach may be useful to determine the phylogeny of ovulate scale features by correlating scale development in living Pinus species (the only known extant genus with an umbo on the scale) with extinct pinacean cones. This study is of umbo and apophysis development within one species of Pinus and it may suggest a pattern of development for other conifer species. In P. elliottii Engelm. (Section Diploxylon, Pinaceae), the umbo forms from the scale tip. To date, the sequence and pattern of differential growth of the apophysis and umbo on scales has not been examined. Over thirty years ago, Mergen and Koerting (1957) remarked that “a detailed morphological descrip- tion” of pollen and seed cone developmental stages had not been made for pines. They further commented that for P. elliottii (slash pine) this is especially regrettable, because that pine is of paramount economic importance world- wide. Their work with P. elliottii addressed pollen cone morphology (cone initiation to just before spore mother cell formation) and ovulate cone mor- phology from initiation to the stage of ovular swelling, i.e., sporogenous tissue formation. The present study extends knowledge of ovulate cone morphology in this species beyond Mergen and Koerting’s stages to the mature seed cone. MATERIALS AND MErHops—AIll seed cones of P. elliottii Englem. were collected from seven trees on the campus of Eckerd College (elev. 3 feet), St. Petersburg, Florida. Collections were made at a six month interval, April 21 and October 24, 1988 as part of an on-going study. Cones were obtained from the lower one-third branches of the trees, divided into developmental stages according to apophysis/umbo morphology, and measured for length and width. All cone and scale measurements were taken at right angles to the surface measured. Fifty measurements for every feature were obtained for each of the six stages. An average is reported (Table 1). Because this is an initial study and some cone measurements are based on fewer than 10 samples, standard deviations were deter- mined only for data that included 10 or more cones. No. 4, 1990] MENTE AND BRACK-HANES— SCALE DEVELOPMENT 278 Aa = ’ . Ee Fic. 1—(A) Pinus elliottii seed cone Stage 1 with incipient umbo area all of the exposed part of scale (arrow) (2.2X), (B) Stage 2 with incipient umbo area (arrow) (1.5X), (C) Stage 3 with umbo area (arrow) and incipient apophysis (ap) (1.5X), (D) Stage 4 with umbo (arrow) and apophysis (ap) (1.2X), (E) Adaxial (ad) and abaxial (ab) surfaces on scale of Stage 1 with incipient umbo area (arrow) (1.7X), (F) Adaxial (ad) and abaxial (ab) surfaces on scale of Stage 2 with incipient umbo area (arrow) 1.7X), (G) Adaxial (ad) and abaxial (ab) surfaces on scale of Stage 3 with umbo area (arrow) and incipient apophysis (ap) (1.7X), (H) Adaxial (ad) and abaxial (ab) sur- faces on scale of Stage 4 with umbo (arrow) and apophysis (ap) (1.7X), (I) Stage 6 open mature cone with umbo (arrow) and apophysis (ap) (0.5X), (J) Adaxial (ad) and abaxial (ab) scale sur- faces of Stage 6 with umbo (arrow), apophysis (ap), bract (B), seed (S), and seed wing (W) (1.5X). 276 FLORIDA SCIENTIST [Vol. 53 ResuLts—Cones and features of cones described herein include the scales and two other morphological scale parameters; umbo and apophysis (Fig. lJ). Other features not included in this study are the bract on the abaxial surface, and the seed and seed wing on the adaxial surface (Fig. 1J). The umbo is the most distal end of the scale that is green at first, tapering to a hair-like tip, and becomes brown at maturity. The apophysis is the enlarged, exposed portion of the scale contiguous with the distal umbo. As with the umbo, it also is green at first and becomes brown as it matures. The exposed scale at maturity includes the apophysis and umbo areas. This study began with the first stage of obvious differentiation in the cones and ovulate scales of a Diploxylon species, i.e., P. elliottii Engelm. Ovulate scale differentiation begins about nine months after seed cone initiation, just after pollination (Mirov, 1967; Foster and Gifford, 1974). At the time, the green scales begin to swell and the cone closes (Fig. 1A). This is the first of the six stages in the study and is notable for the second greatest cone length-to-width ratio (Table 1). The thickness-to-width ratio of the exposed scales appears to remain con- stant for the remaining growth stages. The green, exposed part of the scale is the incipient umbo area (Fig. 1A, 1E). Stage 2 is characterized by tan/green- ish, succulent scales with a hair-like tip (incipient umbo), and resin deposits. A bloom is on the exposed part of the scale (incipient umbo) (Fig. 1B, 1F). The bloom appears as a powder-like deposit that may be similar to the waxy coating of polymeric esters that occur on conifer needles (Robinson, 1967). While the cone has grown in all aspects from the first to second stages, the length-to-width ratio of the cone has decreased and continues to decrease for the remaining stages, except for Stage 6 closed cones. Stage 2 and closed cones of Stage 6 have the greatest variability in length-to-width ratios as reflected in their standard deviations (Table 1). In Stage 3, the umbo area is tan, larger (swollen), and more distinctive; some areas have green bases (incipient apophyses) and resin deposits are present (Fig. 1C, 1G). Although the incipi- ent apophysis is evident, it is not a cone-wide characteristic at this stage and, therefore, not statistically valid. When the scale is broken, four resin canals are observed adaxial to five vascular bundles. Stage 4 has scales without a bloom, but apophyses and umbo areas are developing (Fig. 1D, 1H). Both structures are darker than those of Stage 3 and the umbo area is darker brown than the apophysis. Collapsing of the cells delimiting the umbo has not yet occurred; that is a feature apparent by Stage 6. As a result of this, measure- ments of umbo thickness and width in Stage 4 are greater than in Stage 6. The brown distal end of the scale (umbo) exhibits splitting. Resin deposits are present. The apophysis T/W ratio appears to be smaller in Stage 4 than Stage 6, while the umbo T/W ratio appears to remain constant. The umbo AD/AB ratio appears to decrease slightly in further developmental stages. Stage 4 is characterized by differential growth of the apophysis and displacement of the umbo. In Stage 4, growth of the apophysis in width appears greater than its growth in thickness. Due to collection times (6 months apart) and cone di- mensions, it is apparent that at least one stage is missing between Stage 4 and No. 4, 1990] MENTE AND BRACK-HANES— SCALE DEVELOPMENT PAT, the mature cone in Stage 6. Since this is an on-going study, no data is recorded in Table 1 for Stage 5, but it is included in the table for continuity and to acknowledge that there is at least one obviously missed stage. Perhaps this indicates that the missing stage(s) is of very short duration. In Stage 6, the mature cone, the closed cone has the greatest length to width ratio of all the stages. Apophysis growth appears greatest in the horizontal (width) direction than in the vertical (thickness) direction and its T/W ratio increases, indica- ting substantial growth in both directions. Umbo thickness and width mea- surements decrease from Stage 4 to Stage 6 and its T/W ratio appears to be constant from Stage 4 to Stage 6. The AD/AB ratio of the umbo is the smallest of all the stages (Table 1). In Stage 6, the open cone expression shows a L/W ratio distinctively below the closed cone (Fig. 11, 1J). Because this final devel- opmental form of Stage 6 (open) is not a growth stage, it is not compared with other stages. TABLE 1. Measurements (mean average in mm except where noted) with standard deviations (s) for ratios of closed seed cone features in developmental stages (1-6) in Pinus elliotti. The final expression of Stage 6 is the open cone. ! Features Stages i 2 3 4 5 6 Number 4 13 18 10 _— Te Cone Length (L) (em) 14.0 19.8 21.4 26.8 — 8.3 Width (W) (cm) 7.5 14.1 16.0 22.3 = 4.0 L/W Ratio eS, 1.40 1.34 1.20 — 20 S — oll 1G +.16 — ae DAL Thickness (T) 18.4 2358 23.9 leo _ 80.6 Exposed Width (W) 37.1 44.4 46.6 73.2 _ 133.9 Scales T/W Ratio 0.49 0.54 0.51 0.42 — 0.60 s + .05 + .05 + .03 ell — ae i Thickness (T) — - — 4.4 — Olen, Apophysis Width (W) = - — ES = US T/W Ratio — — — 0.29 — 0.74 s + .09 — + .06 Thickness (T) — — — 27.8 — 22.9 Width (W) — — — 60.4 — 50.7 Umbo T/W/ Ratio _ — —_ 0.40 — 0.42 S + .08 — + .06 Adaxial (ad) — _ — 22.6 — 24.1 Adaxial (ab) — — — 26.0 — ThE) ad/ab Ratio _ _ — 0.87 _ 0.79 S =20i0 — + 0.07 1Number 16, length 9.5 cm, width 7.0 cm, L/W Ratio 1.37, s= +0.09. Discussion—In P. elliottii Engelm. (Diploxylon Section, Pinaceae), the abaxial side of the umbo seems to have an initial growth rate that is greater than the adaxial side. This is changed by later adaxial growth and the ratio becomes nearly equal by the mature stage. Overall, differential growth of the umbo may be displaced to a greater or lesser degree depending on pine spe- cies. This has potential significance for interpreting the fossil record. De- 278 FLORIDA SCIENTIST [Vol. 53 pending on cone, apophysis, the umbo growth ratios, as well as preservation conditions, one may ascertain not only the species, but the stage of growth in the fossil cone. Hills and Ogilvie (1970) demonstrated with living and fossil species of Picea, that they may be classified by scale ratio measurements. The same may apply to Pinus. The present study focused on a set of seed cone/ scale variables that may be used for classification of pines; albeit, one of many characteristics commonly used to divide the genus into either Haploxy- lon or Diploxylon sections. Previously, these sections have been defined mainly on wood and anatomical features (vascularture and resin canals) of scales and seeds (Jeffrey, 1905; Bailey, 1910; Miller, 1976; Alvin, 1988). A review of pertinent literature, however, disclosed that the morphological traits of cone shape and umbo position for Pinus species also appear to typify the two sections. Pinus (Haploxylon) species generally have terminal umbos and cylindrical cones with blunt apices, while Pinus (Diploxylon) species have dorsal umbos and mostly ovoid-tapered cones with conical apices. From a paleobotanist’s point of view, such characteristics as these may be of great importance to the understanding of plant organ phylogeny or even the inter- pretation of ancestry within a family. Pinus elliottii may statistically suggest that all mature cones of Pinus Haploxylon and Diploxylon species have a predictable and very narrowly defined length to width ratio for ovulate cones, as well as a specific AD/AB ratio for umbo growth. Perhaps an ontogenic study of both umbo positions, as they occur in extant species, would reveal a more clearly defined ancestry for Pinus and other pinacean genera as the fossil record becomes known. This method of species differentiation may also apply to Pityostrobus. Pi- tyostrobus, and extinct pinacean genus of seed cones, is known from the Juras- sic (Nathorst, 1897, 1899; Fleiche and Zeiller, 1904; Toyama and Oishi, 1935) into the Oligocene (Crabtree and Miller, 1989). It is recognized as a collection of rather diverse cones with or without umbos. In those species with umbos, both dorsal and terminal ones have been reported; this may be useful for classification. For instance, Alvin (1957) described Pityostorobus villerotensis from numerous specimens of cones. By taking cone length-to-width ratios (from Alvin’s figures), individuals of his collection appear to vary from 1.25 to 6.0. Perhaps this species represents several varieties, species, or even onto- genetic stages. Juvenile stages of cone features, when discovered, may be assigned to individual species or perhaps grouped together into unnatural taxa. This question has been raised with some fossil seed studies such as Pinus wheeleri. Pinus wheeleri is a species of fossil seeds that Chaney and Axelrod (1959) established for a collection of species described previously (Penhallow, 1908; Berry, 1929; Dorf, 1936; Smith, 1941; MacGinitie, 1953). Information for Pinus, as noted by Mergen and Koerting (1957), is still lacking. The statistical data for mature and developmental stages of cone features (length to width ratios) presented in this paper for one Pinus species, is needed for other living species. Moreover, this work suggests that ontoge- netic studies of extant pinacean species is imperative for defining not only No. 4, 1990] MENTE AND BRACK-HANES— SCALE DEVELOPMENT 279 extant species, but also fossil ones. One cannot statistically define a range of features in a fossil if the statistical characteristics of related extant species are unknown. As Hutton formulated (1788), and is now popularly paraphrased, “the present is the key to the past.” LITERATURE CITED Atvin, K. L. 1957. On the two cones Pseudoaraucaria heeri (Coemans) Nov. Comb. and Pityos- trobus villerotensis Nov. Sp. from the Wealden of Belgium. Inst. Roy. Sci. Nat. Belgique Mem. 135:1-33. . 1988. On a new specimen of Pseudoaraucaria major Fleiche (Pinaceae) from the Cretaceous of the Isle of Wight. Bot. J. Linn. Soc. 97:159-170. Bal.ey, I. W. 1910. Anatomical characters in the evolution of Pinus. Amer. Nat. 44:284-293. Berry, E. W. 1929. A Revision of the Flora of the Latah Formation. U.S.G.S. Prof. Pap. 154-H, Pp. 225-264. Cuaney, R. W. ann D. I. AxELRop. 1959. Miocene Floras of the Columbia Plateau. Carnegie Inst. Wash. Pub. 617. Washington, D.C. CraBTREE, D. R. ANDC.N. MILLER, Jr. 1989. Pityostrobus makahensis, a new species of silicified pinaceous seed cone from the Middle Tertiary of Washington. Amer. J. Bot. 76:176-184. Dorr, E. 1936. A Late Tertiary Flora from Southwestern Idaho. Carnegie Inst. Wash. Pub. 476, II, Pp. 73-124. Washington, D.C. FLEICHE, P. AND R. ZEILLER. 1904. Note sur une Florule Portlandienne des Environs de Boulogne- sur-mer. Bull. Soc. Geol. de France 4:787-811. Fiorin, R. 1944. Die Koniferen des Oberkarbons und des Unteren Perms. Paleontographica 85B, Pp. 365-654. Foster, A. S. AND E. M. Girrorp, Jr. 1974 (2nd ed.). Comparative Morphology of Vascular Plants. W. H. Freeman and Co., San Francisco. Hiuts, L. V. anp R. T. Ocitvie. 1970. Picea banksii n. sp. Beaufort Formation (Tertiary), north- western Banks Island, Arctic Canada. Can. J. Bot. 48:457-464. Hutton, J. 1788. Theory of the Earth. Trans. Roy. Soc. Edinburgh. Edinburgh, Scotland. Jerrrey, E. C. 1905. The Comparative Anatomy and Phylogeny of the Coniferales. Pt. 2—The Abietineae. Mem. Boston Soc. Nat. Hist. 6(1). KarrFaLT, E. E. anp D. A. Eccert. 1977. The Comparative Morphology and Development of TIsoetes L. II. in I. tuckermanii A. Br. and I. nuttallii A. Br. Bot. Gaz. 138 (3):357-368. MacGinitie, H. D. 1953. Fossil Plants of the Florissant Beds, Colorado. Carnegie Inst. Wash. Pub. 599. Washington, D.C. MERGEN F. Anp L. E. Koertinc. 