€) , ISSN: 0098-4590 EY “Florida Scientist Volume 67 Spring, 2004 Number 2 CONTENTS Mass Occurrence of the Jellyfish Stomolophus meleagris and an Associated Spider Crab Libinia dubia, Eastern Florida .................. Bjorn G. Tunberg and Sherry A. Reed 93 Uptake of Phosphate and Nitrate Using Laboratory Cultures of Lemna NEE ete IN IE A eR ULL ealas Seda ean na sbonsdcusuceelusehesss vee Daniel P. Smith, Matthew E. McKenzie, Craig Bowe, and Dean F. Martin 105 The Sand Pine Scrub Community: An Annotated Bibliography, erm rt ee ee dls icc sae ino lsu'edies dela Locided sata ss Senn adacnoetie Donald R. Richardson 118 Effects of Social Environment in Early Life on Cortical Depth, Locomotor Activity, and Spatial Learning in the Golden Mouse, STE SAD PEG ean OU tO ee rey Fred Punzo 144 First Report of Aplidium antillense (Gravier, 1955), (Tunicata, Pee emermeiiata) (ON FIOTIGA...:)...25......6.s001cceassecesseaededencedessecenes Thomas Stach 154° A Brief Description of the Courtship Display of Male Pike Killifish PRE APLC SAO Noes hia Ne es los cannes esse bed dad devas davedolndestnceses ere ety OL SCICNCES Medalists o.........06.. 0500 fiiecessoveelacceoessccceoeess 166 PIE MMMIMCTIS TOP THES SCICHCES 200 1)..........5006..sccccccdooesessccesceessessceenes 168 FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright © by the Florida Academy of Sciences, Inc. 2004 Editor: Dr. Dean F. Martin Co-Editor: Mrs. Barbara B. 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Printing by Allen Press, Inc., Lawrence, Kansas Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES DEAN F. Martin, Editor BARBARA B. Martin, Co-Editor ~ Volume 67 Spring, 2004 Number 2 Biological Sciences MASS OCCURRENCE OF THE JELLYFISH STOMOLOPHUS MELEAGRIS AND AN ASSOCIATED SPIDER CRAB LIBINIA DUBIA, EASTERN FLORIDA BJORN G. TUNBERG AND SHERRY A. REED Smithsonian Marine Station, 701 Seaway Drive, Fort Pierce, Florida 34949 ABSTRACT: The jellyfish Stomolophus meleagris was collected both randomly and selectively in the Fort Pierce Inlet area of the Indian River Lagoon, eastern Florida on 26 and 28 March 2003. The total number of S. meleagris randomly sampled was 382 of which 16.5% carried an associated spider crab, Libinia dubia. Two S. meleagris carried two crabs each. The male/female ratio of the crab was 0.82. The mean carapace width (CW) of the males was 22.9 mm and the females 20.0 mm. The difference in size was significant between the sexes. Crabs were only found on jellyfish with a bell diameter between 80 mm and 110 mm, while the total size range of the jellyfish was 70-130 mm. More than twice as many females than males were found on jellyfish with a bell diameter of SO mm, but otherwise the sex distribution was similar regardless of the size of S. meleagris. There was no significant difference between the sexes concerning the relationship between size (CW) and live wet weight. Key Words: _Stomolophus meleagris, Libinia dubia, Indian River Lagoon, Florida STOMOLOPHUS meleagris is one of the most abundant species of scyphome- dusae along the southeastern and Gulf coasts of the United States (Mayer, 1910; Kraeuter and Setzler, 1975; Burke, 1976; Calder and Hester, 1978). According to Corrrington (1927) S. meleagris is by far the most abundant scyphozoan of the South Carolina coast, and one of the more conspicuous planktonic organisms of the littoral zone. The life history has been described by Calder (1982) and the feeding habits by Larson (1991). The spider crab Libinia dubia is found on almost all types of bottom in shallow ocean waters and saltier estuaries from nearshore to ca 50 m depth (Williams, 1984). The known range is from Cape Cod to southern Texas, Bahamas and Cuba. Tabb and Manning (1961) reported that L. dubia is common in Florida Bay and 23 94 FLORIDA SCIENTIST [VOL. 67 Dragovich and Kelly (1964) reported that it is the most common spider crab in Tampa Bay. The larval development of L. dubia has been described by Sandifer and Van Engel (1971). Jellyfish commonly harbor commensal forms, with certain symbionts being characteristic. For instance, some brachyuran crabs exhibit protective and transpor- tive forms of commensalism with jellyfish (Trott, 1972). The association between S. meleagris and L. dubia was first reported by Corrington (1927). Williams (1984) also describes the association between L. dubia and S. meleagris. However, the information on this association is very limited and no detailed observations have been reported. In an earlier study, Weymouth (1910) described the association between Cancer gracilis and jellyfish (species not determined), and Trott (1972) described the relationship between Stomolophus nomurai and the portunid crab Charybdis feriatus. Young blue crabs, Callinectes sapidus, have frequently been observed clinging to the umbrellas of the sea nettle, Chrysaora quinquecirrha (Jachowski, 1963). A mass occurrence of S. me/eagris, some with crabs clearly visible on the body surface, was observed in the Fort Pierce inlet area, eastern Florida in March 2003 that allowed analysis of the association. MetTHOps—Random sampling of the S. me/eagris was performed on 26 and 28 March 2003 from a small boat in the Fort Pierce Inlet area of the Indian River Lagoon (IRL) by using fine mesh dip nets (Figure 1). Each jellyfish was transferred to a container filled with saltwater. The bell diameter was measured with a plastic ruler to the nearest cm, and each specimen was thoroughly examined for L. dubia. If a crab was found, it was measured (carapace width) to the nearest mm and sexed. All crabs were then transferred to another container filled with saltwater for further treatment in the laboratory. On 28 March, selective sampling was also performed, 1.e., only jellyfish with visible crabs associated with them were collected. On 31 March, all crabs sampled on 28 March were weighed to the nearest 0.1 g in the laboratory. The relationship between the bell diameter of the jellyfish and the crab carapace width and the bell diameter and the sex distribution of the crabs were calculated from the random sampling data. RESULTS—Twenty-eight males, 34 females and 1 specimen, presumably a juvenile, of L. dubia were found when random collections of S. meleagris (382 specimens) were performed on 26 March, which corresponds to a male/female ratio of 0.82, and an association rate of 16.5%. The size distribution of L. dubia of the randomly collected S. meleagris is presented in Figure 2. The mean carapace width of the males was 22.9 mm (SD = 4.8 mm) and the females 20.0 mm (SD = 4.0 mm). The size difference was significant between the sexes (p = 0.004, Mann Whitney Rank Sum test). The crab size distribution of the selectively collected jellyfish is presented in Figure 3. The mean carapace width of the males was 23.2 mm (SD = 4.8 mm) and the females 23.5 mm (SD = 4.0 mm). The male/female ratio of the crab on the selectively collected jellyfish was 2.14 (62 males and 29 females). The size difference was not significant between the sexes (p = 0.72, unpaired t-test). The selectively sampled females were significantly larger (CW) than the randomly sampled ones (p = 0.003, Mann Whitney Rank Sum test), but there was no significant size difference between the males (p = 0.561, unpaired t-test). No. 2 2004] TUNBERG AND REED—JELLYFISH AND SPIDER CRABS 95 27°29'N fi Atlantic Ocean af Fort Pierce ) Inlet yi cA ft Dope 1 @ } Indian River | Lagoon aT) 80°19°W oe a Fic. 1. The Fort Pierce Inlet area of the Indian River Lagoon, eastern Florida. Sampling of Stomolophus meleagris was performed on 26 and 28 March 2003 within the area marked with a rectangle. SMS = Smithsonian Marine Station. The male/female ratio of all crabs combined was 1.43 (90 males and 63 females). During the selective sampling on 28 March two specimens of S. meleagris had two crabs attached, one (bell diam. = 100 mm) with two males (CW = 30 mm and 19 mm) the other (bell diam. = 90 mm) with two females (CW = 28 mm and 19 mm). The relationship between carapace width and wet weight for males and females is presented in Figure 4. The best fit was achieved by using polynomial regressions. Both regressions were highly significant: p < 0.001 (power of performed test with alpha = 0.050: 1.000). There was no significant difference in size or weight between the females and males (p = 0.114 and p=0.081, respectively) (Mann-Whitney Rank Sum Test). The relationship between the carapace width of males and females of L. dubia and the bell diameter of S. meleagris is presented in Figure 5. There was no sig- nificant correlation between bell diameter of the jellyfish and the carapace width for females (p = 0.115) and males (p = 0.469) (Pearson Product Moment Correlation). The relationship between bell diameter of the jellyfish and the sex distribution of the crabs is presented in Figure 6. More than twice as many females than males 96 FLORIDA SCIENTIST [VOL. 67 Ml females males IXAN juveniles NUMBER OF INDIVIDUALS @) 5) 10 15 20 295 30 oD) CARAPACE WIDTH (mm) Fic. 2. Size distribution of Libinia dubia from Stomolophus meleagris collected randomly on 26 and 28 March 2003. were found on jellyfish with a bell diameter of 80 mm, but otherwise the sex distribution was similar regardless of the size of S. meleagris. Figure 7 shows the number of crabs found on different size classes of S. meleagris. As presented earlier, crabs were only found on jellyfish within size No. 2 2004] TUNBERG AND REED—JELLYFISH AND SPIDER CRABS 97 12 MM females n= 62 [<=] males 10 15 20 25 30 35 40 CARAPACE WIDTH (mm) Fic. 3. Size distribution of Libinia dubia from Stomolophus meleagris collected selectively on 28 March 2003. 10 NUMBER OF INDIVIDUALS classes 80-110 mm. The highest percentage of crab occurrence was recorded on 80 mm jellyfish (22.6%) and the lowest (10.9%) on those measuring 110 mm. Discussion—Mayer (1910) reported that mature individuals of S. meleagris were abundant in winter and spring off the coast from Florida to South Carolina. 98 FLORIDA SCIENTIST [VOL. 67 @ fem width vs fem weight ——- plot females (1) ~=male width vs male weight —— plot males Females Coefficients: b[0O] 12.23 bit 4875 b[2] 0.09 b[3] - 9.01 e-4 r?- 0.981 Males Coefficients: b[0] -23.02 Dit. <3.12 b[2] -0.14 b[3] 2.38 e-3 eee OSI LIVE WET WEIGHT (g) 10 15 20 PBS) 30 35 40 CARAPACE WIDTH (mm) Fic. 4. The relationship between carapace width (CW) and live wet weight of Libinia dubia. It was rarely seen in brackish harbors, and was mostly confined to ocean water off the coast. However, Kraeuter and Setzler (1975) performed studies on S. meleagris in Georgia estuaries and concluded that this species does occur offshore in the winter, but that it also spends much of its early life in sounds and estuaries. Large indi- viduals appear offshore in March and apparently move in nearshore in May and June. Small individuals were collected in early July in the estuary. These populations No. 2 2004] TUNBERG AND REED—JELLYFISH AND SPIDER CRABS 99 CARAPACE WIDTH (mm) 28 [] bell diam. 2 vs males 26 24 22 20 18 16 14 12 10 80 90 100 Vo) BELL DIAMETER (mm) Fic. 5. The relationship between bell diameter of Stomolophus meleagris and carapace width of males and females of Lihinia dubia. decreased in numbers by August. After mid-October all individuals disappeared until March. According to Larson (1991) in a study performed between June 1986 and October 1987, in the north-eastern Gulf of Mexico, S. meleagris was abundant from June to October. Burke (1976) reported that §. meleagris was found throughout the year in Mississippi Sound, with the highest abundance during 100 FLORIDA SCIENTIST [VOL. 67 Ml Females (7) Males NUMBER OF CRABS 80 90 100 110 BELL DIAMETER (mm) Fic. 6. The relationship between the bell diameter of Stomolophus meleagris and the sex distribution of Libinia dubia. midwinter. Specimens collected ranged in size from 3 to 380 mm bell diameter, suggesting that a few of these medusae may survive for longer than a year. In the Fort Pierce Inlet area, S. me/eagris is usually most abundant during late summer and early fall (pers. obs.). Even though we observed a few jellyfish far from the inlet in the IRL, the highest abundances were near the Fort Pierce Inlet. No. 2 2004] TUNBERG AND REED—JELLYFISH AND SPIDER CRABS 101 140 MM with crabs (“7 without crabs 120 100 80 60 NUMBER OF INDIVIDUALS 40 20 70 80 90 100 110 120 130 BELL DIAMETER (mm) Fic. 7. The relationship between the bell diameter of Stomolophus meleagris and the Libinia dubia association rate. Noteworthy is that during this time, many S. meleagris were found even in an IRL mosquito impoundment (David, 2003). Large amounts of stranded jellyfish were also recorded on the ocean beaches in St. Lucie and Indian River counties during the sampling period and the following week. The sampling on 28 March started early during the incoming tide, with few specimens found near the surface. However, 102 FLORIDA SCIENTIST [VOL. 67 massive amounts of jellyfish appeared very rapidly approximately an hour later during the incoming tide. The jellyfish were either transported in large numbers into the estuary with the incoming tide or migrated vertically toward the surface (or a combination of both). Since few specimens were found at the actual mouth of the inlet at the beginning of the incoming tide on 28 March, a vertical migration possibly occurred. O’Brien and co-workers (1999) sampled L. dubia over a seven-year period in the Great Bay estuary (New Jersey). They found a male/female ratio of 1.33, which was very different from the ratio (0.82) of the randomly collected jellyfish in our study. However, when combining our random and selective samples, the male/ female ratio was 1.43, which is similar to the one found by O’Brien and co-workers. O’Brien and co-workers (1999) also reported a male maximum size (CW) of 75 mm, and males were slightly larger than females (not significantly). The males also had a greater CW size range than the females. Therefore the mean carapace width of the males of 22.9 mm found during our study clearly suggests that only small L. dubia are found on S. meleagris. Interestingly, Weymouth (1910) suggested that the relationship between Cancer gracilis and jellyfish may be obligatory until the crab reaches a length of 15 to 20 mm. Gutsell (1928) reported an association between S. me/eagris and L. dubia in a study performed at Beaufort, North Carolina. In one plankton tow, 19 jellyfish and 9 spider crabs were collected, indicating that more than 47% of the jellyfish had an associated crab, a much higher incidence of jellyfish/crab association than that of our study. Gutsell (1928) also reported that with one exception, the jellyfish and crabs were taken in surface tows. The size (carapace lengths) of the crabs during this study varied between 3 and 37 mm. Corrington (1927) collected 17 S. meleagris between Sullivan’s Island and the Isle of Palms, near Charleston, South Carolina; 16 concealed a L. dubia. Corrington (1927) never found more than one crab attached to each medusa. However, in our samples, we observed two jellyfish, each with two crabs. This may be due to our larger sample size. According to Gutsell (1928), the crab can also enter the subumbrellar space on the jellyfish. We also observed crabs using this space (sex unknown). Consequently, crabs hiding in this space may not have been detected by us when sampling jellyfish selectively. The percentage of females using the subumbrellar space may possibly therefore be higher than for the males. It is unknown how this benthic crab gets associated with the jellyfish. Corrington (1927) hypothesized: “‘Since the crab is absolutely confined to the bottom in so far as its own efforts are concerned, there remain but two alternatives: either the medusa must descend to the substratum at least occasionally, and for an obscure purpose, or else one of the larval stages of the crab must seek shelter within the umbrella and then remain attached during a long period of its mature life, for a reason equally difficult to conjecture.”’ According to Sandifer and Van Engel (1971), the duration of the larval life in L. dubia is short, approximately 9 days, compared to other species of Libinia. In the Chesapeake Bay area planktonic larvae of L. spp. were found from June to October, and were most abundant in July and September (Sandifer, 1973). This is the period No. 2 2004] TUNBERG AND REED—JELLYFISH AND SPIDER CRABS 103 when swarms of S. me/eagris usually occur in the Fort Pierce Inlet area, and may possibly be the time when the larvae of L. dubia attach to the jellyfish. Jachowski (1963) hypothesized that L. dubia attaches to the jellyfish Aurelia aurita by chance contact of the jellyfish with the shallow bottom or with aquatic plants where this non-swimming crab usually occurs. Jachowski (1963) further stated that the habits of these two animals are so different that an association in which one of them is dependent upon the other is considered to be unlikely. Because as many as 16.5% of the jellyfish collected during our study were carrying crabs, we presume this to be an important association, most likely of benefit to the crab. Corrington (1927) concluded that it is unquestionable that the crab receives both shelter and transportation by being attached to the jellyfish, but that it seemed unlikely that the crab preyed upon the medusa. Trott (1972) suggested that the relationship is symbiotic, but was uncertain if the relationship is truly commensal. Jachowski (1963) found numerous small L. dubia on the jellyfish Aurelia aurita. Two individuals of the crab had penetrated into the mesoglea and were feeding upon the medusan tissue. Several crabs also appeared to be feeding on the living medusa when later observed in the laboratory. The crabs pulled fragments of tissue from the exumbrella with their chelipeds and ate them, and none showed ill effect from eating or living in medusae. In the laboratory, we observed L. dubia attacking and eating live S. meleagris. However, not having witnessed this behavior in situ, it is uncertain whether this is a laboratory artifact or not. Several of the jellyfish collected during our study were damaged both on the bell and on the tentacles, but it is unknown if these injuries had been caused by the crabs. ACKNOWLEDGMENTS—Funding and resources for this study were provided through the Smithsonian Marine Station, Fort Pierce, Florida. This is Smithsonian Marine Station at Fort Pierce contribution number 567. LITERATURE CITED Burke, W. D. 1976. Biology and distribution of the macrocoelenterates of Mississippi Sound and adjacent waters. Gulf Res. Rep. 5:17—28. Catper, D. R. 1982. Life history of the cannonball jellyfish, Stomolophus meleagris L. Agassiz, 1860 (Scyphozoa, Rhizostomata). Biol. Bull. 162:149-162. AND B. S. Hester. 1978. Phylum Cnidaria. Pp. 87-93 in R. G. ZINGMARK (ed.). An Annotated Checklist of the Biota of the Coastal Zone of South Carolina. University of South Carolina Press, Columbia, South Carolina. CorRINGTON, J. D. 1927. Commensal association of a spider crab and a medusa. Biol. Bull. 53:346—350. Davip, J. R. 2003. St. Lucie County Mosquito Control District, Fort Pierce, Pers. Comm. Dracovicnu, A. J. AND J. A. KELLY, Jr. 1964. Ecological observations of macro-invertebrates in Tampa Bay, Florida. Bull. Mar. Sci. 14:74-102. GuTsELL, J. S. 1928. The spider crab, Libinia dubia, and the jellyfish Stomolophus meleagris, found associated at Beaufort, North Carolina. Ecology 9:358—359. JACHOWSKI, R. 1963. Observations on the moon jelly, Aurelia aurita, and the spider crab, Libinia dubia. Chesapeake Sci. 4:195. KRAEUTER, J. N. AND E. M. Setzier. 1975. The seasonal cycle of scyphozoa and cubozoa in Georgia estuaries. Bull. Mar. Sci. 25:66—74. 104 FLORIDA SCIENTIST [VOL. 67 Larson, R. J. 1991. Diet, prey selection and daily ration of Stomolophus meleagris, a filter-feeding Scyphomedusa from the NE Gulf of Mexico. Estuar. Coast. Shelf. Sci. 32:511-525. Mayer, A. G. 1910. Medusae of the world. Scyphomedusae. Carnegie Institute, Washington 3:499—735. O’BrIEN, S. B., M. LANDAU, AND K. W. ABLE. 1999. Sex ratios of two species of spider crabs, Libinia dubia H. Milne Edwards, 1834 and L. emarginata Leach, 1815 in the area of Great Bay, New Jersey. Crustaceana 72:187—-192. SANDIFER, P. A. 1973. Distribution and abundance of decapod crustacean larvae in the York River estuary and adjacent lower Chesapeake Bay, Virginia 1968-1969. Chesapeake Sci. 14:235—257. AND W. A. VAN ENGEL. 1971. Larval development of the spider crab, Libinia dubia H. Milne Edwards (Brachyura, Majidae, Pisinae), reared in laboratory culture. Chesapeake Sci. 12:18—25. Tass, D. C. AND R. B. MANNING. 1961. A checklist of the flora and fauna of northern Florida Bay and adjacent brackish waters of the Florida mainland collected during the period July, 1957 through September, 1960. Bull. Mar. Sci. 11:552-649. Trott, L. B. 1972. The portunid crab Charybdis feriatus (Linnaeus) commensal with the scyphozoan jellyfish Stomolopus nomurai (Kishinouye) in Hong Kong. Crustaceana 23:305—306. WeymoutH, F. W. 1910. Synopsis of the true crabs (Brachyura) of Monterey Bay, California. Stanford University Publications 4:1—64. WILLIAMS, A. B. 1984. Shrimps, Lobsters, and Crabs of the Atlantic Coast of the Eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D. C. 550 pp. Florida Scient. 67(2): 93-104. 2004 Accepted: July 25, 2003 Environmental Chemistry UPTAKE OF PHOSPHATE AND NITRATE USING LABORATORY CULTURES OF LEMNA MINOR L. DanieL P. Smite’?, MatrHew E. McKenzie, 9) CRAIG Bowe”, AND DEAN F. Martin “Department of Civil and Environmental Engineering Institute for Environmental Studies, Department of Chemistry, University of South Florida, 4202 East Fowler Avenue, Tampa, FL 33620 Asstract: The use of the Lemna minor L. species of duckweed is an emergent technology that may be effective for the removal of nitrogen and phosphorus from enriched waters. The goal of this project is to find a cost-effective method of removing nitrogen and phosphorus from fresh water resources that have received loadings of these nutrients in stormwater runoff. This research examined the uptake of nitrogen and phosphorus by cultures of duckweed, L. minor L., cultivated in the laboratory under controlled environmental conditions. L. minor L. was grown in Plexiglas reactors (1600-mL) using modified Hillman growth medium. The effect of Lemna on phosphorus and nitrogen uptake and plant growth was measured over a two-week period . Two approaches were used: a batch and continuous flow method. For the latter method, a mass balance calculation was performed using measurements of the mass of phosphorus and nitrogen added in the influent, the mass taken up in the growing Lemna biomass, and the mass exiting in the effluent. A mass balance for high nitrogen, low phosphorus media (650 ppm N and 150 ppm P) indicated that 7% of the nitrogen and 10% of the phosphorus was removed by the plant uptake over the 14-day period of operation. Key Words: Duckweed, phytoremediation, nutrients, nitrate, phosphate, reactor, aquatic treatment system, storm water, wastewater treatment POLLUTION by elevated nitrogen and phosphorus levels in fresh water sources is a problem that has plagued the unique biosphere of the state of Florida in recent years. The increases in the population in the state of Florida, extensive agricultural production, and the advent of heavy industry have exacerbated this problem. Technology has proven to be both a benefit and a detriment for the state water resources. The challenge faced by scientists is the control of chemical constituents added to the environment and the control of nutrients that cause impairment to beneficial uses of surface waters when present at elevated concentrations. The use of biotechnology is a solution to the problem that is cost-effective and practical. There is an increased interest in using aquatic plants, such as duckweed, in the treatment of contaminated surface waters and wastewaters. The use of this technology is of significant importance because it is a relatively cost- effective alternative to methods currently being used in the field of water treatment. Integrated algal and duckweed ponds have proven to be an effective means of controlling nutrient levels while not doing further damage to the environment (Van der Steen et al., 1998). 105 106 FLORIDA SCIENTIST [VOL. 67 Duckweed cultivation systems also offer an approach to sustainable nutrient recovery and reuse. In Asia, duckweed has been a valuable source of nutrition. The larger species, L. minor wolffia, is harvested from Asian farmers’ runoff ponds. Then it is dried and sometimes mixed in with other feed, which is fed to ducks, chickens, livestock, and fish. Lemna minor wolffia contains about forty percent protein (dry weight), similar to soybean protein content, and also contains high levels of all essential amino acids except methionine (Landolt, 1986). The use of duckweed in the removal of nitrogen is also a technology that has proven to be an effective use of this macrophytic aquatic plant (Bonomo et al., 1997). Five species of duckweed have proven to be effective in wastewater treatment and one of these, L. minor L., is the most common species in the state of Florida. The performance of common duckweed species on wastewater has been studied, and the results indicated that two species of duckweed, L. minor gibba and Spirodela polyrhiza, proved to be the most effective in controlling nutrient levels (Vermaat and Hanif, 1998). Duckweed has proven to be important in the removal of nitrogen and phosphorus in domestic water systems (KOrmer and Vermaat, 1998). The focus of this project is to find a relatively inexpensive and environmentally friendly means of removing nitrogen and phosphorus from surface waters, includ- ing industrial and municipal wastewater effluents, and storm water. The use of a plant species commonly found in Florida provides a practical and cost-effective alternative to methods that are currently being explored in government and industry. Species of duckweed, particularly L. minor L., are commonly found within the state of Florida (Long and Lakela, 1976), and they offer an extensive and renewable source for usage on a large scale. There are obvious financial advantages in utilizing a plant species to uptake these nutrients. The availability and accessibility of the plant species for use in pilot studies and full-scale treatment of surface water, storm water, and municipal and industrial effluents is also an advantage. The contributions of this present work are advancements in process-based knowledge that can be applied to aquatic treatment systems treating wastewater effluents. Secondly, the use of an aquatic plant that is readily available within the state of Florida has additional potential applications in cost-effective removal of toxic chemicals and excess nutrients from surface waters. This project addresses aquatic treatment of high contents of nitrogen and phosphorus in industrial and municipal effluents and storm water such as are found in the Tampa Bay area. The goals of nutrient removal from wastewater effluents are to reduce nutrient loading and algal growth in the receiving surface waters. Therefore the goals of these experiments are to monitor the growth of the L. minor L. with the increased levels of phosphate and nitrate nutrients in order to elucidate a maximum level of each nutrient that the plant can withstand without detriment to the plant physiology and to ascertain duckweed’s ability to remove nitrogen and phosphorus from increased nutrient media. MATERIALS AND MetHops—Growth chambers (reactors)}—The approach used in this investigation involved use of bench-scale reactors that emulate larger models that are found in a wastewater treatment No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCK WEED 107 facility. The growth chambers were constructed at the Engineering Shop at the University of South Florida Department of Civil and Environmental Engineering from a single sheet of Plexiglas obtained from GE Polymershapes, Tampa, Florida. The four reactors accommodated the duckweed and the flow rate was controlled using peristaltic flow pumps connected to each individual Plexiglas container (Fig. 1). The total volume of medium in each reactor was 1600 mL. It was necessary to encase the sidewall and bottom of each reactor with light eliminating paper to limit the growth of algae over the period of the study. The paper limited the penetration of light to the inside of the reactors and at the bottom of the reactors and limited the growth of Chlamydomonas gloegama Korschikoff, a freshwater algae, which otherwise tended to proliferate. (The algae were identified by Professors Clinton Dawes and Bruce Cowell). In aquatic growth reactors, such as ponds and basins in which a duckweed mat has achieved high degree of surface coverage, shading by the surface layer of duckweed would reduce solar penetration and limit the growth of algal populations. Sidewall light penetration would not occur in such systems. Culture history—All experiments and stock duckweed cultures were kept in the Phytotron, controlled environment room (Environmental Growth Chambers, Chagrin Falls, OH) in the Department of Biology. Conditions in the chamber were as follows: constant temperature of 26 °C, 80% relative humidity, and a twelve-hour photoperiod with a light intensity of 190 jtE/m*/sec measured by LiCor model LI-185A photometer. The light intensity measured in the Phytotron room was equivalent to 33000 kJ/m?