Biological Services Program FWS/OBS-81/24 January 1982 THE ECOLOGY OF THE MANGROVES OF SOUTH FLORIDA: A COMMUNITY PROFILE **" ?lE !•»-> 03 Q) •O c cr. -■■» OJ M II +J -o E ■•- c ^Q. .c ra o a. 2 aiz co ii oj •» * 1. C3 w 03 o >i •r— *• oJ oi +-> a> "o oj o > 3 3 O >> r-13 1. C 03 o en c > 5- C 3 Q. 03 00 r- E >- >i c < 1-TI O 03 ai E i- OJ • •!- S_ itj i- ii a) -o a. 2 •t- or i. to r-~ O ul « i— o ro "a u- s- c cn x os c •i- II r— lO r— CO- « Ul O a. M- c ■f- 1. •!- 3 • +j m T3 S- +-> > O >>••- i. s_ ^.— oj Q. oj ui SZ f— J3 m\ mo > W 3 o c c S- o c • 01 4-> oj 0J C > 03 (J II O E •■- S- s- . en <♦- +J u. c O CD -03 E — I E Ul 0J II - ^ ■!-> CO 03 O- CD 03 E a- .,-,— •t- Z 4-> JD • +-> U l/l Mr- 3 II >> Ul 03 XJ 03 3 O CO ■o C S_ • c n. - >» 03 03 OJ -a l— >1 > =3 +-> s_ o O CD Q. 03 S. ■— 0) E cn U -Q CJ 1- C 03 X S- o3 c 1— 0) Q. E o oj cj c s_ a> m- ai or ai 0) cn Ol c c ro 03 sz sz -a o o o X X SZ aj ai +-> ai Ul Ul s: OS o3 C3 CO Q. Q_ a. 03 cd a. 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Gross primary production (GPP) at different salinities (Hicks and Burns 1975). Mangrove type Average surface sal i ni ty (ppt ) GPP (gC/m2/day) Red Red Red Bl ack Bl ack Black White White 7.8 21 .1 26.6 7.8 21 .1 26.6 21 .1 26.6 8.0 3.9 1 .6 2.3 5.7 7.5 2.2 4.8 21 HIGH COMMUNITY NET PRIMARY PRODUCTION LOW LANDWARD SEAWARD WATERWAY POSITION Figure 6. The hypothetical relationship between waterway position and community net primary production of Florida mangrove forests (based on Carter et al . 1973). 22 radiation within the ambient range. Gross photosynthesis per unit leaf area was greater at the top of the tree canopy than at the bottom, although the middle levels had the greatest production. Miller (1972) concluded by suggesting that the canopy distribution of red man- grove leaves is nearly optimal for ef- ficient water utilization rather than production. This indicates that the cano- py is adapted to maximizing production under conditions of saturated water sup- ply. The mangrove ecosystem model reported by Lugo et al. (1976) provides hypotheses on succession, time to arrive at steady state conditions (see section 3.2), and several aspects of productivity. The model output suggests that the relative amount of tidal amplitude does not affect GPP significantly; instead, GPP appears to be extremely sensitive to inputs of ter- restrial nutrients. It follows that loca- tions with large amounts of nutrient input from terrestrial sources (riverine man- grove communities) have high rates of mangrove production (see section 3.3). All simulation model -generated hypotheses need to be field tested with a particular- ly critical eye, since the simplifying assumptions that are made in constructing the model can lead to overly simplistic answers. Mangrove productivity research re- mains in an embryonic stage. Certain preliminary tendencies or hypotheses have been identified, but much work must be done before we can conclude that these hypotheses cannot be falsified. 2.6 HERBIVORY Direct herbivory of mangrove leaves, leaf buds, and propagules is moderately low, but highly variable from one site to the next. Identified grazers of living plant parts (other than wood) include the white-tailed deer, Odocoileus virginianus, the mangrove tree crab, Aratus pi soni i , and insects including beetles, larvae of lepidopterans (moths and butterflies), and orthopterans (grasshoppers and crickets). Heald (1969) estimated a mean grazing effect on North River red mangrove leaves of 5.1% of the total leaf area; values from leaf to leaf were highly variable ranging from 0 to 18%. Beever et al. (1979) presented a detailed study of grazing by the mangrove tree crab. This arboreal grapsid crab feeds on numerous items including beetles, crickets, cater- pillars, littoral algae, and dead animal matter. In Florida, red mangrove leaves form an important component of the diet. Beever et al. (1979) measured tree crab grazing ranging from 0.4% of the total leaf area for a Florida Keys overwash forest to 7.1% for a fringing forest at Pine Island, Lee County, Florida. The researchers also found that tree crab grazing rates are related to crab density. Low densities (one crab/m ) resulted in low leaf area damage (less than 1% of total leaf area). High densities (four crabs/m ) were accompanied by leaf area damage ranging from 4% to 6% (see section 6.2). Onuf et al. (1977) investigated in- sect herbivory in fringing and overwash red mangrove forests in the Indian River estuary near Ft. Pierce, Florida. They found six major herbivorous insect species, five lepidopteran larvae and a beetle. Comparisons were made at a high nutrient site (input from a bird rookery) and a low nutrient site. Both red man- grove production and leaf nitrogen were significantly higher at the high nutrient site. This resulted in a four-fold greater loss to herbivores (26% of total leaf area lost to grazing); this increased grazing rate more than offset the in- creased leaf production due to nutrient input. Calculations of leaf area damage may underestimate the impact of herbivores on mangroves. For example, the larvae of the olethreutid moth, Ecdytol opha sp., develops within red mangrove leaf buds and causes the loss of entire leaves. All stages of the beetle, Poeci 1 i ps 23 rhizophorae, attack mangrove propagules while still attached to the parent tree (Onuf et al. 1977). 2.7 WOOD BORERS Many people have the mistaken idea that mangrove wood is highly resistant to marine borers. While this may be true to a limited extent for certain mangrove species in other parts of the world, none of the Florida mangroves have borer- resistant wood. Southwell and Boltman (1971) found that the wood of red, black, and white mangroves has no resistance to Teredo, Pholad and Simnorid borers; pieces of red mangrove wood were completely de- stroyed after immersion in ocean water for 14 months. An interesting controversy surrounds the ability of the wood boring isopod, Sphaeroma terebrans, to burrow into the living prop roots of the red mangrove. Rehm and Humm (1973) were the first to attribute apparently extensive damage of red mangroves stands within the Ten Thousand Islands area of southwestern Florida to an isopod, Sphaeroma. They found extensive damage throughout southwest Florida, some infestation north to Tarpon Springs, and a total lack of infestation in the Florida Keys from Key Largo south to Key West. The destruction process was described as follows: the adult isopod bored into the prop roots (5- mm diameter hole); this was followed by reproduction within the hole and develop- ment of juveniles within the root. This process, combined with secondary decompo- sition from fungi and bacteria, frequently results in prop root severance near the mean high tide mark. These authors attributed loss of numerous prop roots and, in some cases, loss of entire trees during storms to isopod damage. The extent of damage in the Ten Thousand Islands region led Rehm and Humm (1973) to term the phenomenon an "eco- catastrophe" of possibly great importance. They further stated that shrinking of mangrove areas appeared to be occurring as a result of Sphaeroma infestation; this point was not documented. Enright (1974) produced a tongue-in- cheek rebuttal, on behalf of Sphaeroma and against the "terrestrial invader", red mangroves. Snedaker (1974) contributed a more substantial argument in which he pointed out that the isopod infestation