1957. Initiation and development of flower primordia in slash pine. For. Sci. 3(2):145-155. MILter, C. N. 1976. Early Evolution in the Pinaceae. Rev. Palaeobot. Paly. 21:101-117. Mirovy, N. T. 1967. The Genus Pinus. The Ronald Press Co., New York. Natuorst, A. G. 1897. Zur Mesozoischen flora Spitzbergens. K. Svensk. Vetenskapsakad. Hand. 30:1. . 1899. Fossil plants from Franz Josefs Land, Norwegian North Polar Exped. 1893- 1896. Stockholm. PENHALLOw, D. P. 1908. Report on Tertiary Plants from British Columbia. Can. Dept. Mines, Geol. Surv. Branch, No. 1013, Ottawa. Rosinson, T. 1967 (2nd ed.). The Organic Constituents of Higher Plants. Burgess Pub. Co., Minneapolis, Minnesota. SmiTH, H. V. 1941. A Miocene Flora from Thorn Creek, Idaho. Amer. Mid. Nat. 25:473-522. ToyaMA, S. AND S. Otsu1. 1935. Notes on Some Jurassic plants from Chalai-Nor, Prov. North Hsingan, Manchoukuo. J. Fac. Sci., Hokkaido Imp. Univ., Sapporo, Ser. 4:61-75. Florida Sci. 53(4): 274-279. 1990. Accepted: October 3, 1989. Biological Sciences DISTRIBUTION OF THE EASTERN CHIPMUNK (TAMIAS STRIATUS) IN FLORIDA JEFFERY A. GORE Florida Game and Fresh Water Fish Commission, Nongame Wildlife Program, 6938 Highway 2321, Panama City, Florida 32409 Asstract: Reports of sightings of the eastern chipmunk in Florida were solicited from the public and used to locate areas where chipmunks occur. Field observations and collected speci- mens were used to outline the chipmunk’s known distribution, which now includes portions of five northwestern Florida counties (Escambia, Santa Rosa, Okaloosa, Walton, and Holmes) and the watersheds of the Escambia, Blackwater, Yellow, and Choctawhatchee rivers. The species may have been present here since the Pleistocene, but the historical range in Florida is not known. Chipmunks in Florida are associated with hardwood or mixed hardwood-pine (Pinus spp.) forests, but they appear to be unevenly distributed within their range. Because the size of the population in Florida is unknown and the range is small, the chipmunk’s current legal status as a Species of Special Concern should be retained. THE eastern chipmunk occurs across much of eastern North America, but it is conspicuously absent from most of the coastal plain of the southeastern United States (Hall, 1981). Chipmunks were first reported from Florida in 1962, when their known range comprised only a few square kilometers in Okaloosa County near the border with Alabama (Stevenson, 1962). Subse- quent surveys produced additional local specimens including one from Ala- bama (Jones and Suttkus, 1979), but they did not greatly extend the chip- munk’s known range in Florida. These surveys also did not determine whether the Florida population was geographically isolated from chipmunk populations in central Alabama (Jones, 1978). Here I present results of a survey of the distribution of the chipmunk across northwest Florida, with comments on the chipmunk’s range in south- ern Alabama. The objectives of the survey were to document the range of the chipmunk within Florida and to qualitatively assess the status of the Florida population. METHops—Information on sightings of chipmunks was solicited from persons throughout northwest Florida and verified with specimens or my own observations from nearby locations. Because chipmunks are diurnal, often live in close proximity to humans, and look unlike any other animal in Florida, I assumed that the presence of chipmunks in an area could be reliably determined by questioning local residents. More than 200 people across northwest Florida were interviewed during this study. In 1986 interviews were confined to Okaloosa County north of Crestview. In 1987 and 1988 persons in all six Florida counties that border Alabama were questioned. The survey concentrated on locations north of Interstate Highway 10 (approximately latitude 30° 40’N) and, in particular, along the major streams. Three outlying areas were surveyed: along the Chipola River in Calhoun County, east of the Apalachicola River in northern Liberty County, and along Lake Talquin in Leon County. Most persons interviewed lived or worked near what appeared to be suitable habitat for chipmunks, i.e. hardwood or mixed hardwood-pine forests with nearby streams (Stevenson, 1962; Jones and Suttkus, 1979). No. 4, 1990] GORE—EASTERN CHIPMUNK IN FLORIDA 281 Each person questioned was shown a nearly life-size drawing of a chipmunk and asked if they had seen the animal. Most persons were able to respond quickly and confidently. Those who claimed to have seen a chipmunk were asked to recall the date and location and to describe the animal’s behavior and the circumstances under which they had seen the animal. If a person gave an atypical account of chipmunk behavior or ecology (Snyder, 1982), their observation was disre- garded. All persons questioned were also given a printed request for sighting information and asked to pass it to others that might know about chipmunks. Requests for information were also placed in local newspapers and newsletters. The number of persons these indirect requests reached is not known. In 1987, a drawing of a chipmunk was sent along with a map of northwest Florida to approxi- mately 75 biologists, technicians, and wildlife officers working for the Florida Game and Fresh Water Fish Commission (FGFWFC) in northwest Florida. Each was asked to report locations where they had seen chipmunks. Sighting reports that represented range extensions were verified by observations of chipmunks at or near the reported location. If persons had a specimen of a chipmunk, no further attempt was made to corroborate the presence of the species. Collected specimens were deposited in the Florida Museum of Natural History, Gainesville (accession number A-288). Field observations were employed instead of trapping because chipmunks in Florida seem particularly secretive and difficult to trap (Stevenson, 1962 and Jones and Suttkus, 1979). Ideally, all potential chipmunk habitat in Florida would have been systematically surveyed, but that was not practical. In 1986, when the chipmunk’s known range in Florida was extremely small, I began a systematic survey by walking along transects across suitable habitat while listening and watching for chipmunks. Although some animals were found in this way, negative results from the transect surveys were unreliable and walking transects proved to be inefficient for surveying a large area. ResuLtts— More than 30 of the persons interviewed reported seeing chip- munks in Florida. With the help of their reports, I was able to find chip- munks at seven locations (Fig. 1). Other FGFWFC personnel reported obser- ESCAMBIA Escambia Reon WASHINGTON 8) oe tieaeenetandta Rs Fic. 1. Range of the eastern chipmunk in Florida based upon the location of animals collected (4), observed by the author (MM), or observed by other personnel of the Florida Game and Fresh Water Fish Commission (L)). Boundary extrapolations were determined by the location of suit- able habitat and unconfirmed reports of chipmunk sightings. 282 FLORIDA SCIENTIST [Vol. 53 vations from eight additional locations. Four specimens were provided by individuals who live-trapped the chipmunks (two trapped at the same loca- tion in Escambia County) or whose pets had killed the animals in their yards (Fig. 1). I observed or collected chipmunks within 10 km of all but one of the locations reported by FGFWFC personnel. I did not observe chipmunks at Lake Jackson in Walton County, but Paul Moler saw one there (Moler, 1988) and several local residents also reported that chipmunks had been in the area for many years. All of the 18 locations where chipmunks were recorded (Fig. 1) were in hardwood or mixed hardwood-pine forests having oaks (Quercus spp.) as dominant species. Twelve (66%) of the locations were within 100 m of a permanent stream or lake. Although the range depicted (Fig. 1) encompasses areas dominated by long-leaf pine (Pinus palustris), chipmunks were not ob- served or reported in that habitat type. Only two apparently valid reports of chipmunks in Florida came from outside the range shown in Fig. 1. Both those reports were from Okaloosa County between the Yellow River and the town of Holt, but I was unable to verify either. Field observations and interviews with persons along the entire length of the Yellow River in Florida convince me that chipmunks, if present, are very rare below the mouth of the Shoal River. Approximately 20 persons reported seeing chipmunks in southern Alabama, but I did not attempt to verify those reports with field observations. Discussion— Although reports of animal sightings by untrained observers are often not reliable, sighting reports proved to be very useful in identifying areas occupied by chipmunks. By obtaining specimens or making corroborat- ing field observations, I was able to verify range extensions initially suggested by reports from the public. Unfortunately, it was not possible to determine whether areas that produced no reports of chipmunks were, in fact, unoccu- pied. Thus, the range depicted (Fig. 1) should be considered a conservative estimate of the current range of the chipmunk in Florida. The range documented in this study represents a substantial increase over that previously reported for the chipmunk in Florida (Stevenson, 1962; Jones, 1978). Chipmunks are now known to occur at least 20 km farther south on the Yellow River than Stevenson (1962) found them. They are also known from the watersheds of 3 other rivers (Escambia, Blackwater, and Choc- tawhatchee) and 4 new counties, i.e. Escambia, Santa Rosa, Walton, and Holmes (Fig. 1). Surprisingly, no chipmunks were observed or reported along the Apalachicola or Chattachoochee rivers in Florida. This river system pene- trates north through the coastal plain and within Florida contains many large ravines harboring other northern plants and animals (Wolfe et al., 1988). Several reports were received of chipmunks from along the Chattahoochee River in Columbia, Alabama, 30 km from Florida. However, the resident park manager at Chattahoochee State Park on the Alabama-Florida border had never seen chipmunks in the park. Apparently, the current southern limit to the chipmunk’s range along the Chattahoochee and Apalachicola rivers is No. 4, 1990] GORE—EASTERN CHIPMUNK IN FLORIDA 283 between these two points. The paucity of specimens from southern Alabama and Florida (Lindzey, 1970; Jones and Suttkus, 1979) has left unanswered the question of whether the Florida population of chipmunks is isolated. The most common assump- tion is that the chipmunk’s range extends into Florida from central Alabama, but only along a narrow peninsula (Hall, 1981; Snyder, 1982). Although I collected no specimens from Alabama, several biologists and local residents reported seeing chipmunks in Baldwin, Clarke, Covington, Escambia, Ge- neva, and Houston counties. These include all the Alabama counties along the border with Florida. The number and distribution of these reports, com- bined with the locations across northwest Florida (Fig. 1), indicate that the chipmunk is distributed, although perhaps unevenly, across all of southern Alabama. More records are needed from Alabama to confirm the chipmunk’s range there. It is not known how long chipmunks have been in Florida. Several older residents along the Yellow River in Okaloosa County recalled seeing chip- munks in the area more than 60 years ago. These reports, coupled with the presence of the animals at several widely-spaced locations (Fig. 1), suggest that the Florida population did not result from recent transplanting by hu- mans. Fossil specimens of the chipmunk have not been found in Florida, al- though several other northern mammals were present here during the Pleisto- cene (Neill, 1957; Webb, 1974; Kurten and Anderson, 1980). Pleistocene seas flooded major streams up to the current Alabama border, but apparently did not cover the higher lands (greater than 30 m above present sea level) of northwest Florida (Wolfe et al., 1988). Thus the chipmunk potentially could have occupied most of its current range throughout the Pleistocene. Changes in the distribution of hardwood forests, both natural and hu- man-related, may have had a significant effect on the distribution of the chipmunk. During the late Pleistocene, hardwood forests were common across the Gulf coastal plain (Delcourt, 1980) and, therefore, suitable habitat for chipmunks is assumed to have been present. As recently as 5,000 years ago, oaks and hickories (Carya spp.), which are typical components of chip- munk habitat, were much more common than pines over southern Alabama (Delcourt, 1980) and, presumably, northwest Florida. Within the past 5,000 years long-leaf pine communities have occupied much of the upland habitats within the Gulf Coastal plain (Delcourt, 1980; Wolfe et al., 1988). If chip- munks were present in the upland hardwood forests prior to this time, their distribution likely became restricted, along with the hardwood forests, to mesic hammocks, ravines, and river valleys. At the time of settlement by Europeans, much of the coastal plain was apparently dominated by either hardwood (Magnolia-Fagus) forests or long- leaf pine forests (Delcourt and Delcourt, 1977; Delcourt, 1980; Wolfe et al., 1988). European settlers converted much of the long-leaf pine forests in northwest Florida to agricultural land (Delcourt, 1980; Wolfe et al., 1988). 284 FLORIDA SCIENTIST [Vol. 53 When these lands were left untilled, mixed pine-oak-hickory forests suc- ceeded them, particularly where fires were absent (Wolfe et al., 1988). These forests are common today along the northern edge of the Florida panhandle (Wolfe et al., 1988) and some are occupied by chipmunks (Stevenson, 1962; Jones and Suttkus, 1979; this study). Thus, humans may have converted some mesic long-leaf pine sites to mixed pine-oak-hickory sites suitable for chip- munks. On the other hand, humans probably destroyed much chipmunk habitat when clearing hardwood forests. Whether such activities signifi- cantly affected the chipmunk’s distribution or abundance in Florida remains unknown. The present survey indicates that chipmunks occur unevenly across their range and that much of what appears to be suitable hardwood forest habitat in northwest Florida is not occupied by chipmunks. Many explanations for the chipmunk’s uneven distribution are possible, including inaccurate survey methods, inability of the chipmunks to disperse widely (Snyder, 1982) and closely track habitat changes, and inaccurate assessment of suitable habitat. Unfortunately, too little is known about the chipmunk’s ecological require- ments in its southern range, its past distribution, or the vegetational history of northwest Florida, to be able to explain the chipmunk’s current distribution in Florida. I made no quantitative estimate of the density or abundance of chip- munks within Florida. The field observations and interviews with residents indicate that chipmunks are uncommon throughout most of their Florida range, but the animal’s secretive behavior makes even a qualitative estimate difficult. There is, however, no evidence that chipmunk numbers are declin- ing. In fact, Stevenson (1962) believed that the number of chipmunks in- creased during his study. Most of the 30 residents who reported seeing chip- munks noted that the animals had been in their area for many years, but not one person reported that chipmunks were now absent from places where they formerly occurred. Thus, the chipmunk population appears at least stable in number and distribution; however, quantitative monitoring is obviously needed to accurately estimate density levels and determine trends in popula- tion size. In the absence of more data, the chipmunk population in Florida is prob- ably best described as locally or unevenly distributed and not declining in size. The Florida Committee on Rare and Endangered Plants and Animals considers the chipmunk rare in the state (Jones, 1978) and the FGFWFC lists it as a Species of Special Concern (Wood, 1989). These designations should be retained, despite the larger distribution I found, because the species’ entire range in Florida is still small and its population size remains unknown. ACKNOWLEDGMENTS—I thank the many people who kindly responded to my inquiries or oth- erwise assisted in the search for chipmunks, especially R. Carr, B. Klug, R. Klug, T. Magaha, J. McMahon, G. Mausbaum, T. White, and the R. Walker family. C. Jones, H. Stevenson, and R. Suttkus supplied useful advice from their study of Florida chipmunks and S. Humphrey, C. Jones, B. Millsap, and T. O’Meara provided helpful comments on the manuscript. This study was funded by the Florida Game and Fresh Water Fish Commission. No. 4, 1990] GORE— EASTERN CHIPMUNK IN FLORIDA 285 LITERATURE CITED Detcourt, H. R. anp P. A. Detcourtr. 