/day (16,500 kJ/m?/day for the 12-hour photoperiod). This is similar to the measured solar radiation of the months March and October (approximately 15,000 kJ/m?/day) in the southeastern United States (Reifsnyder and Lull, 1965). Duckweed (Lemna minor L.) was obtained from Carolina Biological. Stock duckweed was grown in plastic trays in a 100% Hillman growth medium, Table | (Hillman, 1959a,b). Growth medium was changed every three days to protect against loss of nutrients and the proliferation of algae. Reactor studies—A 15 L Pyrex carboy was autoclaved at 60 psi and a temperature of 115 °C for 90 minutes. After the medium was made, it was autoclaved using the same conditions. The autoclaved medium was allowed to cool to room temperature before the study began. When the medium was brought to the Phytotron, a black plastic bag covered the carboy during the study to prevent the growth of algae. The reactors were autoclaved at a pressure of 60 psi for 90 min, and the autoclaved medium was pumped into the reactor using a peristaltic pump (Cole-Parmer Model 07554-80) with a pump head (07518-12) using Tygon (LFL L/S® 25) tubing. Then the duckweed was transferred from the plastic trays and placed in Plexiglas growth reactors (Fig. 1). In all the studies, the medium was monitored for changes in concentration of nutrients and chlorophyll content. The biomass in each growth reactor was determined at the start of the study and at the completion of the experiment. A 30-mL plastic measuring cup, modified by drilling five holes into the bottom, was used to collect and strain the duckweed when removing it for biomass measurements, or harvesting. This cup had a surface area of 13.8 cm*. We shoved the cup into the duckweed of the reactor, and waited a few seconds for the duckweed to come back together on the surface. The retriever was then lifted up above the waterline. The water was drained out of the holes at the bottom of the retriever. Each time the reactor was initially filled with duckweed, the reactor surface was mixed to spread the duckweed fronds or plant bodies uniformly over the reactor surface area, and a separate scoop of stock duckweed was collected and analyzed to determine the starting duckweed biomass per reactor surface area. Fresh and dry weights were determined for a scoop sample and the values were multiplied by the number of scoops it took to fill the surface of the reactor of that particular study. Batch and continuous flow studies—This study consisted of two experimental methods. The experimental setup, length of study, starting media, chlorophyll a analysis, and water sampling did not change between the two different methods. In the batch method, the effluent from the reactor was recycled back into the carboy . Here the flow rate was 25 mL/min with a water replacement time of 64 minutes. The Continuous Flow method did not have the media being recycled, and the effluent hose led to the drain. Fresh medium was made daily to restore the medium stock. The water replacement time was 229 minutes with a flow rate of 7 mL/min. 108 FLORIDA SCIENTIST [VOL. 67 Top View Side View S Fic. 1. Schematic representation of L. minor growth reactors. Reactor dimensions were 15.2 cm X 8.26 cm X 12.7 cm. On a daily basis for two weeks, water samples were taken from the systems to find the amount of nitrogen and phosphorus in the medium. In the batch method, the water samples were taken from the 15 L carboy, which contained the recycled media. The continuous flow system had two 40 mL water samples, one sample was taken from the 15 L carboy (the influent) and the other was collected from the dripping effluent hose before the water was allowed to drain out of the room. Phosphorus and nitrogen analyses—A Hach total phosphorus kit (model PO-24, Hach Company, P.O. Box 389, Loveland, CO) and Hach kit (Model PI-14) for nitrate analysis were used for phosphorus and nitrogen analyses respectively. The Hach methods for nitrate and phosphorus analyses follow EPA guideline numbers 353.3 and 365.2, respectively. The Hach kit instructions were followed when analyzing the 40 mL water samples except the nitrate analysis procedure was modified, i.e., the agitation of the sample was performed in a Cole-Palmer ultra sonic cleaner for three minutes, instead of the Hach method of shaking the flask for the same amount of time. Water samples (40 mL) were analyzed in triplicate and the mean and standard deviation were recorded. The total aqueous phosphate and nitrate values were converted to total phosphorus and nitrogen. Modified nitrate analysis—The Hach nitrate analysis kit was used, as described above, yet a few modifications were made. A sample (0.125 mL) in a 5-mL Erlenmeyer flask was diluted with 4.875 mL deionized water, and the Hach procedure was followed, except that agitation in a Cole Palmer ultra sonic cleaner for 3 min. was done instead of the Hach method of shaking the flask for three minutes. The Hach procedure was repeated in triplicate, and the mean and standard deviation were calculated. No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCKWEED 109 TABLE |. Summary of experiments performed. Total Total Duration Nitrogen Phosphorus Experiment Designation Flow Mode Media (days) (mg/L) (mg/L) 1 Bl Batch HM! 14 406 155 2 B2 Batch HN/LP? 14 OM 155 3 B3 Batch LN/HP* 14 A06 228 4 Cl Continuous § HN/LP? 14 Wi 155 5 (eH) Continuous = LN/HP® 14 406 228 : Hillman medium. ~ High nitrogen, low phosphorus media. > Low nitrogen, high phosphorus media. Nutrient analysis in biomass—TYo analyze for nitrogen or phosphorus in the biomass, the plant matter was first dry- weighed and recorded. Then it was digested with 7.5 M sulfuric acid and let stand for a day. This material was neutralized with 7.5 M sodium hydroxide. The mixture was then filtered using Whatman GF/A filter paper to remove any undigested plant particles. The filtrate volume was recorded and the filtrate was analyzed for nitrogen and phosphorus, and the values were used to calculate nutrient content of the duckweed. Determining fresh weight—The duckweed samples were placed between two layers of absorbent paper. The two layers were gently pressed together on other paper. The paper was removed after blotting several times and the duckweed was transferred to unused, dry paper. The blotting procedure was repeated and the newly dried L. minor L. sample was transferred to the analytical balance and the fresh weight was determined. Dry weight—Afiter the fresh weight was determined, the L. minor L. sample was placed in a test tube and transferred to an oven (56° C) for a period of 24 hr. Then, the dried L. minor L. sample was allowed to cool to room temperature and was weighed on an analytical balance to get the mass of the dried duckweed. The weight loss was calculated to give the water content. The relationship of dry weight to fresh weight is based on 15 samples of stock duckweed. Microsoft Excel’s trendline was used to calculate this correlation (Eqn. 1). D.W. = 0.0566 + (F.W.) +. 0.0015 (N = 15; R*? = 0.91) (1) Here, D.W. = Dry weight (in g.) and F.W. = Fresh weight (in g.) Fresh weight was related to the frond count, using appropriate data as indicated (Eqn. 2) Fresh weight (in g) = —0.024 + 0.006 « (frond count); N—=17, R* =0.97 (2) The fresh weight of scooped duckweed was then determined as described above. After weighing, the duckweed was returned to the L. minor growth reactor. It took five scoops to cover the surface of the each of the reactors completely, where the duckweed formed a green mat. It was necessary to calculate the fresh weight for seeding the reactors and then converting it into dry weight (Eqn. 1, 2). Percent recovery—Values were obtained for phosphorus and nitrogen by adding known amounts (spikes) of orthophosphate or nitrate to samples and analyzing the sample and the spiked sample. Percent recovery was calculated (Eqn. 3). % Recovery = {(sample + spike) — spike/sample] < 100. (3) Here “sample” and “spike” refers to the concentration of the untreated sample and the concentration of the added increment or “‘spike.”’ 110 FLORIDA SCIENTIST [VOL. 67 Chlorophyll a analyses—A known sample of duckweed fronds were removed from the L. minor growth reactor after each study, and similar samples were taken from the stock duckweed. Chlorophyll- a content was determined spectrophotometrically as described elsewhere (Gallardo et al., 1998). Nutrient-uptake experiments—Five experiments were performed as summarized in Table 1. The three batch experiments were designated as B1 (HM), B2 (HN/LP), and B3 (LN/HP). Two continuous- flow experiments were conducted. Experiment Cl used HN/LP medium while LN/HP medium was used in experiment C2. The three batch duckweed growth experiments were performed with identical starting biomass cultures and identical growth conditions, but each one having a different modified growth media. These tests were designed to compare the effects of the different nitrogen and phosphorus concentrations and N/P ratios and on the coupled processes of duckweed growth (increase in biomass) under non- harvesting conditions and decrease in nutrient concentration in the media. It was anticipated that the decline in media concentrations of nitrogen and phosphorus would be stoichiometrically linked to the net increase in duckweed biomass, and that the decrease in each nutrient would be related to their concentration in dry lemna biomass. The intent of these studies was to document the coupled processes of duckweed growth (biomass increase) and the decline of nutrient concentrations in reactors subject to continuous flow conditions. Such a continuous flow experiment, if operated with biomass harvesting for a sufficiently long period of time, could approach an operational regime with steady state biomass production and nutrient removal rates. RESULTS AND DiscussioN—General conditions of duckweed growth—The results from all the experiments showed nitrate and phosphate levels decreased over the period of the investigation (Figs. 2 and 3). Hillman growth medium and the spiked Hillman growth medium were successful in maintaining and culturing L. minor. Analyses—Nutrient (orthophosphate and nitrate) and chlorophyll analyses were routinely performed. Mean and relative standard deviations were calculated as a means of evaluating precision. For example, for phosphate the relative standard deviation of the mean was 2.5%, while the corresponding value for nitrate was 2.3%, and for chlorophyll a was 1.3%. In addition, the percent recovery was measured for phosphate and nitrate and was found to be 103 and 90%, respectively, for the Cl study (Table 1); for the C2 study the percent recovery values were 97.8% (P) and 99% (N). Dry and fresh weight determinations—Weight is an apparent measure of plant growth. While fresh weight is a useful means of estimating biomass, dry weight is a more precise measurement of biomass because it is not affected by the amount of water on the surface of the fronds or in the plant itself. Table 2 shows the start and end of each study’s biomass in fresh and dry weights. The starting dry weight biomass was calculated (using Eqn | and 2). Furthermore, measuring the fresh weight requires removing the plants from the medium and blotting them with a paper towel. Weighing duckweed plants could be a potential problem because they are delicate and small, and exhibit thigmotropism (the response of plants to mechanical force and vibration) even with gentle handling. Thigmotropism is known to cause changes in growth rate or the pattern of growth in many plants (Riehl and Jaffe, 1984). Thus, whenever the duckweeds were taken out of their medium, it was done quickly and gently. To lessen the stress on the duckweed, fresh and dry weight No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCKWEED iplal 700 - 500 = == BA —*— B2 RA ——B3 300 - Sa : —a— C1 Influent hae nl —2— C1 Effluent : + —a— C2 Influent SS , ee 0 -0 C2 Effluent Nitrogen Concentration, ppm 0 sli T = Faas 123 4 5 6 7 8 9 10 11 12 13 14 Time, days Fic. 2. Nitrogen concentration in effluent as a function of versus time for all experiments. standards were made in order to extrapolate any potential future harvesting experi- ment and biomass composition. Duckweeds were reported to contain between 86% and 97% water by weight (Landolt and Kandeler, 1987). In this study, the percent water in the duckweed stock samples was 92.6 + 1.6% (N = 14), which was within the reported range (Landolt and Kandeler, 1987). Mass balance calculation—High nitrogen/low phosphorus—A mass_ balance calculation was performed for the HN/LP media, or the C1 study, with an N/P molar ratio of 10.3/1 (Table 1). In the Cl study (Table 4), the cumulative phosphorus input was 490.5 g and the sum of the cumulative P output (Table 4) was 441.3 g, leaving a net difference of 49.2 g. The last value may be compared with the total phosphorus content 46.6 g (Table 4), which was obtained for all biomass, except for the biomass corresponding to 71 fronds removed for analyses. The difference (49.2 vs 46.6 g) is 2.6 g, and the value calculated for 71 fronds was 3.9 g, which leaves an excess of 1.3 g, which is ascribed to the uncertainty associated with the analytical errors. In a similar manner, the biomass calculation performed for nitrogen gave a difference between input and exiting value of nitrogen of 152.1 g (Table 3). The nitrogen content of the biomass (71 fronds) was 140.2 g, and the amount calculated for the 71 fronds was another 10.9 g. This resulted in an overall net difference between 152.1 g and 151.2 g of 0.9 g; again, the error is ascribed to the sum of experimental errors. Using the 7 mL/min flowrate entering the reactor, 7% of the nitrogen [(152.1/ 2061.3) X 100; Fig. 3] and 10% of the phosphorus [(49.2/490.5) X 100; Fig. 4] was removed by the lemna plants in the reactors during the Cl run (1.e., 10 N/I P molar ratio). 112 FLORIDA SCIENTIST [VOL. 67 — Bt —*— B2 —-—B3 —«— C1 Influent —4— C1 Effluent on —® C2 influent —o- C2 Effluent Phosphorus Concentration, ppm 0 - T T T T T T T nn omens Maestros ss |) 1:2 3 4 °5°6 °7 8° 94109 isl2e13h14 Time, days Fic. 3. Phosphorus concentration in effluent versus time for all experiments. Mass-balance_ calculation—Low_nitrogen/high phosphorus—A mass_ balance calculation was performed (a N/P molar ratio of 3.9/1;Table 1). The cumulative phosphorus input of the C2 run was 778.1 g (Table 4) and the sum of the cumulative P output (Table 4) was 527.1 g, leaving a difference of 251.0 g. The last value may be compared with the total phosphorus content 200.7 g (Table 4), which was obtained for all biomass except for the biomass corresponding to 80 fronds removed for chlorophyll analysis. The difference (251.0 vs 200.7 g) is 50.3 g, and the value calculated for 80 was 48.3 g, which leaves an excess of 2.0 g, which might be ascribed to the uncertainty associated with the analytical errors. In a similar manner, the biomass calculation performed for nitrogen gave a difference between input and exiting value of nitrogen of 306.3 g (Table 3). The nitrogen content of the biomass (80 fronds) was 293.7 g, and the amount calculated for the 80 fronds was another 11.4 g. This resulted in an overall net dif- ference between 306.3 g and 305.1 g (293.7 + 11.4 g) of 1.2 g; again, the error is ascribed to the sum of experimental errors. Using the same flowrate as was used in the continuous flow method, which produced a 229 minute water residence time, 23% of the nitrogen [(306.3/1335.0) x 100; Table 3] and 32% of the phosphorus [(251.0/778.1) X 100; Table 4] was removed by the lemna plants in the reactors during the C2 run. Chlorophyll a measurements—The chlorophyll a concentrations measured at the end of four lemna growth experiments compared favorably with those in the controls. This is an indication that the duckweed grew well during both batch and continuous flow experiments. In addition, the quantity of nitrogen and phosphorus No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCKWEED 113 TABLE 2. Total plant biomass for experiments described. Experiment Calculated Biomass per Fresh Weight* Dry Weight* Surface Area (g) (g) (mg/cm”) B2 (start) 6.25 O),35)5 2.43 B2 (end) 7.20 0.409 2.80 B3 (start) 28 0.300 JAWS) B3 (end) 10.8 0.613 4.20 Cl (start) 6.10 0.347 2.38 Cl (end) 7.68 0.436 299 C2 (start) 5.65 0.211 1.45 C2 (end) 7.46 0.428 7293 =e : : 5 * Biomass is normalized to the reactor per surface area of 146 cm’. assimilated into lemna biomass increased both as a function of time and as a function of the increase of plant biomass (Table 2). Extrapolated mass removals—We calculated the amount of nitrogen and phosphorus removed under various conditions for two different systems (Batch and Continuous Flow). The surface area of the lemna reactor was calculated to be 146 cm* (=161 cm” total area minus 15 cm? of the outflow shield area). For the continuous flow systems, the amount removed per reactor surface area (mg/146 cm”) was calculated, and this was converted to other units as shown (Eqn. 4-6). g/m> = (mg/146 cm’) X 0.068 (4) kg/ha = (g/m*) x 10 (5) Ib/acre = (kg/ha) /1.12 (6) For the batch system, an additional assumption was made, 1.e., that essentially the medium was cycled through each day of the 14-day study and the medium would be exhausted in a 24-hour period. Consequently, the total nitrogen or phosphorus removed was divided by 14 to give the mean surface area. The results are presented in Table 5. TABLE 3. Summary of nitrogen mass balances in the continuous flow systems’. HN/LP, LN/HP, Calculated Quantity g g Sum of Cumulative N;,, mass 2061.3 1335 Sum of Cumulative N,,,; mass 1909.2 1028.7 Difference of Cumulative N;,, and Cumulative N,.; S21 306.3 Amount of N in N-* fronds (remaining) Moe 298 view Amount of N in * fronds (estimated) 10.97 11.48 Total Estimated N in biomass SIA 305.1 Difference between Cum. N;, and N,,, and N in biomass —0.9 ED ' N-* is the amount of fronds used to find the nitrogen content; here * is the amount of fronds used to find chlorophyll a concentration 114 FLORIDA SCIENTIST [VOL. 67 TaBLE 4. Summary of phosphorus mass balances in the continuous flow systems’. HN/LP, LN/HP, Calculated Quantity g g Sum of Cumulative P;,, mass 490.5 778.1 Sum of Cumulative P,,, mass 441.3 527.1 Difference of Cumulative P;, and Cumulative Poy; 49.2 251 Amount of P in N-* fronds (remaining) AGG) ae 20017 Amount of P in * fronds (estimated) 3.971 48.38 Total Estimated P in biomass 50.4 249 Difference between Cum. P;,, and P,,, and P in biomass 1.3 2, ' N-* is the amount of fronds used to find the phosphorus content; here * is the amount of fronds used to find chlorophyll a concentration. Examining the results (Table 5) gives a comparison of removal efficiencies for the two different approaches. For example, in the batch systems, a greater removal of nitrogen (353 Ib/acre) was obtained using the B2 system, but a lesser amount of phosphorus was removed (76 Ib/acre). Those two values do not give a complete picture; in the system involved, 93.5% of the available nitrogen and 84.5% of the available phosphorus was removed. And the percentage removal for both nitrogen and phosphorus was good. A lesser amount of nitrogen, but a greater amount of phosphorus (168 Ib/acre N, 163 lb/acre P) was predicted to be removed using the B3 medium in the batch system (Figs. 2 and 3). Closer examination indicates that though more phosphorus was removed, the percentage removal of the influent was about the same as the B2 study (84%), but the removal of nitrogen was significantly less, t.e., 72%. Thus, for effective removal of both nitrogen and phosphorus, the B2 medium was superior, which is consistent with the concept of the stoichiometric relationship between the nitrogen and phosphorus requirement of lemna organisms (vide infra). Fewer data are available for continuous flow systems, but for the Cl system, 82 and 267 lb/acre were removed for nitrogen and phosphorus, respectively, though the percent removal was understandably low (7—10%) being a single-pass system. The approach was an experimental advantage since it permits a valid mass balance to be calculated with fewer assumptions. TABLE 5. Extrapolated removal of nitrogen and phosphorus for L. minor L. aquatic treatment systems. manent Removed Experiment Mass N/P Element g g/m? kg/ha Ib/acre B2 4.65 N 9.09 395 395 Bj) P LOW Se 85 76 B3 1.78 N 4.32 18.8 188 168 P 4.20 18.2 182 163 Cl 4.65 N li52 OF 92 82 P 49.2 29.8 298 267 C2 1.78 N 306 18.5 185 165 P. Ds) Sy 1520 1362 No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCKWEED TS In the B3 and C2 experiments, which were the low nitrate/high phosphate media, the duckweed grew the most out of all the studies (Table 2). The dry weight biomasses increased 0.313 g for the B3 study and 0.217 g for the C2 study. Looking into these two studies the removal of nitrogen and phosphorus were the highest out of all the studies performed (Figs. 2 and 3). Since nitrogen is essential for growth, the duckweed in the reactor was more aggressively competing for nitrogen than those of the higher phosphate and standard phosphate medias (B1, B2, and C1). Furthermore, the duckweed was removing the nitrogen at a higher rate; therefore the process was removing a higher rate of phosphorus. The investigation of the use of L. minor L. in the uptake of essential plant nutrients would have practical applications in wastewater treatment facilities. The basic inference as to the amount of nitrogen and phosphorus that could be removed comes from a review by Redfield (1958). He summarized studies of the uptake of these elements by plankton, and concluded that phosphorus and nitrogen appear to be the constituents of the sea in limiting quantities. The Redfield atomic ratios were phosphorus-nitrogen-carbon of 1:15:105. However, as noted in Martin (1970), plants have the ability to remove excess amounts of phosphorus and/or nitrogen. In addition, examining the atomic ratios of plankton, Strickland (1965) indicated that the values of the type Redfield may have used in his review were an average that had, as might be expected, significant variation. For example, for phytoplankton, the % nitrogen reported for about 20 species ranged from 2.7—9.1; % phosphorus ranged from 0.6—2.7% (Strickland, 1965; Table III). Some examples of deviation from Redfield ratios are evident from the present study. Most obvious is the fact that the Hillman’s medium is low with respect to the nitrogen-phosphorus ratio (5.8/1 molar ratio; Table 1) in contrast with the Redfield N/P molar ratio of 15. For the B2 and C1 studies (N/P = 10.3), the atomic ratio for removal was 6.9 N/P, which may indicate that for lemna the optimum for growth was a ratio of about 6, and that a greater enhancement of nitrogen did not result in a significantly greater removal ratio. In a closer examination of the Continuous Flow experiments (studies C1 and C2), there was a surprise in the N/P uptake (Table 6). Obviously, the N/P ratio of the influent would remain constant over time; hence the slope would be zero. The slopes near zero are HN/LP influent, LN/HP influent, and LN/HP effluent (—0.0011, —0.0002, and —0.0155 respectively, Table 6). The duckweed in the lower nitrogen media was more aggressively absorbing the nitrogen; it absorbed more phosphorus consequently, keeping the same ratio. On the other hand, the Cl experiment, or HN/LP, had an increasing slope of 0.0617 (Table 6). Nitrogen is necessary for plant growth, with enough nitrogen present the duckweed grew strong. Furthermore, this increasing slope shows duckweed has the potential of removing more nitrogen with time. This research has been concerned with the uptake of nitrogen and phosphorus by Lemna minor L. We have examined three different nitrogen/phosphorus ratios under conditions that are environmentally representative of Hillsborough County, Florida. We also demonstrated that successful mass balances could be established. The nitrogen and phosphorus concentrations used for optimum growth of Lemna minor L., as used in all three media, were higher than would be expected for a storm 116 FLORIDA SCIENTIST [VOL. 67 TABLE 6. N/P ratios of the influent and effluent during the Continuous Flow studies. The slope of the line is from a plot of Day vs. N/P ratio. Note: The N/P ratio’s slope are close to zero besides the HN/LP effluent N/P ratio. HN/LP LN/HP N/P ratio N/P ratio Day Influent Effluent Influent Effluent 1 4.15338 4.12218 Le7Alisat 0.02652 2 4.21364 4.19127 1.73801 0.12246 3 4.181 4.11793 1.70736 0.54448 4 4.17598 4.23061 1.71274 0.30416 5) 4.2036 4.34485 1.70999 0.23818 6 4.27876 4.39541 1.71439 0.30023 7] 4.2488 4.48729 1.71316 0.27764 8 4.21867 4.69578 75129, 0.23852 9 4.20109 4.64869 1.69644 0.35786 10 4.16342 4.69111 72536 0.40122 1] 4.25548 4.64914 1.69304 0.39374 12 4.18853 4.74932 1.7343 0.06904 13 4.16091 4.7831 1.69637 —0.0196 14 4.14584 4.90942 1.72908 —0.1781 Slope —0.0011 0.0617 —0.0002 —0.0155 water stream, but they seem likely to be similar to concentrations to be found in storm water runoff retention ponds, so that the clean-up potential seems worthy of consideration. ACKNOWLEDGMENTS—We are grateful to the Storm Water Management Section of Hillsborough County Public Works Department for funding this project. We also thank the University of South Florida engineering shop for the construction of the Plexiglas reactors. We thank also the Department of Biology at the University of South Florida for the use of the Phytotron room and the autoclave units. Finally, we are grateful to Dr. Bruce Cowell and Dr. Clinton Dawes, Department of Biology, for their assistance in identifying the algal species Chlamydomonas gloegama Korschikoff. Mrs. Barbara Martin served as editor. LITERATURE CITED Bonomo, L., G. PASTORELLI, AND N. ZAMBON. 1997. Advantages and limitations of duckweed-based wastewater treatment systems. Wat. Sci. Tech. 35(5):239-246. GaLLarpo, M. T., B. B. Martin, AND D. F. Martin. 1998. Inhibition of Salvinia minima by cattail (Typhia domingensis) extracts and by 2-chlorophenol and salicylaldehyde. J. Chem. Ecol. 24: 1483-1490. HILLMAN, W. S. 1959a. Experimental control of flowering in L. minor. I. General Methods. Photo- periodism in L. pepusilla 6746. Amer. J. Bot. 46:466—-473. . 1959b. Experimental control of flowering in L. minor. II. Some effects of medium composition, chelating agents and high temperatures on flowering in L. pepusilla 6746. Amer. J. Bot. 46: 489-495. KOrNER, S. AND J. E. VERMAAT. 1998. The relative importance of L. minor gibba L., bacteria and algae for the nitrogen and phosphorus removal in duckweed-covered domestic wastewater, Wat. Res. 32(12):3651—-3661. LANDOLT, E. 1986. The family of L. minorceae—a monographic study. Vol 1. Jn: LANDOLT, E. (ed.) Biosystematic Investigations in the Family of Duckweeds (L. minorceae). Veroff. Geobot. Inst. ETH, Stiftung Rubel. No. 71. No. 2 2004] SMITH ET AL.—UPTAKE OF NUTRIENTS BY DUCKWEED 1G AND R. KANDELER. 1987. The family of Lemnaceae—a monographic study. Vol. 2, Phyto- chemistry, physiology, application, bibliography. Veroff. Geobot. Inst. ETH, Zurich, 638 pp. Lona, R. W. AND O. LAKELA. 1976. A Flora of Tropical Florida. Banyan Books, Miami, FL. Martin, D. F. 1970. Marine Chemistry, Vol 2.: Theory and Applications, Marcel Dekker, Inc., New York, NY. REDFIELD, A. C. 1958. The biological control of chemical factors in the environment. Amer. Scient. 46: 205-221. RriewL, T. E. AND M. J. JAFFE. 1984. Physiological guides on pea tendrils. Plant Physiol. 75:679-687. REIFSNYDER, W. E. AND H. W. LULL. 1965. Radiant Energy in Relation to Forests. Tech. Bull. No 1344ed. U.S. Department of Agriculture, Washington, DC. STRICKLAND, J. D. H. 1965. Production of organic matter in the primary stages of the marine food chain. Pp. 478-610 (Chapter 12). In: Ritey, J. P. AND G. SkiRROW (eds.) Chemical Oceanography (1*' ed) Vol. 1. Academic Press, New York, NY. VAN DER STEEN, P., A. BRENNER, AND G. Oron. 1998. An integrated duckweed and algae system for nitrogen removal and renovation. Wat. Sci. Tech. 38(1):335-348. VERMAAT, J. E. AND M. K. Hanir. 1998. Performance of common duckweed species (L. minorceae) and the waterfern Azolla Filiculoides on different types of wastewater. Wat. Res. 32(9):2569-2576. Florida Scient. 67(2): 105—117. 