might be an example of a long-term eco- system control process. Further arguments against the "ecoca- tastrophe" theory were advanced by Estevez and Simon (1975) and Estevez (1978). They provided more life history information for Sphaeroma and suggested a possible ex- planation for the apparently destructive isopod infestations. They found two species of isopods inhabiting red mangrove prop roots, S^. terebrans and a sympatric congener, S. quadridentatum. The latter does not appear to be a wood borer but utilizes S. terebrans burrows. Neither species appeared to utilize mangrove wood as a food source. Estevez and Simon (1975) found extensive burrowing into seedlings in addition to prop root damage. In general, infestations appeared to be patchy and limited to the periphery of mangrove ecosystems. In areas with the highest density of burrows, 23% of all prop roots were infested. There appeared to be more colonization by ^. terebrans in regions with full strength sea water (30 to 35 ppt). The most important finding by Estevez and Simon (1975) and Estevez (1978) was that periods of accelerated activity by ^. terebrans were related to periods of fluc- tuating and slightly increased salinity. This suggests that fluctuations in isopod burrowing may be related to the magnitude of freshwater runoff from the Everglades. These authors agree with Snedaker (1974) and suggest that root and tree loss due to Sphaeroma activity may be beneficial to mangrove ecosystems by accelerating pro- duction and root germination. Simberloff et al. (1978) amplified this last sugges- tion by showing that root branching, which is beneficial to individual trees, is stimulated by isopod activity. 24 This ecocatastrophe versus beneficial stimulus argument is not completely re- solved. Probably, Sphaeroma root destruc- tion, in areas of low isopod density, can be a beneficial process to both the in- dividual tree and to the entire mangrove stand. Whether changes in freshwater runoff have accelerated this process to the point where unnatural and widespread damage is occurring is not clear. The data and research perspective to answer this question do not exist. As a result, we are reduced to providing hypotheses which cannot be tested with available knowledge. 2.8 MANGROVE DISEASES Published research on mangrove diseases is rare. The short paper by Olexa and Freeman (1975) is the principal reference for diseases of Florida man- groves. They reported that black man- groves are affected by the pathogenic fungi , Phyl losti eta hi bi scina and Nigro- spora sphaerica. These authors found that P. hi bi sci na caused necrotic lesions and death of black mangrove leaves. They felt that under conditions of high relative humidity coupled with high temperatures, this fungus could pose a serious threat to individual trees, particularly if the tree had been weakened by some other natural agent, such as lightning or wind damage. Nigrospora sphaerica was considered to be of little danger to black mangroves. Another fungus, Cyl i nrocarpon didymum, appears to form galls on the prop roots and stems of red mangroves. Olexa and Freeman (1975) noted mortality of red mangroves in areas of high gall infesta- tions, although a direct causation link was not proven. Further research on mangrove diseases is badly needed. Viral disease must be investigated. The role of pathogens in litter production and as indicators of mangrove stress may be very important. 25 CHAPTER 3. ECOSYSTEM STRUCTURE AND FUNCTION 3.1 STRUCTURAL PROPERTIES OF MANGROVE FORESTS Published information about the structural aspects of Florida mangrove forests is limited; most existing data have been published since the mid-1970's. This lack of information is unfortunate since quantitative structural data greatly aid understanding of processes such as succession and primary production. Even more important, the response of mangrove forests to stress, both climatic and man- induced, can be followed quantitatively with this type of data. Ball (1980) contributed substantially to understanding the role of competi- tion in mangrove succession by measuring structural factors such as basal area, tree height, and tree density. Lugo and Zucca (1977) monitored the response of mangrove forests to freezing temperatures by observing changes in structural proper- ties of the trees. Baseline studies of forest structure have been published by Lugo and Snedaker (1975), and Pool, Snedaker and Lugo (1977). For example, Lugo and Snedaker (1975) compared a fringing mangrove forest and a basin forest at Rookery Bay, near Naples, Florida. They found the fringing forest, which was dominated by red man- groves, to have a tree diversity of H = 1.48, a basal area of 15.9 mz/ha, an aboveground biomass of 17,932 g/m , and a non-existent litter layer. The nearby basin forest was dominated by black man- groves, had a tree diversity of H = 0.96 and a basal area of 23.4 m /ha. The lit- ter layer in the basin forest averaged 550 dry g/m . Tree diversity in a hurricane disturbed section of the Rookery Bay forest was 1.62. Similar data were pre- sented for mangrove forests in the Ten Thousand Islands area (Table 2). Data of this type are useful for many purposes including impact statements, en- vironmental surveys, and basic scientific questions. Cintron et al. (1978) gave an indication of the direction in which fu- ture research might proceed. Working in a mangrove stand in Puerto Rico, they found tree height to be inversely proportional (r = 0.72) to soil salinity in the range 30 to 72 ppt. Above 65 ppt salinity, dead tree basal area was higher than live tree basal area and above 90 ppt there was no live tree basal area. It should be possible to investigate the relationship between a variety of mangrove structural properties and factors such as flushing frequency, soil depth, nutrient availability, pollution stress, and other measures of human impact. Ulti- mately, this should lead to an ability to predict the form and structure of mangrove forests resulting from various physical conditions or artificial impacts. One example of this potential tool is Ball's (1980) documentation of structural changes in mangrove forests resulting from altera- tions in the hydrological conditions of south Florida. 3.2 Z0NATI0N, SUCCESSION AND "LAND- BUILDING" Much of the world's mangrove litera- ture consists of descriptive accounts of zonation in mangrove forests and the spe- cies composition within these zones. Al- thouqh general agreement has been lacking, various hypotheses have been put forth concerning the possible connection between zonation, ecological succession, competi- tion, and the role of physical factors such as soil salinity and tidal amplitude. In this section we review briefly the dominant ideas about mangrove zonation and succession and present our interpretation of the current status of knowledge. Davis (1940), working in south Flori- da, was one of the first investigators to describe distinct, almost monospecific, zones within mangrove ecosystems. In what has become the classical view, he argued that mangrove zonation patterns were equivalent to serai stages in succession. The most seaward zone, dominated by red mangroves, was regarded as the "pioneer stage". More landward zones were dominated by white mangrove, black mangrove, buttonwood and, finally, the climatic climax, a tropical forest. Since 26 CD CD • s_ ra sz cn -^ -o en c ^ ra 1 — >> to s- 1-1 53 -o C c •i— rO l/l -a 3 ai o LO -C i/i r- CD s- c a. ai X l— 0) a> a> .c 5- +-> 1a ai =3 s- CJ o 4- s- 03 CD <1J . — > <_> O • S-CM - — . en E LT> c r-~ ID LO en E 4- CVI 1 O o CD C/l -a -ii Ul 0) ra ra i/i ■a E CO CD o .a c: •r— jQ CD oo i. "O X> 03 c tz ro 3 in O CD O s- =i en en < — r> CD ra _i > > O E -Q o < ■ S- ra 4- ■a • ■i — CD CVJ S- i- o ro CD 1 — i — u. ro .a 4-> rO n- ro 1— o Q CD i- O CD i. > cn ••- c or: ra E in CD cn o c i- i- cn i- c U_ ra t/> J= CD in > ra o 3 s- s- cn ra O E en -O. O =J S- s- cn o c OO ra E ai E rO Q. o O 10 cn o o O 1 , — ro 10 ro I CO ro o cn i 00 •• •* • i •» r— ro CO 1 r-» LO ro i o CVI o co o o CD 1 CO t — ^> CM •3- LO LO 00 r— LO LO cn r— * - *• •* » CO cn <3- CM f— r-^ i— «3" "3" r-^ ,— O O O 00 ro CO r— ct< i — f~. O f — LCI i — *r CM n it •* * * r-. 