1977. Presettlement magnolia-beech climax of the Gulf Coastal Plain: quantitative evidence from the Apalachicola River Bluffs, north-central Florida. Ecology 58:1085-1093. Detcourt, P. A. 1980. Goshen Springs: Late quaternary vegetation record for southern Ala- bama. Ecology 61:371-386. Hatt, E. R. 1981. The Mammals of North America. John Wiley and Sons, New York. 600 pp. Jones, C. 1978. Eastern chipmunk. Pp. 35-36. In Layne, J. N. (ed.). Rare and endangered biota of Florida, Volume One: Mammals. Univ. Presses of Florida, Gainesville. 52 pp. AND R. D. Sutrxus. 1979. The distribution and taxonomy of Tamias striatus at the southern limits of its geographic range. Proc. Biol. Soc. Washington. 91:828-839. Kurten, B. AND E. ANDERSON. 1980. Pleistocene mammals of North America. Columbia Univ. Press, New York. 442 pp. Linpzey, D. W. 1970. Mammals of Mobile and Baldwin counties, Alabama. J. Alabama Acad. Sci. 41:64-99. Moter, P. E. 1988. Florida Game and Fresh Water Fish Commission, Gainesville, Pers. Comm. NEILL, W. T. 1957. Historical biogeography of present-day Florida. Bull. Florida State Mus. 2:175-220. SNYDER, D. P. 1982. Tamias striatus. Mammal. Species 168:1-8. STEVENSON, H. M. 1962. Occurrence and habits of the eastern chipmunk in Florida. J. Mammal. 43:110-111. Wess, S. D. 1974. Chronology of Florida Pleistocene mammals. Pp. 5-31. In: Webb, S. D. (ed.). Pleistocene mammals of Florida. Univ. Presses of Florida, Gainesville. 270 pp. Wo rt, S. H., J. A. REIDENAUER, AND D. B. Means. 1988. An ecological characterization of the Florida Panhandle. U.S. Fish and Wildl. Serv., Biol. Rep. 88(12). 277 pp. Woop, D. A. 1989. Official lists of endangered and potentially endangered fauna and flora in Florida, 1 July 1989. Fla. Game and Fresh Water Fish Comm., Tallahassee. 19 pp. Florida Sci. 53(4): 280-285. 1990. Accepted: October 20, 1989. Biological Sciences A REVIEW OF THE FLORIDA CRAYFISH FAUNA, WITH COMMENTS ON NOMENCLATURE, DISTRIBUTION, AND CONSERVATION RICHARD FRANZ AND SHELLEY E.. FRANZ Florida Museum of Natural History, University of Florida, Gainesville, FL 32611. ABSTRACT: A review of the Florida crayfish fauna is presented as an update of Hobbs’ 1942 monograph “The Crayfishes of Florida.” As currently understood, this fauna includes 50 species belonging to six genera. Distributional information and recent nomenclatural changes are sum- marized. Hosss (1942) provided a comprehensive treatment of the Florida crayfish fauna in “The Crayfishes of Florida” and this volume continues to be widely used as the primary source for information about these crustaceans in the state. However, caution must be employed when using this source, since there have been many adjustments in scientific names and distributions. There has also been the addition of at least 11 previously undescribed taxa to the state’s faunal list in the interim. This paper attempts to provide a preliminary up- date for this very valuable and still important reference. We refer readers to specific pages in Hobbs’ volume so that they can make necessary notations. Florida has one of the richest crayfish faunas in North America, compris- ing 50 species in six genera (Cambarellus [2 spp.], Cambarus [5 spp.], Falli- cambarus [2 spp.], Faxonella [1 sp.], Procambarus [39 spp.], and Troglocam- barus [1 sp.]) (Table 1). According to Hobbs (in press), this fauna is exceeded in number only in Alabama (73), Tennessee (70), Georgia (69), Mississippi (62), and Arkansas (58). Additionally, two cavernicolous crayfishes from Orange and Putnam counties are probably new and will warrant descriptions when Form I males become available. Another species, Procambarus capilla- tus Hobbs, as yet unrecorded from the state, has been collected near the Florida-Alabama state line in Escambia and Conecuh counties, Alabama (Hobbs, 1971a), and will presumably be found in the flatwoods areas of northern Escambia and Santa Rosa counties, Florida, in the future. With these additions, the state’s fauna could include 53 species. Early publications listed the species affinis, fallax, lecontei, alleni, clarkii, versutus, acherontis, barbatus, evermanni, and clarkii paeninsulanus (all as- signed to the genus Cambarus) from the state (Hagen, 1870; Faxon, 1884, 1885, 1890, 1898, 1914; Lonnberg, 1895; Harris, 1903; Ortmann, 1905a). No new species were added to the faunal list until Hobbs (1938) described Cam- barus rogersi from northwest Florida. Between 1940 and 1942, Hobbs (1940, 1941a, 1942a) described 10 new taxa and elevated Ortmann’s subgenus Pro- cambarus (Ortmann 1905b) to generic rank. These early studies in Florida culminated with the publication of Hobbs’ Florida monograph in 1942 in which he listed 39 species (42 taxa including subspecies) as naturally occur- No. 4, 1990] FRANZ AND FRANZ—FLORIDA CRAYFISH FAUNA 287 ring in the state (Hobbs, 1942b). In this volume, Hobbs described 15 new taxa; reported that there was no evidence for affinis and lecontei in Florida and deleted them from the state’s list; elevated Faxon’s paeninsulanus to full species, and showed that the records of barbatus and clarkii were based on undescribed taxa (i.e., escambiensis and okaloosae, respectively). More recent studies on the systematic relationships of Holoarctic cray- fishes have necessitated changes in the taxonomic interpretations presented in Hobbs’ original volume. Hobbs (1974a) elevated the subfamily Cambarinae to familial rank, in recognition of the many distinctive attributes that sepa- rate them from their western American and old world counterparts, and employed Cambarellinae and Cambarinae as subfamilies. He restricted the use of Astacidae to include only those species that occur on the American Pacific basin (Pacifastacus) and in Europe and Western Asia. Taxonomic changes at the generic and species levels have also necessitated alterations in the names of several Florida species, as reflected in Table 1. Certain Cambarus-like species were split from Cambarus and placed in the genus Fallicambarus (Hobbs, 1969); the subgeneric name Faxonella (Creaser, 1933) was elevated by Hobbs (1942b, by implication) to generic rank for the distinctive little Cambarus clypeatus Hay (1899) (a position supported by Fitzpatrick [1963]); and subgenera were established for Cambarellus (Fitzpa- trick, 1983), Cambarus (Hobbs, 1969), Fallicambarus (Hobbs, 1973), and Procambarus (Hobbs, 1972). The unidentified crayfish, listed by Hobbs (1942b:168) as “Cambarus spe- cies incertis’, from the Escambia River floodplain, has recently been shown to represent the species, Fallicambarus fodiens, based on additional materials from near Jay, Santa Rosa County, Florida (Mansell, 1989). Bouchard (1978) synonymized Cambarus floridanus Hobbs with Cambarus striatus Hay and described the orange crayfish mentioned by Hobbs (1942b; 159, 162) as Cam- barus pyronotus. Procambarus advena was recently split into two taxa (Hobbs, 1981). The name advena, as presently understood, applies only to those crayfishes from the lower coastal plain between the Savannah and Oco- nee-Altamaha rivers in Georgia. Florida records for advena (Hobbs, 1942b:77) are now considered to be for Procambarus talpoides. Hobbs (1967) split Procambarus blandingii and acutus and designated the Mexican Pro- cambarus blandingii cuevachicae (Hobbs, 1941b) as a subspecies of acutus. He referred Florida specimens (Hobbs, 1942b:95) to Procambarus acutus acutus (Hobbs, 1967). Additionally, Hobbs (1981) recognized two subspecies of Procambarus pubischelae and listed the Florida specimens (Hobbs, 1942b:43-44) as belonging to the nominate race. Procambarus clarkii, de- leted by Hobbs (1942b), was later confirmed from the state, based on speci- mens from Escambia County (Florida) (Hobbs, 1974b) and by reports of recent introductions in North and South Florida (Miltner, 1988). There are efforts by state agencies to discourage further attempts to introduce this spe- cies east of the Appalachicola River and to use Procambarus alleni and Pro- cambarus paeninsulanus as alternatives in aqua-culture (Shafland, 1977). 288 FLORIDA SCIENTIST [Vol. 53 TABLE 1. Current list of Florida crayfishes. Nomenclatural changes since Hobbs (1942) are noted in parentheses following species name. Page numbers following the species name refer to pages in Hobbs’ 1942 text. Taxa not listed in Hobbs (1942) are shown with *. (Nomenclatural history, synonymies, range, and habitats for each taxon are recorded in Hobbs, in press). Family Cambaridae Hobbs Subfamily Cambarellinae Laguarda Genus Cambarellus Ortmann Subgenus Pandicambarus Fitzpatrick *blacki Hobbs, 1980 schmitti Hobbs 1942b (p.149) Subfamily Cambarinae Hobbs Genus Cambarus Erichson Subgenus Depressicambarus Hobbs latimanus (LeConte) 1856 (p.158) *oyronotus Bouchard 1978 (= orange specimens, pp. 159,162) *striatus Hay 1902 (= Cambarus floridanus Hobbs 1941a, p.161) Subgenus Jugicambarus Hobbs cryptodytes Hobbs 1941a (p.162) Subgenus Lacunicambarus Hobbs diogenes diogenes (Girard) 1852 (p.164) Genus Fallicambarus Hobbs Subgenus Creaserinus Hobbs byersi (Hobbs) 1941a (= Cambarus byersi, p.167) *fodiens (Cottle) 1863 (= Cambarus species incertis, p.168) Genus Faxonella Creaser clypeata (Hay) 1899 (= Orconectes clypeata, p.154) Genus Procambarus Ortmann Subgenus Hagenides Hobbs Advena Group geodytes Hobbs 1942b (p.80) pygmaeus Hobbs 1942b (p.83) *talpoides Hobbs 1981 (=P. advena, p.75) Rogersi Group *rogersi expletus Hobbs and Hart 1959 rogersi campestris Hobbs 1942b (p.90) rogersi ochlocknensis Hobbs 1942b (p.89) rogersi rogersi (Hobbs) 1938 (p.89) Subgenus Leconticambarus Hobbs Alleni Group alleni (Faxon) 1884 (p.69) *milleri Hobbs 1971b Barbatus Group apalachicolae Hobbs 1942b (p.55) econfinae Hobbs 1942b (p.49) escambiensis Hobbs 1942b (p.46) latipleurum Hobbs 1942b (p.52) pubischelae pubischelae 1942b Hobbs (=P. pubischelae, p.41) rathbunae (Hobbs) 1940 (p.59) Shermani Group shermani Hobbs 1942b (p.61) Kilbyi Group kilbyi (Hobbs) 1940 (p.64) Hubbelli Group hubbelli (Hobbs) 1940 (p.67) Subgenus Lonnbergius Hobbs acherontis (Lonnberg) 1895 (p.91) No. 4, 1990] FRANZ AND FRANZ—FLORIDA CRAYFISH FAUNA 289 TABLE |. continued Subgenus Ortmannicus Fowler Blandingii Group acutus acutus (Girard) 1852 (=P. blandingii acutus, p.94) bivittatus Hobbs 1942b (p.96) Evermanni Group evermanni (Faxon) 1890 (p.107) Lucifugus Group lucifugus alachua (Hobbs) 1940 (p.136) lucifugus lucifugus (Hobbs) 1940 (p.134) *erythrops Relyea and Sutton 1975 *franzi Hobbs and Lee 1976 *horsti Hobbs and Means 1972 *leitheuseri Franz and Hobbs 1983 *orcinus Hobbs and Means 1972 pallidus (Hobbs) 1940 (p.139) Pictus Group pictus (Hobbs) 1940 (p.130) youngi Hobbs 1942b. (p.131) Seminolae Group *delicatus Hobbs and Franz 1986 fallax (Hagen) 1870 (p.111) leonensis Hobbs 1942b (p.114) pycnogonopodus Hobbs 1942b (p.117) seminolae Hobbs 1942b (p.142) Subgenus Pennides Hobbs spiculifer (LeConte) 1856 (p.119) *suttkusi Hobbs, 1953 versutus (Hagen) 1870 (p.126) Subgenus Scapulicambarus Hobbs *clarkii (Girard) 1852 okaloosae Hobbs 1942b (p.100) paeninsulanus (Faxon) 1914 (p.104) Genus Troglocambarus Hobbs maclanei Hobbs 1942a (p.146) Eleven new taxa (Cambarellus blacki Hobbs 1980, Cambarus pyronotus Bouchard 1978, Procambarus delicatus Hobbs and Franz 1986, P. erythrops Relyea and Sutton, 1975, P. franzi Hobbs and Lee, 1976, P. horsti Hobbs and Means, 1972, P. leitheuseri Franz and Hobbs, 1983, P orcinus Hobbs and Means, 1972, P. milleri Hobbs, 1971b, P. rogersi expletus Hobbs and Hart, 1959, Procambarus suttkusi Hobbs, 1953) have been described from Florida since 1942 (Table 1). All of these crayfishes have very restricted ranges. Seven are cavernicolous and are confined to karst areas in Dade, Hernando, Jeffer- son, Lake, Leon, Marion, Orange, Pasco, Suwannee, and Wakulla counties (Hobbs et al., 1977; Franz and Lee, 1982, Franz and Hobbs, 1983; Hobbs and Franz, 1986). The burrowing C. pyronotus is apparently restricted to seeps along small streams that flow from bluffs on the eastern side of the Appalachicola River in Liberty County (Bouchard, 1978), and P. r. expletus is found in one seepage area along Ten Mile Creek in Calhoun County (Hobbs and Hart, 1959). C. blacki is known from a creek in Escambia County and P. suttkusi from similar habitats in Holmes and Walton counties (Hobbs, 1953, 1974b, 1980). 