Volume and value of marine ornamentals collected in Florida, 1990–98 CHARLES M. ADAMS1 , SHERRY L. LARKIN2 and DONNA J. LEE2 1

Florida Sea Grant College Program, Department of Food and Resource Economics, Institute of Food and Agricultural Sciences, University of Florida, P.O. Box 110240, Gainesville, FL 32611, USA; 2 Department of Food and Resource Economics, Institute of Food and Agricultural Sciences, University of Florida, P.O. Box 110240, Gainesville, FL 32611, USA Key words: aquarium fish, marine life species, regulations, saltwater fish ABSTRACT The marine life fishing industry in Florida is defined by the state as the non-lethal harvest of marine plants, finfish, and invertebrates that are sold live for commercial purposes (primarily into the saltwater aquarium industry). Approximately 330 different species of finfish and invertebrates are harvested by marine life collectors in Florida, including 180 species of finfish and 150 species of invertebrates and plants. In 1998, the total dockside value of these species was approximately US$1.9 million. The industry is highly regulated via limits on gear, handling methods, harvest sizes, and trip/bag/possession limits. Entry into the industry has been recently curtailed by the implementation of a moratorium on marine life endorsements. The total number of licensed harvesters exceeded 700 in 1998. Approximately 70 wholesales buyers also participated in the market during 1998. The majority of the dockside value is generated in south Florida, with the majority of the remaining value accruing from the region including and north of Tampa Bay. Over 80% of the total value associated with both finfish and invertebrates is generated by only 10 species within each group. Dockside prices vary considerably among the various species landed, with those landed in greater volumes exhibiting the lower prices.

INTRODUCTION

The marine life collection industry represents an important natural-resource based industry in Florida. Marine life collection in Florida, which is defined as the non-lethal harvest of ornamental saltwater plants, invertebrates, finfish, and live rock/sand for non-food commercial purposes (Florida Department of State, 2000, Chapter 68B-42.001), has grown considerably during the last decade. This growth is evidenced by increases in the number of licensed collectors and either the volume or value of landings (Florida Marine Research Institute, 1999). This growth in the Florida industry has coincided with the domestic expansion in the marine aquarium market, which is purported to now be the second most popular hobby in the United States (Pet Industry Joint Advisory Council, 1999). As the industry has continued to increase, however, so has concern over the sustainable use of these resources. As a result, the marine life collection industry in Florida has been confronted with an increasingly stringent regulatory environment. Concern over the sustainable use of the various stocks of finfish and invertebrates harvested by the marine life collection industry in Florida has necessitated a number of recent regulatory measures, which are contained in the Florida Administrative Code (Florida Department of State, 2000) and the Florida Statutes (The Florida Aquarium Sciences and Conservation 3: 25–36, 2001. © 2001 Kluwer Academic Publishers. Printed in the Netherlands.

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Legislature, 1999). A Saltwater Products License (SPL) is required to harvest and sell any marine species. SPLs have been required since 1985 and have provided an accounting of the number of participating harvesters and effort (trips) expended in every commercial saltwater fishery in Florida. A Marine Life Endorsement (MLE) on the SPL has been required since 1992 to participate in the marine life collection industry. During the same year, several species of finfish and invertebrates were designated as ‘restricted species’ in order to restrict the fishery to ‘full-time’ fishers and thereby limit effort. Under this designation, the harvest of certain species is allowed only by those who can demonstrate that at least 25% of their annual income or $5,000 (all dollars cited in this paper are $US) is derived from commercial fishing activity (i.e., including food fish). Initially, only a few marine life species were designated as restricted species. At present, however, over 200 marine life species (129 finfish and 72 invertebrates) are designated as restricted. Additional restrictions for selected marine life species have been implemented as recently as 1995, such as daily bag limits, possession and trip limits, and minimum/maximum size limits. Other legislation applies to virtually all harvested marine life species, such as allowable harvest methods, commercial season designation, and allowable onboard handling methods. The commercial harvest of live rock was prohibited in the state waters of Florida in 1989, with a three year phase-out of harvest in federal waters beginning in 1992. By 1996, all commercial harvest of live rock (which was previously regulated as part of Florida’s marine life industry) was prohibited in both Atlantic and Gulf of Mexico waters adjacent to Florida. The only exception is the harvest of live rock from permitted commercial culture sites approved by the appropriate state and federal agencies. By 1998, there were 7 commercial live rock culture leases off the coast of Florida (Florida Marine Research Institute, 1999). Even though the marine life collection industry was subjected to ever more stringent regulatory measures, the number of participants continued to grow. During the 1990–98 period, the number of MLE holders increased from 159 to 743 (Florida Marine Research Institute, 1999). In addition, the number of licensed wholesale marine life dealers increased from 66 in 1990 to 114 in 1997, however, 40 wholesale dealers exited the industry during 1998. In an attempt to curb the suspected growth of speculative entry by fishers, Senate Bill 1506 established a moratorium on the issuance of new MLEs in July 1998 (Florida Marine Fisheries Commission, 1998). This moratorium is scheduled to expire in July 2002. Senate Bill 1506 also contained language which calls for an assessment of alternative management strategies, such as tiered licenses, individual quotas, vessel/individual harvest restrictions, and others. Of course, the prospect of extending the moratorium exists. Therefore, an assessment of the harvest patterns, current economic rents, and harvester profiles is needed to provide guidance in the development of an economically efficient management regime for the marine life industry. The purpose of this paper is to provide a brief description of the marine collection industry in Florida during the 1990s. Such an overview is needed to provide