00 r-. 00 w— CM i — CTl LO •~ CM CO o o o O CO «3" r~ r» o LO C~. CO CM CM CM CM r~ n *> « * •» Lf) «3" r-^ o r-^ 00 CM LO i-» CM 00 o O O O CO CM LO r~ CO CM cn cn CM r^ cn n «» •» * ■» LO r>- CM CM oo LO CM CM o lO LO o ro o 1 CM «3- co CO CM cy> 1 — CTl CM «3" cn CTV LO « M •» •* •» LO o ■ — CO CO p~ «* 1 — CO CO C3 O o O CO LO CM CO CO 1 — 1 LO CM co cn CO i cn m ■1 * •» i * r^ o , — l~~ 1 LO r-» LO 1— 1 -S3- CM 4-> C31 r- O 1 00 i — ro LO CTl «* 1 o r^ -o CTl r— ■— 1 o in CD > CD in S- 0) o O o o o i- Q. o s_ Q_ i. ■1-J ■u in in i ra O O T- J3 -Q -a ■— c ra 3 ■M O o s- I— m 27 these zones were regarded as progressively later stages in succession, the entire mangrove ecosystem was believed to be moving seaward through a process of sedi- ment accumulation and colonization. Davis based his argument primarily upon the sequence of observed zones and cores which showed red mangrove peat underlying black mangrove peat which, in turn, occurred under terestrial plant communities. Unfortunately, this Clementsian in- terpretation of mangrove zonation was widely accepted, but rarely tested. For example, Chapman (1970) expanded Davis' original successional concept from south Florida to explain zonation in mangrove forests in other parts of the world. Walsh (1974) thoroughly reviewed the man- grove succession/zonation literature. Fortunately, not everyone accepted Davis' point of view. Egler (1952) and later Thorn (1967, 1975) argued that man- grove zonation was a response to external physical forces rather than temporal se- quence induced by the plants themselves. Egler (1952) showed that patterns of sedi- ment deposition predicted by Davis' (1940) theory did not always occur. He also showed that in some cases mangrove zones appeared to be moving landward rather than seaward. Sea level has been rising in south Florida at the rate of 1 ft (30 cm) per 100 to 150 years (Provost 1974). Spackman et al. (1966) emphasized the role of sea level change in determining changes in mangrove zonation, both through sea level rise and land subsidence. Both Egler (1952) and Spackman et al. (1966) along with Wanless (1974) and Thorn (1967, 1975) suggested that mangroves were reacting passively rather than actively to strong geomorphologi cal processes. This implies that mangroves should be regarded as "land-stabilizers" rather than "land- builders". Furthermore, field researchers fre- quently noted that red mangroves were not always the only "pioneer species" on re- cently deposited sediment. It is not unusual to find seedlings of black, white, and red mangroves growing together on a new colonization site. Lewis and Dunstan (1975) found that black mangroves and white mangroves along with the saltmeadow cordgrass, Spartina patens, are often the pioneers on new dredge spoil islands in central Florida. On the northern coast of the Gulf of Mexico, where black mangrove is the only mangrove species present, it may be preceded by marsh grasses such as saltmarsh cordgrass, S. patens, smooth cordgrass, S. al terni fl ora, or the black needle rush, Juncus roemerianus. In Puer- to Rico, we observed that white mangrove often pioneers and dominates sites where oceanic overwash of beach sand has oc- curred. All of these observations detract from Davis' (1940) original contention that red mangroves should be regarded as the initial colonizer of recently de- posited sediments. It appears that under certain conditions, e.g., shallow water depths, substrate type, and latitude, white and black mangroves or marsh grasses can be effective pioneer species. The work of Rabinowitz (1975) added a new perspective to the mangrove zonation debate. Through carefully designed recip- rocal planting experiments in Panamanian mangrove forests using species of Rhi zo- phora , Laguncul a ri a , P e 1 1 i c i e r a and Avicennia, she demonstrated that each species could grow well within any of the mangrove zones. In other words, physical and chemical factors such as soil salinity or frequency of tidal inundation, within each zone, were not solely responsible for excluding species from that zone. To explain zonation, Rabinowitz proposed tidal sorting of propagules based upon propagule size, rather than habitat adap- tation,as the most important mechanism for zonation control. The most recent piece to be added to the zonation/succession puzzle comes from the work of Ball (1980). Based upon re- search of mangrove secondary succession patterns adjacent to Biscayne Bay, Flori- da, she made a strong case for the impor- tance of interspecific competition in controlling zonation. She found that white mangroves, which grow best in intertidal areas, do not occur consis- tently in the intertidal zone of mature mangrove stands. Instead, white mangroves 28 dominate higher, drier locations above mean high water where the red mangrove does not appear to have a competitive advantage. She suggested that competition is not so important during the early stages of succession but becomes critical as individual trees reach maturity and require more space and other resources. Inherent in Ball's concept of zona- tion is the differential influence of physical factors (e.g., soil salinity, depth to water table) on the competitive abilities of the different mangrove species. She concluded that succession proceeds independently within each zone, although breaks in the forest canopy from lightning strikes or high winds may pro- duce a mosaic of different successional stages within a zone. These openings allow species whose seedlings do not com- pete well in shade, such as the white mangrove, to become established, at least temporarily, within solid zones of red mangroves. Zonation of mangrove species does not appear to be controlled by physical and chemical factors directly, but by the interplay of these factors with interspe- cific competition and, possibly, through tidal sorting of propagules. Once succes- sion in a mangrove zone reaches an equili- brium state, change is unlikely unless an external perturbation occurs. These per- turbations range from small-scale distur- bance (lightning strikes) to large-scale perturbations (sea level change, hurricane damage) and may cause succession within zones to regress to an earlier stage. There is some evidence in south Florida that hurricane perturbations occur on a fairly regular basis, creating a pattern of cyclical succession. Except for Ball (1980) and Taylor (1980), the importance of fires as an influence on mangrove succession has been generally ignored. Most fires in the Florida mangrove zone are initiated by lightning and consist of small circular openings in the mangrove canopy (Taylor 