290 FLORIDA SCIENTIST [Vol. 53 TABLE 2. The distribution of genera found in Florida. Species are numerically listed by Flor- ida region. Genera Cambarellus Cambarus Fallicambarus Faxonella Procambarus Troglocambarus TOTAL GADSDEN -7—-.. a NW FL N FL S ie 1 0 0 0 0 NE 20 ] ajoutscoco|y 24 22 =. fnassau ==. LEO a N (JEFFERSON ADISON MT > \ \ ILTON estan : Ry jae OKEECHOBER \\ SARASOTA \ BE soTo ae = Wey Fic. 1. Map of the regions of Florida. West Florida (W), Northwest Florida (NW), North Florida (N), and South Florida (S). No. 4, 1990] FRANZ AND FRANZ—FLORIDA CRAYFISH FAUNA 291 Table 2 summarizes the distributions of the six genera of crayfishes by region, and the boundaries of the Florida regions are defined (Fig. 1). Two genera (Cambarus, Fallicambarus) are restricted to West and Northwest Florida; Faxonella has been found only in Northwest Florida; Cambarellus occurs in West and Northwest Florida and peripherally in North Florida (east to the Suwannee River drainage); and Troglocambarus is restricted to North Florida. Only Procambarus occurs state-wide. The distribution of 50 species are summarized in Table 3. The data pre- sented in this table are based on information from Hobbs (1942b), recent literature, and personal field work, particularly in North and Northwest Florida. Additional surveys are needed to detect further range extensions and to monitor the status of localized species. TABLE 3. Distribution of Florida crayfishes. E = endemic to Florida; E/r = endemic to Florida (restricted to this geographic region); E/p=endemic to Florida (peripheral in this geographic region; E/pos=nearly endemic to Florida (peripheral in other states); X= widespread species; XS = widespread in Southeast; XA = Florida and Alabama only; XG = Florida and Georgia only; X/p = widespread species (peripheral in this geographic region); I = introduced. Species W FL NW FL NFL SFL acherontis — — E/r — acutus x — — — alleni — — E E apalachicolae E/p E — — bivittatus XS — — — blacki E/r — — — byersi XS — — — clarkii x — IP IP clypeata — X — — cryptodytes -- E/pos — -- delicatus — — E/r — diogenes X X/p — — econfinae — E/r — — erythrops — — E/r — escambiensis E/pos — — — evermanni XS — — — fallax — — E/pos E fodiens X — — — franzi — — E/r — geodytes — — E/r — horsti — E/r — — hubbelli E/pos/a E/pos — — kilbyi x E E et leitheuseri — — E/r — latimanus — X/p _ latipleurum — E/r — - leonensis I?/b E E/p -- 292 FLORIDA SCIENTIST [Vol. 53 TABLE 3.— CONTINUED lucifugus — — E/r ss maclanei — = E/r sa milleri = = man E/r okaloosae E/pos == mae ow orcinus — E/r = x! paeninsulanus — XS XS XS/p pallidus — — E/r —~ pictus — — E/r/c — pubischelae -- — XG — pycnogonopodus E/p E 25 iJ pygmaeus — XG/p XG/p — pyronotus — E/r = ees rathbunae E E see = rogerst — E aie a schmitti E E E ab seminolae i. = XG ae shermani Xs ae ms la spiculifer XS XS XS a striatus — X/p was aa suttkusi XA XA i a talpoides es == XG Eis versutus XS XS/p a £2 younge EE le eee TOTAL a/Specimen from Santa Rosa County (Hobbs in litt. to P. Moler, 1986); status unknown. b/Specimen from Okaloosa County thought to be introduced (Hobbs in litt. to P. Moler, 1986); status unknown. c/New records from streams in SE Jacksonville (east of the St. Johns River) in Duval County (J. Pickett pers comm.). The combination of Florida’s strategic position as the most southeastern corner of continental North America, the presence of several major south- flowing rivers (Escambia, Yellow, Choctawhatchee, and Appalachicola) ter- minating along the Gulf coast of the Florida panhandle, and the tumultuous history of the emergences of the Florida panhandle’s coastal strand and the Florida peninsula from the sea has set the stage for the development of a rich crayfish fauna. The complexity of this fauna has, no doubt, been further enhanced by the extension of a large portion of the state from temperate North America into the subtropical zone and the presence of extensive cave systems in Tertiary limestones. Distributional patterns displayed by Florida’s crayfish fauna are summa- rized in Table 4. Of the 50 species in Florida, 30 are endemic, or nearly so. Nineteen species are considered regional endemics, of which 11 are cavernic- olous. An additional eight species have their primary ranges within one re- gion, but also occur peripherally either in other Florida regions or in contigu- ous states. No crayfish species occurs state-wide. The greatest numbers of No. 4, 1990] FRANZ AND FRANZ—FLORIDA CRAYFISH FAUNA 293 TaBLE 4. Summary of distributional data for Florida crayfishes presented in Table 3. FL Regions W FL NW FL NFL S BE FL Regional Endemics 1 6 10 1 FL Endemics/widespread 2 7 3 2 FL Endemics/ peripheral in region 2 0 1 0 FL Near Endemics/ peripheral in other states 3 2 1 0 Widespread/other states 11 9 6 1 Introductions 1 0 1 1 TOTAL 20 24 Zo 5 species occur in Northwest (24), North Florida (22) and West (20); the small- est number is found in South Florida (5) (Table 4). The crayfishes in West Florida include representatives of all genera except Faxonella and Troglocambarus (Table 2). This fauna has its greatest affinities with Alabama, sharing 13 species (Table 4). At least four of these crayfishes are widespread in eastern North America. Nine (excluding the introduced Procambarus leonensis) of the West Florida species occur in other parts of Florida. There is one regional endemic (C. blacki) reported for this area. Northwest and North Florida share only six species, two of which also occur in West Florida. Northwest Florida has eight (including both Procambarus hubbelli and P. leonensis) and North Florida, ten regionally restricted species. The Northwest endemics are associated with the Marianna Karst (1), the Appalachicola Bluffs (1), coastal flatwoods (3), and Woodville Karst (2). The eighth (Procambarus youngi), a somewhat more wide-ranging species, is re- ported from small streams in Gulf County and from the St. Marks River in Leon and Wakulla counties (Hobbs 1974b). In contrast, of the ten regional endemics in North Florida, eight are cavernicolous and are associated with the Northern Peninsula, Lower St. Johns, and Orlando karsts (Franz and Lee, 1982, Franz and Hobbs, 1983, Hobbs and Franz, 1986, Franz unpub- lished data); the remainder include Procambarus pictus from streams in Clay and Duval counties (Franz and Franz, 1978; J. Pickett, 1988) and the bur- rowing Procambarus geodytes from the lower St. Johns and Oklawaha rivers. Three (excluding the introduced Procambarus clarkii) of four species in South Florida also occur in North Florida. Only the cavernicolous Procambarus milleri is endemic to South Florida (Hobbs, 1971b; Franz and Lee, 1982). With 30 species restricted to Florida, there is a compelling need to con- serve this unique assemblage of species. Consequently, the Florida Commit- tee on Rare and Endangered Plants and Animals (FCREPA) suggested the listing of 11 Florida species (Procambarus acherontis as Threatened, Camba- rus cryptodytes, Procambarus erythrops, P. franzi, P. horsti, P. lucifugus, P. milleri, P. orcinus, P. pallidus, P. pictus, and Troglocambarus maclanei as Species of Special Concern) (see Franz, 1982). To date, Procambarus econfi- nae, P. erythrops and P. pictus have been listed by the Florida Game and Fresh Water Fish Commission as Species of Special Concern (Wood, 1989). Additionally, Procambarus acherontis has been listed by the U.S. Fish and 294 FLORIDA SCIENTIST [Vol. 53 Wildlife Service as a species under review for listing, but substantial evidence of biological vulnerability and/or threat is lacking (Wood, 1989). A new FCREPA list is currently being prepared that will recommend action by Flor- ida on an additional five taxa (Procambarus pyronotus, P. youngi, P. delica- tus, P. leitheuseri, and P. rogersi expletus). The status of at least one previ- ously listed crayfish (Procambarus pictus) may need to be upgraded to Threatened in the near future. This may be necessary due to potential changes in land use practices in its native Black Creek habitat: the sale and possible urban development of the 26000-acre Gilman Paper Company tract in the vicinity of the South Fork of Black Creek, the enlargement of State Road 21 to four lanes, and the possible construction of the proposed Jackson- ville-Tampa Toll Road and accompanying bullet train right of way through the heart of this area. The crayfishes of Florida represent an important component of most aquatic food chains in the state, as detritivores, herbivores, and occasional predators, and as prey to a large number of vertebrate predators (see Penn, 1950; Hobbs III and Jass, in press). Studies have shown that some populations of crayfishes can reach sufficient numbers to support populations of carni- vores that feed almost exclusively on them, such as snakes in the genus Regina (Franz, 1977; Godley, 1980). Little is known about the biology of this impor- tant group of Florida crustaceans, and it is hoped that this paper may focus some attention on the needs, not only for continuing studies, but also for efforts to protect this valuable part of Florida’s bio-diversity. ACKNOWLEDGMENTS. We wish to thank Barry Mansell, Paul Moler and Joseph Pickett for sharing data on the distribution of Florida crayfishes; we also want to acknowledge Paul Moler, Carter R. Gilbert, and Horton H. Hobbs, Jr. for their critical reviews of this paper. LITERATURE CITED Boucuarb, R. W. 1978. Taxonomy, ecology, and phylogeny of the subgenus Depressicambarus, with a description of a new species from Florida and redescriptions of Cambarus gray- soni, Cambarus latimanus and Cambarus striatus (Decapoda:Cambaridae). Bull. Ala- bama Mus. Nat. Hist. 3:27-60. Creaser, E. P. 1933. Descriptions of some poorly known species of North American crayfishes. Occ. Pap. Mus. Zool., Univ. Mich. 278:1-8. Faxon, W. 1984. Descriptions of new species of Cambarus, to which is added a synonymical list of the known species of Cambarus and Astacus. Proc. Amer. Acad. Arts and Sci. 20:107- 158. 1885. A revision of the Astacidae. Mem. Mus. Comp. Zool., Harvard Coll., 10(4):1- 186. 1890. Notes on North American crayfishes, Family Astacidae. Proc. U.S. Nat. Mus., 12(758):619-634. 1898. 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The current status of the crayfishes listed by Girard (1852) in his “A Revision of the North American Astaci...” Crustaceana (12)2:124-132. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pp. 93-178. In Hott, P. C., R. L. HOFFMAN, AND C. W. Harr, Jr. (Eds). The Distributional History of the Biota of the Southern Appalachians, Part I: Invertebrates. Virginia Polytechnic Insti- tute, Div. Monogr., 1. 1971a. New crayfishes of the genus Procambarus from Alabama and Texas (Decapoda, Astacidae). Proc. Biol. Soc. Wash., 84(11):81-94. 1971b. A new troglobitic crayfish from Florida. Quart. J. Fla. Acad. Sci. 34(2):114- 124. 1972. The subgenera of the crayfish genus Procambarus (Decapoda, Astacidae). Smithsonian Contrib. Zool. 117:1-22. ________ 1973. New species and relationships of the members of the genus Fallicambarus. Proc. Biol. Soc. Wash. 86(40):461-482. 1974a. Synopsis of the families and genera of crayfish (Crustacea: Decapoda). Smithsonian Contrib. 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Zool. 166:1-161. AND D. S. Lee. 1976. A new troglobitic crayfish (Decapoda, Cambaridae) from penin- sular Florida. Proc. Biol. Soc. Wash. 89(32):383-391. AND D. B. MEans. 1972. Two new troglobitic crayfishes (Decapoda, Astacidae) from Florida. Proc. Biol. Soc. Wash., 84(46):393-409. Hosss, H. H., III anp J. P. Jass, in press. A review of the trophic relationships of crayfishes and shrimps in freshwater ecosystems. Milwaukee Public Museum. LonnBeERG, E. 1985. Cambarids of Florida. Bihang Till Koniglische Svenska Vetenskaps-Akade- miens Handlingar. 20(4):3-14. MANSELL, B. 1989. The occurrence of the crayfish Fallicambarus (Creaserinus) fodiens in Flor- ida. Florida Scient., 52 (3):177-178. MiLtnER, M. 1988. A survey of the range and distribution of Procambarus clarkii in Florida. Aquaculture Market Development Aid Program, Fla. Dept. Agricul. Consu. Serv., Tallahas- see, 24 pp. OrtMANnNN, A. E. 1905a. The mutual affinities of the species of the genus Cambarus, and their dispersal over the United States. Proc. Amer. Philos. Soc. 44(180):91-136. ____—s«di1905b. Procambarus, a new subgenus of the genus Cambarus. Ann. Carnegie Mus. 3(3):435-442. Penn, G. H., Jr. 1950. Utilization of crawfishes by cold-blooded vertebrates in the eastern United States. Amer. Midl. Natur. 44(3):643-658. PickeTT, J. 1988. Jacksonville, FL. Pers. Comm. RELYEA, K. anv B. Sutton. 1975. A New Troglobitic Crayfish of the Genus Procambarus from Florida (Decapoda: Astacidae). Tulane Stud. Zool. Bot. 19(1-2):8-16. SHAFLAND, P. 1977. A Brief Survey with Recommendations Regarding the Commercial Produc- tion of Two Species of Louisiana Crayfish (Genus Procambarus, Family Cambaridae). Un- publ. Report, Florida Game Fresh Water Fish Commn., 10 pp. SHAFLAND, P. 1977. Florida Game and Fresh Water Fish Commission, Boca Raton, Pers. Comm. Woop, D. 1989. Official Lists of Endangered and Potentially Endangered Fauna and Flora in Florida. Fla. Game & Fresh Water Fish Commn. 19 pp. Florida Sci. 53(4): 286-296. 1990. Accepted: December 12, 1989. Environmental Chemistry MEANINGFUL ENVIRONMENTAL DATA: NOT JUST THE LABORATORY’S RESPONSIBILITY DONALD C. ANNE Hart Environmental Management Corporation, 28 Madison Avenue Extension, Albany, New York 12203 Asstract: The increase in the number of environmental regulations have created a new awareness in the validity of laboratory data. What was once thought of as being only the respon- sibility of the laboratory has now been reevaluated. The five aspects of environmental data, comparability, representativeness, precision, accuracy, and completeness, should now be consid- ered the joint responsibility of the laboratory and consultant. In the past twenty years, federal legislation has been enacted in order to help protect the environment. This legislation includes, but is not limited to: 1) Clean Water Act, 40 CFR 100 et seq., 1987; 2) Resource, Conservation, and Recovery Act (RCRA), 40 CFR 260, et seq., 1987; 3) Comprehensive Environmental Response, Compensation and Liability Act (CERCLA), 40 CFR 300, et seq., 1987. During the same time period, the states have passed parallel legislation. In New York State, examples of parallel legislation are: 1) Water Quality Regulations: Surface Water and Groundwater Classifi- cations and Standards (NYSDEC, 1986); 2) Hazardous Waste Management System (NYSDEC, 1987); 3) Solid Waste Management Facilities (NYSDEC, 1988). One of the impacts from increased federal and state environmental legis- lation is an increase in environmental investigative, monitoring and clean-up programs. As a result, more and more consultants and analytical laboratories are entering the environmental market to meet the demand for assistance in answering questions and solving problems. Obtaining samples, maintaining sample integrity, and accurate sample analysis are critical to any environmental program that generates analytical data. The ultimate goal of sample collection and analysis is the extraction of valid information about the composition of the sample. Credible environ- mental data is contingent on both proper laboratory methods and correct sampling and sample preservation procedures. Failure of a laboratory to yield valid, reliable data may jeopardize an entire sampling effort. Con- versely, if the samples were improperly collected or were not representative, the laboratory data, no matter how valid and reliable, will be of little value. There are five aspects of environmental data that must be addressed in order to judge quality, validity, and reliability of both the sampling effort and the laboratory analytical work. The five aspects of meaningful environmental data are comparability, representativeness, precision, accuracy and com- pleteness. In this paper, each aspect of environmental data will be defined 298 FLORIDA SCIENTIST [Vol. 53 and discussed to help persons envolved in environmental