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background information for future management decisions, especially since the only other attempt to characterize the industry was conducted in 1978 (Hess and Stevely, 1978). Focus will be on the existing set of harvest regulations, trends in volume and value of landings by major species groups, and harvest by region. Such information will assist resource managers in developing more effective regulatory measures that provide for the sustainability of the resource, thereby ensuring that Florida retains its share of this important and growing market. MATERIALS AND METHODS

The data utilized for this report were derived from the Florida Trip Ticket Program (Florida Marine Research Institute, 1999). These data are collected by wholesale dealers following their purchases from licensed commercial fishers. The data system tracks fisheries landings from 17 regions for all species of finfish, shellfish, and other invertebrates that are used for any commercial purpose. Thus, the Trip Ticket Program contains data on the landings of numerous species of marine life that are harvested live for the marine aquarium industry. Since 1990, the State has collected data on the harvest of approximately 180 marine life finfish and 150 marine life invertebrate species. These are species collected in state and adjacent Federal waters for sale, barter, or trade. Each Trip Ticket records the type and amount of marine life sold by a fisher to an initial buyer at the conclusion of a trip. Information about what species were landed, the volumes of each, and dockside prices are recorded on the form by the initial buyer. Other information may also be collected, such as gear type used, region where trip was made, etc. These data were obtained from the Florida Marine Research Institute, which is a part of the Florida Fish and Wildlife Conservation Commission, in St. Petersburg, Florida. Annual data were obtained for all marine life species for the years 1990-98. Each record contained (at least) information on the species and volume landed, county of initial sale, and price. This data, which includes all marine life collected live in Florida and retained for commercial purposes, is summarized in the following section. RESULTS

General trends and regional distribution At this point it bears repeating that the ‘marine life’ fishery in Florida is defined in the regulations and legislation to include only saltwater species that are collected live and intended for the aquarium industry (i.e., commercial purposes). The total dockside value of marine life landings in Florida increased from $1.4 million in 1990 to $4.3 million in 1994 (Figure 1). The total value of this fishery then decreased to $3.5 million in 1995 and continued to decrease to $1.2 million by 1998. The majority of the decline since 1995 can be accounted for by decreases

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Figure 1. Annual dockside value of commercial marine life landings in Florida, 1990–98.