1980). These openings present an opportu- nity for secondary succession within an established zone. For example, we have frequently observed white mangroves flourishing in small lightning-created openings in the center of red mangrove forests. Fire may also play a role in limiting the inland spread of mangroves. Taylor (1981) pointed out that Everglades fires appear to prevent the encroachment of red and white mangroves into adjacent herbaceous communities. Finally, Lugo and Snedaker (1974), Cintron et al. (1978) and Lugo (1980) suggested that mangrove ecosystems function as classical successional systems in areas of rapid sediment deposition or upon recently colonized sites such as offshore islands. They concluded that in most areas mangrove forests are an example of steady-state cyclical systems. Concep- tually, this is synonymous to E. P. Odum's (1971) cyclic or catastrophic climax. Chapman (1976a, b) suggested the idea of cyclic succession for a variety of coastal ecosystems. If Florida mangrove ecosystems are cyclic systems, then there should be an identifiable perturbation capable of set- ting succession back to an early stage. Lugo and Snedaker (1974) suggested that hurricanes may play this role. They pointed out (without substantiating data) that major hurricanes occur about every 20-25 years in south Florida. Coinci- dently, mangrove ecosystems appear to reach their maximum levels of productivity in about the same period of time (Lugo and Snedaker 1974). This hypothesis suggests that succession within many mangrove eco- systems may proceed on a cyclical basis rather than in the classical fashion. Possibly other physical perturbations may influence mangrove succession including incursions of freezing temperatures into central Florida, periodic droughts causing unusually high soil salinities (Cintron et al. 1978), and fire spreading into the upper zones of mangrove forests from ter- restrial sources. Although understanding of zonation and succession in mangrove ecosystems remains incomplete, a clearer picture is emerging, at least for south Florida. Contrary to early suggestions, mangrove 29 species zonation does not appear to repre- sent serai stages of succession except, perhaps, for locations of recent coloniza- tion or where sediment is accumulating rapidly. The role of mangroves in land-bui'lding seems more passive than active. Geomorphological and hydrological processes appear to be the dominant forces in determining whether mangrove shorelines recede or grow. The role of mangroves is to stabilize sediments which have been deposited by physical processes. 3.3 NUTRIENT CYCLING Current understanding of nutrient cycles in mangrove ecosystems is far from satisfactory. Sporadic field measurements have been made, but a complete nutrient budget has not been published for any mangrove ecosystem in the world. Several pioneering field studies were conducted in Florida (Carter et al. 1973; Snedaker and Lugo 1973; Onuf et al. 1977) and one simulation model of mangrove nu- trient cycling has been published (Lugo et al. 1976). Preliminary measurements of nitrogen fixation were made (Zuberer and Silver 1975; Gotto and Taylor 1976; Zuberer and Silver 1978; Gotto et al. 1981). Based on these studies, we present the following preliminary conclusions. Mangrove ecosystems tend to act as a sink (net accumulator) for various ele- ments including macro nutrients such as nitrogen and phosphorus, trace elements, and heavy metals. As we have discussed in section 1.7, these elements are removed from waters flowing through mangrove swamps by the concerted action of the mangrove prop roots, prop root algae, the associated sediments, the fine root system of the mangrove trees, and the host of small invertebrates and microorganisms attached to all of these surfaces. Al- though the turnover times for these ele- ments in mangrove swamps are not known, it appears that at least a portion may be stored or tied up in wood, sediments, and peat for many years. Although mangrove ecosystems may tend to accumulate nutrients, there is a con- tinual loss through export of particulate and dissolved substances. If significant nutrient storage and resultant high pri- mary production are to occur, there must be a continual input of nutrients to the mangrove forest from outside the system (Figure 7). Where nutrient influx to the mangrove ecosystem is approximately balanced by nutrient loss in exported organic matter, then nutrient storage will be minimal and mangrove net primary pro- duction will be low. This appears to occur in the scrub mangrove community type and to a lesser extent in the basin and hammock community types. Carter et al. (1973) and Snedaker and Lugo (1973) have hypothesized that the greatest natural nutrient inputs for man- grove swamps come from upland and terres- trial sources. Apparently for this rea- son, the most luxuriant and productive mangrove forests in south Florida occur in riverine locations or adjacent to signifi- cant upland drainage. Localized sources of nutrients, such as bird rookeries, can result in greater nutrient storage and higher mangrove pro- ductivity (Onuf et al. 1977). If however, large bird rookeries (or artificial nu- trient inputs) occur in poorly flushed sections of mangrove ecosystems, resultant high nutrient levels may inhibit mangrove growth (R. R. Lewis, III, Hillsborough Community College, Tampa, Fla.; personal communication 1981). The output from the simulation model of Lugo et al. (1976) suggests that if nutrient input to a mangrove ecosystem is reduced, then nutrient storage levels within the mangrove ecosystem will be reduced and mangrove biomass and produc- tivity will decline. To our knowledge this hypothesis has not been tested in the field. Nitrogen fixation occurs in mangrove swamps at rates comparable to those measured in other shallow, tropical marine areas (Gotto et al. 1981). Nitrogen 30 SMALL IMPORT LOW STORAGE LOW BIOMASS LOW PRODUCTIVITY SMALL EXPORT LARGE IMPORT MODERATE EXPORT HIGH STORAGE HIGH BIOMASS HIGH PRODUCTIVITY Figure 7. The hypothetical relationship between nutrient input (excluding carbon), biomass, primary productivity, and nutrient export (including carbon) from mangrove ecosystems. Top: small nutrient import. Bottom: large nutrient import. 31 fixation has been found in association with mangrove leaves, both living and dead, mangrove sediment surfaces, the litter layer in mangrove swamps, and man- grove root systems (Gotto and Taylor 1976; Zuberer and Silver 1978; Gotto et al. 1981). In virtually all cases, nitrogen fixation appears to be limited by the availability of labile carbon compounds. Perhaps for this reason, the highest rates of mangrove nitrogen fixation have been measured in association with decaying mangrove leaves; presumably, the decaying leaves act as a carbon source and thus accelerate nitrogen fixation. Macko (1981), using stable nitrogen ratio techniques, has indicated that as much as 25% of the nitrogen associated with black mangrove peat in Texas is derived from nitrogen fixation. Zuberer and Silver (1978) speculated that the nitrogen fixation rates observed in Florida mangrove swamps may be suf- ficient to supply a significant portion of the mangrove's growth requirements. Al- though this hypothesis is impossible to test with present information, it might explain why moderately productive mangrove stands occur in waters which are severely nitrogen depleted. In summary, knowledge of nutrient cycling in mangrove swamps is highly speculative. These ecosystems appear to act as a sink for many elements, including nitrogen and phosphorus, as long as a modest input occurs. Nitrogen fixation within the swamp may provide much of the nitrogen needed for mangrove growth. 