2004 Accepted: August 29, 2003 Biological Sciences THE SAND PINE SCRUB COMMUNITY: AN ANNOTATED BIBLIOGRAPHY, 1989-2001 DONALD R. RICHARDSON Ecological Consultants, Inc., 5121 Ehrlich Road, Suite 103A, Tampa, Florida 33624 ABSTRACT: This annotated bibliography represents a revised 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. The Scrub ecosystem is one of the oldest plant communities in Florida and is rapidly becoming fragmented and isolated due to encroachment of urban areas. As much as 95% or more of this upland plant community has been lost in Palm Beach County alone due to development over the past 20 years. Other areas such as the Lakes Wales Ridge in central Florida, once thought of as a wilderness island, have seen most of the natural scrub being replaced by large scale citrus groves, with estimates of loss ranging from 80-90% of the original scrub. About 40% of the scrub ecosystems in Brevard County are contained within federal government ownership, but recent mapping studies indicate that about 75% or more of this plant community has been eliminated over the past 40 years. Estimates suggest that by the year 2000, as much as 80% of the scrub in southern Florida may have been lost or impacted due to urban sprawl. A growing concern for the disappearance of this relic community has resulted in the Federal listing of many scrub endemics, as well as a rapid increase in land acquisition for nature preserves around the state. This rapid demise has led to a three-fold increase in the amount of research and study of this unique plant community. This bibliography represents an updated effort to include research on plants, animals, and insects endemic to the scrub pine community. Included is the most recent research on management and preservation of the scrub ecosystem. Key Words: Florida sand pine scrub, habitat loss, fragmentation, development, scrub endemics ABRAHAMSON, W. G. 1995. Habitat distribution and competitive neighborhoods of two Florida palmettos. Bull. Torrey Bot. Club 122(1):1-14. [Discussed are two ecologically similar palmettos, saw (Serenoa repens) and scrub (Sabal etonia), which co-occur on the Florida peninsula’s central ridge. Sharing many characteristics of growth form, reproductive strategies, responses to fire, and habitat occurrence, their coexistence suggests differences in micro- habitat distributions and details of life histories. ] . 1999. Episodic reproduction in two fire-prone palms, Serenoa repens and Sabal etonia (Palmae). Ecology 80(1):100—115. [This study examined how fire events affected flowering of two native palms found in flatwoods, scrubby flatwoods, scrub and sandhill communities in Florida.] AND J. R. ABRAHAMSON. 1996. Effects of a low-intensity winter fire on long-unburned Florida sand pine scrub. Nat. Areas J. 16(3):117—183. [A long-unburned stand of sand pine (Pinus clausa) was burned with a low intensity winter burn. The results indicate that this burning regime did not restore the populations of endemic herbaceous species nor did it restore a sand pine canopy. In fact, the community shifted toward a xeric hammock, characterized by the persistence of woody understory species. This study demonstrates the necessity of monitoring the consequences of the 118 No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 119 reintroduction of fire to old stands of sand pine and of recognizing variation in vegetative responses to different fire regimes. ] ALTMAN, B. S. 1990. The scrub jay—Florida’s fragile native. USF Magazine 32(5):4-6. [The article details the study of the Florida scrub jay in its natural environment, its reproductive habits, and its state of near extinction. ] AusTIN, D. F. 1990. Vegetation on the Florida Atlantic University ecological site. Florida Scient. 53(1):13-27. [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, endangerment and introduced or native status given.] , C. F. NAUMAN, AND B. E. TatyeE. 1981. Hobe Sound scrub vegetation study. Unpublished report submitted to Nathaniel Reed. 31 pp. [A vegetation analysis was conducted to determine vegetative cover, species richness, and overall health of a scrub site located in Hobe Sound, FL (T39S, R42E, Sections 26 & 27). Special reference was given to listed plant and animal populations within areas of scrub and successional trends. ] Bauts, P. AND M. Deyrup. 1988. A habitual lurking predator of the Florida harvester ant. Pp. 547-551. In: Adv. Myrmecol. TRAGkR, J., (ed.). Leiden, E. J. Brill. [Discussed is the apparent aggregation of the antlions (Brachynemurus nebulosus) in nest mounds of Florida harvester ants. Antlions normally prey on harvester ants (Pogonomyrmex badius) by lying ambush under a thin layer of soil or sand. This behavior is possibly linked to the unique aposomatic coloration of B. nubulosus.| Bancroft, G. T. AND G. E. WOOLFENDEN. 1982. The molt of scrub jays and blue jays in Florida. Omotholog. Monogr. (29):1—51. [Detailed molt data were collected on live scrub jays and blue jays at Archbold Biological Station, Highlands County, FL, where both species are common permanent residents. Due to an extensive banding program for the Florida scrub jay population at Archbold, numerous jays of known age, sex, and exact breeding status were studied. Age and breeding information for blue jays is less detailed, but still provides valuable information. The pattern of molt by Florida scrub jays and blue jays in Florida is similar to that exhibited by other passerines. This paper suggests that the separation between molt and breeding may be a combination of temperature, water regulation and perhaps flying efficiency. ] BARRENTINE, C. D. 1991. Notes on aggressive behavior of the gopher tortoise. Herp. Review 22(3):87-88. [A first hand account of aggressive behavior was observed for two male gopher tortoise along a fire lane in the sand pine scrub at Archbold Biological Station, Highlands County, FL. Three principal action patterns were used in the encounter, which lasted about 30 minutes. A review of the literature is provided with a discussion on the consequences of overturning. ] BatTrTog, L. E. AND E. F. Lowe. 1991. Acidification of Lake Annie, Highlands County, Florida. Water Air Soil Pollut. 65:69-80. [Lake Annie is a clear-water sinkhole on the property of Archbold Biological Station in Highlands County, FL. The surrounding vegetation is a mixture of sand pine scrub, flatwoods and scrubby flatwoods. A long-term data set of surface water chemistry exists for Lake Annie, which is remote from significant pollution sources. PH and bicarbonate alkalinity have decreased, while specific conductivity and sulfate concentration have increased. The chemical changes are apparently due to acidification from atmospheric deposition. ] BERGEN, S. 1994. Characterization of fragmentation in Florida scrub communities. Masters thesis, Florida Institute of Technology, Melbourne, FL. 71 pp. [Geographic information systems (GIS), historical aerial photographs, and soil maps were utilized to distinguish between historic and current scrub fragmentation in Brevard County, FL.] Berry, D. M. AnD E. S. Mences. 1995. Post-fire changes in resource limitation of Florida scrub plants. Pp. 197-201. In: GREENLEE, J. (ed.) Fire effects on rare and endangered species conference. International Assoc. of Wildland Fire, Coeur d’Alene, Idaho. [Four scrub species were studied to determine which change or changes resulting from fire are responsible for elevated post-fire levels of growth and fruit production. ] Bostater, C. R., C. R. HALL, AND D. R. BREININGER. 1997. High-resolution optical signatures and band selection techniques for endangered species habitat management. Proc. Int. Symp. Spectral 120 FLORIDA SCIENTIST [VOL. 67 Sensing Research 1:556—569. [This paper studies the use of high-resolution spectral reflectance signatures to identify six scrub habitat components for the Kennedy Space Center. ] BoyLe, S. R. 1996. A landscape approach to Florida scrub-jay (Aphelocoma coerulescens) habitat use in Brevard County, Florida. M.S. Thesis, Florida Institute of Technology, Melbourne, FL. 134 pp. [GIS analysis was used to determine the extent of Florida scrub jay use in Brevard County, FL. Of the remaining 13,000+ acres of scrub habitat, less than 2% was determined to be in optimal condition for Florida scrub jays. ] Brancu, L. C., D. G. Hoxir, B. M. StitH, B. W. Bowen, AND A. M. Crark. 1999. Effects of landscape dynamics on endemic scrub lizards: an assessment with molecular genetics and GIS modeling. Florida Game and Fresh Water Fish Comm. Final Rep., Tallahassee, FL. 88 pp + vi. [The effect of landscape dynamics in the Florida scrub was studied on 3 species of endemic lizards. Declines in habitat quality, loss of landscape connectivity, and reduction of patch size in scrub all pose problems for scrub lizards. ] AND D. G. Hokir. 2000. A comparison of scrub herpetofauna on two central Florida sand ridges. Florida Scient. 63(2):108—117. [A herpetological survey of the Avon Park Air Force Range and the Arbuckle Tract of the Lake Wales State Forest was conducted using visual surveys, trap arrays, trapping grids, and coverboards. Forty species were captured on the Bombing Range Ridge and 15 species on the Arbuckle Tract. Further ecological studies will be required for other scrubs as a basis for managing this rare ecosystem, especially as more and more of the scrub becomes fragmented due to urban sprawl.] BRAND, K. B. 2001. Body mass variation in breeding Florida scrub-jays (Aphelocoma coerulescens). Master of Science, University of South Florida, Tampa, FL. 56 pp. [The loss of body weight in breeding birds has been explained by several hypotheses. Predictions of these hypotheses were tested using Florida scrub jay data gathered from Archbold Biological Station in 1999 and 2000.] BREININGER, D. R. 1981. Habitat preferences of the Florida scrub jay (Aphelocoma coerulescens coerulescens) on Merritt Island National Wildlife Refuge, Florida. Masters Thesis, Florida Institute of Technology, Melbourne, FL. [The Florida scrub jay was found to occupy three plant communities (i.e., coastal scrub, coastal strand, and flatwoods) at Merritt National Wildlife Refuge. A brief analysis of the correlation of Florida scrub jay abundance with such factors as low open areas, open space, percentage of vegetation, height of shrub layer, and winter insect supply was studied. | . 1989. A new population estimate for the Florida scrub jay on Merritt Island National Wildlife Refuge. Fl. Field Nat. 17(2):25—5S2. [A variable circular plot method was used to sample avifauna within different vegetation types identified from aerial imagery. The Florida scrub jay population was estimated to range between 1400 and 3600 birds within the MINWR, suggesting that the state population number may be much lower than believed. ] . 1999. Florida scrub-jay demography and dispersal in a fragmented landscape. Auk 116(2):520— 527. [Population dynamics and cooperative breeding was studied in an isolated population in southern Brevard County between 1992 and 1998. The results indicate that the population declined due to poor reproductive success. ] AND R. B. Situ. 1992. Relationships between fire and bird density in coastal scrub and slash pine flatwoods in Florida. Am. Midl. Nat. 127:233—240. [Bird densities within coastal scrub and slash pine flatwoods were compared with time since fire, mean shrub height, number of snags and per- cent burn. The results suggest that fires every 4 years or less are likely to have a negative influence on shrub dwelling birds. ] AND P. A. SCHMALZER. 1990. Effects of fire and disturbance on plants and birds in a Florida oak/ palmetto scrub community. Am. Midl. Nat. 123(1):64-74. [The vegetation and avian communities of a site located on the Kennedy Space Center consisting of oak/palmetto scrub was intentionally burned. The prescribed fire altered avian community composition for at least 6 months where nearly all vegetation burned. Twenty years after clearing of an oak/palmetto scrub, the regenerated oak-scrub differed from adjacent oak/palmetto scrub in exhibiting more bare ground, a taller shrub layer, and more herbaceous species. The data suggests that disturbed areas can represent important wildlife habitat. ] No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 121 . P. A. SCHMALZER, AND C. R. HINKLE. 1991. Estimating occupancy of gopher tortoise (Gopherus polyphemus) burrows in coastal scrub and slash pine flatwoods. J. Herp. 25(3):317—321. [One hundred and twelve plots were established in scrub and flatwoods on the Kennedy Space Center to evaluate relationships between number of burrows and densities. The standard correction factor (0.614) was not suitable for estimating the number of tortoises from burrow counts for the Kennedy Space Center. ] , P. A. SCHMALZER, AND C. R. HINKLE. 1994. Gopher tortoise (Gopherus polyphemus) densities in coastal scrub and slash pine flatwoods in Florida. J. Herp. 28(1):60-65. [Densities of gopher tortoises were compared with habitat characteristics in scrub and in flatwoods on the Kennedy Space Center. The relationship between densities of tortoises and time since fire classes were inconsistent. ] , P. A. SCHMALZER, D. A. RYDENE, AND C. R. HINKLE. 1988. Burrow and habitat relationships of the gopher tortoise in coastal scrub and slash pine flatwoods on Merritt Island, Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl., Program Final Report. 238pp. + x. [Field plots were established within scrub and pine flatwoods on Kennedy Space Center to study gopher tortoise habitat relationships. A census was conducted over a one-year period to determine time since last fire, canopy cover, shrub cover, oak cover, herbaceous cover, soil type, burrow activity and density. A census was done by bucket trapping or stick method in the fall, camera system or pulling hook in the winter, and camera system in the spring. The results indicate that tortoise densities remained similar season to season. These two habitat types may actually contain as many or more tortoise than sandhills. ] , M. J. PROVANCHA, AND R. B. Smitu. 1991. Mapping Florida scrub jay habitat for purposes of land-use management. Photogram. Eng. Rem. Sensing 57(11):1467—1474. [GIS applications were used to map areas of primary and secondary Florida scrub jay habitat on Kennedy Space Center using vegetation and soil maps. ] —, M. A. BUuRGMAN, AND B. M. StitH. 1999. Influence of habitat quality, catastrophes, and population size on extinction risk of the Florida scrub-jay. Wildlife Soc. Bull. 27(3):810—822. [A population risk model was developed to examine influence of population size, catastrophes (epidemics and hurricanes), and habitat management scenarios on Florida scrub jay populations. ] , V. L. Larson, B. W. DUNCAN, AND R. B. Smitu. 1998. Linking habitat suitability to demographic success in Florida scrub jays. Wildlife Soc. Bull. 26(1):118—128. [Spatial patterns in habitat suit- ability (HSI), reproductive success, and survival were quantified at different scales from attributes of habitat patches, nest sites, and territories of the Florida scrub jay using GIS and remote sensing on the Kennedy Space Center. ] , V. L. Larson, D. M. Oppy, AND R. B. Situ. In press. How does variation in fire history influence Florida scrub-jay demographic success? Jn: GREENLEE, J., (ed.) Fire effects on rare and endangered species conference. International Assoc. of Wildland Fire, Fairfield, Washington. [The Florida scrub jay is threatened with extinction because of habitat loss, ecosystem fragmentation, and disruption of natural fire patterns. The objectives of this study were to determine how Florida scrub jay demography varies with fire on the Kennedy Space Center.] , V. L. Larson, D. M. Oppy, R. B. SmitH, AND M. J. Barkaszi. 1996. Florida scrub jay demography in different landscapes. Auk 113(3):617-625. [Florida scrub jay demography and cooperative breeding were measured from 1988 to 1993 at two sites on Kennedy Space Center. ] , V. L. Larson, B. W. DuncaAN, R. B. SmitH, D. M. Oppy, AND M. F. GooDcHiLb. 1995. Landscape patterns of Florida scrub jay habitat use and demographic success. Conser. Biol. 9(6):1442—1453. [Comparisons of habitat types with measures of density and demographic performance showed that open oak was the optimal habitat type for Florida scrub jays on the Kennedy Space Center using GIS and remote sensing techniques. | , V. L. Larson, R. ScHaus, B. W. Duncan, P. A. SCHMALZER, D. M. Oppy, R. B. Situ, F. ADRIAN, AND H. HILL, Jr. 1996. A conservation strategy for the Florida scrub jay on John F. Kennedy Space Center/Merritt Island National Wildlife Refuge: An initial scientific basis for recovery. National Aeronautics and Space Administration Technical Memorandum 111676. Kennedy Space Center, FL. [Population trends were predicted using population modeling and field 122 FLORIDA SCIENTIST [VOL. 67 data on reproduction and survival of Florida scrub jays collected from 1988-1995 on the Kennedy Space Center. | Burcu, J. N. 1992. Notes on southern Gulf Coast scrub. Florida International University, Miami Shores, FL. [This study provides information from several sites in southern coastal Lee County through the southernmost limit of xeric habitats on the southern gulf coast of Florida, in Collier County. ] BuRGER, J. AND G. E. WOOLFENDON. 1999. Metal concentrations in the eggs of endangered Florida scrub- jays from central Florida. Arch. Environ. Contam. Toxicol. 37:385—388. [Metal levels in the eggs of the Florida scrub jay were studied between 1997 and 1998 for a population at Archbold Biological Station. On the basis of trophic-level consideration, the concentrations in the eggs should have been low, however, they were variable. Metal levels in these eggs may be the result of feeding on insects that are at a higher trophic level than seed or grass eating species. ] Buroman, M. A., D. R. BREININGER, B. W. DUNCAN, AND S. FERSON. 2001. Setting reliability bounds on habitat suitability indices. Ecological Applications 11(1):70—78. [This paper discusses alternative approaches to dealing with uncertainty in setting reliability bounds on HSI calculations and outlined an example application to the Florida scrub jay habitat model.] CARRINGTON, M. E. 1997. Soil seed bank structure and composition in Florida sand pine scrub. Am. Mid. Nat. 137(1):39-47. [The soil seed bank of selected sand pine scrub sites was characterized for herbaceous and some woody species, especially as it relates to recovery after fire. ] . 1999. Post-fire seedling establishment in Florida sand pine scrub. J. Veg. Science 10:403-412. [This study looks at quantification of pre- and post-fire seedling establishment and microsite characteristics in two Florida sand pine scrub sites burned in May 1993. Post-fire seedling establishment in sand pine scrub was sparse relative to other communities. | CHRISTENSEN, N. L. 1988. Vegetation of the southeastern coastal plain. Pp. 322-331. Jn: BARBouR, M. G. AND W. D. BILLINGS, (eds.), North American Terrestrial Vegetation. [This paper discusses the weather of the southeastern coastal plain, the various xeric upland plant communities (1.e., northern pine barrens, sandhill pine forests, and sand pine scrub) and vegetation responses to fire.] CHRISTMAN, S. P. AND W. S. Jupp. 1990. Notes on plants endemic to Florida scrub. Florida Scient. 53(1):52-73. [A three-year field survey of Florida interior sand pine scrubs has resulted in better understanding of many Central Florida scrub endemic plants. The status of 21 species of plants that are endemic to central peninsular Florida are reported, with habitat notes, range extensions, and corrections of erroneous literature records documented. | Cox, A. C. 1998. Comparative reproductive biology of two Florida pawpaws Asimina reticulata Chapman and Asimina tetramera Small. Ph.D. Dissertation, Florida International University, Miami Shores, FL. 119 pp. [Two closely related pawpaws were studied in the sand pine scrub on the Atlantic Coastal Ridge to determine if differences contribute to the rarity of Asimina tetramera. | Cox, J., D. INKLEY, AND R. Kautz. 1987. Ecology and habitat protection needs of gopher tortoise (Gopherus polyphemus) populations found on lands slated for large-scale development in Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Technical Report No. 4. Tallahassee, Fl. 75 pp. [This paper offers criteria to use in deciding when the protection of gopher tortoise habitat becomes a biologically important issue in the review of large projects. Life-history characteristics, general biology, population viability, minimum area requirements, estimating densities, and management plans are discussed for long-term survival of this complex species. ] , R. Kautz, M. MACLAUGHLIN, AND G. TERRY. 1994. Closing the gaps in Florida’s wildlife habitat conservation system. Florida Game and Freshwater Fish Commission. Tallahassee, FL. 239 pp. [This report describes habitat areas in Florida that should be conserved if key components of the state’s biological diversity are to be maintained. The project employed a computerized Geographic Information System to manipulate geographic data sets and create distribution maps for selected species of wildlife, threatened species of plants, and rare plant communities. Satellite imagery was utilized to note over 25,000 geographically referenced points documenting known occurrences of rare animals, plants, and communities. ] CRUMPACKER, D. W., E. O. Box, AND E. D. Harpin. 2001. Potential breakup of Florida plant communities as a result of climatic warming. Florida Scient. 64(1):29-43. [A climatic envelope model was No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 123 utilized to predict potential changes of 13 major plant communities (including sand pine scrub) in Florida over the next 100 years.] Curry, R. L. 1989. Florida scrub jay kills a mockingbird. Condor 92:256—257. [Detailed description of a Florida scrub jay attack, killing, and then feeding on a healthy adult northern mockingbird. Florida scrub jays are aggressive, omnivorous birds, which will feed on small vertebrates including frogs, lizards, snakes, rodents, and fledgling birds. ] DELANEY, K. R. AND R. P. WUNDERLIN. 1989. A new species of Crotalaria (Fabaceae) from the Florida central ridge. SIDA 13(3):315-324. [Crotalaria avonensis is described from the xeric white sand scrub habitat of Highlands County, FL. This species was a new discovery for the Lake Wales Ridge region of Highlands and Polk counties. ] . R. P. WUNDERLIN, AND B. F. HANSEN. 1989. Rediscovery of Ziziphus celata (Rhamnaceae). SIDA. 13(3):325-—330. [The rare Florida endemic Ziziphus celata, last collected in 1955 and presumed extinct, was rediscovered in Highlands and adjacent Polk counties. | DEPARTMENT OF THE INTERIOR. 1992. Endangered and threatened wildlife and plants; proposed endangered status for three Florida plants of the Genus Conradina. Fed. Register 57(98):21369-21374. [The Service proposes to list three plant species belonging to the genus Conradina (minty rosemaries) as endangered species, Conradina glabra (Apalachicola rosemary), Conradina brevifolia (short- leaved rosemary), and Conradina etonia (Etonia rosemary). All three of these species are found in xeric scrub uplands. ] Deyrup, M. 1989. Arthropods endemic to Florida scrub. Florida Scient. 52(4):254—270. [An annotated list of 46 species endemic to the scrub community is detailed for the central ridges, especially the Lake Wales Ridge. ] . 1990. Arthropod footprints in the sands of time. Florida Entomol. 73(4):529-538. [Arthropod distribution in Florida’s sandy uplands shows some obvious patterns. Arthropods endemic to Florida scrub are concentrated on the Lake Wales Ridge, apparently a long-term refuge. While scrub areas to the north apparently lost diversity, some species that must have migrated through there from southwestern North America are now only in southern scrubs. ] , C. JOHNSON, G. C. WHEELER, AND J. WHEELER. 1989. A preliminary list of the ants of Florida. Florida Entomol. 72(1):91—101. [A preliminary list of ant species from Florida is provided. Ants from xeric uplands are included, but not identified. ] AND E. S. MENGES. 1997. Pollination ecology of the rare scrub mint Dicerandra frutescens (Lamiaceae). Florida Scient. 60(3):143—157. [Natural populations of this rare scrub endemic were examined on the Lake Wales Ridge to determine if pollinator specificity or poor service could be factors contributing to this species restricted range and decline. ] AND R. E. Wooprurr. 1991. A new flightless Psammodius from Florida’s inland dunes (Coleoptera: Scarabaeidae). Coleopterists Bull. 45(1):75—80. [Psammodius relictillus is described from specimens taken from relict inland dunes in Florida. This blind and flightless species occurs in Florida scrub habitat on the southern Lake Wales Ridge and on the highly disjunct Atlantic Coastal Ridge. ] Diemer, J. E. 1987. Gopher tortoise status and harvest impact determination. Final report submitted to the Florida Game and Fresh Water Fish Commission, 4005 South Main Gainesville, FL. 150 pp. [A 6.5 year study of gopher tortoise status, harvest levels, population dynamics, and movements was finished in June 1987. Questionnaires and field surveys revealed that the gopher tortoise remains widely distributed in Florida. The report discusses the occurrence of tortoise in a wide variety of xeric habitats, including oak scrub and sand pine scrub communities. Marking and tracking with radio telemetry were utilized to provide insight on tortoise use of clear cuts, windrows, and flooded burrows, as well as to gain information on burrow defense, usurpation, and cohabitation. ] . 1987. The status of the gopher tortoise in Florida. Pp. 72-83. In: Opum, R., K. RIDDLEBERGER, AND J. OzieR (eds.). Proc. Third Southeast. Nongame and End. Wildl. Symp., Ga. Dept. Nat. Resour., Game and Fish Div., Atlanta, GA. [Field surveys were conducted between 1981-1987 to determine the status and distribution of the Florida gopher tortoise. All xeric uplands and dry prairies were surveyed. Gopher tortoise decline, coupled with the species’ low fecundity, necessitates further evaluation of tortoise harvest regulations. | 124 FLORIDA SCIENTIST [VOL. 67 . 1989. An overview of gopher tortoise relocation. Pp. 1-5. Jn: Diemer, J., D. R. JACKSON, J. L. LANpers, J. N. LAYNE, AND D. A. Woop (eds.) Gopher Tortoise Relocation Symp. Proc., Nongame Wildlife Prog. Tech. Rep. #5. Florida Game and Fresh Water Fish Commission. Tallahassee, FL. [Previous studies have provided minimal or conflicting data regarding the fate of relocated gopher tortoises (Gopherus polyphemus). Suggestions on maintaining gopher tortoise populations are given with biological considerations associated with translocating tortoises. ] Do an, R. W., R. YAHR, E. S. MENGES, AND M. D. HALFHILL. 1999. Conservation implications of genetic variation in three rare species endemic to Florida rosemary scrub. Am. J. Bot. 86(11):1556—1562. [Fire suppression and land alteration over the past 50 years has greatly reduced and altered much of Florida’s scrub vegetation. Patterns of isozyme variation in three rare perennial plants endemic to the rosemary scrub of the Lake Wales Ridge were studied to provide management procedures that may optimize conservation of these and other scrub endemics.] Doren, R. F., R. E. ROBERTS, AND D. R. RICHARDSON. 1989. Sand pine scrub burning: Some positive steps for management. Proc. 17th Tall Timbers Fire Ecology Conference. 2 pp. Tallahassee, FL. [Notes on the importance of fire as an ecological factor in species distribution, composition, and productivity of the sand pine scrub community are discussed. Due to narrowly adapted fauna and flora, increasingly reduced and fragmented habitat, and isolation from other scrub types, the modified chaparral fuel model (NFFL Model 4) was utilized as a basis for modifying and testing a prescription for managing the sand pine scrub. ] Douc ass, J. F. 1990. Patterns of mate-seeking and aggression in a southern Florida population of the gopher tortoise (Gopherus polyphemus). Proc. Symp. Desert Tortoise Council. Pp. 155-199. [Studies on the courtship and combative behaviors of the Florida gopher tortoise were observed as part of a long-term study at Archbold Biological Station on the southern end of the Lake Wales Ridge. This study observed tortoise in low flatwoods, scrubby flatwoods, sand pine scrub, sandhills, and cultivated areas. ] Doy.e, T. W., L. GORHAM, AND B. PLatr. 1992. The effect of gulf coast hurricanes on the growth and stand dynamics of Florida panhandle sand pine communities. Unpublished report submitted to Natl. Wetlands Research Center. Miami, FL. [A dendroecological study of sand pine growth and stand development in the Florida panhandle was conducted to investigate the effect of hurricanes on coastal plant communities. ] DrescHEL, T. W., R. B. SmitH, AND D. R. BREININGER. 1990. Florida scrub jay mortality on roadsides. Florida Field Nat. 18(4):82—83. [Four Florida scrub jay carcasses were found between May and June 1989 on two roadsides in Brevard County. Road mortality may be significant for small populations where it may contribute to the extirpation of small local populations. Data on habitat features are needed to develop strategies to mitigate the problem throughout the range of this species. | Duncan, B. W., D. R. BREININGER, P. A. SCHMALZER, AND V. L. Larson. 1995. Validating a Florida scrub jay habitat suitability model, using demography data on Kennedy Space Center. Photogr. Eng. Rem. Sensing 61(11):1361—1370. [A habitat suitability index model for the Florida scrub jay was tested for the Tel-4 study site on Kennedy Space Center. The model used suitability graphs that quantify habitat preference with respect to a given variable to produce spatial estimates of Florida scrub jay habitat suitability. ] , S. Boye, P. A. SCHMALZER, AND D. R. BREININGER. 