programs produce meaningful environmental data. In performing an environmental sampling program, a clear understand- ing of the goals of the program must be established in order to integrate the aspects of environmental data. This involves the preparation of a program specific sampling plan. The sampling plan must include information on de- contamination procedures for equipment, sequence of sample collection, nec- essary field measurements, and frequency and type of field QA/QC samples. The sampling plan must also include the types of matrices to be sampled, the number of samples to be obtained for each matrix, and the analytical para- meters for each sample. Comparability—The first aspect of environmental data is comparability. Comparability becomes involved early in the environmental program. Con- sultants will first review historical information that pertains to a specific environmental problem. Historical environmental data may need to be used in conjunction with the data to be generated during current or future sam- pling events. The data from all current, past and future activities must be comparable. Comparability essentially means keeping the variables to a min- imum. Historical information will help in determining the analyses required, the frequency of each sampling event, the matrices to be sampled, and the number of samples for each of the matrices. Keeping data comparable helps prevent the apples vs. oranges syndrome. After reviewing historical data, the following questions may be asked: First, should the analytical parameters be changed? Second, how many sampling rounds are needed to define the problem and what should the sampling frequency be? Third, how is the data going to be used, such as regulatory review or internal client information? This must be understood before analytical pa- rameters and level of QA/QC are chosen so that the program is cost effective; i.e., additional information, not just additional numbers. Once a program has been established, the variables should be kept to a minimum. The same laboratory should be contracted for the entire duration of the project. The consultant should dedicate personnel to a project in order to maintain familiarity with the project. Finally, sampling and analytical procedures should remain consistent throughout the history of the project. Comparability is the responsibility of the consultant. The consultant must work with the laboratory in order to obtain the best comparable data. Representativeness—The second aspect of environmental data, represen- tativeness, is the unbiased representation of the population of interest. When obtaining environmental samples, the objective is to remove a small portion of an environment that is representative of the entire body and maintain the sample integrity until the time of analysis. Every measurement made and every sample collected should be recorded in a field log book with the time, date, and person or persons in the field team. Information on the sample’s initial physical condition (color, texture, No. 4, 1990] ANNE— MEANINGFUL ENVIRONMENTAL DATA 299 clarity, odor, phase, etc.), sample location and weather conditions at the time of sample collection should also be adequately documented in field notes. The information in the field log book will be used to assess the environmental data and the rationale for collecting the samples. After the samples are collected, placed in the appropriate sample contain- ers, and labeled, the samples are ready to be prepared for shipment to the laboratory. Preparation involves preserving the samples, documenting the sampling event, (e.g., chain-of-custody procedures), and packaging the sam- ples for shipment. At a minimum, preservation will involve cooling the samples to a temper- ature of 4°C or less. Certain analyses of specific matrix types require the addition of chemical preservatives. Preservation requirements are docu- mented in analytical protocols or may be found in references such as SW-846, 3° Edition (USEPA, 1986), Standard Method, 17° Edition (APHA, et al., 1989), Methods for Chemical Analysis of Water and Wastes (USEPA, 1983) and Method for Organic Chemical Analysis of Municipal and Industrial Wastewater (USEPA, 1982). The preservatives may be added to the samples in the field or the laboratory may add the preservative to the appropriate bottle before sampling. The consultant and laboratory must work together to be sure it is clearly understood who will be preserving the samples properly. Further documentation beyond notes in the field log are required for environmental samples. A properly prepared chain-of-custody form must ac- company all environmental samples collected. The chain-of-custody form provides an accurate written record which can be used to trace possession and holding of samples from the time of collection through sample analysis and data reporting. The following information will be specified for each sample on the chain-of-custody form: 1) Sample ID number; 2) date and time; 3) matrix; 4) number of sample containers; 5) analyses requested; 6) samplers names; and 7) final disposition of the samples (name and address of labora- tory doing the analytical work). All samples must be carefully packed before being transported to the lab- oratory. Samples should be packed in such a way as to prevent breakage and to ensure the integrity of the samples. Samples must be shipped or delivered to the laboratory in a timely fashion in consideration of preservation and to ensure compliance to holding times. Before samples are delivered or shipped to the laboratory, the laboratory should be notified. This will help the laboratory to prepare for the samples and at this time any samples with unusual handling characteristics should be discussed (i.e. known toxic materials). These steps will ensure both legal and physical representativeness of the environmental data. Precision and Accuracy—The third and fourth aspects of environmental data, precision and accuracy, are typically used to describe analytical data and are the primary responsibility of the laboratory. Accuracy is defined as the nearness of a measurement, or the mean of a set of measurements to the true value. Accuracy is routinely assessed by means of spiked samples and 300 FLORIDA SCIENTIST [Vol. 53 reference materials. Precision is defined as the agreement between a set of replicate measurements without assumption or knowledge of the true value. Precision is assessed by means of duplicate/replicate analyses. In addition to results from duplicate analysis and spike sample recoveries, the laboratory has other analytical information to help determine precision and accuracy. The laboratory maintains instrument tuning information as well as calibration data. If the instrument is not tuned and calibrated prop- erly, the data it generates is worthless. The laboratory also analyzes other internal QA/QC samples such as method blanks, interference check samples, serial dilution samples, and others in order to assist in tracing the origin of a problem. Problems can be caused by the matrix, instrument, analyst, analyti- cal method, reagents, sampler or any combination. The laboratory QA/QC information along with information on the sampling event can be used de- ductively to determine the problem areas. Before the analytical data is used by the consultant, the data should un- dergo a review to confirm or establish usability of the data. The degree to which data is reviewed will be dependent upon the intent of the program. If the program is only to survey for the absence or presence of contamination, the review may only involve checking sample holding times, method blank results, and methods used (historical precision and accuracy for that method). The other end of the spectrum would be a remedial investigation where specific compounds need to be defined in quantitative terms that are both accurate and precise. A review of data generated for a remedial investigation could involve evaluating all the supporting data that is required by the Contract Labora- tory Program. In either case, the data is first reviewed in order to determine the data usability. Some useful references for evaluating the precision and accuracy of envi- ronmental data are: Quality Assurance of Chemical Measurements (Taylor, 1987); Functional Guidelines for Evaluating Inorganic Analyses (USEPA, 1988); Functional Guidelines for Evaluating Organics Analyses (USEPA, 1988); SW-846, 3” Edition (USEPA, 1986); Standard Methods, 17" Edition (APHA, et al., 1989); and Handbook for Analytical Quality Control in Water and Wastewa- ter Laboratories (USEPA, 1979). Completeness—The fifth aspect of environmental data ties all steps in- volved in obtaining the environmental data. Data completeness involves en- suring that all the necessary sampling and analytical requirements are met. Completeness is the minimum amount of valid and reliable environmental data necessary to achieve the goals of the program. This involves working closely with the laboratory in order to ensure that data completeness is achieved. The laboratory needs to know if there are any particular samples considered critical to the program. In addition, steps such as reanalysis, re-extraction, and resampling to bring data for out-of-specifica- tion or unacceptable critical sample data back into specification need to be No. 4, 1990] ANNE— MEANINGFUL ENVIRONMENTAL DATA 301 discussed. In addition, the laboratory and consultant must discuss whose re- sponsibility it will be to implement any corrective actions. CoNcLusIoN—In summary, the purpose of collecting and analyzing envi- ronmental samples is to extract information. Therefore, the resultant infor- mation must be reliable and valid in order to define the extent and severity of environmental problems. For environmental data to be reliable and valid, it must be precise, accurate, comparable, complete, and representative. The reliability and validity is the responsibility of all those involved in a project. Environmental data can become meaningless if data validity and reliability cannot be determined. LITERATURE CITED AMERICAN PuBLiC HEALTH ASSOCIATION, AMERICAN WATER WorkKS ASSOCIATION, AND WATER POoL- LUTION CONTROL FEDERATION. 1989. Standard Methods for the Examination of Water and Wastewater, 17th Edition. American Public Health Association, Washington, D.C. Cob OF FEDERAL REGULATIONS. 1987. Protection of Environment, Title 40, Parts 100, et seq., US Government Printing Office. Washington, D.C. , 1987. Protection of Environment, Title 40, Parts 260 et seq., US Government Print- ing Office. Washington, D.C. , 1987. Protection of Environment, Title 40, Parts 300 et. seq., US Government Print- ing Office. Washington, D.C. New York STATE DEPARTMENT OF ENVIRONMENTAL CONSERVATION. 1986. Water Quality Regula- tions; Surface Water and Groundwater Classifications and Standards, 6NYCRR, Chapter X, Parts 700-705. Albany, New York. , 1987. Hazardous Waste Management System, 6NYCRR, Parts 370-375. Albany, New York. ____—«, 1988. Solid Waste Management Facilities, 6(NYCRR, Part 360. Albany, New York. Tay or, J. K. 1987. Quality Assurance of Chemical Measurements. Lewis Publishers, Inc. , Chel- sea, Michigan. Unirep STATES ENVIRONMENTAL PRoTEcTION AcENCy. 1979. Handbook for Analytical Quality Control in Water and Wastewater Laboratories. EPA-600/4-79-019. Environmental Mon- itoring and Support Laboratory, Cincinnati, Ohio. , 1982. Methods for Organic Chemical Analysis of Municipal and Industrial Wastewa- ter, EPA-600/4-82-057. Environmental Monitoring and Support Laboratory, Cincinnati, Ohio. , 1983. Methods for Chemical Analysis of Water and Wastes. EPA-600/4-79-020. Envi- ronmental Monitoring and Support Laboratory, Cincinnati, Ohio. , 1986. Test Methods for Evaluating Solid Wastes, (3rd ed.) Volumes IA, IB, IC and II. Office of Solid Waste and Emergency Response, Washington, D.C. , 1988. Laboratory Data Validation Functional Guidelines for Evaluating Organics Analysis. February, Washington, D.C. , 1988. Laboratory Data Validation Functional Guidelines for Evaluating Inorganics Analyses. July, Washington, D.C. Florida Sci. 53(4): 297-301. 1990. Accepted: February 7, 1990. Biological Sciences ICHTHYOFAUNAL EVALUATION OF THE PEACE RIVER, FLORIDA THOMAS R. CHAMPEAU Florida Game and Fresh Water Fish Commission, 3900 Drane Field Road, Lakeland, FL 33811 ABSTRACT: Fishes were collected from six locations along the freshwater portion of the Peace River from 1983 through 1988. Analysis of fish community characteristics such as species rich- ness, diversity, abundance, biomass and species composition indicated the fish community of the Peace River has been significantly impacted by human activities. Degraded water quality and invasion of exotic fishes have altered species composition, diversity and fish production. Effluent from headwater lakes and streams increased turbidity and biological oxygen demand in the upper 35-km section of the river that was dominated by pollution-tolerant fishes. Dilution from tribu- taries exhibiting higher water quality improved biotic conditions downstream as evidenced by higher species diversity and more complex fish community structure. Tidal influence on the lower 35-km portion of the river reduced flushing and stressed conditions were indicated by lower species richness and diversity. Fish community structure at the lower river was dominated by large piscivores and omnivores. THE Peace River begins in Polk County at Lake Hancock in south-central Florida and flows southwest through Hardee and DeSoto counties before discharging into Charlotte Harbor estuary in Charlotte County (Fig. 1). The river is 211 km in length, descending 30 m at an average gradient of 0.2 m/km with an average annual discharge of 32.7 m*/s (Estevez et al., 1981). The Peace River valley is a pinnate system, draining 5959 km’. Micoene deposits rich in phosphate ore were mined from the river bottom in the late 1800’s and from lands within the valley from 1900 to the present. Most strip mine operations have occurred in the upper valley, drastically altering natural hydrology and degrading river water quality (Hand et al., 1988; Estevez, et al., 1981). Mining operations have caused catastrophic fish kills, most recently occurring in 1967 and 1971 (Ware, 1969; Chapman, 1973). These were caused by massive discharges of wastewater from clay- filled settling basins which created extremely high turbidity and resulted in high mortality of fishes and invertebrates. Public outcry and legislative actions during the late 1970’s resulted in stringent regulations that signifi- cantly reduced the incidence of these events. The Peace River receives a large amount of pollutants from municipal/ industrial effluent and urban/agricultural runoff. Most cultural inputs occur near the headwaters; as a result, the upper river exhibits poor water quality (Hand et al., 1988; Estevez et al., 1981; Florida Department of Environmen- tal Regulation, 1980; Florida Board of Health, 1965). Lake Hancock is a hypereutrophic