3.4 LITTER FALL AND DECOMPOSITION Unless otherwise stated, litter fall refers to leaves, wood (twigs), leaf scales, propagules, bracts, flowers, and insect frass (excrement) which fall from the tree. Mangrove leaves are shed con- tinuously throughout the year although a minor peak occurs during the early part of the summer wet season in Florida (Heald 1969; Pool et al. 1975). Sporadic litter fall peaks may follow periods of stress from cold air temperatures, high soil salinities, and pollution events. Litter fall typically can be partitioned as 68% to 86% leaves, 3% to 15% twigs and 8% to 21% miscellaneous; the latter includes flowers and propagules. Litter fall is an important ecosystem process because it forms the energy basis for detritus-based foodwebs in mangrove swamps (see sections 3.5 and 3.6). The first measurements of litter fall in man- grove swamps were made by E.J. Heald and W.E. Odum, working in the North River estuary in south Florida in 1966-69. This was subsequently published as Heald (1969), Odum (1970), and Odum and Heald (1975a). They estimated that litter pro- duction from riverine red mangrove forests averaged 2.4 dry g of organic matter/m /day (or 876 g/m /year or 8.8 metric tons/ha/year). Subsequent studies agreed with this early estimate (Table 3), although varia- tion clearly exists between different types of communities. Scrub forests with scattered, very small trees have the smallest amount of leaf fall. Basin and hammock forests, which appear to be nutrient limited, have intermediate leaf fall values. Not surprisingly, the highest values occur in the highly produc- tive fringing, overwash, and riverine forests. Odum and Heald (1975a) suggested that the relatively uniform litter fall values from productive mangrove forests around the world result from the shade intolerance of the canopy leaves and the tendency for the canopy size to remain the same in spite of increasing height. If detailed information is lacking, red man- grove forests of south Florida, which are not severely limited by lack of nutrients, can be assumed to produce litter fall of 2.0 to 3.0 g/m /day of dry organic matter. Pure stands of black mangroves usually have a lower rate of 1.0 to 1.5 g/nr/day (Lugo et al. 1980). Decomposition of fallen Florida man- grove leaves has been investigated by a number of researchers including Heald (1969), Odum (1970), Odum and Heald (1975a), Pool et al. (1975), Lugo and Snedaker (1975), Twilley (1980) and Lugo et 32 ,— 3 te • 4-> 01 t- > i — o S- ■— o> ro c +J ii to OJ on J- o t- • ^ OJ s_ > rO o -Q t- en -a c c rO ra E n C t/> •p- {. ai i — 3 ■ — o rO f— >+- <4- s- « . oj 00 0J *-> en > ■t-> i- o •n- 3 s- . — *-> en 00 •r— ^ 01 3 CJ +-> s- rO ro 4- 1 — E „ -Q +-> ol II 00 ia CO cu * • »— OJ OO > 00 o ai OJ s- r— T3 CT1 JD 3 C rO i — ro 1— O E 0J u c aj OJ on i- >, ro S_ -C OJ -^ +-> oo 4-> C •i- o r— O fO T- 3 S- C +-> C 0J < E rO ra OJ ^ ■MCM ra i— o ro I— >»- , — >> « — rO fO -o <4- ' — . CM 4- E ro ^■^ O) en u a> a. 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For example, using the litter bag method, they found that only 9% of the original dry weight remained after 4 months in sea water. By comparison, 39% and 54% re- mained at the end of comparable periods in brackish water and freshwater. Under dry conditions, 65% remained. Higher decompo- sition rates in sea water were related to increased activity of shredder organisms, such as crabs and amphipods. Heald (1969) and Odum (1970) also found increases in nitrogen, protein, and caloric content as mangrove leaves pro- gressively decayed. The nitrogen content of leaves decaying under brackish condi- tions (on an AFDW basis) increased from 1.5% (5.6% protein) to 3.3% (20.6% protein) over a 6-month period. Subse- quent information (Odum et al. 1979b) suggested that the protein increase may not have been this great since some of the nitrogen increase probably included non- protein nitrogen compounds such as amino sugars. Fell and Master (1973), Fell et al. (1980), Fell and Newell (1980), and Fell et al. (1980) have provided more detailed information on red mangrove leaf decomposition, the role of fungi in decom- position (see section 4), and nitrogen changes and nitrogen immobilization during decomposition. Fell et al . (1980) have shown that as much as 50% of weight loss of the leaf during decomposition is in the form of dissolved organic matter (DOM). Heald et al. (1979), Lugo et al. (1980) and Twilley (1980) discovered that black mangrove leaves decompose more ra- pidly than red mangrove leaves and ap- parently produce a higher percentage of DOM. Pool et al. (1975) have shown that mangrove litter decomposes and is exported most rapidly from frequently flooded riverine and overwash forests. These communities have little accumulation of litter on the forest floor. Communities which are not as well-flushed by the tides, such as the basin and hammock forests, have slower rates of decomposi- tion and lower export rates. 3.5 CARBON EXPORT Research from Florida mangrove swamps forms a small portion of the larger con- troversy concerned with the extent to which coastal wetlands export particulate organic carbon (reviewed by Odum et al. 1979a). Available evidence from Florida, Puerto Rico and Australia (Table 4) sug- gests that mangrove swamps tend to be net exporters. The values in Table 4 should be regarded as preliminary, however, since all five studies are based upon simplistic assumptions and methodology. Golley et al. (1962) based their annual estimate of particulate carbon export from a Puerto Rican forest upon a few weeks of measurements. Odum and Heald's estimates were derived from two or three measurements a month. All investi- gators have ignored the importance of bed load transport and the impact of extreme events. All investigators except Lugo et al. (1980) have failed to measure DOC flux. It seems relatively clear that man- grove forests do export organic carbon to nearby bodies of water. The magnitude of this export has probably been underesti- mated due to ignoring bedload, extreme events, and DOC. The value of this carbon input to secondary consumers in receiving waters is not clear. As shown in section 3.6, food webs based primarily upon mangrove carbon do exist. The relative importance of mangrove carbon to Florida coastal ecosys- tems remains speculative. We suspect that mangrove-based food webs are dominant in small bays, creeks and rivers within large mangrove ecosystems such as the North River system studied by Heald (1969) and Odum (1970). In intermediate-sized bodies of water, such as Rookery Bay near Naples, Florida, mangroves are probably important but not dominant sources of organic car- bon. Lugo et al. (1980) estimate that mangroves supply 32% of the organic carbon 34 Table 4. Estimates of particulate carbon export from mangrove forests. Lugo et al . (1976) estimated export from a theoreti- cal, steady state forest using a simulation model. Lugo et al . (1980) measured export from an inland black mangrove forest. Investigators Location Export g/m /day tonnes/ha/yr Golley et al . (1962) Heald (1969), Odum (1970)' Lugo and Snedaker (1975) Lugo et al . (1976) Boto and Bunt (1981) Lugo et al. (1980)b ^Estimate only includes carbon of mangrove origin. Estimate includes dissolved and particulate carbon. Puerto Rico 1.1 4.0 Florida 0.7 2.5 Florida 0.5 2.0 Florida 1.5-1.8 5.5 - 6.6 Australia 1.1 4.0 Florida 0.2 0.7 35 input to Rookery Bay. In very large sys- tems, such as Biscayne Bay near Miami, Florida, mangroves are clearly less impor- tant than any other sources such as algae and sea grasses, although mangrove carbon may be important in localized situations such as the immediate vicinity of fringing and overwash forests. The magnitude of mangrove carbon export to unenclosed coastal waters and offshore remains a mystery. 