1996. Spatial quantification of historic landscape change within two study sites on John F. Kennedy Space Center. 21 p. Proc. Sixteenth Ann. USRI Conference. [Aerial images from 1943-1989 photography were measured to determine historical changes in scrubby flatwoods and scrub landscapes. Land use categories were mapped and digitized into ARC/INFO. The data indicates that since the 1940’s, there has been a dramatic decrease in the amount of open scrub with a concomitant increase in forest cover. These data provide an initial check of existing management practices, particularly as it might relate to listed species. ] , S. Boye, D. R. BREININGER, AND P. A. SCHMALZER. 1999. Coupling past management practice and historic landscape change on John F. Kennedy Space Center. Landscape Ecol. 14:291—309. [Historic land cover dynamics in scrubby flatwoods and scrub were measured using aerial images No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 125 from several years beginning in the 1940’s through 1989. This study highlights the importance of sound management practices if we are to perpetuate these valuable pine ecosystems. | EISNER, T., K. D. McCormick, M. SAKAINO, M. EISNER, S. R. SMEDLEY, D. J. ANESHANSLEY, M. DEYRUP, R. L. Myers, AND J. MEINWALD. 1990. Chemical defense of a rare mint plant. Chemoecology 1:30- 37. [Analyses of leaf extracts from Dicerandra frutescens, (a highly aromatic mint plant from scrub community in Central Florida), revealed presence of 12 closely related monoterpenes. The terpenes produced serve for defense against insects, and are also released when the leaf is injured. ] EncE, K. M. 1997. Habitat occurrence of Florida’s native amphibians and reptiles. Tech. Rep. No. 16. Florida Game and Fresh Water Fish Commission. Tallahassee, FL. 44 pp + vi. [Data are provided that give the relative abundance of various native amphibian and reptile species in 31 Florida habitats (i.e., xeric uplands, scrub and sandhill) based on drift-fence surveys, literature, observations, and educated guesses. ] AND K. Dopp, Jr. 1986. A bibliography of the herpetofauna of Florida. Smithsonian Herp. Info. Serv. 72:68 pp. [This is a directory of the scientific literature, popular articles, theses and dissertations and much of the “‘gray”’ literature about Florida’s amphibians and reptiles, inclusive of the scrub community. ] AND K. Dopp, Jr. 1992. An indexed bibliography of the herpetofauna of Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Prog. Tech. Rep. No. 11. 231 pp. [This bibliography includes standard scientific references, popular articles, newsletter articles (including photocopied newspaper articles), theses and dissertations, and much of the so-called “gray” literature dealing with contracted-for but unpublished government and privately funded research on amphibians and reptiles endemic to the scrub ecosystem. ] Evans, J. K., A. J. PARKER, K. C. PARKER, AND D. S. LEIGH. 1996. Edaphic properties and foliar elemental concentrations from sand pine (Pinus clausa) populations throughout Florida. Phys. Geogr. 17(3):219-241. [Soil physical and chemical properties, plus foliar elemental concentrations, are reported for surface and subsurface soil and foliar tissue samples from 20 sand pine populations, 9 from the Choctwhatchee variety in the Florida panhandle and 11 from the Ocala variety. Foliar elemental concentrations did not reflect soil elemental patterns. ] Evans, M. E., R. W. DoLan, E. S. MENGEs, AND D. R. Gorpbon. 2000. Genetic diversity and reproductive biology in Warea carteri (Brassicaceae), a narrowly endemic Florida scrub annual. Am. J. Bot. 87(3):372-381. [The mating patterns, isozyme variation, and effective population sizes of Carter’s mustard were examined to better understand its population biology as it might relate to preserve design and management. | FERNALD, R. T. AND B. TOLAND. 1991. The Florida scrub jay. Florida Game and Fresh Water Fish Commission, Office of Environmental Services, Tallahassee, FL. 7 pp. [This pamphlet briefly summarizes the range and habitat of the Florida scrub jay, its food, nesting preferences, social and family lifestyles, and management practices. ] FiscHerR, N. H., G. B. WILLIAMSON, J. D. WEIDENHAMER, N. TANRISEVER, A. DELA PENA, E. JORDAN, AND D. R. RICHARDSON. 1989. Allelopathic mechanisms in the Florida scrub community. Pp. 183-193. In: CHou, C. H. AND G. R. WALLER (eds.), Phytochemical Ecology: allelochemicals, mycotoxins and insect pheromones and allomones. Academia Sinica Monograph Series No. 9. [The hypothesis that allelochemicals released from members of the Florida scrub community deter the invasion of fire-prone sandhill grasses was investigated. The germination and growth of grasses is reduced in soils from beneath the scrub perennial, Polygonella myriophylla (Small) Horton, supporting the hypothesis that this shrub chemically interferes with the growth of other species. ] , N. TANRISEVER, AND G. B. WILLIAMSON. 1988. Allelopathy in the Florida scrub community as a model for natural herbicide actions. Pp. 233-249. In: CuTLer, H. (ed.) Natural Products: Potential in Agriculture. Amer. Chem. Soc. Symp. Ser. 380. American Chemical Society, Washington, D.C. [The possibility that allelochemicals released from plants of the Florida scrub community deter the invasion of fire-prone sandhill grasses was investigated. Three scrub species, Ceratiola ericoides, Conradina canescens, and Calamintha ashei, were tested for effects on the germination and radicle growth of lettuce (Lactuca sativa) and little bluestem (Schizachyrium scoparium), a native grass of the Florida sandhill community. ] 126 FLORIDA SCIENTIST [VOL. 67 , G. B. WILLIAMSON, J. D. WEIDENHAMER, AND D. R. RICHARDSON. 1994. In search of allelopathy in the Florida scrub: the role of terpenoids. J. Chem. Ecol. 20(6):1355—1380. [The hypothesis was tested that allelopathic agents released from fire-sensitive plants of the Florida scrub community deter the invasion of fire-prone sandhill grasses. ] , T. HEEKYUNG, AND G. B. WILLIAMSON. 1992. Photochemical allelopathic activation in the Florida scrub. Unpublished report submitted to Louisiana St. Univ., 6 pp. [The hypothesis was tested that during the long fire-free periods in the Florida scrub, allelopathic action of the endemic false rosemary (Ceratiola ericoides; Empetraceae) inhibits germination and growth of fire-facilitating graminoids (Schizachyrium scoparium and Leptochloa dubia) and pines (Pinus palustris and P. elliotii) of the sandhill. ] , G. B. Witiiamson, N. TANRISEVER, A. DE LA PENA, J. D. WEIDENHAMER, E. D. JORDAN, AND D. R. RICHARDSON. 1989. Allelopathic actions in the Florida scrub community. Biolog. Plant. 31(6):471-478. [The hypothesis that allelochemicals released from members of the Florida scrub community deter the invasion of fire-prone sandhill grasses was investigated. Constituents of the endemic scrub members, Ceratiola ericoides, Conradina canescens and Calamintha ashei, were examined for their phytotoxic activity. Effects of the plant natural products on the germination and radicle growth of lettuce (Lactuca sativa), as well as little bluestem (Schizachyrium scoparium) and green sprangletop (Leptochloa dubia), two native grasses of the Florida sandhill community, were tested. ] Fitzpatrick, J. W. 1991. Florida’s vanishing scrubland. Field. Bull. Field Mus. Nat. Hist. Jan./Feb. Pp. 9-10. [A brief geologic history of the scrub is discussed with emphasis on the delicate balance of this system and its continued existence. ] . 1992. Vanishing Florida scrub. Florida Nat. 65(1):8—9. [A brief overview of the Florida scrub environment with special reference to endemic species, endangerment, protection, and restoration goals. | . 1992. Florida scrub jay. Florida Nat. 65(1):13. [A brief article on the Florida scrub jay and its habitat, adaptations, and ecological specializations. ] AND G. E. WOOLFENDEN. 1988. Components of lifetime reproductive success in the Florida scrub jay. Pp. 305-320. In: CLutroNn-Brock, T. H. (ed.). Reproductive Success, University of Chicago Press, Chicago, IL. [Patterns of lifetime reproduction among Florida scrub jays were analyzed for one hundred jays whose complete lifetime breeding histories have been documented. ] , G. E. WooLFENDEN, AND M. T. Kopeny. 1991. Ecology and development-related habitat requirements of the Florida scrub jay (Aphelocoma coerulescens coerulescens). Nongame Wildlife Program, Tech. Rep. (8):1-49. [The statewide population of the Florida scrub jay was recently estimated at about 7,000 to 11,000 birds. Over half of the remaining jays in the state occur on and around two large, federally owned tracts of land: the Ocala National Forest and Merritt Island National Wildlife Refuge. Procedures for determining population size of Florida scrub jay families and territories are provided in the context of establishing preservation sites within developments throughout Florida. Measures for implementing suitable jay habitat buffer zones and corridors are also discussed. This publication establishes a basis for Florida scrub jay preservation techniques on a statewide level. ] AND G. E. WOOLFENDEN. 1989. Florida scrub jay. Pp. 201-218. Jn: Newton, I. (ed.) Lifetime reproduction in birds. Academic Press Ltd., Orlando, FL. [The results of a 14-year survey of a wild population of the Florida scrub jay at the Archbold Biological Station were used to determine patterns of lifetime reproductive success. Also discussed are the habitat, social system, and demographic attributes of this co-operative breeding bird.] , G. E. WOOLFENDEN, AND K. J. McGowan. 1988. Sources of variance in lifetime fitness of Florida scrub jays. Pp. 876-891. Jn: OUELLET, H. (ed.) Congressus Internationalis Ornithologicus. Univ. Ottawa Press. Ottawa, Canada. [The lifetime fitness of 67 known-age Florida scrub jays (Aphelocoma c. coerulescens) from seven cohorts now absent from the breeding population was analyzed. The sexes show similar patterns of variance in fitness. Certain breeders simply outperform others in most or all respects, exaggerating variance in lifetime reproduction. ] No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 127 Fieck, D. C. AND J. LAYNe. 1990. Variation in tannin activity of acorns of seven species of central Florida oaks. J. Chem. Ecol. 16(10):2925—2934. [Acorns of seven sympatric species of oaks (Quercus spp.) occurring in central Florida were compared for protein-precipitating ability (PPA). Six of the seven species studied occurred in sandhills, pine flatwoods, scrubby flatwoods, and sand pine scrub communities. The other species, laurel oak, occurred in mesic forest communities at Archbold Biological Station. ] FLEISCHER, A. L. Jr. 2000. The influence of time budget and rate of food handling and consumption on the timing of breeding of female Florida scrub-jays (Aphelocoma coerulescens): a comparison between populations in natural and suburban habitats. Masters thesis, Univ. of South Florida, Tampa, FL. 82pp. [The relationship between pre-laying behavior and food handling and con- sumption rates on the timing of breeding of female Florida scrub jays in natural and suburban scrub were studied in 1996 and 1997. The results suggest variation in food handling rates may explain annual variation in timing of breeding in natural scrub whereas suburban birds showed little varia- tion due to supplemental food supplies.] FLEISCHER, T. L. 2000. Reactions of Florida scrub-jays (Aphelocoma coerulescens) to eggs and shams added to their nests. Master thesis, Univ. of South Florida, Tampa, FL. 40pp. [Brood parasites may pose another threat to the Florida scrub jay. The reactions of Florida scrub jays to nest additions of brown-headed cowbird eggs and eggs of five other species were studied. ] FLORIDA Dept. OF AGRICULTURE AND CONSUMER SERVICES. 1989. Protecting Florida homes from wildfire: A guide for planners, developers and fire services. Fl. Dept. of Ag. and Consumer Serv. 21 pp. [A brief explanation on preventing homes from being destroyed by wildfire; due to the increasing number of homes near areas where fire could damage. |] FLORIDA GAME AND FRESH WATER FISH COMMISSION. 1992. The scrub ecosystem. Wild Florida Publication. 1:10. [Endemic wildlife, unique flora and fauna, fire ecology, and the typical traits known to scrub communities are discussed. | . 1992. Recognizing scrub. Florida Nat. 65(1):10. Reprinted and edited from: The Wild Florida Publication, Vol. 1. [Brief article, which identifies the various types of scrub, its fire history, and its unique flora and fauna. |] FLORIDA NATURAL AREAS INVENTORY. 2000. Tracking list of rare, threatened, and endangered plants and animals and exemplary natural communities of Florida. Tallahassee, FL. [Identifies listed plant and animal species found in scrub and other plant communities in Florida. ] FRANK, P. A. AND J. N. LAyne. 1991. Nests and daytime refugia of cotton mice (Peromyscus gossypinus) and golden mice (Ochrotomys nuttalli) in South-Central Florida. Am. Midl. Nat. 127:21—30. [Daytime refuge selection, nest construction, and patterns of refuge use of cotton mice and golden mice were studied at the Archbold Biological Station in south-central Florida. The habitat was scrubby flatwoods with some mention of Florida mice and their occurrence in scrub habitats. ] AND K. Lips. 1989. Gopher tortoise burrow use by long-tailed weasels and spotted skunks. Florida Field Nat. 17(1):20-22. [Approximately 60 species of vertebrates are known to occur in association with burrows of the gopher tortoise (Gopherus polyphemus) as accidental, occasional, or obligate residents. Spotted skunks were observed using gopher burrows in the scrub habitat at Archbold Biological Station, Highlands County, FL.] FRANZ, R. AND L. L. Smitu. 1999. Distribution and status of the striped newt and Florida gopher frog in peninsular Florida. Florida Fish and Wildl. Conserv. Comm. Final Rep. 46 pp + vi. Tallahassee, FL. [The distribution of the Florida gopher frog was documented based on museum records, published literature, field notes and fieldwork. The Florida gopher frog is known from 258 sites in 45 counties and primarily occurs in xeric uplands (i.e., scrubby flatwoods, sandhills, and scrub), but breeds in isolated wetland ponds. | FRONCZEK, F. R., N. TANRISEVER, AND N. H. FiscHer. 1987. Structure of 2’,4”-Dihydroxychalcone. Acta Crystallogr. C43:158—160. [Ceratiola ericoides (false rosemary) is an endemic, dominant shrub of the Florida Scrub. Preliminary investigations showed C. ericoides releases allelopathic constituents, which inhibit the germination and growth of plants of the Florida sandhill community. Chemical analysis of rosemary was performed leading to isolation of a number of flavonoids. | 128 FLORIDA SCIENTIST [VOL. 67 GREENBERG, C. H., D. G. NEARY, AND L. D. Harris. 1994. Effect of high-intensity wildfire and silvicultural treatments on reptile communities in sand-pine scrub. Conserv. Biol. 8(4):1047—1057. [This paper tested whether the herpetofauna response to clear cutting followed by site preparation was similar to high intensity wildfire followed by logging in sand pine scrub.] , D. G. Neary, L. D. Harris, AND S. P. LInDA. 1995. Vegetation recovery following high-intensity wildfire and silvicultural treatments in sand pine scrub. Am. Midl. Nat. 133:149-163. [This paper hypothesized that clear-cutting mimics natural high-intensity disturbance by wildfire followed by salvage logging in sand pine scrub, and tested whether vegetation adapted to recovery from fire would respond similarly to other types of biomass removal. The results suggest that many scrub species responded similarly to aboveground biomass removal. ] HALL, J. 2002. Sand pine scrub fragments in Pinellas County, Florida-species richness and mapping using GIS. Master thesis, Department of Geography, Univ. of South Florida, Tampa, FL. S55pp. [Geographic information systems (GIS) was used to identify and examine landscape features of fragmented patches of Florida scrub in Pinellas County.] Hammirt, Y. C. 1992. Florida’s ancient archipelago. The Stuart News/Port St. Lucie News. Pp. 1-8. [A series of articles dealing with many aspects of the Florida scrub ecosystem.] Hawkes, C. V. AND E. S. MENGEs. 1994. Density and seed production of a Florida endemic, Polygonella basiramia, in relation to time since fire and open sand. Am. Midl. Nat. 133:138-148. [Density and reproductive output in relation to fire, open sand, and other site factors were determined for this federally endangered scrub endemic. The open sand habitat is critical in the life history strategy of this obligate-seeding, perennial herb. ] AND E. S. MENGES. 1996. The relationship between open space and fire for species in a xeric Florida shrubland. Bull. Torrey Bot. Club 123(2):81—92. [Fire and open space may both play a role in the development and maintenance of species assemblages in the rosemary phase of sand pine scrub. This study examined individual species densities with time since fire, open space, elevation, ground cover, canopy cover and soil type for several scrub species. ] HERNANDEZ, H. P. AND N. H. FiscHer. 1988. Unambiguous structure determination of a new flavonoid. Spectr. Lett. 21(9&10):927—934. [The structure elucidation of a new flavone, 5,6,4’-trihydroxy- 7,8,3'-trimethoxyflavone, was performed by the use of Insensitive Nuclei Assigned by Polarization Transfer (INAPT), a simple and sensitive NMR technique. This new flavone was isolated from Calamintha ashei, a scrub endemic. ] HERNDON, A. 1999. Life history of Liatris ohlingerae (Asteraceae), an endangered plant endemic to the Lake Wales Ridge, FL. Florida Fish and Wildl. Conserv. Comm. Final Rep. Tallahassee, FL. 46 pp. + vi. [Liatris ohlingerae, an endemic wildflower found only in the lake Wales Ridge of Florida, has low rates of seedling recruitment and adult mortality, and therefore a low rate of population turnover. No new management approaches are suggested based on the data collected. ] AND W. J. PLatr. 1991. Life-history studies on the scrub endemic Liatris ohlingerae. Unpublished report. 4 pp. [Liatris ohlingerae is a narrowly endemic member of the Florida scrub community. Basic information on the requirements for the establishment of new individuals in the population will be obtained through a seed germination study in the field, necessary due to the very few protected populations, which could disappear. ] Hokir, D. G., B. M. StrTH, AND L. C. BRANCH. 2001. Comparison of two types of metatpoulation models in real and artificial landscapes. Conserv. Biol. 15(4):1102—1113. [Patch occupancies and demographic rates for regional populations of the Florida Scrub Lizard (Sceloporus woodi) were measured and compared to observed occupancies with those of both the incidence function and matrix models for a large population located at the Avon Park Bombing Range in Florida. ] Huck, R. B., W. S. Jupp, W. M. WuitTEN, 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 cladistic analysis and discussion of endemism. System. Bot. 14(2):197-213. [Dicerandra christmanii, differing from D. frutescens in anther and corolla color, essential oils, average leaf length, and anther connective glandularity, is described from sclerophyllous oak-sand pine scrub on yellow, well-drained sandy soils near Sebring, in Highlands County, FL.] No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 129 IversON, G. B. AND D. F. Austin. 1992. A regional comparison of Florida scrub sites, Palm Beach County. Unpub. Rept. Boca Raton, FL. [This paper reports on the extent to which Florida Scrub sites in Palm Beach County as of 1987 share a group of plant species characteristic of or endemic to this ecosystem on the sub-tropical Atlantic Coastal Ridge prior to development. ] AND D. F. AusTIN. 1988. Inventory of native ecosystems in Palm Beach County, Phase III Report. Location and evaluation of sites for possible preservation as wilderness island park preserves. Unpublished Report submitted to Florida Atlantic University, Boca Raton, FL. [A listing of possible sites for wilderness preserves in Palm Beach County. ] -JoHNsoN, A. F. AND W. G. ABRAHAMSON. 1990. A note on the fire responses of species in rosemary scrubs on the southern Lake Wales Ridge. Florida Scient. 53(2):138—143. [Three scrub species appeared in post burn samples of plots in rosemary scrubs, that were rare or absent in preburn samples of the same plots and were also rare in unburned scrubs. Changes in cover levels with stand age suggests that these species are displaced by a dense cover of rosemary within 9 to 12 years.] — AND J. W. MULLER. 1993. An assessment of Florida’s remaining coastal upland natural com- munities: Final Summary Report. Unpublished report submitted to Florida Department of Community Affairs. 37 pp. [This study identified, described, and ranked (primarily in terms of their vegetative communities) the remaining undeveloped coastal upland sites in an effort to provide a background for selecting those natural sites most urgently in need of preservation. To this end both publicly and privately owned sites were evaluated. ] AND W. C. ABRAHAMSON. 2001. Stem turnover in the rhizomatous scrub oak, Quercus inopina, from south-central Florida. Am. Midl. Nat. 147:237—246. [In Q. inopina scrubs, median life span of stems was about 4 years, precluding the scrubs it dominates from reaching the 3-4 meter height, which is unfavorable to the Florida scrub jay. It appears that stem lifespan rather than fire return intervals dictates height growth in inopina scrubs on the Lake Wales Ridge in central Florida. ] Jones, C. A. 1989. First record of pawpaw consumption by the Florida mouse. Florida Scient. 52:7. [A Florida mouse was observed collecting a fruit of flag pawpaw in the sand of the Ordway Preserve. ] . 1990. Review of the effects of fire on Peromyscus and Podomys. Florida Scient. 55(2):75—-84. [Details the long-term effects of prescribed burns and lightning strikes on small non-game mammals (mice) with some discussion concerning the Florida mouse. | — AND J. N. Layne. 1993. Mammalian Species: Podomys floridanus. Amer. Soc. Mammal. (427): 1—5. [This paper discusses the general characteristics, distribution, fossil record, form, function, ontogeny and reproduction, ecology, behavior, and genetics of Podomys floridanus. | JORDAN, E. D. 1990. Seasonal changes in concentrations of secondary compounds from foliage, litter and soils of the Florida scrub. Ph. D. Dissertation, Louisiana State University, Baton Rouge, LA. 128 pp. [Chemical constituents from selected scrub species and soils were examined in different times of the year to determine peak activity especially as its may relate to function. ] , [. C.-Y. Hsien, AND N. H. FIscHer. 1992. Volatile compounds from leaves of Ceratiola ericoides by dynamic headspace sampling. Phytochemistry 31:1203—1208. Pergamon Press. [Ceratiola ericoides is a shrub endemic to the Florida scrub community and has been investigated in conjunction with studies of allelopathic interactions that affect members of the adjacent sandhill community. Headspace volatiles of C. ericoides leaves collected in spring, summer, and autumn were absorbed on Tenax TA, thermally desorbed, cryogenically refocused, and identified by GC-MS. In spring leaves hydrocarbons were most prevalent, while alcohols, aldehydes, and ketones were most abundant in summer leaves. Esters were the major components in autumn leaves. | , I. C.-Y. Hsien, AND N. H. Fiscuer. 1993. Volatiles from litter and soil associated with Ceratiola ericoides. Phytochemistry 33:299—302. [The litter of scrub rosemary and soil were analyzed for their volatiles. The major classes of volatiles were aliphatic alcohols, aldehydes and ketones, benzenoids, monoterpenes, and sequiterpenes. , N. H. FiscHer, G. B. WILLIAMSON, AND J. D. WEIDENHAMER. 1988. Determination of the allelopathic hydrocinnamic acid in the soil under the Florida wild rosemary (Ceratiola ericoides). Newsletter Phytochem. Soc. 28(1). Abstract. [Ceratiola ericoides is a shrub endemic to the Florida sand pine scrub community and often grows in open sandy soils devoid of vegetation, especially in the vicinity 130 FLORIDA SCIENTIST [VOL. 67 of mature shrubs. Hydrocinnamic acid (HCA) is biologically active on seeds of grasses native to the neighboring Florida sandhill community. Water extracts of monthly samples of the soil under three individual Ceratiola shrubs were analyzed by reverse-phase HPLC. The extracts contained varying concentrations, with the higher concentrations occurring during the summer rainy season. ] Jupp, W. S. AND D. W. HALL. 1984. A new species of Ziziphus (Rhamnaceae) from Florida. Rhodora 86:386—387. [A new endemic, Ziziphus celata, is described from the xerophytic scrub and pine- lands of the Lake Wales Ridge in peninsular Florida. ] Kautz, R. S. 1998. Land use and land cover trends in Florida. Florida Scient. 61(3/4):171—187. [This paper documents changes in forest cover types between 1936 and 1995. Sand pine plantations have increased from 1.2% to 1.9% of the state.] KEATING, W. J. 1999. Nocturnal roosting behavior of the Florida scrub-jay. Masters thesis, Univ. of South Florida, Tampa, FL. 67pp. [Florida scrub jays were observed at dawn and dusk for one year to determine two aspects of nocturnal roosting behavior; time of initiation and termination of diurnal activity and roost site use. ] KINSELLA, J. M. 1990. Comparison of helminths of three species of mice, Podomys floridanus, Peromyscus gossypinus, and Peromyscus polionotus, from southern Florida. Can. J. Zool. 69: 3078-3083. [The helminths of 102 Florida mice, 86 cotton mice, and 41 old-field mice from the Lake Wales Ridge of southern Florida were studied. Nineteen species of helminths were found, and high indexes of similarity were found among the three host species.] KLEIN, H. G. AND J. N. Layne. 1978. Nesting behavior in four species of mice. J. of Mammalogy 59(1):103—108. [Several aspects of nesting behavior such as nest-height preference, nest and nest- site attachment, and numbers and types of nests constructed were studied for Peromyscus floridanus, P. gossypinus, P. polionotus, and Ochrotomys nuttalli in the laboratory.] KRaAL, R. AND R. B. McCartney. 1991. A new species of Conradina (Lamiaceae) from northeastern peninsular Florida. SIDA 14(3):391—398. [Conradina etonia, a new species from Putnam County, was discovered in the sand pine scrub west of St. Augustine in an area supporting many scrub plant and animal species that appear to have reached their northern geographic limits. ] Kramer, D. A. 2000. Recaching facilitates cache recovery in the Florida scrub-jay: A ‘rolling horizon of short-term memory’ model. Masters thesis, Univ. of Wisconsin-Madison, Madison, WI. 68 pp. [Acorn caching and recovery were studied in a Florida scrub jay population at Archbold Biological Station. ] LAMBERT, B. B. AND E. S. MENGES. 1996. The effects of light, soil disturbance and presence of organic litter on the field germination and survival of the Florida goldenaster, Chrysopsis floridana Small. Florida Sci. 59(2):121—137. [Chrosopsis floridana, a federally endangered perennial herb endemic to the scrub community of west-central Florida was studied to determine the effects of light, disturbance, and litter build-up on germination. Seedling emergence was favored by disturbed soil, by the absence of litter and by their combination. Fire did not affect seed germination or seedling survival, but did increase flowering. ] Layne, J. N. 1989. Comparison of survival rates and movements of relocated and resident gopher tortoises in a south-central Florida population. Pp. 73-80. Jn: Diemer, J., D. R. JAcKson, J. L. LANDERS, J. N. LaYNE, AND D. A. Woop (eds.) Gopher Tortoise Relocation Symp. Proc., Nongame Wildlife Prog. Tech. Rep. #5. Florida Game and Fresh Water Fish Commission. Tallahassee, FL. [Survival and movements of 44 relocated gopher tortoises (Gopherus polyphemus) and 100 randomly selected residents representing the same size classes as the relocated group were compared on a 430-ha study area located on the Archbold Biological Station, Highlands County, FL. The major vegetation associations within the study area include sandhill, sand pine scrub, scrubby flatwoods, flatwoods, and bayheads. ] AND R. J. Jackson. 