waterbody that discharges water with extremely high con- centrations of blue-green algae, resulting in high turbidity, high biological oxygen demand and low dissolved oxygen levels. Peace Creek drains agricul- tural lands and many urban lakes before entering the river 4.4 km below Lake Hancock. Studies compiled by Estevez and co-workers (1981) and Hand No. 4, 1990] CHAMPEAU— PEACE RIVER ICHTHYOFAUNA 303 Lake Hancock Peace Creek -H Whidden Creek -FM POLK 2b Hen ah i eR ih a Arete a Heme Ee aeere Zettai ttertasststersactrestrtctstenscreeetse HARDEE ee ee ee eee ee eee eee ee eer secrete ce ceceseceneseststatata® Matatatatatatate® ae Seen a eae ere a eet wy ae Ue rer ee ee ee 1 EERE REeEEE ee Meee eee ere sree ee ee ee SeSeSeseseseses PE cststetstatetetetatetetets ME Satatetatatetet ats & Matar etetas Mat etetatatatat erate ne ett eter ee Shell Creek CHARLOTTE Charlotte Harbor Fic. 1. Location of the Peace River and fish population sample stations (H-Homeland, FM- Fort Meade, W-—Wauchula, G—Gardner, N-Nocatee, FO-Fort Ogden), 1983-1988. and co-workers (1988) noted that progressive dilution from less disturbed tributaries resulted in improved water quality downstream. Ross and Jones (1979) determined that Charlie, Joshua and Horse Creeks were the least pol- luted of all Peace River tributaries. Groundwater withdrawals for agricultural, mining and potable uses have resulted in a significant decline in the potentiometric surface of the Floridan aquifer (Barcelo et al., 1989). Groundwater contributions to the Peace River water budget have been reduced, most severely at the upper valley (South- 304 FLORIDA SCIENTIST [Vol. 53 west Florida Water Management District, 1988). This groundwater flow re- duction exacerbates water quality problems by eliminating dilution effects and decreasing flow rates. Limited study of Peace River fishes has occurred. First collections of fish were by Woolman (1892), who collected 20 species from the river and several tributaries during the winter of 1890-91. The Florida Game and Fresh Water Fish Commission (FGFWFC, 1963) conducted fish sampling at irregular in- tervals from 1961 through 1963. Investigations of catastrophic fish kills by Ware (1969), Ware and Fish (1969) and Chapman (1973) have provided the most comprehensive information on Peace River fishes. Most recent fish col- lections were conducted in 1976 by Texas Instruments (Estevez et al., 1981). The objective of this study was to update the ichthyofaunal data base through a comprehensive five-year sampling program. The fish community was also evaluated to determine the degree that man’s activities have altered riverine ecology from the expected natural state. The advantages of using fish as indicators of environmental disturbance over water quality or other aquatic organisms were summarized by Karr (1981) and Hocutt (1981). Karr (1981) developed an ecological assessment system (index of biotic integrity or IBI) that incorporates a series of fish community attributes related to species composition and ecological structure. Metrics used to calculate IBI are re- gionally specific (Fausch et al., 1984) and no metrics applicable to peninsular Florida streams have yet been developed. In this study, evaluation of biotic stress was made by integrating the following parameters: fish abundance and biomass, species richness, composition and diversity. MATERIALS AND MErHops— Fish collections were made from six sites along Peace River (Fig. 1). Sample stations and distance downstream from Lake Hancock were: Homeland-21 km, Fort Meade-35 km, Wauchula-70 km, Gardner-123 km, Nocatee-170 km and Fort Ogden-195 km. River width varied between 7 and 12 m at all stations except for Nocatee and Fort Ogden where approximate widths were 30 m and 60 m, respectively. The fish community was sampled during various seasons from 1983 through 1988. Sampling was not conducted during extreme high nor low flow periods to avoid anomalous results. Fish were collected using an electrofishing boat, operating on pulsating direct current varying from 6.0 to 7.0 amps. Electrical output was regulated by a Smith-Root model VI-A electrofisher, utilizing the aluminum boat hull as the cathode and a pair of bow-mounted boom electrode arrays as anodes. Two people at the bow of the boat captured stunned fish with dip nets. Stand- ardized sampling technique was used in all samples. Available habitat types (brush, macro- phytes, pools and eddies) at each station were sampled for 30 to 60 minute time periods. Time periods were measured by the electrofisher as “pedal down time” or the amount of time electric- ity was actually applied to the water. A total of 57 samples was conducted for a total electrofish- ing effort of 2866 “pedal-down” minutes. Species richness is the number of species collected per sample. Species diversity (H’) was calculated by the Shannon-Weaver index (Shannon and Weaver, 1949). Numerical abundance per station is expressed by catch-per-unit-effort (CPUE) as the number of fish collected per “pedal-down” minute. Biomass indices are expressed as total weight (kg) of fish collected per “peda-down” minute. Fish community composition for each station is the median percent com- position of CPUE for all samples. 305 CHAMPEAU— PEACE RIVER ICHTHYOFAUNA No. 4, 1990] C3 GG 6G Ts 60 0°0 snjeydao [isn , T0 0°0 0°0 0'0 0'0 0'0 snsojns sniqosoi1py , 9 TT cal e9 6S ve 0'°0 sIjeutoepun snwiodonua’), 9°0 v0 T0 60 0'0 0'0 B7e1)SO1 BT[INSuy, T0 9°0 cl 6G v0 20 sn}e[NoeUL S9}D9UTIT, , 0°0 0'0 l0> 10 0'0 0'0 snuvdes sniopopo1ydy 00 10 0°0 0'°0 0'°0 0'0 QUIIOJISNJ BUIO}SO9Y} A G0 9°0 8G ST L‘0 G0 sn[NdoIs say sepiqe’] 10 10 T0 T0 6S VT SBONIOSAIO SNUOSIWII}ON 0°0 0°0 0°0 0'°0 0'°0 T0o> snjzelnoew sidon0oN 60 ith VI SL jen, L‘0 tuosiayed stdo10N 9G OV Lh Se eV 0's SI[OUIWas sn[npuNn] L‘0 T'0 cat 6 0 10 £G STUTJFe BISNGUIB St) 00 0'°0 9°0 10 €'0 O'T euUldyey] eiplos0g 00 00 0'°0 l0> 0'0 3'0 asusuejzed BUIOSOIOG 60 £0 U0 T0> Ll el wmnuetpedso euloso10q 0°0 00 £0 8'T £0 ¢'0 eyeoons uOZAWIIY 9'T vi £0 er eal v's BATeO BIUIY (nal LG SL CT G0 ¢'0 snasso snaysostda'T C'sT 691 Lael Cac 8°ET 83'°SSG snoutyiAze]d snoaysostda'T 0'0 0'0 0'°0 l0o> 0'0 0°0 snuIIAS sninjoN 0°0 60 T0 10> 0'0 0'0 sIjejeU sn.Ny]e,] T0 £0 0'°0 v'0 @ 0 r'0 snsoynqeu sn.inye,o] OL 9°0 L‘0 9°0 G0 r'0 snjevo sninyey] LG! 9°8 v6 8°S 80 90 snjzejound sninye zd] 00 00 0°0 10 0'°0 PI snjze[NOVUIOISIU STXOUIOg 60 60 v0 v0 €'0 c‘0 snso[ns stwioda'] 0'0 (0) €'0 20 0'0 roar) sn}eulsieur stuoda'T VI BL SIT 6 FI G13 Vil snjeyound stwoda'T] 8'V v8 G9 LS Il 9°6 snydoyororur sturoda'T 66 v8 0's aa 9°31 6'°OI sniroor1oew sturodaT jit VIl v6 6 IT L’8 VP saploulyes sna} do101jy sa[dures jo ueipayy—uolztsoduio7) yUs010g satoeds uspsO "17 990}8900N ioupiey emnyone oproy ‘Ww pueswopyy ‘88-E861 JOATY cova OUR BUOTP SUOT}E}S XIS LIOIJ P9}D9T[OO saTdures SuTYsTjo1{09]9 JO UOTISOduIOO soloed ‘| ATAV], FLORIDA SCIENTIST [Vol. 53 306 GG 0°0 0°0 6G 8°0 6 0 Te 0°0 T'0 8°9 0°0 0°0 £6 Soe a ee Sooncosd satoeds o1oxy, , satoods ouliey , P9}09][00 satoads jo Joquinu [e}0], B]Jopt uoposuAreydous}), , snyorieq Sele), voine eIdeil]L, , snuIO}SOY}ST]O sn.19}deIq, snuueih} &IOOAIIg, penunuoD— | aTAvL, No. 4, 1990] CHAMPEAU—PEACE RIVER ICHTHYOFAUNA 307 ResuLts—A total of 37 species of fish were collected from the river proper (Table 1). Indigenous freshwater species found in the main channel totaled 27 and represented 12 families. Along with the freshwater forms, seven marine species and three non-indigenous species were collected. No endangered or threatened species were documented. Centropomus undecimalis, a marine species, was the only fish collected that was of special concern status. No catastrophic events such as phosphate waste spills, severe drought or flooding occurred during the study period. Seasonal hydroperiod changes and periodic release of water from Lake Hancock resulted in inconsistent environ- mental conditions which caused variable data. Since the assumption of nor- mality could not be made, comparisons between stations (Table 2) were ana- lyzed non-parametrically (Kruskal-Wallis Test). TABLE 2. Peace River sample station comparisons of species richness, diversity (H’), numerical abundance (CPUE) and biomass, 1983-88. Number Median Median Median CPUE Median Biomass Station of Samples _ Richness H’ (fish/minute) (kg/minute) Homeland 9 15.0 2.05* OFOr | 1.4 Ft. Meade 3 17.0 Dp’) 3.4 1.0 Wauchula 14 15.5 2.20* 3.2 1.0 Gardner 12 13.0 ghee Mey) Oe Nocatee 9 14.0 2ASs 2.9 1.0 Ft. Ogden 10 11.0** Lee) G= 1.1 *Significant difference between other stations to p<0.05. **Significant difference between other stations to p<0.025. Total number of species collected at all stations ranged from 23 to 31 (Table 1) and species richness per sample varied from 9 to 20. Significantly lower richness occurred at the Fort Ogden station (p<0.025). Median rich- ness was not significantly different among the other stations (Table 2). Species diversity, H’, varied from 1.09 to 2.55 and was not correlated with species richness (R’=0.43), and H’ was significantly different between stations (p<0.05). Median H’ was low at Homeland (2.05, increased downstream (Ft. Meade-2.29, Wauchula-2.20, Gardner-2.28) and decreased as the river became tidally influenced (Nocatee-2.18, Fort Ogden-1.99). Median CPUE was highest at Homeland, 5.8 fish/minute, and lowest at Fort Ogden, 1.6 fish/minute. CPUE at the middle four stations varied from 2.1 to 3.4 fish/ minute and were not significantly different. Biomass was not significantly different between stations except for Gardner which was lowest at 0.70 kg/ minute. Biomass for the other stations ranged from 1.0 to 1.4 kg/minute. Fishes abundant at all stations were Lepisosteus platyrhincus, Tilapia aurea, Micropterous salmoides and several Lepomis spp. (Table 1). Abun- dance of Lepisosteus osseus, Centropomus undecimalis and Ictalurus puncta- tus were higher at the three downstream stations. Ictalurus catus was abun- dant only at the Fort Ogden station. The presence of Notropis petersoni, Notropis maculatus and Labidesthes sicculus appeared to be related to water quality and were considered indicative of favorable biotic conditions. Notro- 308 FLORIDA SCIENTIST [Vol. 53 pis petersoni was most abundant at Fort Meade, Wauchula and Gardner. Abundance of Labidesthes sicculus was higher at Wauchula and Gardner than at other stations. Discussion— The freshwater fish fauna of the Peace River is depauperate in species richness; which is primarily a result of the natural zoogeography of peninsular Florida (Gilbert, 1987). Review of earlier Peace River studies in- fers that anthropogenic influences may have reduced species richness during recent times. Records from FGFWFC (1963), Ware and Fish (1969), Chap- man (1973), Texas Instruments (Estevez et al., 1981 and Layne et al., 1977) documented 11 freshwater species not collected in the main channel during this study. Several of these species were collected by Champeau and co-work- ers (1988) in tributaries exhibiting higher water quality than the Peace River proper. It is probable that habitat requirements for these species are not cur- rently being met in the main channel and these species have been either reduced to less impacted tributaries or extirpated from the system. Besides cultural degradation of aquatic habitat, human introduction of non-indigenous fishes has also affected the ichthyofauna. Historical records (FGFWFC, 1963; Buntz and Manooch, 1968; Ware and Fish, 1969) infer that Tilapia aurea, an African cichlid, invaded the Peace River between 1963 and 1967. Buntz and Manooch (1968) documented the rapid expansion of Tilapia aurea in several Polk County lakes from 1961 to 1968. Migration to the Peace River most likely occurred from these upper-basin lakes. Tilapia aurea has become well established throughout the river and is a significant component of the community. These omnivorous filter-feeders exploit areas of Peace River where algal and detrital biomass are high. Tilapia aurea is opportunistic and capable of surviving dissolved oxygen levels that are intol- erable to most indigenous species. Dominance of Tilapia aurea in other Flor- ida systems has been associated with poor water quality and low biotic integ- rity (Foote, 1977). The ability of Tilapia aurea to exploit severely polluted habitats is well-documented; however, the degree of displacement of native fishes by Tilapia aurea in higher quality habitats is poorly understood (Noble et al., 1975; Noble and Germany, 1985; Shafland and Pestrak, 1983; Shafland and Metzger, 1986). Competition, predation by piscivores and al- teration of energy flow thereby changing trophic structure are possible im- pacts of Tilapia aurea on native ichthyofauna. Clarias batrachus became established in the Peace River between 1973 and 1977, although the route of initial river invasion is not clear. This species was not abundant in the samples and probably does not have a significant impact on the fish community. One specimen of Ctenopharyngodon idella was collected that probably escaped from a headwater lake where stocking was utilized for aquatic plant control. Several marine species were found in the Peace River. Centropomus unde- cimalis and Mugil cephalus were most common and were collected at all but the uppermost station. Centropomus undecimalis ranged from 1 to 12 kg in weight, and this piscivore is a competitor with Micropterus salmoides, Lepi- No. 4, 1990] CHAMPEAU— PEACE RIVER ICHTHYOFAUNA 309 sosteus spp. and Amia calva. Centropomus undecimalis was most abundant in the lower river and was commonly collected from habitats incorporating deep water, flow and dense cover. Trinectes maculatus was abundant at all stations where sandy substrates existed. Other marine species appeared to be transients with the exception of Anguilla rostrata, a catadromous eel. Two components of degraded water quality that appeared to affect the fish community most were high turbidity and unstable dissolved oxygen. High concentrations of phytoplankton in Lake Hancock effluent results in elevated levels of suspended organic matter and biological oxygen demand loading of the upper river (Hand et al., 1988; Estevez, 1981). High turbidity and low dissolved oxygen directly impact fish communities by displacing in- tolerant species. Tilapia aurea, Lepisosteus spp. and Amia calva became dominant since these species were able to exploit stressed conditions. Seasonal rainfall patterns and variable discharge from Lake Hancock re- sulted in fluctuations in diversity and species composition; however, these data indicate a pattern of differentiated fish assemblages from upper to lower stations. These differences are a result of pollution-loading of the upper river rather than a function of natural longitudinal zonation. Consistently poor water quality at the Homeland station resulted in low diversity; however, this area supported a high density of fish. Lepisosteus platyrhincus and Amia calva were dominant predators supported by a prey base consisting of Tilapia aurea and Lepomis spp. High fish density