3.6 ENERGY FLOW At least seven sources of organic carbon may serve as energy inputs for consumers in mangrove ecosystems (Figure 8). The pathways by which this energy containing material is processed and made available to each consumer species is indeed complex. Not surprisingly, current understanding of energy flow in Florida mangrove ecosystems exists largely in a qualitative sense; quantitative data are scarce and piecemeal. A variety of inves- tigators have contributed information over the past decade including, but not limited to, Heald (1969), Odum (1970), Odum and Heald (1972), Carter et al. (1973), Snedaker and Lugo (1973), Heald et al. (1974), Lugo and Snedaker (1974, 1975), Odum and Heald (1975a, b), and Pool et al. (1977). Probably, the most complete study to date is the investigation of energy flow in the black mangrove zone of Rookery Bay by Lugo et al. (1980). It is possible at this time to pre- sent a series of hypotheses concerning the relative importance of these energy sources. First, the relative importance of each source can vary from one location to the next. As will be shown in the following discussion, the consumers in certain mangrove forests appear to depend primarily upon mangrove-derived carbon while in other locations inputs from phy- toplankton and attached algae are probably more important. Our second hypothesis is that energy flow based upon phytopl ankton is most important in overwash mangrove forests and other locations associated with large bodies of clear, relatively deep water. Conversely, phytoplankton are hypothesized to be relatively unimportant to the energy budgets of the large riverine forest com- munities along the southwest coast of Florida. It should be remembered, how- ever, that even where phytoplankton are quantitatively unimportant, they poten- tially perform an important function as the basis of phytopl ankton-zoopl ankton- larval fish food webs (Odum 1970). As a third hypothesis, Iver Brook (Rosensteil School of Marine and Atmos- pheric Sciences, Rickenbacker Causeway, Miami, Fla.; personal communication 1979) has suggested that both sea grasses and benthic algae serve as an important energy source for fringing mangrove communities adjacent to large bodies of water such as Biscayne Bay and Whitewater Bay. Although little evidence exists to test this hypo- thesis, observations of extensive deposits of sea grass and macroalgal detritus with- in mangrove forests suggest intuitively that Brook's hypothesis may be correct. In regions where mangrove shading of the prop roots is not severe, our fourth hypothesis suggests that carbon origina- ting from prop root epiphytes may be sig- nificant to community energy budgets. Lugo et al. (1975) have measured net pro- duction of periphyton in mangroves fringing Rookery Ba^ and found average values of 1.1 gC/m /day. Hoffman and Dawes (1980) found a lower value of 0.14 gC/mVday. Because these values are roughly comparable to average exports of mangrove leaf carbon (section 3.5), its potential importance is obvious. The fifth hypothesis states that mangrove organic matter, particularly leaf material, is an important energy source for aquatic consumers. This hypothesis was first espoused by Heald (1969) and Odum (1970), who worked together in the riverine mangrove communities between the Everglades and Whitewater Bay. Clearly, mangrove carbon is of great importance within the riverine and basin communities all along the southwest coast of Florida (Odum and Heald 1975b); Carter et al . (1973) and Snedaker and Lugo (1973) 36 to >> >,*♦- as nj O 3 3 x: >, o •(-> c c tO o <0 t- ac » •r- as .— cox: ■— -r- a. 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OJ IO O IO r— UEIi-r • m c i— c i— - OJ r— O •>- ■— J= > +j i>- in 3 Oj 5 « t- "1-j S- C O .£= «f- 3 2 r- O OJ II cn io r— s- ■— •■- j- a) oj i- o u. -a >> Q.**- CM 51 herons, ibis and the wood stork (Heald et al. 1974). 7.2 RIVERINE FORESTS Tidal streams and rivers, fringed largely by red mangroves, connect the freshwater marshes of south Florida with the shallow estuarine bays and lagoons (Figure 12). Few of these streams have been studied thoroughly. The exception is the North River which flows into White- water Bay and was studied by Tabb (1966) and Odum (1970). Springer and Woodburn (1960) collected fishes in a bayou or tidal pass connecting Boca Ciega Bay and Old Tampa Bay. Carter et al. (1973) reported on the fishes of two tidal streams entering Fahkahatchee and Fahka Union Bays. Nugent (1970) sampled fishes in two streams on the western shore of Biscayne Bay. Characteristics of these areas and sampling gear used by the inves- tigators are summarized in Appendix A. These tidal streams and associated riverine mangrove forests exhibit extreme seasonal variability in both physical characteristics and fish community compo- sition. Salinity variations are directly related to changes in the make-up of the fish assemblage. During the wet season (June - November), salinities fall throughout the water courses and, at some locations in certain heavy runoff years, become fresh all of the way to the mouth (Odum 1970). Opportunistic freshwater species, which are normally restricted to the sawgrass and black needle rush marshes of the headwaters, invade the mangrove zone. These include the Florida gar, Lepi sosteus pi a ty r h i nc us; several centrarchid sunfishes of the genus Lepomis and the largemouth bass, Micropterus sal moi des ; the freshwater catfishes, Ictalurus natalis and Noturus gyrinus; and the killifishes normally considered freshwater inhabitants such as Lucani a goodei and Ri vulus marmoratus. During the dry season (December to early May) salinities rise as a result of decreased freshwater runoff and continuing evaporation. Marine species invade the tidal streams primarily on feeding forays. Examples include the jewfish, Epinephelus ita jara, the stingrays (Dasyatidae), the needlefishes (Belonidae), the jacks (Carangidae), and the barracuda, Sphyraena barracuda. Other seasonal movements of fishes appear to be temperature related. Tabb and Manning (1961) documented move- ments of a number of species from shallow inshore waters to deeper water during times of low temperature stress. The lined sole, the hogchoker, the bighead searobin, and the striped mullet, for example, are much less frequently caught in winter in shallow inshore waters. A third type of seasonality of fish populations in the tidal rivers is related to life cycles. Many of the fish which utilize the tidal stream habitat do so only as juveniles. Thus, there are peaks of abundance of these species following offshore spawning when larval or juvenile forms are recruited to the mangrove stream habitat. In general, recruitment occurs in the