in the landings of live rock and sand, which fell from approximately 1.2 million pounds (544,311 kg) in 1995 to 218,000 pounds (98,883 kg) in 1998. The relative share of landings value attributed to fish (i.e., finfish) and invertebrates (e.g., crustaceans, mollusks, echinoderms, plants, live rock and sand, and all others) has fluctuated slightly over time (Figure 1). During the 1990–98 period, fish and invertebrates accounted for an average of 37% and 63% of the total value, respectively. Fish accounted for 55% of the total in 1990, but this share decreased to 40% by 1994 and was 40% in 1998. Of the total value associated with fish, angelfish (which is comprised of 6 individual species) accounted for approximately 52% in each year. Of the total value associated with invertebrates, live rock (which, like angelfish, is also comprised of 6 individual species) accounted for 40% in 1990. The live rock share increased to almost 60% in 1995 before declining to 19% in 1998 due to the phase-out of non-cultured live rock landings mentioned earlier. The majority of the landings value is concentrated in the south Florida region, primarily in Monroe (Florida Keys) and Dade Counties region (Figure 2). During the 1990–1998 period, $18 million, or 70% of the total marine life dockside value, was reported in Monroe and Dade Counties. Less than $500,000 was reported north of Dade County on the Atlantic coast. The Gulf region, excluding the Keys, accounted for the remaining total value during the same period, with 26% of the total value reported from the region including and north of Tampa Bay. Trends in the volume and value of finfish landings Landings and value of marine ornamental finfish increased to peak levels in 1994, then decreased through 1998 (Figure 3). Landings increased from 245,000 individual fish in 1990 to 426,000 in 1994, then declined to approximately 200,000 in 1998. Dockside value followed the same pattern, increasing from $766,000 in

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Figure 2. Geographical distribution of the total value of marine life landed in Florida, 1990–98.

Figure 3. Annual landings and dockside value of marine fish collected in Florida, 1990–98.

1990 to $1.6 million in 1994, then declining to $759,000 in 1998. Note that in 1992, landings increased 35% while the total value of landings remained stable. The increased landings were due, primarily, to a five-fold increase in the collection of seahorses (from approximately 14,000 harvested in 1991 to 83,700 harvested

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in 1992), primarily Hippocampus zosterae (i.e., Dwarf seahorses). In addition, the increased landings of seahorses resulted in a lower market price; the average price paid by dealers for seahorses fell from $1.10 in 1991 to just $0.17 in 1992, a decline of nearly 84%. During the 1990–98 period, over 180 individual species of finfish were harvested. For simplicity, these species were grouped into 66 categories using their common name as defined by the Florida Marine Research Institute. The Institute uses a three digit code for each species and associated with this code are: (1) a common name, (2) genus and species, and (3) family. The common name is most closely associated with the family. For example, the data set contains three genus and species of ‘cowfish’ including Lactophyrs polygonia, Lactophyrs quadricornis, and family Ostraciidae, which are listed (in the common name field), respectively, as honeycomb cowfish, scrawled cowfish, and other cowfish. Although each species has its own unique code, each is member of the Ostraciidae family and data from all three are aggregated and included under the common name ‘cowfish.’ Note that not all codes are associated with a unique genus and species and, thus, fall into an ‘other’ category. Consequently, the number of individual species should be considered as conservative. The 66 aggregate finfish groups are listed in Table 1. If a group consists of multiple species, parentheses are used to indicate the number of individual species that are included in the common name groupings. Of these groups, ten accounted for nearly 84% of the total dockside value (Table 2). The five most economically important species groups, in descending order of importance, were angelfish, hogfish, damselfish, jawfish, and wrasses. The most important single species group was angelfish, which represented 54% of the total value. Hogfish accounted for 7.5% of the total, while the other eight groups accounted for approximately 22% of the total dockside value of live marine finfish collected from 1990 to 1998. These same ten species groups represented 80% of the total dockside value for 1998. While the total value of these species groups remained virtually unchanged in 1998 as compared to the 1990–98 average, the relative importance of the groups Table 1. Fish species groups collected by the commercial marine life industry in Florida, 1990–98∗ Angelfish (6) Balloonfish Barracuda Bass (8) Batfish Bigeye Blenny (8) Brotula Burrfish Butterflyfish (6) Cardinalfish (3) ∗

Catfish Chub Clingfish Coronetfish (3) Cowfish (3) Cusk-Eel Damselfish (14) Drum (4) Filefish (6) Flounder Frogfish (2)

Goatfish (2) Goby (3) Grouper (5) Grunt (5) Hamlet (6) Hawkfish Hogfish (3) Jack (2) Jawfish (4) Lizardfish Minnow

Mojarra Moray (5) Parrotfish (9) Perch Pilotfish Pipefish Porgy Puffer (3) Ray (4) Razorfish Remora (2)

Scorpionfish (2) Seahorse (3) Searobin Soapfish Soldierfish Spadefish Squirrelfish (3) Shark (3) Sheephead Skate Snapper (3)

Stargazer (2) Stingray (2) Surgeonfish (4) Sweeper Tilefish Toadfish Triggerfish (3) Tripletail Trumpetfish Trunkfish (2) Wrasse (8)

Species groups listed in alphabetical order. Parentheses contain the number of individual species that comprise each group.