resulted from high algal biomass exhibited in the upper river. Low diversity and dominance of fishes tolerant to unstable dissolved oxygen demonstrated poor water quality in the upper river created biotic disturbances that greatly affected the fish community. Physical and biotic conditions became less stressful downstream of Home- land. The Fort Meade station is 35 km downstream of Lake Hancock, (14 km below Homeland) and improved conditions were evidenced by increased di- versity and higher abundance of Notropis petersoni and Micropterus sal- moides. This area does not receive dilution from major tributaries, and it is probable that biotic conditions are occasionally unstable. Reduced stress was evident at the Wauchula station 35 km downstream of Fort Meade. Increased diversity and more complex community structure indicated that habitat con- ditions were consistently more favorable than upstream. Four major and several minor tributaries enter the river between Fort Meade and Wauchula. Dilution from these inputs improves water quality which allows for more fish species to exist. Downstream 53 km, the Gardner station was influenced by input of relatively unpolluted water from Charlie Creek and seven smaller tributaries. Biomass estimates indicated that carry- ing capacity was lowest in this area; however, species diversity was high. Dilution of upstream loadings of chemical nutrients and suspended organic matter resulted in lower fish production while higher quality habitat pre- vented dominance and allowed for good equitability of species. Tidal influence on water flow at the Nocatee and Fort Ogden stations created biotic conditions dissimilar from observations upstream. Species rich- 310 FLORIDA SCIENTIST [Vol. 53 ness, H’, and CPUE decreased while fish production (biomass) was un- changed. Insectivorous and planktivorous species were scarce from the lower river while omnivores were abundant. Large piscivores (Lepisosteus spp., Centropomus undecimalis, and Micropterus salmoides) that could utilize a prey base consisting primarily of Tilapia aurea and Ictalurus spp. dominated the community. This community structure resulted in decreased fish density while maintaining higher biomass. ConcLusions— Cultural impacts have significantly affected biotic condi- tions of the Peace River system. Analysis of fish communities indicates the upper (35 km) and lower sections (35 km) showed greater impact than the middle section (125 km). These impacts are believed to be expressions of pollution from headwater lakes and streams. Dilution from less polluted trib- utaries improved biotic conditions in the middle section. Agricultural activi- ties and phosphate mining within the drainage basins of these streams is in- creasing and wise land management practices must be mandated for their future protection. Tributaries already affected by mining should be re- claimed to re-establish flow of high quality water to the main channel. Treat- ment of effluent from Lake Hancock and Peace Creek by diversion to created wetlands would improve biotic conditions throughout the river’s course. The Peace River fish community recovered from catastrophic events in the past (Ware and Dequine, 1967; Ware, 1969; Chapman, 1973) and re- sponds favorably to improved water quality. A holistic, long-term solution to improve the Peace River involves many factors such as ecologically-sound phosphate land reclamation, municipal and industrial wastewater manage- ment, urban and agricultural stormwater management, flood control, lake restoration, and regulation of land use. Protection and enhancement of the river’s aquatic resources are possible if these factors are incorporated into a comprehensive basin management program. LITERATURE CITED Barce.o, M. D., D. L. SLoNENA, S. C. Camp ann J. D. Watson. 1989. Ridge II: A hydrogeolo- gic investigation of the Lake Wales ridge. Southwest Fl. Water Manag. District. Brooks- ville, Florida. Buntz, J. AND C. S. Manoocu III, 1968. Tilapia aurea, a rapidly spreading exotic in south central Florida. Southeast. Assoc. of Game and Fish Comm., Proc. 22nd Ann. Conf. 495- 501. CHAMPEAU, T. R., K. W. DENSON, AND K. G. Garpner. 1988. Peace River fish population moni- toring and sunshine bass stocking evaluation. Completion Report. Fl. Game and Fresh Water Fish Comm., Tallahassee, Florida. Cuapman, P. G. 1973. Pollution investigation— Peace River and Whiddon Creek, Cities Service Company incident, December 3, 1971. Fl. Game and Fresh Water Fish Comm., Tallahas- see, Florida. Estevez, E. D., J. MILLER, AND J. Morris. 1981. Charlotte Harbor estuarine ecosystem complex and the Peace River. Vols 1 and 2. Mote Marine Lab. for Southwest Fl. Reg. Plan. Coun- cil. Ft. Myers, Florida. Fauscu, K. D., J. R. Karr, AND P. R. Yant. 1984. Regional application of an index of biotic integrity based on stream fish communities. Tran. Am. Fish. Soc. 113:39-55. FLoriwa Boakp oF HEALTH. 1965. Interim Report: Water quality of the Peace River 1959-1964. Tallahassee, Florida. No. 4, 1990] CHAMPEAU— PEACE RIVER ICHTHYOFAUNA 311 FLoRIDA DEPARTMENT OF ENVIRONMENTAL REGULATION. 1980. Water quality inventory for the State of Florida. Tallahassee, Florida. FLoripA GAME AND FRESH WATER FisH Commission. 1963. Peace River fish population survey. Tallahassee, Florida. Foote, K. J. 1977. Annual Performance Report. Blue tilapia investigations. Fl. Game and Fresh Water Fish Comm., Tallahassee, Florida. Gi.berT, C. R. 1987. Zoogeography of the freshwater fish fauna of southern Georgia and penin- sular Florida. Brimleyana No. 13:25-54. Hann, J., V. TAUXE, AND M. FRIEDMANN. 1988. Water quality assessment for the State of Florida. Dept. Environ. Reg., Tallahassee, Florida. Hocutt, C. H. 1981. Fish as indicators of biological integrity. Fisheries (Bethesda) 6(6):28-31. Karr, J. R. 1981. Assessment of biotic integrity using fish communities. Fisheries (Bethesda) 6(6):21-27. Layne, J. N., J. A. StaLtucup, G. E. WooLFENDEN, M. N. McCautey, Anp D. J. Wor ey. 1977. Fish and wildlife inventory of the seven-county region included in the central Florida phosphate industry area-wide environmental impact study. U.S. Dept. Interior, Contract No. 14-16-0009-77-005. Washington, D.C. Nosxe, R. L., R. D. GERMANY, AND C. R. Haut. 1975. Interactions of blue tilapia and large- mouth bass in a power plant cooling reservoir. Southeast. Assoc. Game and Fish Comm., Proc. 29th Ann. Conf.:247-251. NoBLE, R. L. AND R. D. Germany. 1985. Changes in Fish Populations of Trinidad Lake, Texas, in response to abundance of blue tilapia, Tilapia aurea. Proc. Fish Culture and Mang. Symp., American Fisheries Society, Bethesda, Maryland. Ross, L. T. ann D. A. Jones. 1979. Biological aspects of water quality in Florida. Pt. III. Withlacoochee, Tampa Bay, Peace and Kissimmee Drainage Basins. Fl. Dept. Envirn. Reg., Techn. Ser. 4(3):1-291. SHAFLAND, P. L. AnD R. Merzcer. 1986. Annual Performance Report. Associations of blue tilapia with native fishes. Fl. Game and Fresh Water Fish Comm., Tallahassee, Florida. SHAFLAND, P. L. anp J. M. Pestrak. 1983. Suppression of largemouth bass production by blue tilapia in ponds. Southeast. Assoc. Game and Fish Comm., Proc. 37th Ann. Conf.:441- 446. SHANNON, C. E. anp W. Weaver. 1949. The Mathematical Theory of Communication. Univ. Illinois Press. Urbana, Illinois. SouTHWEST FLoRIDA WATER MANAGEMENT Districr. 1988. Groundwater resource availability inventory: Polk County, Florida. Brooksville, Florida. Wake, F. J. anv J. F. DEQuine. 1967. Joint report on the recovery and current status of the Peace River for fishes and other aquatic life. F1. Games and Fresh Water Fish Comm., Tallahas- see, Florida. Wake, J. F. anp W. V. Fisu. 1969. Peace River Study: Pollution investigation. Mobil incident. F1. Game and Fresh Water Fish Comm.., Tallahassee, Florida. Ware, F. J. 1969. Effects of phosphate clay pollution on the Peace River, Florida. Southeast. Assoc. Game and Fish Comm.., Proc. 23rd Ann. Conf.:359-0373. Wootman, A. J. 1892. Report on the rivers of central Florida tributary to the Gulf of Mexico, with lists of fish inhabiting them. Bull. U.S. Fish Comm. 10 (1890):293-302. Florida Sci. 53(4): 302-311. 1990. Accepted: February 23, 1990. Anthropological Sciences BEHAVIORAL CHARACTERISTICS OF SQUIRREL MONKEYS AT THE BARTLETT ESTATE, FT. LAUDERDALE RyAn J. WHEELER Department of Anthropology, Florida Atlantic University, Boca Raton, FL 33431 ABsTRACT: Several colonies of exotic primates are known to exist in Florida. This study was undertaken to report on the existence of the semi-wild colony of squirrel monkeys at the Bartlett Estate in Ft. Lauderdale, Florida and examine the behavioral characteristics of these animals. The Bartlett monkeys are compared with wild troops in South America and with provisioned animals at Monkey Jungle, a tourist attraction in Miami, Florida. Many similarities were ex- pected since squirrel monkeys are known to perform a number of rather stereotypical activities (i.e. penile display). These types of activities were observed, but there were also deviations from the reports given for other troops. THE purpose of this study is to report on the squirrel monkey troop at the Bartlett Estate, Ft. Lauderdale. Goals include determining the population size, population composition with regard to age and sex, existence of sub- groups, origin of the troop, and the species of squirrel monkey in question. Descriptions of subsistence, reproduction, communication, grooming, play, and aggression are included and compared with reports on wild and other semi-wild populations. A possible explanation for the remarkable adaptation of a tropical species to a sub-tropical environment is also presented. Stupy AREA—The Bartlett Estate is bounded by State Road AIA to the East, the Intracoastal Waterway to the West, Vistamar St. to the South, and N.E. 9th St. to the North. Several interesting ecosystems combining native and exotic vegetation can be found on the 35 acre, Ft. Lauderdale estate. The western portion of the property contains tall-canopied Australian pines and red mangroves which border the Intracoastal Waterway and a shallow natu- ral channel. The main road through the center of the estate is lined with Australian melaleuca trees and several large Ficus fig trees. The area nearest AIA is a dense forest predominated by large fig trees, Gumbo Limbos, Sea Grapes, various fruiting trees, and, in some parts, thick scrub. A large spring- fed pond is located in front of the Bonnet House, which is the focal point of the estate. Summers are normally hot and wet while winters are cool and dry. MernHops—This study is based on about 100 hours of direct observation of the animals made by the author during May, June, July, August, and October, 1988. The methods utilized follow those described by Baldwin (1967) and Thorington (1968). Observations were made at all hours of the day with a special focus on early morning and early evening when the animals were most active. The animals were watched with a 7-15 X 35, zoom binoculars and with the unaided eye. All observations were logged into a field journal and on several occasions the animals’ activities were recorded on video tape. The animals were easily followed as they traveled along the tree- lined paths; they were rarely agitated by the presence of a human observer. The animals were occasionally provisioned in order to census them. All animals were identified by the author and many were assigned names to aid in evaluating intra-animal behavior. Notes on weather condi- tions were entered along with behavorial data into the field journal. No. 4, 1990] WHEELER— SQUIRREL MONKEY BEHAVIOR 313 TABLE 1. Bartlett Estate subgroup composition (May 1988). Subgroup Females Males Total Adults Juveniles Adults Juveniles Red’s 2 4 — — 6 Silver’s 4 3 — — 7 Louise’s 2 5 — -- 7 Sasha’s 2 3 -— -- 5 Males — — 3 15 18 N=43 ResuLts— Observations of the Bartlett Estate squirrel monkeys yielded information regarding the species of monkey present, the size of the colony, intra-animal behavior, and adaptation to the environment. Colony History—The squirrel monkey colony that inhabits this territory is of the Brazilian variety and are the descendants of two pairs of animals that were released 25 years ago from a nearby social club. Napier and Napier (1967) suggest that Saimiri sciureus, with a blue-grey crown, is the common South American squirrel monkey and Saimiri oerstedii, with a black crown, is the Costa Rican/Panamanian variety. Accepting this classification, the au- thor has concluded that the Bartlett Estate animals are Saimiri sciureus. Population Dynamics— There were 42 monkeys in the colony when obser- vations began in May, 1988; five of the animals were pregnant at the time and gave birth during the study. Accurate counts revealed that there were 47 squirrel monkeys present in October, 1988 when the observations ended. Census information was collected when the animals gathered to accept grapes, and when they traveled along a fence or powerline (Table 1). Each female subgroup has a matriarch which seems to be an older animal that will typically lead during foraging and may initiate subgroup move- ment. It appears that subgroups are an excellent means by which to exploit trees with widely scattered fruit. These subgroups have fixed membership and animals within each subgroup are typically seen sleeping and foraging together. The young males travel together with the troop and can be observed playing and resting with each other. It seems that as the young males reach one year of age they are turned out of their family subgroup and they join this play unit. The three adult males do not associate with any particular sub- group and do not assume any leadership role for the colony as a whole; how- ever, the males interact with all the animals and occasionally engage in play with the juveniles. Subsistence—Thorington (1968) states that due to their wide range, squir- rel monkeys have adapted to many botanically and zoologically different habitats. This capability has clearly allowed them to live and reproduce in southern Florida. Studies indicate that Saimiri are diurnal foragers that spend most of their time searching for insects (Klein and Klein, 1979). The Bartlett Estate monkeys typically awake at dawn and move into a fruiting tree where they feed intensively. Ficus fig, Sea Grape, Black Cherry, Surinam Cherry, Sapodilla, Mango, and Avocado trees are all utilized, but 314 FLORIDA SCIENTIST [Vol. 53 examination of fecal remains suggests that figs are the monkeys’ stable food during the summer and fall. As the animals move through the property in their subgroups they occasionally find lizards, tree-crabs, and Cuban tree snails, all of which seem to be prized foods. More common fare includes large carpenter ants which are collected by peeling tree bark, spiders which can be found by unrolling leaves, orb-weaving spiders, wasps, bees, dragonflies, cat- erpillars, and butterflies. Unrolling leaves and peeling park are apparently two methods of foraging shared by the Bartlett and