late spring or early summer fol- lowing late winter and spring spawning offshore or in tidal passes (Reid 1954). Numerous species are involved in this life cycle phenomenon including striped mullet, grey snapper, sheepshead, spotted sea trout, red drum, and silver perch. The only estimate of fish standing crop from tidal stream habitats is that of Carter et al. (1973). They recorded 27 species weighing 65,891 g (wet wt.) from an area of 734 m or about 90 g/m . This is probably an overestimate since an un- known portion of the fish community had moved from the flooded lowlands to the stream on the ebb tide; sampling occurred at low tide in October. Nonetheless, this is an indication of the high fish standing crop which this mangrove-associated habi- tat can support. The number of species reported from individual tidal streams annually ranges from 47 to 60 and the total from all tidal streams in southwest Florida is 111 species (Appendix B). The food webs in these riverine man- grove ecosystems appear to be predomi- nantly mangrove detritus-based, although the Biscayne Bay stream studied by Nugent 52 Figure 12. Aerial photograph of the mangrove belt of southwest Florida near Whitewater Bay. Note the complex system of pools and small creeks which connect with the tidal river system. 53 (1970) may be an exception. The basic link between the mangrove leaf and higher order consumers is provided by micro- organisms (fungi, bacteria, Protozoa) which colonize the decaying leaf and con- vert them into a relatively rich protein source (Odum 1970; Odum and Heald 1975a). These decaying leaf fragments with asso- ciated microorganisms are fed upon by a group of omnivorous detritivores including amphipods, mysids, cumaceans, ostracods, chironomid larvae, harpacticoid and calanoid copepods, snapping shrimp, caridean and penaeid shrimp, a variety of crabs, filter-feeding bivalves, and a few species of fishes (Odum 1970; Odum and Heald 1972; Odum and Heald 1975b). These detritivores, in turn, are consumed by a number of small carnivorous fishes, which in turn, are consumed by larger piscivorous fishes. The concept of man- grove trophic structure is also discussed in section 3.6. See Appendix B for species specific dietary information. The tidal creeks studied by Nugent (1970) on the western shore of Biscayne Bay differ from the previously discussed streams in the Everglades estuary. The mouths of the Biscayne Bay creeks have dense growths of sea grasses which con- tribute sea grass detritus. The salini- ties are considerably greater and the streams are located only a few kilometers from coral reefs, which are largely absent on Florida's west coast, at least close to shore. As a result, 23 species listed in Appendix B were captured by Nugent (1970) and are not recorded from riverine man- grove habitat on the west coast of Florida. Examples include several of the grunts (Pomadasyidae), the gray trigger- fish, Balistes capriscus, the barbfish, Scorpaena brasiliensis, the scrawled box- fish, Lactophrys quadricorni s, and the snappers, Lutjanus apodus and L. synagris. Riverine mangrove communities and associated tidal streams and rivers are typified by the following families of fishes: killifishes (Cypri nodontidae), livebearers (Poeci 1 i idae), silversides (Atherinidae), mojarras (Gerreidae), tar- pon (Elopidae), snook (Centropomidae), snappers (Lut jani dae) , sea catfishes (Ariidae), gobies (Gobi idae), porgys (Sparidae), mullets (Mugilidae), drums (Sci aenidae), and anchovies (Engraulidae). The mangrove-lined streams and associated pools are important nursery areas for several marine and estuarine species of gamefish. The tarpon, Megalops atlantica, snook, Centropomus undecimalis, and lady- fish, Elops saurus, utilize these areas from the time they reach the estuary as post-larvae, having been spawned offshore. Gray snapper, Lutjanus g r i s e u s , sheepshead, Archosargus probatocephalus, spotted seatrout, Cynoscion nebulosus, and red drum, Sciaenops ocellata, are re- cruited to grass beds of shallow bays and lagoons as post-larvae and enter the mangrove-lined streams for the next sever- al years (Heald and Odum 1970). Of these species, only the spotted seatrout prob- ably spawns in the estuary (Tabb 1966). Other species of commercial or game impor- tance which use the riverine fringing habitat include crevalle jack, gafftopsail catfish, jewfish, striped mojarra, barra- cuda, Atlantic thread herring, and yellow- fin menhaden (Odum 1970). 7.3 FRINGING FORESTS ALONG ESTUARINE BAYS AND LAGOONS Mangrove-fringed estuarine bays and lagoons are exemplified by the Ten Thousand Islands area and Whitewater Bay. Quantitative fish data are available from Fahkahatchee Bay (Carter et al. 1973; Yokel 1975b; Seaman et al. 1973), Fahka Union Bay (Carter et al. 1973), Rookery Bay (Yokel 1975a), the Marco Island Estuary (Weinstein et al. 1977; Yokel 1975a), and Whitewater Bay (Clark 1970). Individual site characteristics are summarized in Appendix A. All except Fahka Union Bay contain significant amounts of sea grasses. Macroalgae domi- nate the benthic producers of Fahka Union Bay. Studies by Reid (1954) and Kilby (1955) near Cedar Key, Florida, were not included in our summary because mangroves are sparse in this area and no mention of mangrove collecting sites were made by these authors. Studies of Caloosahatchee Bay (Gunter and Hall 1965) and of Charlotte Harbor (Wang and Raney 1971) 54 were omitted because the areas studied have been highly modified and because data from many habitats were pooled in the final presentation. All of the bays reviewed in our sum- maries are fringed by dense growths of red mangroves and all contain small mangrove islets. Carter et al. (1973), in their studies of Fahkahatchee and Fahka Union bays, estimated that 57% to 80% of the total energy budget of these two bays is supported by exports of particulate and dissolved organic matter from the man- groves within the bays and inflowing tidal streams. Lugo et al. (1980) estimated that the mangroves surrounding Rookery Bay provide 32% of the energy base of the heterotrophic community found in the bay. Salinities in these bays tend to be higher than in the tidal streams and rivers and the fish assemblages reflect both this feature and the added habitat dimension of sea grass and macro algae beds. Truly freshwater species are rare in these communities and a proportionally greater percentage of marine visitors is present. The dominant fish families of the benthic habitat include drums (Sci aenidae), porgys (Sparidae), grunts (Pomadasyidae), mojarras (Gerreidae), snappers (Lutjanidae), and mullet (Mugili- dae). Other familes with sizeable contri- butions to the benthic fauna include pipe- fishes (Syngnathidae), flounder (Bothi- dae), sole (Soleidae), searobins (Trigli- dae), and toadfishes (Batrachoididae). Numerically abundant fishes of the mid and upper waters include anchovies (Engraul idae), herrings (Clupeidae) and needlefishes (Belonidae). At all loca- tions studied, the benthic fauna was domi- nated by the pinfish, Lagodon rhomboides, the silver perch, Bairdiella chrysura, the pigfish, Orthopristis chrysoptera. and the mojarras, Euci nostomus gula and E. argenteus. The most common midwater and surface species include the two anchovies, Anchoa mitchilli and A_. hepsetus, and two clupeids, Brevoortia smithi and Harengula pensacol ae. The total number of species recorded in the individual