1994. Burrow use by the Florida mouse (Podomys floridanus) in south-central Florida. Am. Midl. Nat. 131:17—23. [The pattern of burrow use by the Florida mouse was investigated at ABS in sandhill and sand pine scrub habitats. ] LEENHOUTS, B. 1995. Number of Florida scrub jays detected along census routes in habitats with different fire severities. Pp. 77-79. In: GREENLEE, J. (ed.) Fire effects on rare and endangered species conference. International Assoc. of Wildland Fire, Coeur d’Alene, Idaho. [Florida scrub jays were No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 131 censused along transects in two fire regime community types (flatwoods and coastal scrub) with differing fire return intervals in the Merritt Island National Wildlife Refuge. ] Lewis, P. O. 1992. Allozyme variation in geographically restricted and widespread species of Polygonella: Biogeographical Implications. Unpublished report submitted to North Carolina State University, Raleigh, N.C. [A survey of allozyme variation in Polygonella revealed that four of the five restricted endemics have as much or more gene diversity within their populations as do their closely related widespread congeners. ] Lips, K. R. 1991. Vertebrates associated with tortoise (Gopherus polyphemus) burrows in four habitats in south-central Florida. J. of Herp. 25(4):477-481. [Burrows located in sand pine scrub, slash pine- turkey oak, and scrubby flatwoods were surveyed for tortoise activity. Habitat, size, status of the burrow, and season were studied to see how these factors affected the species composition and abundance of vertebrate associates of gopher tortoise burrows. ] LOVESTRAND, E. 1990. Living treasures of the Florida scrub. Wild Florida. Florida Game and Fresh Water Fish Commission 1:1—6. [Details the flora and fauna of the scrub ecosystem with special emphasis on the xeric or desert-like conditions and the requirement for long-term wildfires. ] Macias, F. A., F. R. FRONCZEK, AND N. H. FiscHer. 1989. Menthofurans from Calamintha ashei and the absolute configuration of desacetylcalaminthone. Phytochemistry 8(1):79—82. [Water washes from the aerial parts of Calamintha ashei, a scrub endemic shrub, provided, besides the known monoterpenes, menthofuran, (+)-epievodone and (—)-calaminthone, the new (+)desacety]l- calaminthone. The molecular structure and absolute configuration of desacetylcalaminthone was determined by singe crystal X-ray analysis of its p-bromobenzoate derivative. ] Main, K. N. AND E. S. MENGES. 1997. Archbold Biological Station, Station Fire Management Plan. Land Manag. Publ. 97—1. 94 pp. [A fire management plan has been designed to provide five fire-return intervals for the vegetation associated with ABS. The plan creates flexibility, increases hetero- geneity, and provides research opportunities for both lightning and prescribed burns. The plan includes a burning program for scrub, sandhill, flatwoods, cutthroat flatwoods, seasonal ponds, scrubby flatwoods and bayheads. | Marois, K. C. 1999. Tracking list of rare, threatened, and endangered plants and animals and natural communities of Florida. Florida Natural Areas Inventory, Tallahassee, FL. 74 pp. [This document is an annotated list of plant and animals that occur within the scrub and other plant communities in Florida with special reference to their degree of endangerment.] Martin, D. 1992. The Lake Wales Ridge National Wildlife Refuge: A new endangered species refuge for Florida’s scrub. Florida Nat. 65(1):12. [A proposal to create a National Wildlife Refuge for scrub plants and animals within the Lake Wales Ridge system is discussed. Due to the high degree of endemic species, land conservation is an important tool for long-term preservation. ] McCartTan, L. 1992. Evolution of landforms on the Florida platform with special emphasis on the Lake Wales Ridge. U.S. Geo. Surv. Bull. 2122. Pp. 1-26. [Discusses the geological beginnings of Florida, mainly the Lake Wales Ridge area, and discusses such factors as long-term crustal arches and basins which have focused deposition, other major factors that controlled marine deposition are climate, sea level position, and the presence or absence of the Panama land bridge. ] McCay, D. H. 2000. Effects of chronic human activities on invasion of longleaf pine forests by sand pine. Ecosystems 3:283—292. [Studies at Eglin Air Force Base in the Florida panhandle indicate that the amount of sand pine has nearly doubled in acreage following the extraction of turpentine from longleaf pine forests and subsequent fire suppression in the 1940’s.] . 2001. Spatial patterns of sand pine invasion into longleaf pine forests in the Florida panhandle. Landscape Ecology 16:89—98. [GIS analysis using 1949 and 1994 aerial photographs showed sand pine expansion and increased canopy cover in the pine forests of Eglin Air Force Base in the panhandle of Florida. ] McCormick, K. D., M. A. Deyrup, E. S. MENGES, S. R. WALLACE, J. MEINWALD, AND T. EISNER. 1993. Relevance of chemistry to conservation of isolated populations: the case of volatile leaf components of Dicerandra mints. Proc. Natl. Acad. Sci. 90:7701—7705. [Chemical analysis of the essential oils of four congeneric species of mint plant (Dicerandra spp.) endemic to the Florida 132 FLORIDA SCIENTIST [VOL. 67 scrub revealed a pattern of chemical similarity and dissimilarity that would not have been predicted on morphological or geographic criteria. ] McCoy, E. D. AND H. R. Musuinsky. 1992. Rarity of organisms in the sand pine scrub habitat of Florida. Conserv. Biol. 6:537—548. [Plants, amphibians, and reptiles of the Florida scrub were categorized based on several schemes of rarity. The distribution of taxa among categories of rarity were different for plants compared to amphibians and reptiles and those of previously published studies of rarity.] AND H. R. MusHINSKyY. 1994. Effects of fragmentation on the richness of vertebrates in the Florida scrub habitat. Ecology 75(2):446-457. [Vertebrates inhabiting fragments of the severely threatened sand pine scrub habitat of interior peninsular Florida were surveyed for structure, composition, distance to nearest scrub, distance to nearest larger scrub, habitats between scrubs, corridors, distance to permanent water, cover of surrounding habitats, and area reduction over time. The results suggest that the need for single large scrub reserves for resident vertebrates is not founded and that small reserves could support at least as many taxa as large ones.] McNas, W. H., K. W. OUTCALT, AND R. H. BRENDEMUEHL. 1985. Weight and volume of plantation-grown Choctawhatchee sand pine. USDA Forest Service, Res. Pap. SE-252. 44p. Southeast For. Exp. Stn., Asheville, NC. [The above ground green weight of the total tree and its major components were determined in eight stands of planted Choctawhatchee sand pine ranging in age from 7 to 27 years. | MENGES, E. S. 1992. Habitat preferences and response to disturbance for Dicerandra frutescens, a Lake Wales Ridge (Florida) endemic plant. Bull. Torrey Bot. Club 119(3):308—313. [A survey of all known populations revealed that this species is virtually restricted to excessively drained yellow sands in areas that support evergreen oak-dominated scrub vegetation. The sensitivity of this narrow endemic to loss of aboveground tissue is consistent with its recently described chemical defense. | . 1994. Fog temporarily increases water potential in Florida scrub oaks. Florida Scient. 57(3):65— 74. [The effects of fog on predawn water potential of two scrub oaks species were studied on the Lake Wales Ridge of central Florida during two winter dry seasons. | . 1999. Ecology and conservation of Florida scrub, Pp. 7-22. In: ANDERSON, R. C., J. S. FRALISH, AND J. M. BASKIN (eds.), Savannas, Barrens, and Rock Outcrops plant communities of North America. Cambridge University Press, Cambridge, U.K. [This paper provides a general overview of the Florida scrub and discusses the need for continued research especially as it relates to land management and conservation efforts. | AND N. P. GALLo. 1991. Water relations of scrub oaks on the Lake Wales Ridge, Florida. Florida Scient. 54(2):69—79. [Seasonal water relations in three scrub oaks species were studied on the Lake Wales Ridge. Results suggest that site and species differences are present in water relations and the potential exists for different drought adaptations among the oak species. ] AND V. T. SALZMAN. 1992. Archbold Biological Station plant list. Plant list of the Archbold Biological Station-reprint. 74pp. [The habitat, life history, and distribution of the vascular plants known to occur at Archbold Biological Station are discussed. This plant list is an expansion, reorganization, and update of a previous list authored by Sam Vander Kloet.] , W. G. ABRAHAMSON, K. T. Givens, N. P. GALLO, AND J. N. LAyne. 1993. Twenty years of vegetation change in five long-burned Florida plant communities. J. Veg. Sci. 4:375—386. [Rates and directions of change over a 20-year interval in five long-unburned plant communities (1.e., sandhills, scrub, scrubby flatwoods, flatwoods, and bayheads) were studied using multivariate analyses and compositional vectors. The study sites were located in fire and summer-drought adapted, xerophytic vegetation with many endemics on acidic, nutrient-poor, sandy soils in south- central peninsular Florida. | AND D. R. Gorbon. 1996. Three levels of monitoring intensity for rare plant species. Nat. Areas J. 16(3):227—237. [A three-level, hierarchical approach of increasing intensity for monitoring rare plant species was conducted on the Tiger Creek Preserve for several rare scrub species. Level 1 monitoring looked at the distribution of populations, level 2 measured population size and condition, and level 3 measured life history traits.] No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 133 AND J. Kimmicu. 1996. Microhabitat and time since fire: effects on demography of Eryngium cuneifolium (Apiaceae), a Florida scrub endemic plant. Am. J. Bot. 83(2):185—191. [Survival, growth and fecundity of 1287 individuals over a 4-year period were studied. The results suggest that this species is dependent on an open habitat maintained by periodic fires. Below ground competition or allelopathy from shrubs probably restricts this species to recently burned open patches. ] AND K. KouHFELDT. 1995. Life history strategies of Florida scrub plants in relation to fire. Bull. Torrey Bot. Club 122:282—297. [Scrub species were classified from two types of scrub (scrubby flatwoods and rosemary scrub) into guilds based on mechanisms of post-fire recovery and post-fire abundance. Management strategies to conserve scrub species are discussed in relation to fire intervals. | AND C. V. Hawkes. 1998. Interactive effects of fire and microhabitat on plants of Florida scrub. Ecological Applications 8(4):935—946. [Studies were conducted in rosemary scrub and scrubby flatwoods to examine the response of each community and associated species to alterations in fire frequency, with special reference to seeders and resprouters. ] , P. J. McIntyre, M. S. Finer, E. Gross, AND R. YAuR. 1999. Microhabitat of the narrow Florida scrub endemic Dicerandra christmanii, with comparisons to its congener D. frustescens. J. Torrey Bot. Soc. 126(1):24-31. [Microhabitat characters of this rare scrub mint were studied along an old sand road and in adjacent undisturbed scrub at the Flamingo Villas site, a USFWS National Wildlife Preserve. Plants of this species occurred in microhabitats with significantly more open canopies, shorter shrubs, and lower litter cover and depth than those in the undisturbed scrub. ] Miprorp, P. E. 1999. Social learning and tradition in the Florida scrub-jay (Aphelocoma coerulescens). Ph.D. Dissertation, University of Wisconsin-Madison, Madison, WI. 195 pp. [To understand the operation of traditions in wild animal populations, 210 free-living Florida scrub jays in 49 families received training to perform a simple foraging task.] Miter, H. A. 1993. A new species of Dicerandra (Lamiaceae) from Florida. Phytologia 75(3):185—189. [Dicerandra thinicola, a new species, was found associated with old dune in scrubs of yellow sands. ] Minno, M. C. AND M. Minno. 1992. The natural history of Asclepias curtissii Gray (Apocynaceae), a rare scrub endemic from Florida. Unpublished report submitted to University of Florida, Gainesville, FL. [Discusses the Curtiss’ milkweed (Asclepias curtissii Asa Gray) as a state endangered species endemic to Florida and limited to the scrub community. Natural history observations were made to conclude that some factors that limit the plant’s population are habitat specificity, germination site specificity, low seed production, and herbivory. ] . 1992. Lepidoptera of the Archbold Biological Station, Highlands County, Florida. Florida Entomol. 75(3):297—329. [A total of 1,112 species of Lepidoptera is reported from the Archbold Biological Station based on literature citations, specimens in collections, and new surveys. Surveys were conducted within the sand pine scrub and other plant communities located on the station grounds. This paper serves to pool all records together, update nomenclatural changes, and provide a baseline to which new faunal studies may be compared. ] MOoHLENBROCK, R. H. 1988. Big scrub, Florida. Nat. Hist. 97(7):22—24. [Discussion of the scrub and sandhill communities in the Ocala National forest, their soils, origins of, and the plant and animal life indigenous to each.] MuLter, J. W., E. D. Harpin, D. R. Jackson, S. E. GATEwoop, AND N. Carre. 1989. Summary report on the vascular plants, animals, and plant communities endemic to Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Tech. Rep. No. 7. 113 pp. [The objectives of this report are to provide current lists of terrestrial and freshwater animals and vascular plants endemic or nearly endemic to Florida, as well as a consideration of plant communities restricted to Florida; to annotate these lists with available information on distribution and abundance for all vertebrate species, all vascular plant taxa, and all infraspecific taxa listed as endangered or threatened by the United States Fish and Wildlife Service. | Moumwe, R. L. ANnp T. H. BeLow. 1995. Relocation as a management technique for the threatened Florida scrub jay. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Project Rep. 134 FLORIDA SCIENTIST [VOL. 67 48 pp + vii. Tallahassee, FL. [This study suggests that relocation of the Florida scrub jay is a viable technique for management and restoration of this threatened subspecies. A total of 18 jays were removed from Archbold Biological Station and relocated to suitable habitat at Rookery Bay National Estuarine Research Reserve in Collier County. About 50% of the relocated birds had established territories and at least two of the four pairs nested successfully. ] , 8. J. SCHOECH, G. E. WOOLFENDEN, AND J. W. Fitzpatrick. 2000. Life and death in the fast lane: demographic consequences of road mortality in the Florida scrub jay. Conserv. Biol. 14(2):501— 512. [The survival and reproductive success of a small Florida scrub jay population along a two- lane roadway at Archbold Biological Station was studied for 9 years. High roadway mortality presents a difficult challenge for the management and conservation of this threatened and declining species. ] MUSHINSKY, H. R. AND E. D. McCoy. 1995. Vertebrate species composition of selected scrub islands on the Lake Wales Ridge of central Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Project Rep. 325pp + xiv. Tallahassee, FL. [Vertebrate species composition of 16 patches of scrub habitat of three different size categories along the Lake Wales Ridge of Central Florida were sampled. ] Myers, R. L., S. E. BOETTCHER, AND H. A. Tuck. 1987. Sand pine (Pinus clausa) seeding response following fire. ASB Bull. 34(2):68-69. Abstract. [Seed rain in a 60-year-old sand pine stand with 73% serotiny was monitored for one-year pre and post burn. ] . 1990. Scrub and high pine. Pp. 150-193. Jn: Myers, R. L. AND J. J. EWEL (eds.) Ecosystems of Florida. University of Central Florida Press, Orlando, FL. 765 pp. [An excellent overview of the sand pine scrub ecosystem in Florida.] OSTERTAG, R. AND E. S. MENGES. 1994. Patterns of reproductive effort with time since last fire in Florida scrub plants. J. Vegetation Sci. 5:303—310. [Fire frequency may be an important selective pressure for the evolution of plant reproductive allocation patterns. This hypothesis was studied using scrub plants by developing three models of reproductive effort with time since last fire.] OutTcaLt, K. W. 1986. Stand density affects growth of Choctawhatchee sand pine. South. J. Appl. For. 10:128—131. [Choctawhatchee sand pine was grown for 20 years at densities of 400, 600, and 800 trees per acre. Growing the Choctawhatchee sand pine at 400 trees per acre resulted in significantly larger trees, but less wood was produced per acre than at the higher densities. ] . 1986. Sand pine for dry sites. Forest Farmer 45:34,37. [Planting Choctawhatchee sand pine is recommended for acid sand areas in North Florida at densities of 500 tree per acre.] . 1987. The effect of seed treatment and sowing method on germination of Ocala sand pine. Pp. 81-83. Jn: Fourth Biennial Southern Silvicultural Research Conf. Proc., Atlanta, Ga. Nov. 4-6, 1986. USDA For. Ser. Gen. Tech. Rep SE-42, Southeast For. Exp. Stn., Asheville, NC. [Germination of Ocala sand pine seeds were tested at different moisture levels coupled with or without presoaking. ] . 1987. Re-establishment of sand pine: An example of how the harvest system effects regeneration. Pp. 39-42. In: Second Plantation Establishment Workshop Proc. Feb. 17-19, 1987. Georgetown, SC. [Mechanized harvesting systems offer opportunities for integrating harvest and regeneration as part of total stand management rather than requiring separate operations. The tree-length windrow system seems to be the best because it yields high quality chips, retains site nutrients and produces a site not difficult to regenerate. ] . 1988. An efficient system for harvesting sand pine biomass. Pp. 164—166. Jn: Ninth Ann. South. For. Biomass Workshop Proc. June 8-11, 1987. Biloxi, MS. [A harvesting system using feller- bunchers, grapple skidders, and an on-site chipper is discussed for sand pine. ] . 1988. Establishing Choctawhatchee sand pine using strip site preparation. South. J. Appl. For. 12:178-181. [A long-term (>10 years) study in the Georgia sandhills shows that strip site preparation is a beneficial-cost effective method in establishing sand pine on partially treated scrub hardwood sites. | . 1990. Operational trials of a scarifier-seeder for regenerating Ocala sand pine. South. J. Appl. For. 14(2):85—88. [Most harvested stands of the Ocala sand pine have been regenerated by direct seeding. An integrated system using a scarifier-seeder can reduce site disturbance, control spacing, No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 135 and conserve seed. Further research is needed to determine the most effective method for different site characteristics. ] . 1990. A method for increasing the germination of Ocala sand pine (Pinus clausa var. clausa D. B. Ward) seed in stressful environments. Pp. 57—58. In: Proc. 11th North American Forest Biology Workshop. June 13-15, 1990. Athens, GA. [Direct seeding of Ocala sand pine often fails due to high seed predation. Experiments were conducted to determine if pre-soaking, or the use of moisture retaining gels, and/or planting depth would increase germination in the field. ] . 1991. Aerated stratification improves germination of Ocala sand pine seed. Tree Planters’ Notes, winter 22—26. [A stratification system of soaking in water with and without aeration followed by chilling was tested for its effect on the germination of Ocala sand pine seed.] . 1991. Effect of pesticides and number of seed per spot on seedling establishment from direct- sown Ocala sand pine seed. Jn: Sixth Biennial Southern Silvicultural Research Conf. Proc., Memphis, TN, October 30-November 1, 1990. Gen. Tech. Rep. SE-70 Vol. 1, Pp. 47-51. Southeastern For. Exp. Stn., Asheville, N.C. [Seed treatment with pesticides and the number of seeds per spot were studied to determine if treatment reduces predation. The results suggest that seed predation if sown at the proper time of year is not severe enough to justify repellants and five well-placed seeds per spot were adequate to obtain a well-stocked stand.] . 1992. Effect of cold storage on germination of stratified Ocala sand pine seed. Jn: Proc. 12th North American Forest Biology Workshop. August 17—20. 1992. Sault Ste, Marie, Ontario, Can. Pp. 182. [The objectives of this study were to determine if stratified seed retained their attribute of improved germination under stressful conditions following redrying and differing periods of cold storage. | . 1993. Southern pines performance on sandhills sites in Georgia and South Carolina. Southern J. Appl. For. 17(2):100-102. [Various pines were grown on sandhills to determine yield. Choctawhatchee sand pine grew the fastest and yielded the most volume after 28 years. Longleaf pine was also a viable species on the sandhill sites, especially with longer rotations. ] . 1997. An old-growth definition for sand pine forest. USDA For. Serv. Gen. Tech. Rep. SRS-12. Asheville, NC. 8pp. [Selected stands of old sand pine were sampled on the Ocala National Forest and near Niceville, FL, to determine the characteristics of old growth. Old growth stands had an overstory of sand pine with some trees at least 14.0 inches in diameter and more than 55 years old. ] AND C. A. GREENBERG. 1998. A stand replacement prescribed burn in sand pine scrub. Proc. of 20th Tall Timbers Fire Ecology Conf. on July 23, 1997. Pp. 141-145. [This paper describes fire characteristics and the immediate effects of a prescribed, high-intensity burn on a 12.2-hectare portion of a stand of Ocala sand pine scrub. ] PACKER, W. C. AND J. N. Layne. 1990. Foraging site preferences and relative arboreality of small rodents in Florida. Am. Midl. Natl. 125:187—194. [Foraging microhabitat preferences of Ochrotomys nuttalli, Peromyscus gossypinus, P. polionotus, Podomys floridanus and Sigmodon hispidus in sandhill and sand pine scrub vegetation associations in south-central Florida were studied by placing bait containing different-colored plastic markers on the ground in structurally open and closed sites and in trees and recovering the markers from feces of live-trapped individuals. | PARKER, A. J., K. C. PARKER, AND H. W. Brown. 2000. Disturbance and scale effects on southern old growth forests (USA): The Sand Pine example. Nat. Areas J. 20(3):273—-279. [Sand pine often is a monotypic dominant in the Florida scrub assemblage and does not conform to the traditional image of old growth forest. This research questions the conceptual basis and management perspective implicit in developing a stand-scale definition of old-growth sand pine. |] PARKER, K. C. AND J. L. Hamrick. 1996. Genetic Variation in sand pine (Pinus clausa). Can. J. For. Res. 26:244—254. [The genetic diversity of sand pine was surveyed for both varieties from 21 different locations throughout Florida. ] , A. J. PARKER, R. M. Beaty, M. M. FUuLLer, AND T. D. Faust. 1997. Population structure and spatial patterns of two coastal populations of Ocala sand pine (Pinus clausa (Chapm. ex Engelm) Vasey ex Sarg. var. clausa D. B. Ward). J. Torrey Bot. Soc. 124(1):22-33. [Two coastal populations of sand pine at Jonathan Dickinson State Park were compared for effects of disturbances occurring at different scales on spatial dispersion and age structure. ] 136 | FLORIDA SCIENTIST [VOL. 67 PLatTT, B. 1992. Development of scrub ecosystems in Florida. Unpublished report submitted to Louisiana State University, Baton Rouge, LA. [Study to determine whether or not persistence of sand pine in peninsula stands is likely to depend on lightning-initiated fires originating in adjacent savannas, and burning into scrub during drought years. ] Potts, W., D. McDoNnaLp, J. W. FitzPATRICK, AND G. E. WOOLFENDEN. 1992. Genetic population structure of Florida scrub jays across the peninsula: preliminary findings. University of Florida College of Medicine; Archbold Biological Station; University of South Florida. [These preliminary data show differences in allelic frequencies between comparisons of both distant and closely spaced populations by use of molecular genetic techniques called microsattelite loci.] PROVENCHER, L., B. J. HERRING, D. R. Gorpon, H. L. RopGers, G. W. TANNER, L. A. BRENNAN, AND J. L. HARpesty. Restoration of northwest florida sandhills through harvest of invasive Pinus clausa. 2000. Restor. Ecol. 8(2):175—185. [Much of the longleaf pine sandhills of Elgin Air Force Base in the Florida panhandle have been invaded by sand pine (Pinus clausa) following fire suppression and logging. Silvicultural removal of sand pine with minimal soil disturbance followed by fire and longleaf pine (Pinus palustris) planting appears to be a promising and cost-effective restoration approach to P. clausa invaded sandhills. ] Putz, F. E. 1991. Microclimate, plant community structure and fruit production on edges of clearcut sand pine scrubs in Ocala National Forest. Pp. 1—2. [Sand pine scrub which currently is an endangered community containing over 40 endemic species has the largest existence in Ocala National Forest, which at present is being intensively managed primarily for sand pine (Pinus clausa) pulpwood production, and less concerned with wildlife. The tremendous amounts of clear-cutting is resulting in an extreme increase in the forest’s edge/interior ratio, the effects of which are of great concern. | AND M. MInNNo. 1995. The pollination biology and ecology of Curtiss’ milkweed (Asclepias curtissii). Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Project Rep. 121 pp. + x. Tallahassee, FL. [This project was conducted to determine the nature and causes of rarity for the Curtiss milkweed, a scrub endemic in Florida. ] Quinn, J. S., G. E. Woo.LFENpoN, J. W. Fitzpatrick, AND B. N. White. 1999. Multi-locus DNA fingerprinting supports genetic monogamy in Florida scrub-jays. Behav. Ecol. Sociobiol. 45:1—10. [Extensive behavioral and pedigree data for a Florida scrub jay population at Archbold Biological Station suggested that this cooperatively breeding species is monogamous, with extremely rare exceptions in which males have two mates. | QUINTANA-ASCENCIO, P. F. AND E. S. MENGES. 1996. Inferring metapopulation dynamics from patch-level incidence of Florida scrub plants. Conserv. Biol. 10(4):1210—1219. [A metapopulation model was used to analyze the distribution of extinction rates and rate of migration among patches in the rosemary scrub. The results suggest stronger effects of patch size and patch isolation on extinction probabilities of herbs, compared with woody species] AND E. S. MENGES. 2000. Competitive abilities of three narrowly endemic plant species in experimental neighborhoods along a fire gradient. Am. J. Bot. 87(4):690—699. [Field experiments involving three endemic scrub species were studied to assess seed germination, survival, biomass and fecundity along a time since fire gradient. The results indicate that fire is a useful tool in controlling the competitive effects of large shrubs and lichens, especially on the three species studied in the Florida scrub. ] AND M. Mora.es-HERNANDEZ. 1997. Fire-mediated effects of shrubs, lichens and herbs on the demography of Hypericum cumulicola in patchy Florida scrub. Oecologia 112:263—271. [Hypericum cumulicola is a narrowly adapted scrub endemic restricted to open areas of well- drained white sand in the Florida rosemary scrub. Survival, growth and fecundity among 1214 individuals in 14 rosemary scrub patches were studied under different fire histories. H. cumulicola and other herbaceous species depend on sporadic fires to decrease interference of shrubs and ground lichens. Understanding the role of fire may be an important factor in ecosystem manage- ment of endangered species in the Florida scrub.] , R. W. DoLaN, AND E. S. MENGES. 1998. Hypericum cumulicola demography in unoccupied and occupied Florida scrub patches with different time since fire. J. Ecology 86:640—651. [The patchy pattern of site occupancy by Hypericum cumulicola is probably due to limited dispersal and No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 137 periodic extinction, especially associated with long fire-free intervals. Management practices should include measures to protect unoccupied patches for long-term survival. | Rep, G. K. 1991. The gopher tortoise-landlord of the sandhills. Florida Nat. Spring. Pp. 3-5. [Briefly describes a generalized account of the gopher tortoise, its occurrence in sandhill and scrub vegetation, breeding habits, social behavior, diet, burrows and concerns on their preservation. ] RICHARDSON, D. R. 1990. The sand pine scrub community: General aspects. Unpublished report submitted to Rookery Bay National Estuarine Research Preserve, Naples, FL. 10 pp. [Stresses the importance of considerable research to determine sound management methodology and goals for preserving remaining fragmented scrubs. ] . 1989. The sand pine scrub community: an annotated bibliography. Florida Scient. 52(2):65—93. [Detail listing of 316 references related to the Florida sand pine scrub community. ] , 1. J. Stout, R. E. Roperts, D. F. AUSTIN, AND T. R. ALEXANDER. 