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Table 2. The ten most economically important groups of fish collected in Florida, 1990–98 Fish group

Total value 1990–98 ($)

Percentage of total fish value (%)

Cumulative percentage of total fish value (%)

1. Angelfish 2. Hogfish 3. Damselfish 4. Jawfish 5. Wrasse 6. Butterflyfish 7. Seahorses 8. Parrotfish 9. Surgeonfish 10. Drum

4,891,917 676,696 316,368 293,857 289,019 273,876 238,631 233,147 201,162 174,865

54.0 7.5 3.5 3.2 3.2 3.0 2.6 2.6 2.2 1.9

54.0 61.5 65.0 68.2 71.4 74.4 77.0 79.6 81.8 83.7

Table 3. Data (1998) and change in landings of the economically important groups of fish collected in Florida Fish group

1998 Value ($)

1. Angelfish 2. Hogfish 3. Damselfish 4. Jawfish 5. Wrasse 6. Butterflyfish 7. Seahorses 8. Parrotfish 9. Surgeonfish 10. Drum

396,765 62,647 25,298 13,886 22,712 15,402 13,664 17,205 26,728 14,278

Average price per specimen ($) 8.12 8.44 1.19 2.36 1.68 2.35 0.80 5.74 3.47 2.11

Landings

Percentage change in landings, 1990–1998 (%)

48,839 7,419 21,225 5,894 13,512 6,551 16,977 2,998 7,702 6,781

−32 −13 −34 −7 −42 −48 +184 −39 +18 −43

changed in 1998 (Table 3). In 1998, the five most economically important species groups were angelfish (48,839 fish valued at $396,765), hogfish (7,419 fish valued at $62,647), surgeonfish (7,702 fish valued at $26,728), damselfish (21,225 fish valued at $25,298), and wrasses (13,512 fish valued at $22,712). Jawfish dropped out of the top five and surgeonfish jumped from ninth to third in economic importance. With the exception of seahorses and surgeonfish, all species groups exhibited a decline in landings volumes from 1990 to 1998. The largest species group decline was reported for butterflyfish (48%), while seahorses (184%) were the species group with the largest increase. Average per unit prices varied considerably across species. For example, in 1998 the average unit price for angelfish and hogfish both exceeded $8, while the unit price for damselfish, jawfish, wrasse, butterflyfish, and drum were less than $3 (Table 3). The average price for seahorses was less than $1. With the exception of angelfish, the species exhibiting the highest landings volume (i.e., damselfish, wrasse, and seahorses) also exhibited the lowest average unit price. The average unit price for angelfish varied considerably during the 1990–98 period (Figure 4),

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Figure 4. Annual landings and dockside price of angelfish collected in Florida, 1990–98. Table 4. Average annual landings, value, and dockside prices for angelfish species (family Pomacanthidae) collected in Florida, 1990–98 Species

Landings

Value ($)

Unit price ($)

Blue (Holacanthus bermudensis) Queen (Holacanthus ciliaris) Rock Beauty (Holacanthus tricolor) Other (Cherubfish, French, and Grey)∗ Total

19,962 (28%) 8,853 (12%) 22,218 (31%) 20,820 (29%) 71,853 (100%)

146,519 (27%) 127,252 (23%) 100,560 (19%) 168,499 (31%) 542,830 (100%)

7.43 14.41 4.52 8.09

∗ Cherubfish is species Centropyge argi, French is species Pomacanthus paru, and Grey is species Pomacanthus arcuatus.

increasing from $5.60 in 1990 to $9.10 in 1993, before declining to $6.90 in 1995. The unit average price for angelfish then increased to $8.10 in 1998. Prices also varied across individual species of angelfish (Table 4). Six species of angelfish have been landed commercially in Florida. The three most important species of angelfish were the blue, queen and rock beauty. Blue angelfish comprised the largest share of the value (27%) during the 1990–98 period, with an average price of $7.43. Rock beauty angelfish comprised the largest share of the volume (31%), but had an average price of just $4.52 over the same period. Queen angelfish, which represented only 23% of the value and 12% of the volume, had the highest average price of $14.41. Price trends for the individual species were similar to the overall trend exhibited by the angelfish group. Trends in the volume and value of invertebrate landings The approximate 150 individual species of invertebrates collected by the marine life industry in Florida from 1990 to 1998 were grouped, using the same procedure