Columbian squirrel mon- keys (Thorington, 1968). Leaves are chewed and licked to obtain moisture; the monkeys also retrieve water from knot holes and dripping spigots. Reproduction— Actual coitus was observed only six times, suggesting that the mating season had already ended. The sexual encounters recorded at the Bartlett Estate were similar to the “group activities” described by DuMond (1968) in which other males rush to the coital pair to smell their genitals and touch their lips. Squirrel monkey birth seasons coincide with the period of greatest rain- fall, placing the Florida birth season between June and August and the mat- ing season between December and March, analogous to the situation at Mon- key Jungle (DuMond and Hutchinson, 1967). DuMond and Hutchinson (1967) suggest that seasonal spermatogenesis occurs in male squirrel monkeys and causes several temporary secondary sex- ual characteristics to appear. These include a heaviness in the arms and torso, giving the animals a “fatted” appearance, and an increase in aggressive social interactions during the height of the mating season (DuMond and Hutchin- son, 1967). The “fatted” condition was observed in Jim, one of the Bartlett males, at the beginning of the study, but seemed to be subsiding. Communication—Communication in Saimiri includes an interesting set of behavioral activities, most of which involve vocalizations. Winter (1968) catalogs a host of these vocal signals, but Thorington (1968) notes that it is difficult to distinguish many of these in the wild. The following is a list of calls commonly heard at the Bartlett Estate: Isolation peeps, “eeeeee”, are heard when an animal becomes separated from the group. Isolation calls always elicit a similar call from other animals and allow the separated monkey to maintain contact with the group. Alarm peeps are very short calls and spread throughout the Saimiri group quickly. This call is heard in response to avian or terrestrial predators, and the animals will move up or down in the canopy depending on the origin of the threat. Twittering is used during feeding situations and as a reaction to other squirrel monkeys. Purr is an affectionate “rrrrt” call with a true purr or humming sound given by mothers to infants and by males during coitus. In agreement with Winter’s study (1968), the most common calls heard were “peep calls”. Visual signals are another means of communication in Saimiri and most of these have been described in other sections of this paper. No. 4, 1990] WHEELER— SQUIRREL MONKEY BEHAVIOR 315 Play— Among the activities observed at the Bartlett Estate, play was the most interesting and diverse. Non-social activity play involving running, jumping, and swinging was observed in the five infant squirrel monkeys and conformed to descriptions made by Rosenblum (1968). DuMond (1968) de- scribes two types of social play at Monkey Jungle; contact play which involves two animals wrestling and scuffling on the ground, and distance play in which the play pair chase each other. Social play also takes on sexual connota- tions with older individuals; this type of play may include pursuit, mounting, and an interesting “invitation posture” in which one animal will look be- tween its legs or over its shoulder at its play partner (Baldwin and Baldwin 1974). The Bartlett Estate squirrel monkeys engage in all of the above types of play frequently and in a type of group game which is not described in the literature. This game typically begins when two or three juvenile males wres- tle near the ground. They are soon joined by the rest of the juvenile males and by some of the young females. Running, chasing, hopping, wrestling, head grasping, tail pulling, mounting, and the sexual invitation stance are all com- ponents of this game. The most striking difference between group play and other social play is that it involves the constant exchange of partners and is focused primarily on the ground. Group play was observed nine times and lasted as long as an hour and as short as fifteen minutes. Aggression— Physical aggression was not observed during the study, but an aggression display was used by the animals to signal dominance. Winter (1968) describes this as the “open” penile display in which one animal directs its genitalia at a partner; displays of this nature were seen on five occasions and occurred between the adult males. Penile displays and head grasping were common components of the group game described above. DiscussiloN— Population dynamics of other squirrel monkey groups ap- pear to be very similar to that of the Bartlett animals. Klein and Klein (1979) and Thorington (1968) also observed subgroups within Saimiri colonies. Evi- dence from the Bartlett colony partially confirms Thorington’s suspicion (1968) that these subgroups represent family units inasmuch as the subgroups at the Bartlett Estate have fixed membership. DuMond (1968) notes groups of pre-adult males at Monkey Jungle, but these tend to live separately from the rest of the animals. There does seem to be a difference between the Bartlett colony and the Monkey Jungle colony regarding aggression. DuMond (1968) describes vio- lent outbursts by adult males which occasionally were fatal or left lasting scars, and attacks made by adult females on members of the pre-adult male group. These activities were not reported for wild troops or for the Bartlett animals and may be due to the relatively large size of the Monkey Jungle population and the small habitat area provided. Comparison with other accounts of squirrel monkey subsistence indicate that the Bartlett monkeys prefer the same fare and strategies as their counter- parts at Monkey Jungle and in South America. Hill (1960) notes that protein from insects—including flies, butterflies, mosquitoes, and spiders—is an im- 316 FLORIDA SCIENTIST [Vol. 53 portant part of their diet, but he states that Saimiri subsist primarily on vegetable matter. DuMond (1968) indicates that the provisioned colony of Monkey Jungle squirrel monkeys collect flowers, fruits, berries, the growing tips of a variety of native and introduced plants, and also search for flying insects under wood and leaves. Vegetable matter does seem to be the staple upon which all squirrel monkeys rely, with protein from vertebrates and in- vertebrates providing a necessary and desired supplement. This study indicates that many of the types of play described by Baldwin and Baldwin (1974), DuMond (1968), and Winter (1968) are, in fact, very stereotypical activities which are performed by all Saimiri. However, when considering the group game observed among the Bartlett monkeys, there does seem to be considerable room for adaptation. CoNCLUSIONS— Many similarities exist between the Bartlett Estate squir- rel monkeys and other squirrel monkeys in wild and semi-wild environments; however, the Bartlett animals’ behavior differs in two ways. Intense group play on the ground and insignificant amounts of aggression are two interest- ing behavioral characteristics which have allowed the colony to grow and persist in sub-tropical Florida. Elaborate play may allow the animals to es- tablish cohesive social bonds, in turn, these bonds eliminate aggression as the animals mature. Any fighting resulting in the death of a reproductively active animal would adversely affect the integrity of such a small colony. ACKNOWLEDGMENTS—I would like to thank Mrs. Bartlett, the staff of the Bonnet House, and John Fletemeyer. LITERATURE CITED BaLpwin, J. D. 1967. A study of the social behavior of a semi-freeranging colony of squirrel monkeys (Saimiri sciureus). Ph.D. dissertation, Johns Hopkins Univ., Baltimore. AND J. I. BaLpwin. 1974. Exploration and social play in squirrel monkeys (Saimiri). Am. Zoolog. 14:303-314. DuMonp, F. V. 1968. The squirrel monkey in a seminatural environment. Pp. 88-144. In: Ro- SENBLUM, L. A. AND R. W. Cooper (eds.). The Squirrel Monkey. Academic Press, New York. AND T. C. Hutcuinson. 1967. Squirrel monkey reproduction: The ‘fatted’ male phe- nomenon and seasonal spermatogenesis. Science 158: 1067-1070. HILL, W. C. O. 1960. Primates: Comparative Anatomy and Taxonomy. Edinburgh Univ. Press. Edinburgh. Kuen, L. L. AND D. J. Kie1n. 1979. Social and ecological contrasts between four taxa of neo- tropical primates. Pp. 107-131. In: Sussman, R. W. (ed.). Primate Ecology. John Wiley & Sons, New York. Napigr, J. R. anp P. H. Napier. 1967. A Handbook of Living Primates. Academic Press, New York. . 1985. The Natural History of the Primates. MIT Press, Cambridge. RosENBLUM, L. A. 1968. Mother-infant relations and early behavioral development. Pp. 207- 233. In: RosENBLUM, L. A. AND R. W. Cooper (eds.). The Squirrel Monkey. Academic Press, New York. Tuorincton, R. W. 1968. Observations of squirrel monkeys in a Columbian forest. Pp. 69-84. In: RosENBLUM, L. A. AND R. W. Cooper (eds.). The Squirrel Monkey. Academic Press, New York. Winter, P. 1968. Social communication in the squirrel monkey, Pp. 235-252. In: RosENBLUM, L. A. AND R. W. Cooper (eds.). The Squirrel Monkey. Academic Press, New York. Florida Sci. 53(4): 312-316. 1990. Accepted: April 3, 1990. Florida Scientist QUARTERLY JOURNAL of the FLORIDA ACADEMY OF SCIENCES VOLUME 53 DEAN F. MakrrTIN Editor BARBARA B. MARTIN Co-Editor Published by the FLORIDA ACADEMY OF SCIENCES, INC. Orlando, Florida 1990 The Florida Scientist continues the series formerly issued as the Quarterly Journal of the Florida Academy of Sciences. The Annual Program Issue is published independently of the journal and is issued as a separately paged Supplement. Copyright© by the FLoripa AcADEMy OF SCIENCES, INc. 1990 CONTENTS OF VOLUME 53 NUMBER 1 Monitoring Florida’s Riverine Fish Communities ...................004. D. Gray Bass, Jr. Vegetation on the Florida Atlantic University Ecological STUBS o wero ciateeaheeabete SOE ACRE A nee Ca rs ee Daniel F. Austin Nonpoint Source Phosphorus Control By a Combination Wet Detention/Filtration Facility in Kissimmee, Florida .................. Jeffrey Dee Holler Survival of Florida Bay Fish Tagged with Internally Anchored SHPEEAISO I PGS acca oe > Cee en, ee rE Gerald M. Ludwig, Jorgan E. Skjeveland, and Nicholas A. Funicelli Postures Associated with Immobile Woodland Salamanders, Penns Od ONS se Le TY Sse elaine Fee, C. Kenneth Dodd, Jr. exmmowiedement Of RCVIEWEFS 2s. 05 ke ce ee eo edhe BE wine ae meals Note on the feeding behavior of the Common Atlantic Marginella Prunum Apicinum (Gastrododa, Marginellidae) ..................... Thomas M. Baugh OGLE ROVE Gee eee en nn neg ocean Cn Rey Ae Clinton J. Dawes Steven P. Christman and Walter S. Judd Landfills—A Thing of the Past? ................. Richard C. Johnson, Sr. NuMBER 2 Environmental Influences on the Distribution of Rice Rats (Oryzomys palustris) in Coastal Marshes .................+.-- James L. Wolfe Recent Changes in the Distribution of Calulerpa prolifera MMeMeuGlan WIVER LAGOON, FlOFIGa, oy. .ccys cy < Gseate Bon, enced niebeya 0i6 ae ogee aon ont Conrad White and Joel W. Snodgrass Vascular Plants of Fakahatchee Strand State Preserve ...............00-. Daniel F. Austin, Julie L. Jones, and Bradley C. Bennett eh vosonptionmimblorida Springs .. .u.c: es cs oe et we eo eee eee ene Carlos M. Duarte and Daniel E. Canfield, Jr. Food Selection by Early Life Stages of Blue Tilapia, Oreochromis aureus, in Lake George, Florida: Overlap MALMO VIMIpPAatric SiiaduGaATae.-4 si g.t heh ee cca teat SoG rkeee efeitos hoses hawt ae Alexander V. Zale and Richard W. Gregory Development and Application of a Soil-Air Radon Analysis pee lanai ule pine El On Ua recap tae ce sires a Pet ca lutte A ate hewslid Gadi quieres Robert S. Braman and Robert L. Sutton A Note on the Fire Responses of Species in Rosemary Scrubs on the Somenenileawe Wales RiGGe\s 5 aniateskaceda nr a wis ieeirecnd on00Sonteahe® 4 0a, 0,0 Ann F. Johnson and Warren G. Abrahamson SOG RGSS Aan age a ener fee nen er ee Richard P. Wunderlin NUMBER 3 OREWOLG er oe Po ek ae Ned P. Smith and Richard L. Turner Numerical Modeling of Tidal Hydrodynamics and Salinity ianspore inthe indian River Wagoony *. (Ge ee ek Y. Peter Sheng, S. Peene, and Y. M. Liu 28 38 52 74 81 85 89 118 123 130 138 144 145 147 Groundwater Seepage into the Indian River Lagoon at Port St. TEuicie 5 ones oe eek Aes eee Ashok Pandit and Clovis Clovis El-Khazen Water Quality Changes Associated with Rotary Ditched and Breached Mosquito Control Impoundments in Mosquito Eagoon’.. ooo. eh eon nen a J. C. Gamble, J. P. Stewart, and W. R. Ehrhardt Precipitation Chemistry: Atmospheric Loadings to the Surface Waters of the Indian River Lagoon Basin by Riamnrall se oP FA eae ete eee Thomas W. Dreschel, Brooks C. Madsen, Lee A. Maull, C. Ross Hinkle, and William M. Knott III Salt Marsh Mitigation: An Example of the Process of Balancing Mosquito Control, Natural Resource, and Development IGE ROSES Sinan wa tema o sea ae eee Peter D. O’Bryan, Douglas B. Carlson, and R. Grant Gilmore Holocene Evolution of Indian River Lagoon in Central Brevard County, Florida .... 0.2.0. .c00s cesses cce sob eee ee Sharon F. Bader and Randall W. Parkinson An Introduction to the Tides of Florida’s Indian River Lagoon. II. UIE MUS? 6. sy crscctv'nn to oi ce S.A won ae as dei MR ee Ned P. Smith Ichthyofauna Associated with Spoil Islands in Indian River agoon. Flotida sco. cc tee Ces eet teu Nancy Brown-Peterson and Ross W. Eames Adaptive Specializations of the Cyprinodont Fish RAOUL US MATIMOTAIUS, hae a $540 9 cas Pines ee D. Scott Taylor The Large Spatial and Temporal Biological Variability of Imeian River Lagoon. 6... 4sc-sc0eur soem etedannay wy Robert W. Virnstein NUMBER 4 Novel Marine Steroids from the Florida Keys. I. Isolation and Structure Elucidation of A®-9, 11-seco-Gorgostene-3, 11,246- triol-9-one from the Gorgonian (Pseudopterogorgia americana (Gmelin))) 2.55306 ehs 4d dower s ooh eee Cte Re ee M. J. Musmar and Alfred J. Weinheimer BOOK REVIEW 22 s4i2n clio bead hd SE ee eee Richard Franz BOOkKIREVICW 5 sco Aon Oe se ee eee Richard Wunderlin Hoodsot the Key Deer ..<22352:60: - yee eee eee W. D. Klimstra and Allan L. Dooley Scale Development in Ovuliferous Cones of Pinus Elliottii Engelm., Pinaceae seat. 07 8005 Sn eo ee eee R. F. Mente and S. D. Brack-Hanes Distribution of the Eastern Chipmunk (Tamias striatus) in | 2 (c) oc FN reed NI ry he ee na ac reese es renee eo 3c Jeffrey A. Gore A Review of the Florida Crayfish Fauna, with Comments on Nomenclature, Distribution, and Conservation .............000eeeees Richard Franz and Shelley E. Franz Meaningful Environmental Data: Not Just the Laboratory's Responsibility ..........:...2.5205- eee Ichthyofaunal Evaluation of the Peace River, Florida ................... Thomas R. Champeau Behaviorial Characteristics of Squirrel Monkeys at the Bartlett Estate; Ft. Lauderdale... 2.2.42 .- =F w” AMM = 7) an NOILNLILSNI NVINOSHLINWS S ”) > ye (7? sai 7) a: - 5 & oc ey o ao ea o — oO = ai z - - = am) o = OG ke >on > FS ‘om rad ay eS Ae z Oo NOILNLILSNI NVINOSHLINS S$ NWINOSHLIWS \. 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