1986. Design and management recommendations for a sand pine scrub preserve: the Yamato scrub. Unpublished report submitted to City of Boca Raton, Boca Raton, FL. 142pp. [This report discussed the fate of the ““Yamato Scrub” which lies along the eastern edge of the Atlantic Coastal Ridge in Palm Beach County. The uncertain fate of the scrub is stressed, as a portion of the 163 acres of sand pine scrub has been slated for development as a multiuse commerce park. ] Roserts, R. E. AND A. C. Cox. 2000. Sand pine scrub vegetation response to two burning and two non- burning treatments. Pp. 114-124. In: Moser, W. K. AND C. F. Moser (eds.) Fire and forest ecology: innovative siliviculture and vegetation management. Tall Timbers Fire Ecology Conf. Proc. No. 21, Tall Timbers Research Station, Tallahassee, FL. [Florida’s sand pine scrub association tends to burn with catastrophic crown fires of high intensity under sometimes extreme weather conditions, thus exhibiting uncontrollable and unpredictable behavior. To provide a better understanding of prescription and non-burning techniques for various size parcels of scrub, prescribed fire methods combined with several mechanical alteration treatments were applied to numerous plots of sand pine at Jonathan Dickinson State Park, Hobe Sound, FL.] AND I. J. Stour. 1992. Response of Florida mice to prescribed burning at Jonathan Dickinson State Park. Division of Sponsored Research, University of Central Florida, Orlando, FL. [Report of the demography and behavior of a Florida mouse (Podomys floridanus) population inhabiting sand pine scrub burned in 1982 at Jonathan Dickinson State Park, Hobe Sound, FL.] Rockwoop, D. L., B. YANG, AND K. W. OuTcaLt. 1997. Stand yield prediction for managed Ocala sand pine. USDA For. Serv., Res. Pap. SRS-3, Asheville, NC. 16 pp. [The purpose of this study was to develop the site index and stand-level growth and yield equations for managers to make informed decisions by tree component. Actual yields were compared to predicted yields by sampling 22 sand pine plantations. ] , K. W. Reppy, C. W. Comer, W. H. McNaps, AND K. W. OurTcALt. 1987. Weight and volume prediction equations for sand pine trees in Florida. Fla. Ag. Exp. Stn. Tech. Bull. 869. 16 pp. University of Florida, Gainesville, FL. [An intensive sampling of sand pine stands was initiated in 1982 to determine tree weight and volume prediction equations for both varieties of sand pine. Selected trees were harvested in various age stands and from different densities. ] SANGER, M. B. 1991. The big scrub. Young Naturalists 64(1):1—4. [Details the origin of the Ocala National Forest from the ice age approximately 10,000 years ago to present, and stresses the unique balance of the scrub forest and its importance to the endemic species that thrive there, and whose survival is linked to the forest. ] ScHaAus, R., R. L. MummMe_, AND G. E. WoOLFENDEN. 1992. Predation on the eggs and nestlings of Florida Scrub Jays. Auk 109(3):585-593. [Nest predation was investigated in a population of Florida scrub jays (Aphelocoma c. coerulescens) at Archbold Biological Station, Lake Placid, FL.] SCHMALZER, P. A. AND C. R. HINKLE. 1985. A brief overview of plant communities and the status of selected plant species at John F. Kennedy Space Center, FL. Unpublished report, Kennedy Space Center, FL. [Brief description of various plant communities including scrub is provided with discussion of various species. ] AND C. R. HINKLE. 1990. Flora and threatened and endangered plants of John F. Kennedy Space Center, Florida. NASA Tech. Memo. 102791. Kennedy Space Center, FL. 68 pp. [The vascular 138 FLORIDA SCIENTIST [VOL. 67 flora of the Kennedy Space Center is provided with special reference to endemic and listed plant species. ] AND C. R. HINKLE. 1990. Geology, geohydrology and soils of Kennedy Space Center: a review. NASA Tech. Memo. 100313. Kennedy Space Center, FL. 46 pp. [Discusses the geology and soils of the region, many of which support scrub vegetation. | AND C. R. HINKLE. 1991. Dynamics of vegetation and soils of oak/saw palmetto scrub after fire: Observations from permanent transects. NASA Tech. Memo. 103817. 149 pp. [Two burned and two unburned areas within an oak/saw palmetto scrub on the Kennedy Space Center were sampled to determine structural features of the vegetation 5 years post-burn. ] AND C. R. HINKLE. 1992. Recovery of oak-saw palmetto scrub after fire. Castanea 57(3):158—173. [Previously established permanent line-intercept transects (15m in length) in oak-saw palmetto scrub on Merritt Island were sampled at 6, 12, 18, 24, 36, 48, and 60 months after a 1986 fire to determine patterns of recovery.] AND C. R. HINKLE. 1992. Species composition and structure of oak-saw palmetto scrub vegetation. Castanea 57(4):220—251. [In 1983, four stands of oak-saw palmetto scrub vegetation were sampled using permanent line transects that were 2, 4, 8, and 25 years since the previous fire. These transects were re-sampled two years later to determine post fire recovery of the dominant shrubs and shifts in the vegetation that might have occurred. Soils were sampled and analyzed from the 0— 15cm and 15—30cm layer at each transect.] AND C. R. HINKLE. 1996. Biomass and nutrients in aboveground vegetation and soils of Florida oak-saw palmetto scrub. Castanea 61(2):168—193. [Four stands of oak-saw palmetto scrub vegetation were sampled at 2, 4, 8, and 25-year intervals since the previous fire by harvesting 1- meter plots. Retention of nutrients in soils and re-growing vegetation after fire may be important to the persistence of scrub on low nutrient soils. ] , D. R. BREININGER, F. W. ADRIAN, R. SCHAUB, AND B. W. Duncan. 1994. Development and implementation of a scrub habitat compensation plan for Kennedy Space Center. NASA Tech. Memo. 109202. Kennedy Space Center, FL. 55 pp. [Kennedy Space Center, located on Merritt Island on the east coast of central Florida, is one of the three remaining major populations of the Florida scrub jay (Aphelocoma coerulescens coerulescens), listed as threatened by the U.S. Fish and Wildlife Service since 1987. Over the next five years construction of new facilities by NASA has the potential to impact up to 193 ac (78.1 ha) of Florida scrub jay habitat. NASA has agreed to a compensation plan for loss of Florida scrub jay habitat, and that plan is discussed. ] AND S. R. BoyLe. 1997. Restoring long-unburned Florida oak-saw palmetto scrub using mechanical cutting and prescribed burning. Abstr. Soc. Restor. Ecology 9™ Annual Intern. Conf., Fort Lauderdale, FL. P. 108. [Mechanical alteration techniques can be used to re-establish shrub structure, which can then be maintained by periodic burning practices. ] , 5. R. BoYLe, AND H. M. Swain. 1999. Scrub ecosystems of Brevard County, Florida: A regional characterization. Florida Scient. 62(1):13-47. [The geology of the scrub ridges in Brevard County is discussed in reference to the distribution of the vegetation, soils, rare or listed species, and conservation measures. | , B. W. Duncan, V. L. Larson, S. BOYLE, AND M. GimMonpb. 1996. Reconstruction of historic landscapes of the Indian River Lagoon basin. Pp. 849-854. In: Pro. Eco-Informa’ 96: Global Networks for Environmental Information, Vol. 11. Environmental Research Institute of Michigan, Ann Arbor, Michigan. [Historic changes in landscape composition were documented for the Florida scrub and other community types by comparing historic aerials and other information. ] ScHMutTz, D. D. 1997. Translocation and microhabitat distribution of Podomys floridanus on native upland and reclaimed mined sites. Masters thesis, University of South Florida, Tampa, FL. 162 pp. [Florida mice were captured from scrubby flatwoods donor sites and relocated into both natural and phosphate-mined reclaimed xeric oak scrub recipient sites at Noralyn Mine sites, Polk County Jal SCHOECH, S. J. 1999. Florida scrub-jay nestlings preyed upon by eastern coachwhip. Florida Field Nat. 27(2):57—58. [Considerable indirect evidence implicates diurnal snakes as major predators on eggs No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 139 and nestlings of the Florida scrub jay. This paper describes predation of two offspring by eastern coachwhip. ] SmiTH, R. B., D. R. BREININGER, AND V. L. LARSON. 1997. Home range characteristics of radio tagged gopher tortoises on Kennedy Space Center, Florida. Chelo. Conserv. Biol. 2(3):358—362. [Data from 14 radio tagged gopher tortoises on the Kennedy Space Center were analyzed to determine home range sizes, the number of burrows used by each tortoise, and the preferred use of habitats within the home ranges. Oak scrub and oak palmetto scrub were compared to 5 other vegetation types. ] ~StirH, B. M. 1999. Metapopulation dynamics and landscape ecology of the Florida scrub-jay, Aphelocoma coerulescens. Ph.D. Dissertation, University of Florida, Gainesville, FL. [The Florida scrub jay is rapidly disappearing throughout much of its range. To characterize the spatial structure and vulnerability of this species, GIS was used to describe the species metatpopulation structure. ] StouT, I. J. 1992. Long-term studies of east-central Florida scrub: structure and dynamics. University of Central Florida, Orlando, FL. [Sand pine scrubs of central Florida were typically fire-derived and exhibited a high similarity among stands in the tree (76%), shrub (62%), and ground layers (40%). Typical stand metrics were provided. ] . 2001. Rare plants of the Florida scrub, USA. Nat. Areas J. 21(1):50—60: [Thirty-eight plants found in the Florida scrub have been recognized as endangered or threatened with extinction under the Endangered Species Act or comparable state regulations. Life history strategies are discussed for species found in different regions of the state. ] AND W. R. Marion. 1993. Pine flatwoods and xeric pine forests of the southern (lower) coastal plain. Pp. 373-446. Jn: Biodiversity of the Southeastern United States/Lowland Terrestrial Communities. John Wiley & Sons, Inc. [A review of the pine communities of the southeastern coastal plain is discussed with emphasis on their classification, structural attributes, dynamics and management. | , T. J. DOONAN, R. E. RoBerTs, AND D. R. RICHARDSON. 1987. Comparison of results of three gopher tortoise relocations in central and southeast Florida. Pp. 15-42. Jn: Diemer, J. E. et al. (eds.) Proc. Gopher Tortoise Relocation Symp. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Tech. Rept. #5, Tallahassee, FL. [This paper presents the results of gopher tortoise relocation efforts at three sites in central and southeast Florida. Methods, season of release, and conditions of release were different in each case; however, radio tracking was used in each study to evaluate the relocation strategy. ] , D. R. RICHARDSON, AND R. E. RoBerts. 1987. Response of resident and relocated gopher tortoises to a prescribed burn in a sand pine scrub community. Pp. 84-85. /n: Diemer, J. E. et al. (eds.) Proc. Gopher Tortoise Relocation Symposium. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Tech. Rept. #5, Tallahassee, FL. [This paper reports the response of tortoise to a prescribed burn of a 4 ha recipient site in Boca Raton, FL.] , D. R. RICHARDSON, AND R. E. RoBerts. 1988. Management of amphibians, reptiles, and small mammals in xeric pinelands of peninsular Florida. Pp. 98-108. In: SzARo, R. C., K. E. SEVERSON, AND D. R. PATTON (eds.), Management of Amphibians, Reptiles, and small mammals in North America, Proc. of the Symp., USDA Forest Service, General Technical Report RM-166, Flagstaff, AR. [Pineland habitats preserve design, and management strategies are discussed in relation to various vertebrate assemblages in Sandhill and Sand Pine Scrub communities in Florida. ] , D. R. RicHarpson, R. E. RoBerts, AND D. F. Austin. 1987. Design of a nature preserve in a subtropical, urbanizing landscape: application of ecologic and genetic principles. Bull. Ecol. Soc. Am. 68(3):423-424. [Design standards are discussed to maintain a sand pine scrub preserve in the City of Boca Raton, FL.] THE Stuart News/Port Sr. Lucite News. 1992. Florida’s ancient archipelago: scrublands drowning in progress. Newspaper Article. pp. S1—S8. [Details some of the species endemic to the Florida scrub, its inhabitants, the need for long-term survival of the scrub community, and its past and uncertain future. | TANRISEVER, N., N. H. FiscHer, AND G. B. WILLIAMSON. 1987. Menthofurans from Calamintha ashei: effects on Schizachyrium scoparium and Lactuca sativa. Phytochemistry 627(8):2523—2526. 140 FLORIDA SCIENTIST [VOL. 67 [From the aerial parts of Calamintha ashei the new menthofuran, calaminthone, as well as the known terpenoids(+)-evodone, caryophyllene oxide and ursolic acid were isolated and identified by spectroscopic methods (NMR and MS). The possible role of ursolic acid in the release and transport of allelopathic lipids from the source plant into the soil through rain leachates is discussed. | THAXTON, J. E. AND T. M. HINGTGEN. 1992. Effects of habitat fragmentation and urbanization on Florida scrub jay dispersals in Sarasota County. Unpublished report submitted to Department of Recreation and Parks, District 8 Office, Tallahassee, FL. [Dispersal behavior of Florida scrub jays (Aphelocoma c. coerulescens) was compared for birds with territories in fragmented, urbanized habitats designated “‘suburban” and birds with territories in non-fragmented scrub habitat designated “‘preserve’’.] U.S.D.A. Forest Service. 1981. Southeast sandhills can be productive. Management alternatives for pines (slash, longleaf, and sand). SE Forest Experiment Station, Olustee, FL. 5 pp. [Choctawhatchee sand pine is very suitable to sandhill droughty soils and should add substantially to the wood production capacity of the south.] U.S. FISH AND WILDLIFE SERVICE. 1989. Recovery plan for eleven central Florida scrub plant species. U. S. Fish and Wildlife Service, Atlanta, GA. 64 pp. [The eleven plant species covered by this plan occur primarily (or entirely) in Florida scrub vegetation dominated by sand pine and evergreen oaks. In the counties where these plants are concentrated, scrub occurs primarily in small tracts on a series of hills called the Lake Wales Ridge. Recovery objectives are discussed for the eleven species. | . 1987. Technical draft recovery plan for Florida golden aster (Chrysopsis floridana). U.S. Fish and Wildlife Service, Atlanta, GA. 12 pp. [This report discusses the threat to the Florida Golden Aster (Chrysopsis floridana), its systematics and identification, reproductive and population biology, habitat and general distribution and population status. ] . 1986. Recovery plan for three Florida mints: Longspurred mint (Dicerandra cornutissima, Scrub mint (Dicerandra frutescens), and Lakela’s mint (Dicerandra immaculata). U.S. Fish and Wildlife Service, Atlanta, GA. 20 pp. [A recovery plan to protect existing populations of the mints which may require short-term emergency measures, and will require habitat protection and management, including prescribed burning or mechanical disturbance to prevent successional changes. ] . 1992. Notice of availability of the draft environmental assessment and land protection plan; proposed establishment of Lake Wales Ridge National Wildlife Refuge; Highlands and Polk Counties, Florida. Fed. Reg. 57(177):41770. [This notice advises the public that the U.S. Fish and Wildlife Service, Southeast Region, proposes to establish a national wildlife refuge in the vicinity of Highlands and Polk Counties, FL. The purpose of the proposed refuge is to protect and manage up to 12,200 acres of rare Florida scrub habitats and their associated endangered and threatened plants and animals along the Lake Wales Ridge in central Florida. ] . 1992. Endangered and threatened wildlife and plants; proposed endangered or threatened status for seven central Florida plants. Fed. Reg. 57(190):45020-45028. [The Service proposes endangered status pursuant to the Endangered Species Act of 1973 as amended (Act) for the following five plants: Cladonia perforata (Florida perforate cladonia), Crotalaria avonensis (Avon Park harebells), Nolina brittoniana (Britton’s beargrass), Polygala lewtonii (Lewton’s polygala), and Polygonella myriophylla (sandlace). The Service proposes threatened status for two plants: Clitoria fragrans (pigeon wings) and Erigonum longifolium ver. gnaphalifolium (scrub buckwheat). All seven plants are found in Highlands and Polk counties in central Florida (scrub areas). | U.S. DEPARTMENT OF THE INTERIOR. 1996. Jupiter Inlet, final coordinated resource management plan. U.S.D.I., Bureau of Land Management, Jackson District, Jackson, MS. 58 pp. plus appendices. [A resource management plan is provided for the 86 acre Jupiter Inlet tract. The plan outlines the management of key resources, including wildlife and vegetation, historic and cultural resources, and recreation and environmental education. | Watts, W. A. 1992. The vegetation history of south-central Florida from 50,000 yr. Bp to the present: evidence from Lake Annie and Tulane. Unpublished report submitted to Trinity College, Dublin, No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 141 Ireland. [Studies were conducted of the vegetation history of Highlands County by pollen analysis of sediments from Lake Annie at the Archbold Station and Lake Tulane at Avon Park, and information on the macrofossil studies. | AND B. C. S. HANSEN. 1994. Pre-holocene and holocene pollen records of vegetation history from the Florida peninsula and their climatic implications. Palaeogeog., Palaeoclim., Palaeoecol. 109:163—176. [Pre-holocene sediments dating back to about 50,000 years B.P. have been shown to contain pine, oak and ragweed pollen from lakes in central Florida. | Wess, S. D. 1992. Vertebrate paleontology and the origin of Florida scrub. Unpublished report submitted to University of Florida, Gainesville, FL. [The earliest indirect evidence of open savanna-like habitats on deep sandy soils in Florida comes from the Thomas Farm. The abundant terrestrial species such as gopher tortoise (Geochelone), grazing horses (Parahippus), and granivorous rodents (Proheteromys) suggest a proto-sandhill biota, maintained by periodic lightning fires. ] WEIDENHAMER, J. D. 1987. Allelopathic properties of Polygonella myriophylla. Ph.D. Dissertation, Univ. of South Florida, Tampa, FL. [The chemical properties of Polygonella myriophylla were surveyed in the Florida scrub community with special reference to its allelopathic potential. ] ——. 1994. Distinguishing resource competition and chemical interference: overcoming the methodological impasse. Agron. J. 88:866—875. [Understanding allelopathy may hold the key to new weed management strategies. The difficulty of distinguishing chemical interference from competition has hindered studies of allelopathy in natural and cultivated plant communities. This study consists of testing the feasibility of analytical methods which measure allelochemical flux rates rather than static concentrations. ] , D. C. Hartnett, AND J. T. Romeo. 1989. Density-dependent phytotoxicity: distinguishing resource competition and allelopathic interference in plants. J. Appl. Ecol. 26:613-624. [This paper discusses the experimental procedures done to investigate the potential interacting influences of allelopathy and resource competition on plant response and yield-density relationships. ] AND J. T. RoMEo. 1989. Allelopathic properties of Polygonella myriophylla: field evidence and bioassays. J. Chem. Ecol. 15(7):1957—1970. [Polygonella myriophylla is a perennial shrub endemic to the Florida scrub. Striking bare zones surround mature Polygonella stands. This supports the hypothesis that the bare zones result from chemical interference by Polygonella with the growth of other species. ] , M. MeneLAou, F. A. Macias, N. H. FISCHER, D. R. RICHARDSON, AND G. B. WILLIAMSON. 1994. Allelopathic potential of menthofuran monoterpenes from Calamintha ashei. J. Chem. Ecol. 20(12):3345-3359. [Calamintha ashei is one of several perennial shrubs of the Florida scrub community for which evidence of allelopathic effects on pines and grasses of the sandhills is accumulating. A reversed-phase HPLC analysis was used to separate and quantify five menthofuran monoterpenes in Calamintha ashei leaf soaks and washes. Aqueous solubilities of the menthofurans were determined to exceed concentrations required for growth inhibition. ] AND N. H. FiscHer. 1989. Allelopathic suppression of bahiagrass, Paspalum notatum Fluegge, by the perennial shrub, Polygonella myriophylla. Weed Science Society Abstracts 29:60-61. [Polygonella myriophylla is a perennial shrub endemic to the Florida scrub. Striking bare zones surround mature Polygonella stands. Where areas of scrub border abandoned fields dominated by bahiagrass, sharp boundaries between the scrub and bahiagrass often are evident. Quantitative measurements of root distribution show that few Polygonella roots extend into the bare zones, supporting the hypothesis that the bare zones result from chemical interference by Polygonella with the growth of bahiagrass and other species. | WILLIAMSON, G. B. 1992. Mechanisms of allelopathy in the southeastern coastal plain. Unpublished report submitted to U.S. Department of Agricultural Science and Education. 57pp. [Research is focused on plant interactions in the pine forests of the Southeastern Coastal Plain of the USA, where we have hypothesized that fire-sensitive shrubs of the sand pine (Pinus clausa) scrub community release allelotoxins which inhibit germination and growth of fire-facilitating (sensu Mutch 1970) graminoids (Schizachyrium and Aristida spp.) and pines (Pinus palustris and P. elliottii) of the sandhills. | , N. H. FiscHer, D. R. RICHARDSON, AND A. DELA PENA. 1989. Chemical Inhibition of fire-prone grasses by fire-sensitive shrub, Conradina canescens. J. Chem. Ecol. 15(5):1567—1577. [In an 142 FLORIDA SCIENTIST [VOL. 67 investigation of potential chemical activity of fire-sensitive shrubs in Florida’s sand pine community, bioassays of foliar washes of Conradina canescens showed significant inhibitory activity on three native grasses that are known to fuel frequent surface fires; inhibition was concentrated seasonally in spring and summer. ] AND N. H. FIscHer. 1992. Mechanisms of allelopathy in forest ecosystems. Unpublished report submitted to School of Forestry and Wildlife Resources, Louisiana State University, Baton Rouge, LA. 24 pp. [Details the study of investigating mechanisms of allelopathy-plant production and release of chemicals that inhibit or stimulate growth and development of other plants in forest ecosystems. Research is focused on plant interactions in the pine forests of the Southeastern Coastal Plain of the USA, where its hypothesized that fire-sensitive shrubs of the sand pine (Pinus clausa) scrub community release allelotoxins which inhibit germination and growth of fire-facilitating graminoids (Schizachyrium and Aristida spp.) and pines (Pinus palustris and P. elliottii) of the sandhills. ] , D. R. RICHARDSON, AND N. H. FISCHER. 1992. Allelopathic mechanisms in fire prone communities. Pp. 58-75. In: Rizvi, S. J. H. AND V. Rizvi (eds.) Allelopathy: Basic and applied aspects. Chapman & Hall, London. [Evidence from bioassays with leaf washes suggests strongly that shrubs from the Florida scrub community produce allelochemicals that inhibit the germination and radicle growth of grass species native to the Florida sandhill community. A field transplant experiment, designed to control for alternative competitive mechanisms, implicated allelopathy as the mechanism of interaction. | , E. M. OBEE, AND J. D. WEIDENHAMER. 1992. Inhibition of Schizachyrium scoparium (Poaceae) by the allelochemical hydrocinnamic acid. J. Chem. Ecol. 18:2095—2105. [Bare zones around shrubs in the Florida scrub indicate the possibility of allelopathy by shrubs in controlling the distribution of grasses and other forbs. Hydrocinnamic acid is a breakdown product of ceratiolin from the shrub Ceratiola ericoides. This study indicates that hydrocinnamic acid was shown to have a strong inhibitory effect on shoot and root biomass of the grass Schizachyrium in greenhouse studies and that reduced nutrient levels caused greater inhibition by HCA under low N and P solutions. This may be important in the scrub where levels of N and P are known to be low.] WOOLFENDEN, G. E. 1975. Florida scrub jay helpers at the nest. Auk 92(1):1—15. [Discusses the Florida scrub jays as a close knit society with helpers at the nest to assist in rearing of the offspring, also characterizing them as territorial, permanently monogamous, and single brooded with a short, highly synchronized breeding season. |] AND J. W. Fitzpatrick. 1991. Florida scrub jay ecology and conservation. Pp. 542—565. In: PERRINE, C. M., J. D. LEBRETON, AND G. J. M. Hirons, (eds.) Bird Population Studies, Relevance to Conservation and Management. Oxford Univ. Press, Oxford. 683 pp. [Summarizes aspects of life history, habitat, and demography that are pertinent to protection and management of a species that now exists only in small, isolated populations. ] AND J. W. Fitzpatrick. 1990. Florida scrub jays: a synopsis after 18 years of study. Pp. 241-266. In: STACEY, P. B. AND W. D. Koenic (eds.) Cooperative Breeding In Birds. Cambridge Univ. Press, 1990. [Differentiates between the Florida populations of Florida scrub jays with that of its counterparts in the west. ] AND M. C. Garvin. 2000. Florida scrub-jay disease analysis. Florida Fish and Wildl. Conserv. Comm. Final Rep., E-1 [I-1-6. Tallahassee, FL. 14pp + 11. [The Florida scrub jay is declining in numbers and distribution in Florida due to loss of scrub habitat. Little is known about the frequency and impact of diseases in wild populations for most avian species. This study hopes to determine the relative prevalence and intensity of pathogens present in the jay population at Archbold Biological Station in central Florida. ] WUNDERLIN, R. P. 1988. Endangered and threatened plants of central Florida. Unpublished report submitted to The Bok Tower Garden Foundation. 12 pp. [This paper gives a brief history of the Lake Wales Ridge, with mention to its being one of the best and possibly oldest examples of a scrub community. Discussed are some of the endemics to Florida, and some of the endangered or threatened taxa found in Florida.] No. 2 2004] RICHARDSON—SAND PINE SCRUB COMMUNITY 143 , D. LINDSEY, AND S. UpcHurcu. 1982. Vegetational survey of the Avon Park Air Force bombing range. Unpublished report submitted to MacDill Air Force Base, Tampa, FL. 49 pp. [The purpose of this study is to provide for the APBR a description and a map of the major plant communities (including sand pine scrub), a comprehensive list of plant species, and documentation on all plant species listed in Volume 5 of the ““Rare and Endangered Biota of Florida’’.] Younc, C. C. AND E. S. MENGES. 1999. Postfire gap-phase regeneration in scrubby flatwoods on the Lake Wales Ridge. Florida Scient. 62(1):1—12. [This paper contrasts scrubby flatwoods with rosemary scrub and suggests that periodic fires in Florida scrub communities facilitates the gap-phase regeneration of scrub species. ] ZoNA, S. AND W. S. Jupp. 1986. Sabal etonia (Palmae): systematics, distribution, ecology, and comparisons to other Florida scrub endemics. SIDA 11(4):417-427. [A taxonomic study of Sabal etonia Swingle ex Nash and related taxa (involving field work throughout Florida along with the study of more than 800 herbarium specimens) has shown that this palm is morphologically and ecologically distinctive (endemic to sand pine scrub) and most closely related to S. palmetto.] Florida Scient. 67(2): 118-143. 2004 Accepted: September 9, 2003 Biological Sciences EFFECTS OF SOCIAL ENVIRONMENT IN EARLY LIFE ON CORTICAL DEPTH, LOCOMOTOR ACTIVITY, AND SPATIAL LEARNING IN THE GOLDEN MOUSE, OCHROTOMYS NUTTALLI FRED PUNZO Department of Biology, University of Tampa, 401 W. Kennedy Blvd., Tampa, FL 33606 ABSTRACT: Studies were conducted to assess the effect of group living early in life on brain development (cortical depth), locomotor activity, and spatial learning in males of Ochrotomys nuttalli, as compared to animals (single family groups, SFG) whose mothers had litters comprised of only one offspring and who only had contact with their maternal parent. Animals reared with related and unrelated adults and young (multiple family groups, MFG) were born in nests containing from 6-8 individuals and were thus exposed to a more complex early rearing environment with an increased variety of olfactory, visual, and tactile cues. There was a significant effect of environmental complexity on body weight, brain weight, and cortical thickness (at 3 months of age) between the single and multiple family groups. Subjects from the MFG exhibited a significantly higher level of running wheel activity as compared to SFG subjects. In addition, MFG animals showed a significantly improved level of performance in spatial learning ability in an 8-arm radial maze. Key Words: Brain, locomotor activity, Ochrotomys nuttalli, social interactions, spatial learning THE effects of environmental complexity (EC) occurring early in life on de- velopment of the central nervous system (CNS) and subsequent behavior has been studied extensively in inbred laboratory strains of mice and rats (see review by Rosenzweig and Bennett, 1996). In recent years, many zoos and aquariums have been including environmental enrichment strategies as part of their husbandry programs in view of the beneficial effects of EC to captive mammals including a reduction in aggression (Van Loo et al., 2002) and stereotypical behaviors (Poole, 1992), enhanced exploratory behavior (Poucet et al., 1986) and learning abilities (Nilsson et al., 1999; Van Praag et al., 1999; Punzo, 2003), as well as an increase in their behavioral repetoire (see review by Shepherdson et al. 1998). Factors associated with EC include physical features of the immediate environment (Newberry, 1995), rearing conditions (Harvey et al., 2002; Punzo and Alvarez, 2002), and degree of contact between offspring and their parents and/or siblings (Mellen 1992; Punzo and Ludwig, 2002). In addition to behavior, EC also has profound effects on the CNS including enhanced neurogenesis and synpatogenesis (Nilsson et al., 1999), and levels of neurotransmitter synthesis (Bennett et al., 1964; Buonomano and Merzenich, 1998). 