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as with the finfish, into 32 major species groups (Table 5). Due to the diversity of the invertebrate species, these groups are further aggregated into the following three categories: (1) invertebrate animals (including crustaceans, mollusks, starfish, anemones, sea cucumbers, sponges, nudibranchs, bryozoa, etc.), (2) marine plants, and (3) live rock and live sand. The patterns in invertebrate landings volumes and value during the 1990–98 period varied somewhat across the three major groups (Figures 5 and 6). Landings of invertebrate animals exhibited a steady increase from approximately 850,000 individual animals in 1990 to 3.3 million animals in 1998, an increase of 290%. However, the total dockside value of the animals increased from approximately $376,000 in 1990 to a peak of $1.2 million in 1994, then declined steadily to $896,000 in 1998 as species less valuable on a per unit basis (such as snails, starfish, and sand dollars) garnered an increasing share of the total volume. Landings of plants increased from approximately 31,000 individuals in 1990 to a peak of 37,000 in 1995. Plant landings then declined dramatically (approximately 62%) to 14,000 in 1998. The dockside value of marine plants reached peaks in 1992 and 1995, then declined with landings volumes to $22,000 in 1998. The landings of live rock and live sand mirror the enactment of legislation intended to eliminate the harvest of naturally occurring live rock. Live rock landings increased from approximately 245,000 pounds (110,250 kg) in 1990 to 1.2 million pounds (530,000 kg) in 1995, a 390% increase. Following the moratorium on landings in federal waters, landings decreased to 170,000 pounds (76,500 kg) in 1998. The dockside value of live rock and sand reached equivalent peaks of $1.4 million in 1992 and 1995, then decreased dramatically to $220,000 in 1998 as reported landings were comprised predominantly of live rock cultured on permitted lease sites. Ten species groups accounted for 89% of the total dockside value attributable to invertebrate animals, plants, and live rock and sand during the 1990–98 period (Table 6). The five most important species groups, in descending order of economic importance, were live rock, snails, anemones, crabs, and starfish. The most important single species group was live rock, which accounted for almost 50% of the dockside value accumulated during the 1990–98 period. Snails, anemones,

Table 5. Invertebrate species groups collected by the commercial marine life industry in Florida, 1990–98∗ Anemone (6) Bryozoa Chiton Clam (4) Conch (7) Cowrie (2) Crab (15) Fileclam (2) ∗

Gorgonian (3) Jellyfish (2) Isopod Live Rock (6) Live Sand Lobster (3) Nudibranch (3) Octopus (4)

Oyster Penshell Plant (4) Polychaete (5) Sand Dollar (4) Scallop (2) Sea Biscuit (3) Sea Cucumber (2)

Sea Hare Shrimp (8) Snail (26) Sponge (4) Starfish (8) Tunicates Urchin (5) Whelk (2)

Species groups listed in alphabetical order. Parentheses contain the number of individual species that comprise each group.

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Figure 5. Annual landings of invertebrates collected in Florida by type, 1990–98.

Figure 6. Annual dockside value of invertebrates collected in Florida by type, 1990–98.

and crabs combined accounted for 20% of the value, with the other six species contributing the remaining 30% of the total dockside value. These same ten invertebrate species groups accounted for 83% of the total dockside value during 1998 (Table 7). Live rock continued to represent the most important single species group (in terms of value), though the harvest of live rock was only allowed from approved lease sites during 1998. Snails, crabs, and sand dollars dominated the production volumes. With the exception of live rock and anemones, all of the top ten invertebrate species groups experienced increases in landings

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Table 6. The ten most economically important groups of invertebrate species groups collected in Florida, 1990–98

1. Live Rock 2. Snails 3. Anemones 4. Crabs 5. Starfish 6. Gorgonians 7. Sand Dollars 8. Urchins 9. Sponges 10. Live Sand

Total value 1990–98 ($)

Percentage of total invertebrates value (%)

Cumulative percentage of total invertebrates value (%)

7,357,422 1,262,345 1,128,348 913,848 729,706 685,047 542,991 385,953 349,564 307,662

48.8 8.1 7.2 5.9 4.7 4.4 3.5 2.5 2.2 2.0

48.8 56.8 64.1 70.0 74.7 79.1 82.6 85.1 87.3 89.3

Table 7. 1998 data and change in landings of the economically important invertebrate species groups collected in Florida Invertebrate group