144 No. 2 2004] PUNZO—GOLDEN MOUSE STUDIES 145 In contrast to the extensive literature that exists for laboratory animals, relatively few studies have investigated the effects of early EC on rodents from wild populations (Shepherdson et al., 1998). In this study, I examine the effects of varying degrees of social interaction (group living) on the cerebral cortex and spatial learning in the golden mouse, Ochrotomys nuttalli. This species is found throughout the southern U.S. from northern Kentucky to central Florida, and from eastern Texas to the Atlantic Ocean (Whitaker, 1996). Adults - range in total length from 127-190 mm and weigh 18—24 g (Linzey and Linzey, 1967; Nowak, 1991). This is an arboreal and gregarious species that is an adept climber and can move rapidly on high branches using its prehensile tail for balance and support. It prefers thick woodlands with dense undergrowth as well as swampy areas, and is especially prevalent in some parts of its range in greenbriar (Smilax glauca), red cedar (Juniperus virginiana), or honeysuckle (Lonicera japonica) (Linzey and Packard, 1977). It feeds primarily on acorns and other types of seeds as well as on a variety of invertebrates (Goodpaster and Hoffmeister, 1954; Peles et al., 1995). This mouse nests in shrubs, trees, and on the ground. Ground nests are typically found in association with leaf litter and are often constructed within depressions in the soil or under logs (Frank and Layne, 1992). In shrubs or trees, globular nests are constructed of leaves, bark, and grass, and are typically situated from 1—5 m above the ground (Packard and Garner, 1964; Linzey, 1968). Another structure, called a feeding platform, is a type of retreat where food may be carried and eaten by these animals (Goodpaster and Hoffmeister, 1954). Home ranges can overlap and are from 0.053 to 0.672 ha (Linzey, 1995), and litter sizes range from 1-4 (Blus, 1966), with a mean of 2.2 (Linzey and Linzey, 1967; Pagels, 1999). In a study of O. nuttalli in eastern Florida, Ivey (1949) reported that nests were typically occupied by solitary individuals or a maternal parent with her young (single family group, SFG). However, golden mice are also known to exhibit social behavior with 3-8 adults (mostly females with their sexually mature young) occupying a single nest (Barbour, 1942; Goodpaster and Hoffmeister, 1954; Rose and Walke, 1988). Dietz and Barrett (1992) found that there were no significant differences in the frequencies of group nesting of kin and unrelated mice, and other studies have shown that, given the relatively small litter sizes for this species, nests containing 6-8 in- dividuals indicate that more than one litter is remaining in the nest (multiple family groups, MFG) for an extended period of time (Layne, 1958; Dietz and Barrett, 1992). Although MFG nests can be found throughout the year, it has been reported that they are more common during colder months followed by a more solitary terrestrial nesting habit during the summer (Layne and Dolan, 1975). This led to the sug- gestion that group living may represent a strategy whereby metabolic rates are lowered in response to decreased food supply, thereby increasing energy and water conservation for individuals within the nest (Springer et al., 1981; Peles et al., 1995). However, at several locations in Florida (Gadsden, Jackson, Leon, and Lafayette Counties) I have found O. nuttalli occupying MFG nests throughout the spring and early summer months (Punzo, unpubl. data), when these mice are reproductively active. This suggests that, in addition to thermoregulation and energy and water con- servation, group nesting may contribute to fitness in some other ways. 146 FLORIDA SCIENTIST [VOL. 67 One possible adaptive advantage of MFG nesting may be related to the effects of environmental complexity on developing offspring. Contact with group members, including parents, siblings, and non-kin, can provide the offspring with a greater variety of chemical, olfactory, and visual stimul than would be present under solitary conditions (Kempermann et al., 1997; Tarou et al., 2000; Punzo and Alvarez, 2002; Punzo and Lau, 2003). As stated previously, early EC can significantly affect CNS development and a variety of subsequent behaviors that might contribute to over- all fitness. Young of golden mice that are reared in multiple family groups are presumably exposed to increased stimuli not only due to the increased number of young in the nest, but also to contact with non-related adults. The purpose of the present study was to compare the effects of EC janie extra-familial group living (MFG) versus the maternal parent-sibling unit (SFG) on CNS development (cortical depth), running wheel activity, and spatial learning ability in O. nuttalli. MATERIALS AND METHoDs—Subjects and rearing methods—All\ animals used in this study were observed and/or live-trapped (N = 15 males; 19 females) in Leon and Lafayette Counties, Florida during the period of 10 Mar.—7 Nov., 2001. Animals were placed in plastic cages (20 < 20 X 15 cm) containing sawdust substrate and provided with water via a tubelick water bottle. Arboreal nests and any occupants (N = 22) were collected and placed in a vinyl container (1 m*). Any animals present in the nest were housed together in a plastic rodent cage (60 X 45 X 20 cm) containing sawdust and a water bottle. In captivity, animals collected from each group nest were placed in separate vinyl cages (0.5 m°; Bush Biological Supply, Neodosha, KS). Each of these cages had a clear plastic sliding window panel in the front which allowed access to the interior of the cage, and ventilation holes on the top and sides. Each cage was provided with a substrate consisting of wood shavings plus materials that have been reported from nests in the wild, including leaves of oaks, honeysuckle, maple, hackberry, and greenbriar, strands of shredded inner tree bark, spanish moss, and soft outer parts of milkweed seedpods. Animals collected as solitary individuals in the field were housed separately in plastic containers (25 * 25 * 20 cm) under the same conditions described above. Cages were housed in a room with temperature and relative humidity maintained at 21—22°C, and 62-72%, respectively. Animals were provided with water ad libitum and fed on a diet of rolled oats and the seeds of sumac (Rhus), clover (Trifolium), milkweed (Asclepias), greenbriar (Smilax), and oak (Quercus). I have reared golden mice successfully in captivity using this diet (Punzo, unpubl. data), and all of these seeds represent foods eaten by golden mice in the wild (Goodpaster and Hoffmeister, 1954). Females from group nests and those captured alone that had been impregnated before capture began showing outward signs of pregnancy after 2-3 weeks in captivity. Because some group nests in the wild have been shown to consist of unrelated females and their young (Dietz and Barrett, 1992), 6 groups, each consisting of 2 gravid females, were selected at random and placed in cages as those described above. These groups represented MFGs. The females remained together until they gave birth, and thereafter until their pups were 3 months old. These MFGs ranged in size from 6—8 individuals (2 adult females, and 4-6 young). At 3 months of age, the young were removed from their groups and housed individually in plastic rodent cages for 24 weeks. As these 3.5—4-month-old young had been exposed to a more complex rearing environment (more varied degree of chemical, visual, and olfactory stimuli) as the result of social interactions with both related and unrelated adults and young, compared with their counterparts from single mother-young families (SFGs), they were designated as the multiple family group (MFG) and provided subjects for all subsequent experiments. Offspring (N = 45) obtained from females that had been captured singly in live traps on the ground (N = 17) were housed individually in similar cages and otherwise treated in the same way as the mice from MFGs. At birth, the size of each of these litters was reduced to 1 offspring in order to remove any effect of sibling interaction. Because these offspring had been exposed only to their maternal parent during weaning, they were designated as the single family group (SFG). Because of No. 2 2004] PUNZO—GOLDEN MOUSE STUDIES 147 an inadequate supply of females, all experiments were conducted on male subjects that were 3.5-4 months old. Spatial learning experiments—Six and 10 subjects, respectively, were selected at random from the SFGs and MFGs and tested for spatial learning ability using a radial 8-arm maze (Columbus Instruments, Model 0500-1-D40, Columbus, Ohio) described in detail by Punzo and Lau (2003). To summarize, the maze was constructed of white stainless steel sheeting, with a circular central arena 24.6 cm in diameter. Eight arms, 80 cm long and 12 cm wide, radiated out from the center at equal intervals. The walls of the - central arena and arms were 15 cm high, and the arena and arms were covered with transparent plastic covers that prevented the subjects from jumping out of the maze. A small depression 1 cm in diameter and depth served as a food cup at the end of each arm. The maze was placed in a room with a temperature of 21—22°C, and relative humidity of 60-70%. The room contained abundant extramaze cues on its walls in the form of bookshelves, windows, cabinets, and a cool fluorescent light provided uniform iJlumination. All observations were made behind a one-way mirror to minimize disturbance to the animals. Before each training session, the food cups were baited with a 25 mg Noyes food pellet. Each subject was placed in the center arena in a random orientation, and the sequence of arms visited and the direction of the turn made when exiting each arm were recorded. Subjects were required to move to the end of each arm in order to detect the food pellet which was concealed by the edge of the depression at distances up to 15 cm. A session was ended when the subject had spent 30 min in the maze, had made 16 choices, or had consumed all 8 baits. The maze was washed thoroughly with soapy water after each trial. Each animal was tested daily for 3 weeks, and only data from the last 10 days was used for statistical analyses. The first 11 days of training allowed the animals to adjust to conditions within the apparatus (Magni et al., 1979). Running wheel experiments—The procedure used to assess locomotor activity involved a standard Wahmann running wheel connected to a running wheel activity monitor with automated relay output (Columbus Instruments, Model 1020-D40), which has been described in detail elsewhere (Osborne et al., 1980). Six and 7 subjects, respectively, were selected at random from the MFGs and SFGs and tested individually. Each subject received 6 daily 10-min sessions, and the mean number of rotations/day for all 6 days exhibited by MFG and SFG subjects was compared using a Student t-test (Sokal and Rohlf, 1995). Analysis of brain morphology—At the end of the experiments, mice from the MFG (N = 24) and SFG (N = 22) were sacrificed by administering an overdose of sodium pentabarbitol, and subjected to intracardiac perfusion with 0.9% saline followed by 10% formalin. Their brains were removed and stored in buffered formalin for 7 days, after which the olfactory bulbs were removed and the brain stem severed directly caudal to the cerebellum. The brains were weighed on an electronic balance and then sectioned coronally, starting at the initial appearance of the genu of the corpus callosum and continuing beyond the splenium. A Leitz microtome (Model 1370) was used to prepare tissue sections 30 um in thickness. Every fourth section was saved and stained with metachromatic thionin as described by Walsh (1981). Morphological measurements on the cerebral cortex of the brain were measured as described by Carughi and co-workers (1989) for the rat. The appropriate cortical sections included the frontal area immediately anterior to the first crossing of the corpus callosum; parietal area immediately anterior to the anterior commissure; and occipital area at the crossing of the posterior commissure (see Fig. 1). Camera lucida tracings were made of those sections at 20X magnification. The first measurement of cortical depth was taken 2.2 mm lateral to the median elevation of the corpus callosum. The second was taken 3.1 mm lateral to the first, and all subsequent measurements were taken 5 mm apart. Measurements of the 3 cortical regions were obtained from the left and right cerebral hemispheres. With respect to the frontal lobe, the first 3 measurements from the midline (Fig. 1, upper, A), and the remaining 3 (B) were averaged. In the parietal lobe, the averages were based on groups (Fig. 1, middle, C), the following 3 (Fig. 1, middle, D), and the remaining 3 (Fig. 1, middle, E). Of 18 occipital lobe measurements (Fig. 1, lower), the averages were based on 6 (M), 4 (N), 4(O), and 4 (P) groups of measurements proceding laterally from the midline. Data on cortical depth were analyzed using an analysis of variance (ANOVA) with environmental complexity (MFG vs. SFG), cortical regions (frontal, parietal, and occipital lobes), and lateralization (right vs. left cerebral hemisphere) as factors (Sokal and Rohlf, 1995). 148 FLORIDA SCIENTIST [VOL. 67 Fic. 1. Camera lucida drawings of coronal cross-sections of the brain of Ochrotomys nuttalli, show- ing various measurements of cortical depth. All measurements were made at 20 magnification. The lobes shown are: upper—frontal lobe (F), middle—parietal (Par), and lower—occipital (Occ). See text for details. RESULTS—At 3 months of age, the body weights of MFG animals ranged from 19.4—23.3 g (mean: 21.4 + 1.8 SD), whereas SFG animals ranged from 16.2—20.5 g (mean: 18.3 + 1.3). This difference in body weight was significant (P < 0.05). The degree of social complexity had a significant effect on brain weight in males of No. 2 2004] PUNZO—GOLDEN MOUSE STUDIES 149 O. nuttalli as well. The brains of animals from the MFG weighed between 459.4— 466.5 mg (mean: 462.3 + 8.2 SD) as compared to 432.7-448.2 mg for the SFG nea: 4427 += 7.2) ((=2:18, df= 44, P < 0.05). Social complexity also had a significant effect on the cortical thickness of the frontal, parietal, and occipital lobes of the brain (Table 1). There was a significant effect of rearing condition (F; 4; =4.14, P < 0.05), brain region (Fg 32g = 2.43, P < 0.05), and lateralization (right vs. left hemispheres) (Fj 4; = 4.21, P < 0.05), as well ~ as a significant interaction between rearing condition and brain region (Fg 32g = 2.39, P= 0.05). During the last 10 sessions with the radial maze, the animals from the MFG group required a mean of 9.64 (41.2 SD) choices to remove the food from all 8 arms. On the average, 7.24 of their first 8 choices were correct in the sense that they represented the first visit to that arm during the given training session. This level of performance was significantly better than chance (5.25 choices; see Olton, 1978 for calculations), assuming random, independent choices (t = 19.46, df =9, P < 0.01). In contrast, subjects from the SFG performed more poorly, requiring an average of 14.21 choices per session, achieving a mean of 6.09 correct choices out of the first eight, which was also significantly better than chance (t = 12.05, df =5, P < 0.02). Differences in performance levels of the 2 groups was significant (t = 8.04, df= 14, P < 0.001). Subjects from the MFG exhibited a significantly higher level of locomotor activity, as measured by running wheel activity, as compared to the SEG. MFG subjects averaged 270.43 += 22.6 SD revolutions/min (range: 238-294), as compared to a mean of 207.31 +£15.3 (range: 168-227) for the DE animals (t = 2.97, df = 11, P < 0.05). An analysis of covariance (ANCOVA) showed that the enhanced locomotor activity of the MFG was not the result of differences in body sen, 4 — 143; P-<)0.60). DiscussioN—The results of this study indicate that enhanced group living experienced early in life can have effects on brain development and behavior in O. nuttalli. These data represent the first of their kind for this species. This suggests that, in addition to its impact on thermoregulation and energy conservation, the additional sensory stimulation associated with group living results in a thicker cerebral cortex resulting in increased exploratory activity and greater spatial learning ability. The results of this study, the first for rodents from a wild population, are in agreement with previous research on inbred laboratory subjects demonstrating that both formal training and informal experience in varied environments lead to mea- sureable changes in the neurochemistry and neuroanatomy of the rodent brain (see review by Rosenzweig and Bennett, 1996). Differential experience in early life has been shown to affect neurogenesis and synaptogenesis (Nilsson et al., 1999), brain weight (Diamond et al., 1993), thickness of the cerebral cortex, corpus callosum, and hippocampus (Kempermann et al., 1997), and concentration of neurotransmitters (Thoenen, 1995) in house mice and rats. In addition, those animals reared in environ- ments that allowed contact with parents, siblings and other conspecifics, and/or 150 FLORIDA SCIENTIST [VOL. 67 TABLE 1. Means + SE of measurements (mm) of brain cortical thickness of golden mice subjected to different levels of social environmental complexity in early postnatal development. Different letters in rows indicate significant differences, P < 0.05). Measurements were taken 2—5 mm apart from the midline of the frontal (A, B), parietal (C, D, E), and occipital M, N, O, P) cortices. See text for details. Single Family Groups (N = 22) Multiple Family Groups (N = 24) Brain region Left Right Left Right Frontal cortex A 24.7 + 0.3a 24.9 + 0.2a Di == OMI 25.8 + 0.2b B 24.9 + 0.2a 25.2, 220M 25.58 0b pray pes (5) Parietal cortex € 23°6-+00la 23.8 = 02a 24.7 + 0.2b 24.9 + 0.3b D 23.2 = 0:2a Pi yy = (0.12) 24.5 + 0.1b 24.6 + 0.2b E 22:6; = 0:24 23-2202 24.7 + 0.1b 24.6 + 0.2b Occipital cortex M 13y 2024 1327522, 0Ma 13.6 + 0.2a 13.8 + 0.2a N 13:95 03a 1358 = 02a 14.9 + 0.2b 14.9 + 0.1b O 5.2: 02a 1523: = OFZa 16.4 + 0.1b 16.5 + 0.3b P iis ae0i3.a 1733 = 02a 18.6 + 0.2b 18.7 + 0.3b access to novelty objects (running wheels, multi-colored wooden blocks, tunnels, ladders, etc.), exhibited an enhanced preference for novel environments (Sahakian et al., 1977) and superior learning abilities (Mellen, 1992) when compared to animals reared in isolation in barren cages. The differential effects of early experience on subsequent behavior that were identified through these previous studies have been used to develop more effective environments for domestic livestock (Wemelsfelder et al., 2000) and captive animals in zoos and animal parks throughout the world (see Shepherdson et al., 1998). Young animals housed under conditions lacking in novelty and diversity and that offer few opportunities for interaction frequently exhibit sterotypical patterns of behavior and a decreased tendency to breed and care for young after reaching sexual maturity (Wood-Gush and Vestergaard, 1989; Wemelsfelder et al., 2000). The opportunity for interaction with conspecifics and the placement of novelty objects in enclosures have been shown to reduce aggression in kangaroos (Hohn et al., 2000) and mice (Van Loo et al., 2002), enhance mating activities in pigs (Pearce and Pearson, 1993), red deer (Whittington and Chamove, 1995), and seals (Hunter et al., 2002), increase the use of vertical space and enhance breeding in orangutans (Hebert and Bard, 2000), and enhance maternal care of young in monkeys (Suomi, 1997). It has been suggested that sufficiently enriched early experience may be a requirement for the full manifestation of species-specific brain characteristics and behavioral potential (Rosenzweig, 1984). In this study, the enhanced running wheel activity and spatial learning ability of male golden mice that had been reared under MEG living conditions could increase overall fitness. Increased locomotor activity has been associated with an increased tendency to explore novel objects and places (Sahakian et al., 1977; Prior and Sachser, 1995). This could result in more efficient foraging strategies and subsequent caloric/nutrient intake and contribute to an No. 2 2004] PUNZO—GOLDEN MOUSE STUDIES ily increase in fecundity and survivorship of offspring. Increased cortical depth is associated with an increase in neurogenesis (Nilsson et al., 1999). As the cerebral cortex is involved in learning and memory processes (Rosenzweig and Bennett, 1996), the increase in cortical depth exhibited by O. nuttalli from the MFG could account for their superior performance in the radial maze. This is especially pertinent because the radial maze has been shown to present animals that actively search for resources with an ecologically relevant task for assessing spatial learning abilities (Davey, 1989). The ability to accurately navigate and locate food and escape routes, and to learn to utilize shortcuts and detours using novel pathways, would contribute to survivorship. Indeed, it has been argued that spatial behaviors are essential to the survival of all species of mammals despite the diversity of their morphology or natural habitats (Benhamou and Poucet, 1996; Punzo, 2003). ACKNOWLEDGMENTS—I would like to thank C. Farmer, L. Davila, and J. Trahan for assistance in maintaining the animals in captivity, M. Sarge and K. Smith for assistance in the maze training sessions, B. Garman for consultation on statistical procedures, and J. Layne, D. Martin, B. Martin, C. Hoffman, and anonymous reviewers, for commenting on an earlier version of the manuscript. All animal protocols followed the guidelines for the ethical treatment of animals of the Animal Behavior Society, Washington, D.C. LITERATURE CITED Barsour, R. W. 1942. Nests and habitat of the golden mouse in eastern Kentucky. J. 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Florida Scient. 67(2): 144-153. 2004 Accepted: August 20, 2003 Biological Sciences FIRST REPORT OF APLIDIUM ANTILLENSE (GRAVIER, 1955), (TUNICATA, APLOUSOBRANCHIATA) FROM FLORIDA THOMAS STACH Royal Swedish Academy of Sciences, Kristinebergs Marina Forskningsstation, 45034 Fiskebackskil, Sweden ABSTRACT: The aplousobranch ascidian species Aplidium antillense (Gravier, 1955) was found growing on floating docks in Fort Pierce, Florida in the Indian River Lagoon. Previously described from several locations in the Caribbean and Bermuda, this is the first record from the continental North American coast. The description of A. antillense is augmented; it is concluded that the species can be distinguished from similar ones occurring in Florida by the number of stomach plications (11-14). The present report draws attention to A. antillense and will eventually allow a more rigorous investigation of the taxonomy and biogeography of the diverse genus Aplidium. Key Words: Ascidian, Polyclinidae, biogeography, taxonomy THE ASCIDIAN Aplidium antillense (Gravier, 1955) was found in the vicinity of Fort Pierce inlet in the Indian River Lagoon: Fort Pierce, Florida, 27°26'57"N latitude, 80°19'18”W longitude, floating dock, 25 cm depth, voucher specimens deposited at National Museum of Natural History, Smithsonian Institution, Washington, DC, USNM 1011799, coll. T. Stach. Aplidium antillense (Gravier, 1955) is a small colonial ascidian (sea squirt) species. Colonies are of irregular shape, up to 7 cm in diameter, and up to 10 mm in thickness. The colonies are rounded, the test relatively soft, and the matrix material is colorless. Most Floridian colonies examined are strikingly white in ap- pearance (Fig. 1A and B), though some colonies possess a bluish hue. The coloration stems from numerous small reflective structures in the test (Fig. 1D). Besides these reflective structures that are about 3—5 um in diameter, the test contains larger, irregularly shaped inorganic particles. Zooids are irregularly distributed in the colony (Fig. 1A). The branchial siphon has six lobes (Fig. 2). The postabdomen can easily exceed the length of the branchial and abdominal sections combined. Branchial section and abdomen together measure up to 3 mm in length, whereas the postabdomen can reach a length of up to 4 mm. A slender conspicuous languet arcs over the atrial opening. There are 9—12 rows of stigmata in the branchial basket. The stomach has straight, though occasionally divided, longitudinal plications. The number of these plications is approximately 12 (Fig. 2). This is the most reliable character to distinguish A. antillense (Gravier, 1955) from A. exile (Van Name, 1902) that is also reported from Florida waters (Van Name, 1945, Bingham, 1990). The number of stomach plications in A. exile is 20 or more (Van Name, 1945). The intestine has constrictions and seems to 154 No. 2 2004] STACH—NOVEL ASCIDIAN a5 Fic. 1. .A—Macrophotograph of a colony of Aplidium antillense. Note the irregular arrangement of the zooids. B—Higher magnification to demonstrate distribution of tunic inclusions (arrow). C—Light micrograph of a larva of A. antillense dissected from the parental atrium. D—Higher magnification of tunic. ab—abdomen, sg—irregular inclusions, probably sand grains, wc—small inclusions that give the living colony their white appearance. be divided in three parts as usual in the genus. However, the partition of the intestine is often obscured by the presence of fecal pellets. A flat part that makes a 90° twist as described by Monniot (1972) was not observed possibly due to the presence of fecal pellets. The intestine opens into the atrial cavity. The ovary lies in the anterior part of the postabdomen. Numerous testes are situated in the posterior part of the postabdomen. Testes are in pairs and the sinusoid sperm duct 1s conspicuous. The heart is situated posterior to the gonads. Larvae are commonly found brooded in the atrial cavity. A larva dissected from the atrial cavity is depicted in Fig. 1C and 2. The trunk of a larva before hatching measures about 280 um in length. Its organs are well developed. It possesses epidermal papillae and three funnel-shaped larval attach- ment papillae that are arranged along the sagittal plane at the anterior end. Rarely more than two larvae were observed in the atria. Distribution (Fig. 3}—Aplidium antillense was first described from Martinique (Gravier, 1955) under the name Amaroucium antillense. Monniot described the species from the Bermudas (Monniot, 1972) and Guadeloupe (Monniot, 1983). Goodbody (2000) reported the species from Belize and found it recently in the harbor of Kingstontown, Jamaica (Goodbody, 2003). It was found by the present author in the Indian River Lagoon on the East Coast of Florida in the proximity of an inlet and close to harbor facilities. Field marks—The species is hard to distinguish from A. exile in the field. However, the combination of small size and the white color are good distinguishers for A. antillense in the Indian River Lagoon. 