1. Live Rock 2. Snails 3. Anemones 4. Crabs 5. Starfish 6. Gorgonians 7. Sand Dollars 8. Urchins 9. Sponges 10. Live Sand ∗

1998 ∗∗

Value ($)

Average price per specimen∗ (%)

Landings

175,580 166,310 97,061 138,442 47,116 98,374 65,423 68,105 49,243 42,158

1.93 0.21 0.48 0.18 0.09 2.41 0.08 1.67 2.87 0.56

90,975 lbs 805,210 201,629 788,598 511,297 40,743 771,817 40,900 17,166 75,584 lbs

Percentage change in landings, 1990–98 (%) −63 +791 −26 +755 +1,824 +129 +203 +29 +1 NA

Except for live rock and live sand which are expressed as the average price per pound. Number landed except where noted.

∗∗

volumes during the 1990–98 period, with some being dramatic. For example, starfish, snails, and crabs exhibited increases in landings of 1,824%, 791%, and 755%, respectively, from 1990 to 1998. As with finfish species, dockside prices varied across invertebrate species groups. The highest unit prices during 1998 were associated with sponges ($2.87), gorgonians ($2.41), live rock ($1.93 per pound), and urchins ($1.67). DISCUSSION

The marine life collection industry in Florida has grown during the past decade as the number of licensed collectors (i.e., fishers with MLEs) increased from 159 to 743 and either the volume or value of the primary species increased. The growth is particularly evident in the collection of invertebrate animals. The harvest of live

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rock and sand also increased dramatically during the 1990–95 period, but declined due to a moratorium on the collection of naturally occurring rock and sand in state and federal waters. Although the number of harvesting participants increased dramatically during the 1990–98 period, the implementation of a temporary moratorium on marine life endorsements has limited further entry into the industry. The moratorium extends to 2002. Regulations have also been imposed on certain species (e.g., size limits, bag limits, and trip limits), but most regulations apply to the industry as a whole (e.g., acceptable harvesting methods). The implementation of these regulations reflects concern regarding the sustainability of the marine life resources. The information presented in this report represents the only analysis of harvest data collected by Florida Marine Research Institute since the initiation of data collection efforts in 1990. The reported trends in landings provide some insight into the harvest pressure being exerted on wild stocks of ornamental finfish and invertebrate animals. Although no stock assessments exist for any of the individual species targeted by the marine life collection industry, such information (particularly for the predominant species) could be useful to resource managers as they develop effective management measures for this growing industry. Landings data should next be examined in terms of expended effort (i.e., trips) to better determine if the State’s management goals are being achieved. Without such insight, the assurance of the sustainable use of these marine life resources in the face of growing demand by domestic and international markets cannot be ensured. ACKNOWLEDGEMENT

This article was developed under the auspices of the Florida Sea Grant College Program with support from the National Oceanic and Atmospheric Administration, Office of Sea Grant, U.S. Department of Commerce, Grant No. NA76RG0120. REFERENCES Florida Department of State (2000) Marine life. In: Florida Administrative Code Chapter 68B-42. Tallahassee, FL: Division of Elections, pp. 293–300. Florida Marine Fisheries Commission (1998) Florida marine life – proceedings of the limited entry workshop. December 1998. Tallahassee, FL, 12 pp. Florida Marine Research Institute (1999) Unpublished data. St. Petersburg, FL. Hess, D. and Stevely, J. (1978) The aquarium reef fish collecting industry of Monroe County, Florida. Florida Sea Grant Extension Program. Gainesville, FL, 23 pp. The Florida Legislature (1999) Saltwater fisheries. In: 1999 Florida Statutes, Title XXVIII: Natural Resources; Conservation, Reclamation, and Use. Tallahassee, FL: Law Book Services Office, Chapter 370. Pet Industry Joint Advisory Council (1999) U.S. ornamental aquarium industry. Pet Information Bureau. Washington, DC, 2 pp. Address for correspondence: C. M. Adams, Florida Sea Grant College Program, Department of Food and Resource Economics, Institute of Food and Agricultural Sciences, University of Florida, P.O. Box 110240, Gainesville, FL 32611, USA Phone: +1 352-392-1826; Fax: +1 352-392-3646; E-mail: [email protected]