156 FLORIDA SCIENTIST [VOL. 67 Fic. 2. Line drawings of a single entire zooid, a larva, and across section through the stomach region (plane see arrow) of Aplidium antillense. as—atrial siphon, en—endostyle, fp—fecal pellet, gd—gonoduct, in—intestine, la—larva, lg—languet, Im—longitudinal muscle, lp—tlarval papilla, oc—ocellus, ov—ovary, rst—rudiment of stomach, sc—statocyte complex, sd—sperm duct, st—stomach, ta—tail, te—testes. No. 2 2004] STACH—NOVEL ASCIDIAN 157 Bermudas =F ort Pierce, Florida Kingston, Jamaica ‘Guadeloupe ® Martinique 4 » Fic. 3. Map showing localities of earlier (circles) and present (arrow) findings of Aplidium antillense. The zooids of A. exile are variably colored as described by Van Name (1945), sometimes brown, yellow, or even orange. The living zooids of A. antillense from Florida were of a dull gray color. Only the stomach was brightly colored, yellow to orange. Colonies of A. exile from the same location were brown. Discussion—Aplidium is a genus that contains about 200 described species. Although they are generally clearly distinguished from one another, their small size does not attract attention from many observers. Van Name (1945) lists A. stellatum (Verrill, 1871), A. pellucidum (Leidy, 1855), A. constellatum (Verrill, 1871), A. ber- mudae (Van Name, 1902), A. exile (Van Name, 1902) from Florida, and A. funginum (Sluiter, 1898) from Tortugas Island. In an informal identification key, appended to his doctoral thesis, Bingham lists A. exile and A. constellatum from the Indian River Lagoon (Bingham, 1990). A. antillense has not been reported from the continental United States so far. Several hypotheses could account for this. Taxonomic inconsistency and/or misidentification might be reasons. Van Name in his original description of A. exile mentions that the species possesses 9 stomach folds (Van Name, 1902). In his major work, he corrects this number to 20 or more (Van Name, 1945). It is conceivable that he was dealing with two different species. Gravier (1955) distinguished A. antillense from A. exile by: a smaller number of rows of stigmata (never more than 12) in A. antillense, a smaller number of stomach plications (12-14), the subdivision of the intestine into three parts, and a comparatively longer postabdomen. Monniot, on the other hand realized that 158 FLORIDA SCIENTIST [VOL. 67 most of these characters were too variable and overlapping to be reliable for species identification. This author acknowledged only the number of stomach plications, 11-14 in A. antillense versus more than 20 and up to 35 in A. exile, as a valid distinguishing characteristic to identify A. antillense (Monniot, 1972; Monniot, 1983). On this ground the latter author could discern between the two similar species co-occurring in the same geographical areas. The number of stomach plications also clearly distinguishes A. antillense from A. exile from the Indian River Lagoon. It is 11—13 in the Floridian specimens of A. antillense, whereas in A. exile this number is always higher than 20. In addition, the stomach is slightly longer compared to its diameter in A. antillense versus a more globular shape in A. exile. The fact that A. antillense in the present study was found only in the vicinity of harbor facilities might be a reason to consider another hypothesis. Because ship traffic in the Caribbean is intense, A. antillense might have been transported attached to ships’ hulls or in the bilge water of ships. Finally, it might be also conceivable that the geographic range of the species changed over the last 48 years since its original description. However, the low number of reported observations renders this hypothesis hard to test. ACKNOWLEDGMENTS—I thank the former director Dr. Mary E. Rice and the staff of the Smithsonian Marine Station at Fort Pierce for support during this study, especially S. Reed for guidance in the field. This is Smithsonian Marine Station Contribution No. 573. LITERATURE CITED BinGHAM, B. L. 1990. The ecology of epifaunal communities on prop roots of the red mangrove, Rhizophora mangle. Ph.D. dissert. Florida State Univ. Tallahassee, FL. Goopsopy, I. 2000. Diversity and distribution of ascidians (Tunicata) in the Pelican Cays, Belize. Atoll Res. Bull. 480:304—326. . 2003. University of the West Indies, Jamaica, Pers. Commun. GRAVIER, R. 1955. Ascidies recoltees par le “‘President Theodore Tissier’ (Campagne de printemps 1951). Rev. Trav. Inst. scient. Peches marit. 19:61 1-631. Leipy, J. 1855. Contributions towards a knowledge of the marine fauna of the coasts of Rhode Island and New Jersey. J. Acad. Nat. Sci. Philadelphia. 2(3):135—152. Monniot, F. 1972. Ascidies aplousobranches des Bermudes. Polyclinidae et Polycitoridae. Bull. Mus. nat. Hist. nat., Paris 3e ser. (no. 82):949—962. . 1983. Ascidies littorales de Guadeloupe. II. Polyclinidae. Bull. Mus. nat. Hist. nat., Paris 5(2):413-422. SLUITER, C. P. 1898. Tuniciers recueilli en 1896, par la Chazalie, dans la mer des Antilles. Mém. Soc. Zool. France. 11:5—34. VAN Name, W. G. 1902. The ascidians of the Bermuda islands. Trans. Connecticut Acad. Sci. 11:325— 412, plates 46-64. . 1945. The North and South American Ascidians. Bull. Am. Mus. Nat. Hist. 84:1-475, 31 plates. VERRILL, A. E. 1871. Descriptions of some imperfectly known and new ascidians from New England. Amer. J. Sci. 3(1):54 446. Florida Scient. 67(2): 154-158. 2004 Accepted: September 23, 2003 Biological Sciences A BRIEF DESCRIPTION OF THE COURTSHIP DISPLAY OF MALE PIKE KILLIFISH (BELONESOX BELIZANUS) Lisa Hortu! Kellogg Biological Station, Michigan State University, 3700 East Gull Lake Drive, Hickory Corners, MI 49060 ABSTRACT: The mating behavior of the different live-bearing fish species within the family Poeciliidae varies widely. Several species, such as guppies (Poecilia reticulata), sailfin mollies (Poecilia latipinna) and swordtails (Xiphophorus helleri) are well-studied. However, very little information has been recorded regarding the largest poeciliid, the pike killifish (Belonesox belizanus). This work documents the courtship behavior of male pike killifish, a species which persists within the United States, only in Florida. Unlike males in the non-courting Poeciliidae species, pike killifish males repeatedly conduct particular behavioral acts when in the presence of females. Females do not conduct this suite of behaviors. A courting male will position himself in front of, and perpendicular to a female, and fan his fins and gonopodium (male mating structure), in her direction, prior to attempting to mate. His coloration becomes more vibrant during a display, which is often followed by mating attempts. Pike killifish compose a monotypic genus that has previously been used as an outgroup when studying relationships between live-bearing species. Thus, documenting the courtship behavior of this species is relevant for use in studies where mating-behavior comparisons are being made across genera within this family. Key Words: Mating behavior, live-bearing fish, teleost, Poeciliidae UNDERSTANDING the mating behavior of closely related taxa provides useful information for testing many sorts of hypotheses, such as those involving sexual selection and speciation. Such studies often draw comparisons in which male mating behavior is of prime importance (Seehausen et al., 1997; Uy and Borgia, 2000; Panhuis et al., 2001; Boughman, 2001). Mating behavior data have also been used in cladogram construction. Cladograms are then assessed to determine whether the relationships identified in them reflect molecular phylogenies (Paterson et al., 1995). As an example, such cladistic analyses have been used to determine how well phylogenies account for social organization in primates (Di Fiore and Rendall, 1994). These types of comparisons are relevant in the live-bearing fishes, as well. In swordtails (Xiphophorus helleri), there exists a standing hypothesis that females prefer to mate with males possessing an exaggerated male trait, the sword (an elongated caudal fin), and that this preference arose in swordless species (Basolo, 1990a, b). This hypothesis was recently called into question when character states ' Present address: Section of Evol. And Ecol., Storer Bldg., 1 Shields Ave., University of California at Davis, Davis, CA 95616-5270. 159 160 FLORIDA SCIENTIST [VOL. 67 for male tail-traits were mapped onto molecular phylogenies (Meyer et al., 1994; see also Meyer, 1997). Marcus and McCune (1999) used phenotypic and DNA sequence data to construct a Xiphophorus phylogeny. They found a correlation between increased body size and sword growth rate, and suggest increases in these traits relates to high environmental visibility for females. Their hypothesis is strengthened by phylogeny-wide female preference for the presence of a sword (Basolo, 1995). Marcus and McCune (1999) additionally suggest that selection for small body size in male X. continens is derived, and may contribute to the Joss of the sword despite the presence of female preference for a sword in this swordless species. The mating behavior of many of the fishes that compose the live-bearing family, Poeciliidae, has been well studied (Farr, 1989). Though a complete phylogeny of this family remains to be published, smaller phylogenies of groups within the family have been constructed (Meyer et al., 1994; Morris et al., 2001). Such phylogenies are sometimes created using the pike killifish as an outgroup (Lydeard et al., 1995a, b). However, very little mating behavior data has been gathered for this monotypic genus. In general, mating behavior in the family varies markedly from species to species. Some species exhibit conspicuous male displays (e.g., sailfin mollies, Poecilia latipinna, Travis and Woodward, 1989), and female preferences (e.g., guppies, Poecilia reticulata, Houde, 1988), while others show little to no evidence for such behaviors (e.g. Limia sp., Farr, 1984). For example, in guppies (Poecilia reticulata) there exists a correlation between female preference and males with bright color-spot patterns (Houde, 1987; Houde and Endler, 1990). However, there is also a higher cost of predation for males that display bright spots (Endler, 1980). Within populations of sailfin mollies (Poecilia latipinna), females prefer larger- bodied males, which have larger dorsal fins (that are used in courtship displays), and typically higher rates of display (Ptacek and Travis, 1997). Large differences also occur among populations in size-specific male courtship rates (Ptacek and Travis, 1996). In contrast, gila topminnow (Poecilipsis occidentalis) males conduct very simple courtships displays (Constanz, 1975). Finally, least killifish (Heterandria formosa) males express no obvious color or size polymorphisms and are not known to display to females (Rosen and Tucker, 1961). The pike killifish (Belonesox belizanus) is the largest-bodied (~ 105 mm) of the poeciliids and forms a monotypic genus within the tribe Gambusiini, in the subfamily Poeciliinae (Meffe and Snelson, 1989). This piscivorous species is native to Mexico and Central America (Rosen and Bailey, 1963) and was introduced to Dade County, Florida in 1957 (Belshe, 1961), where it has remained rare, but persisted. Currently, extant pike killifish populations in Dade County are found in a small number of freshwater ditches. The species has not expanded its range outside of south Florida. The limited descriptor of the species’ reproductive behavior includes that used in a broad comparison of the poeciliids (Rosen and Tucker, 1961). Rosen and Tucker (1961) simply indicate that the species has a mating display, including the demonstration of a ‘specific orientation’ by the males, and a lack of mouth/body contact. However, it was not relevant to the message of that publication to describe the mating behavior more thoroughly. A second account states that males demonstrate a simple courtship display only after females give birth (Farr, 1989). No. 2 2004] HORTH—PIKE KILLIFISH COURTSHIP DISPLAY 16] Farr’s reasonable rationale regarding reduced display periods is that these are ‘sit and wait’ predators for whom a high level of sexual activity may deter prey. Belshe (1961) also mentions an extended period of copulatory attempts, but does not describe mating behavior any further. The objective of the present work is to clarify information about the courtship display of pike killifish collected from a field population in south Florida. MeEtTHops—In June 1995, fifteen pike killifish (13 males and two females) were collected via cast-net from canals in Dade County, Florida. Though the fish are distinctive (~3—6” long, slender, swim near the water surface), they are quick and often found in and around vegetation, so several casts were made for each fish that was collected. The fish were transported to Kellogg Biological Station (Hickory Corners, MI) and housed in a 1-KL dark-gray plastic vat, filled with well water and adorned with artificial plants. In August 1995, the fish were moved to a 380-L clear, rectangular Plexiglas aquarium, measuring 2 m long by | m wide by 1 m deep. The aquarium was located directly beneath a 2 m by 1.5 m window and received full sun. Sand was placed on the bottom of the aquarium, and artificial plants were anchored in it. Live guppies (Poecilia reticulata) were added daily, as prey. During autumn, heaters were used to maintain a 22°C water temperature. Observations were made daily, from July to October, on the fish. Fish were observed in the morning and evening for at least 30 minutes, and were checked on several additional times during the day to determine whether displays were occurring throughout the day. Data were recorded daily regarding the sex of the fish observed, whether or not the fish displayed, and what behaviors composed the display. Video recordings were made of the mating behavior and display, as well. The mating ritual was comprised, in part, of several behaviors previously described in other poeciltid species (e.g., intromission: Kadow, 1954; caudal fin spread: Baerends et al., 1955; post-copulatory jerking: Liley, 1966; Parzefall, 1969; 1979; Farr, 1977). RESULTS AND DiscussloN—The activity of the pike killifish when located in the gray vats included simple behaviors: rapid darting from behind plants to capture prey, rapid swimming from behind one plant to another, and occasional swimming parallel, and in close proximity to, another individual. The body-color of fish in the vat was typically dark-gray, occasionally light gray. No other activity was observed. In contrast, within a few days of being transferred to the transparent Plexiglas housing, males were seen displaying to females. Females did not display. Both sexes appeared brighter in color in the clear aquarium, their body-colors being silvery-gray to white abdominally, with the presence of golden hues dorsally. A change in hue from dark to light gray in lower vertebrates (e.g., frogs) often results from a change in the dispersion of melanin, a black pigment present in dermal cells (Hadley, 1996). In general, male displays occur intermittently, several times a day, from early morning until dusk. Displays occurred daily during the entire study period of June through October. Displays last anywhere from a few seconds to many minutes, and may persist, as repeated acts, for the majority of a day. Often, a male displays for several minutes, then leaves the female and swims around for a short period (e.g., 30 sec.), then returns to the female, and continues to display for several more minutes. This was a very typical observation. All males displayed to females frequently. During a mating display, the central portion of a male’s body, inclusive of the dorsal fin and gonopodium (modified fin used for mating), is near the female’s nasal and oral cavities. The male maintains his position in front of the female while moving just the central portion of his body from side-to-side in a sigmoid-motion. 162 FLORIDA SCIENTIST [VOL. 67 He swings his gonopodium (a modified anal fin used to inseminate the female) perpendicular to the long-axis of his body and toward the female, while simultaneously rapidly fanning his erect dorsal fin, in the direction of her snout. Generally, poeciliid males swing the gonopodium forward and backward in a more linear motion, in preparation for, or during mating. The traditional movement does not include distinct sideways thrusts, as occurs during these displays. During a mating-display, a female typically remains motionless and maintains her position in clear view of the male. Sometimes a female will shift her position and change the angle between herself and the displaying male. The male then repositions himself in front of the female, and resumes fanning in her direction. On occasion, the female swims away just prior to, or during the-display. Typically the male follows her and continues to display. When another female swims near the pair, the courting male may re-direct his display to the new female. After fin fanning, the male sometimes positions himself beneath, and parallel to, the female, his snout near her gonopore. Molly (Poecilia latipinna) and gila topminnow (Poecilipsis occidentalis) males conduct gonoporal nibbles of the female urogenital pore (Travis and Woodward, 1989; Farr, 1997; Constanz, 1975) when in this position. I did not see any nibbles or any male contact with a female’s urogenital pore, which is consistent with the observations of Rosen and Tucker (1961). While beneath a female, a male will occasionally open his mouth wide, arc his body slightly, and swing his gonopodium forward, then backward slowly, and in the traditional pattern of live-bearing males (parallel, not perpendicular to the axis of his body). The gonopodial swing and mouth-opening behavior also occurs in males that display for a long period, but that have not moved beneath the female. After displaying, a male occasionally attempts to copulate, and swings his gonopodium directly toward the female’s gonopore, in a manner consistent with other poecillids (more parallel than perpendicular to his own body). If a second male attempts to intrude while a female is being courted, the courting male either chases the potential new suitor away, or jockeys to maintain his position between the intruder and the female. The defending male swims parallel to the female, very near to her, holding his position between the female and the intruding male. During these encounters, and sometimes independent of a female, the two males will swim parallel to one another, displaying their throat-patches at one another. Additional changes in the male’s appearance when courting a female include a deepening in hue of the typical gray to light-gold body-coloration, to a much deeper gold. A small pelvic fin, near the gonopodium, turns deep gold and pink. In addition, three body regions also turn pink: 1) a small area on the head, just posterior to the otoliths, 2) an area on the body immediately ventral to the dorsal fin, and 3) the area on the body anterior to the tail fin. The caudal fin, though typically translucent, displays a black, translucent hue during courtship. The caudal fin-spot (present in all individuals in this collection, but not visible at all times), and horizontal rows of black, body-spots, darken. How the body-spots and caudal fin- spot display is used in these, and other poeciliids (e.g. Heterandria Formosa, Poecilia latipinna), remains to be determined. Pike killifish males also have two pouches of skin, one under the mouth on the throat, and one on the upper abdomen, No. 2 2004] HORTH—PIKE KILLIFISH COURTSHIP DISPLAY 163 which they sometimes distend during a display to a female, or in an aggressive encounter with another male. In contrast to males, females do not display. Their primary mating-behavioral repertoire appears to consist of either remaining motionless while a nearby male displays, or moving away from a displaying male. Females occasionally open their mouths wide when being courted, potentially allowing them to receive chemical cues from a fanning male. However, the actual purpose of this mouth-opening be- havior is not known. Females gave birth at least three times during the study period. On all occasions, the young were cannibalized within two days. Males courted females both before and after parturition. In the collection of 13 males and 2 females that I had, all males conducted mating-displays toward females. I made only one observation out of hundreds, where a male turned upside-down and thrust his gonopodium at a female without displaying first. Lack of a display is consistent with Farr’s observations (1989). In addition, if the larger males were displaying to the females, since there was a male- biased sex ratio, the smaller males sometimes displayed to other males, though when they had the opportunity they would preferentially display to females. Farr (1997) has noted that in guppies (Poecilia reticulata), when the sex ratio is male-biased, males will display to other males. Gambusia holbrooki males also occasionally attempt to mate with other males (pers. obs.). With respect to the lack of mating-displays noted in other studies of pike killifish, it may be that Farr (1989) did not observe the mating-display that I did because he was not using the equivalent of a clear, oversized-housing, exposed to intense sunlight, as was used in this study. Indeed, his behavioral descriptor is a subset of the pattern that I observed. Even in my study, there appeared to be much more activity, both with respect to behavior and color-change, in the brightly sunlit Plexiglas chamber, than in the dark gray vat. However, this was observational study was not designed as an empirical manipulation so confirmation of the importance of light to the display is not possible from this work. Observations on these fish in nature would be most helpful for understanding the significance and frequency of male displays to females. Such information, as well as that documented here, may prove useful to those developing theories regard- ing mating behavior in poeciliids, or mapping characters such as mating behavior onto phylogenies to identify the role of phylogenetic constraints on behavioral development. ACKNOWLEDGMENTS—Thanks to R. Cailteaux, J. Trexler for comments, J. Farr for sharing unpublished observations, W. Loftus and A. Turner for assistance in collecting fish, G. Mittelbach for supplies, and C. Huckins for the hand-made Plexiglas aquarium. This work was supported by NSF Research Training Grant DBI 9602252. MSU AUF 04/94-106-01. LITERATURE CMED BAERENDS, G. P., R. BROUWER AND H. T. WATERBOLK. 1955. Ethological studies on Leibistes reticulates (Peters). I. An analysis of the male courtship pattern. Behaviour 8:249-335. Basoo, A. L. 1990a. Female preference predates the evolution of the sword in swordtail fish. Science Washington. 250:1426—1427. 164 FLORIDA SCIENTIST [VOL. 67 . 1990b. Female preference for male sword length in the green swordtail, Xiphophorus helleri (Pisces: Poecillidae). Anim. Behav. 40:332—338. . 1995. Phylogenetic evidence for the role of a pre-existing bias in sexual selection. Proc. R. Soc. Lond. B 259:307-311. BELSHE, J. F. 1961. Observations of an Introduced Tropical Fish (Belonesox belizanus) in Southern Florida. M.S. 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Social context and courtship flexibility in male sailfin mollies, Poecilia latipinna (Pisces: Poeciliidae). Anim. Behav. 38:1001—1011. Uy, J. A. C. AND G. Borata. 2000. Sexual selection drives rapid divergence in bowerbird display traits. Evolution 54:273—278. Florida Scient. 67(2): 159-165. 2004 Accepted: September 29, 2003 166 1963 1964 1965 1966 1967 1968 1969 1970 LO Fa LOT2 LOFe 1974 es) 1976 LOTT 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 OO 11992 1993 1994 WDD 1996 1997) Dr Florida Academy of Sciences Medalists . Archie Carr . Werner A. Baum . Alex G. Smith . Karl Dittmer . Alfred H. Lawton . Sidney Fox . F. G. Walton Smith . Pierce Brodkorb . Maurice A. Barton . Lloyd M. Biedler . Ruth S. Breen ie: L. Yorker . Alex E. S. Green . Robert N. Ginsburg . Michael Kasha . John Edward Davies . Stanley S. Ballard . Thomas D. Carr . Harold J. Humm . George B. Butler . Karen Steidinger . Yngve Ohrne . William Sears . E. Dwight Adams . Larry Hench . Gene C. Ness . Frank Bradshaw Wood . Martin Uman . Frank Millero Dr. Gregory Choppin D. O. Shah Mrs. Barbara B. Martin Dr Dr Dr Dr Dean F. Martin James N. Layne . Leshe Sue Lieberman . George R. Newkome FLORIDA SCIENTIST University of Florida University of Miami University of Florida Florida State University University of South Florida University of Miami University of Miami University of Florida Mound Park Foundation Florida State University Florida State University University of Florida University of Florida University of Miami Florida State University University of Miami University of Florida University of Florida University of South Florida University of Florida Florida Dept. Natural Resources University of Florida Florida Atlantic University University of Florida University of Florida University of South Florida University of Florida University of Florida University of Miami Florida State University University of Florida University of South Florida University of South Florida Archbold Biological Station University of Florida University of South Florida [VOL. 67 Biology Meteorology Astronomy Chemistry Medicine Biochemistry Marine Science Zoology Medicine- Teaching Physiology- Biophysics Botany Agriculture Physics Geology Molecular Biophysics Public Health Optics Astronomy Marine Biology Chemistry Biology Chemistry Anthropology Physics Engineering Biochemistry Astronomy Electrical Eng’n Oceanography Chemistry Chemical Eng’n Chemistry Chemistry Zoology Anthropology Chemistry No. 2 2004] 1998 1999 2000 2001 2002 ~ 2003 Dr. Joseph J. Krzanowski Dr. Abraham Kandel Richard F. Lockey, M.D. Dr. Clinton J. Dawes Dr. John Trefry Dr. Christian Clausen MEDALISTS University of South Florida University of South Florida University of South Florida University of South Florida Florida Technological Univ 167 Pharmacology Computer Science Medicine Biology Oceanography University of Central Florida Chemistry 168 FLORIDA SCIENTIST [VOL. 67 The Florida Endowment for the Sciences The Florida Academy of Sciences was founded in 1936 with the purpose of advancing the sciences in Florida through educational programs and public outreach. Although the Academy serves as an important means of communicating scientific research conducted by State agencies, as well as by the State University System, it receives no State assistance. Thus, it exists independently by means of membership dues, sale of its publications, occasional gifts or grants, and modest registration fees at meetings. Over the years the Academy has built up an Endowment Fund designed to provide the necessary financial stability for its activities. The assets of this endowment fund now total over $100,000, and the Trustees of the Fund wish to encourage the membership to keep the Endowment Fund in mind as they make charitable donations. The Academy is recognized by the IRS as a non-profit scientific and educational organization. Your gifts to the Endowment Fund are removed from the amount subject to Federal Income Tax when you itemize deductions on your personal income tax return. This can result in substantial tax savings. In addition to making gifts by check, you may also make gifts of other investments. When you do so, you take advantage of their appreciated value. These can include stocks, bonds, mutual funds, insurance policies, real estate, paintings, unwanted jewelry, rare books, etc. You may also want to remember the Endowment Fund in your will. Cash donations should be sent as checks made out to the Florida Academy of Sciences and mailed to Dr. Donald W. Lovejoy at Palm Beach Atlantic University, P. O. Box 24708, West Palm Beach, FL 33416. The Academy will provide you with a receipt to be filed with your Federal Income Tax return. For gifts other than cash, please contact Don at 561-803-2275 or by email at Donald_Lovejoy@pba.edu 168 INSTRUCTION TO AUTHORS This information is available at two web sites: (1) IES site: http:www.chumalcas.usf.edu (click on “Centers and Institutes”, then select “Institute for En- vironmental Studies”, then select “Florida Scientist’). (2) FAS site: http://www.floridaacademyofsciences.org (select “Florida Scientist”). It is possible to search for papers and abstracts of papers for the past six years: Go the FAS site (above), select “Allen Press”. When the link is complete, select “Advanced Search” and follow the directions. The Florida Academy of Sciences Web page, indicated above, allows interested persons to join the Florida Academy of Sciences and pay dues. FLORIDA ACADEMY OF SCIENCES CORPORATE MEMBER Tierra Verde Consulting, Inc. INSTITUTIONAL MEMBERS Archbold Biological Station FFWCC (Gainesville) Army Corps of Engineers Technical Florida Marine Research Institute Library, Jacksonville Hillsborough Community College Disney’s Animal Kingdom NIOZ-Netherlands Institute for Sea Research Duke University Science Library, University of Chicago Florida Community College at Jacksonville South Florida Water Management District Florida Fish and Wildlife Conservation University of North Florida Commission (Panama City) University of Washington FFWCC (Ocala) US EPA Library FFWCC (Olustee) Virginia Institute of Marine Science FFWCC (Lakeland) Virginia Polytechnic University FFWCC (West Palm Beach) WDI Florida FFWCC (Tallahassee) Membership applications, subscriptions, renewals, and changes of address should be addressed to the Executive Secretary, Florida Academy of Sciences, Orlando Science Center, 777 East Princeton St., Orlando, FL 32803. 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PROCEEDINGS OF THE FLORIDA ACADEMY OF SCIENCES (1936-1944) Volumes 1—7 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES (1945-1972) Volumes 8—35 FLORIDA SCIENTIST (1973—) Florida’s Estuaries—Management or Mismanagement?—Academy Symposium FLORIDA 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 Second Indian River Research Symposium—Academy Symposium FLORIDA SCIENTIST 53(3)—$15.00 Human Impacts on the Environment of Tampa Bay—Academy Symposium FLORIDA SCIENTIST 58(2)—$15.00 Please send payment with order. If required, an invoice will be sent on purchases over $20 from a recognized institution. 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