studies ranged from 47 to 89; a total of 117 species was collected in these mangrove-fringed bays and lagoons (Appendix B). In none of these studies were the fishes specifically utilizing the fringing mangrove habitat enumerated separately from those collected in the bay as a whole. The collections were most often at open water stations easily sampled by otter trawl. Carter et al. (1973) had two shore seine stations adjacent to mangroves but the data were pooled for publication. Of the four stations in Rookery Bay sam- pled by Yokel (1975a), one was immediately adjacent to the fringing mangrove shore- line and had moderate amounts of sea grasses. The typical pattern which emerges from many estuarine studies is that rela- tively few fish species numerically domi- nate the catch. This is certainly true in mangrove-fringed estuaries. In Rookery Bay (Yokel 1975a) six species comprised 88% of the trawl -catchable fishes, in Fahkahatchee Bay seven species comprised 97% of the catch from three capture techniques (Carter et al. 1973), and in the Marco Island estuary 25 species com- prised 97% of the trawl -catchable fishes (Weinstein et al. 1977). Like tidal river and stream communi- ties, these shallow bays serve as nur- series for numerous species of estuarine- dependent fishes that are spawned off- shore. Based on the distribution and abundance of juvenile fishes of all spe- cies in six habitats, Carter et al. (1973) ranked the mangrove-fringed bays as the most important nursery grounds; the tidal streams were a close second. Shallow bays and tidal streams provide safe nurseries due to seasonally abundant food resources and the low frequency of large predators (Carter et al. 1973; Thayer et al. 1978). The relative lack of large predaceous fishes is probably due to their general inability to osmoregulate in waters of low and/or fluctuating salinity. As in tidal streams, the peak abun- dance of juvenile and larval fishes in the bays is in spring and early summer (Reid 1954). In general, the highest standing 55 crops and the greatest species richness of fishes occur in the late summer and early fall (Clark 1970). Fish densities decline in the autumn and winter as many fishes move to deeper waters. 7.4 FRINGING FORESTS ALONG OCEANIC BAYS AND LAGOONS Mangrove-fringed "oceanic" bays and lagoons are exemplified by Porpoise Lake in eastern Florida Bay (Hudson et al. 1970), western Florida Bay (Schmidt 1979), southern Biscayne Bay (Bader and Roessler 1971), and Old Rhodes Key Lagoon in eastern Biscayne Bay (Holm 1977). Charac- teristics of these sites are summarized in Appendix A. Compared to the mangrove- fringed bays discussed in the previous section, these environments generally ex- hibit clearer water, sandier substrates, and higher and less variable salinities. Closer proximity to the Florida reef tract, the Atlantic Ocean, and the Gulf of Mexico results in a larger potential pool of fish species. These four locations have produced reports of 156 fish species (Appendix B). Mangrove fringes make up a relatively small proportion of these environments; accordingly, their contribution to the bay food webs is probably not very large. Bader and Roessler (1972) estimated that the fringing mangrove community contrib- utes approximately 1% of the total energy budget of southern Biscayne Bay; they considered only mainland mangroves and did not include the small area of mangrove islands. The main ecological role of the fringing mangroves in this type of en- vironment is probably twofold. First, they increase the habitat diversity within an otherwise relatively homogeneous bay system. Second, they provide a relatively protected habitat for juvenile fishes (and certain invertebrates) that later move to more open water or coral reef communities. The second role is analogous to one of the ecological roles of sea grass communities (see Zieman, in prep.) although the fish species involved may be different. Based primarily on habitat designa- tions of Voss et al. (1969), the fishes of Biscayne Bay can be characterized as to preferred habitat. Of the three main habitat types, (1) rock/coral/seawall, (2) grassbed/tidal flat, and (3) mangrove, the grassbed/tidal flat ranked first in fish species occurrences. One hundred and twenty-two of 156 species (79%) are known to occur in this environment. Rock/coral /seawal 1 habitats were fre- quented by 49 species (32%) and mangroves are known to be utilized by 54 species (35%) of the total fish species recorded from this bay. 7.5 OVERWASH MANGROVE ISLANDS In terms of fish-related research, these communities are the least studied of all mangrove community types in south Florida. They are typified by the low- lying mangrove-covered islands that occur in the Florida Keys and Florida Bay and may be overwashed periodically by the tides. Examples include Shell Key, Cotton Key, and the Cowpens. Islands of this type extend southwest from the Florida mainland through the Marquesas. The Dry Tortugas lack well -developed mangrove com- munities although stunted trees are found (Davis 1942). These islands are the most oceanic of any of the mangrove communities discussed. They are characterized by relatively clear water (Gore 1977) and are largely free of the freshwater inflow and salinity varia- tions which characterize other Florida mangrove communities to varying degrees. Numerous statements exist in the litera- ture acknowledging the frequent proximity of mangrove islands to coral reefs and sea grass beds (McCoy and Heck 1976; Thayer et al. 1978). Olsen et al. (1973) workinq in the U.S. Virgin Islands, found 74% to 93% overlap in the fish species composition of fringing coral reefs and shallow mangrove- fringed oceanic bays. Voss et al. (1969) listed fish species that were collected from all three types of communities: fringing mangroves, coral reefs and sea 56 grass beds in Biscayne Bay, but there appears to have been no systematic survey of the fish assemblage characteristic of the mangrove-covered or mangrove-fringed Florida Keys. No one has quantified the faunal connections which we hypothesize exist between the mangroves and sea grasses and between the mangroves and coral reefs. In the absence of published data from the mangrove key communities, only tenta- tive statements can be made. In general, we expect that while mangrove islands serve as a nursery area for juvenile fishes, this function is limited largely to coral reef and marine inshore fishes and not the estuarine-dependent species that we have discussed previously. The latter (juvenile snook, red drum, spotted seatrout) appear to require relatively low salinities not found in association with most of the overwash islands. Casual observation around the edges of these islands suggests that characteristic fishes include the sea bass family (Ser- ranidae), t ri ggerf i shes (Bal istidae), snappers (Lut jam' dae), grunts (Poma- dasyidae), porgies (Span'dae) parrotfishes (Scaridae), wrasses (Labridae), bonefishes (Albulidae), jacks (Carangidae), damsel- fishes (Pomacentridae), and surgeonf ishes (Acanthuridae); many of these fishes occur on or are associated with coral reefs. We also suspect that considerable overlap occurs in the fish assemblage of these mangrove islands and sea grass communi- ties; examples include puffers (Tetrao- dontidae), pipefishes (Syngnathidae) , go- bies (Gobiidae) and scorpionf ishes (Scor- paenidae). Stark and Schroeder (1971) suggested that juvenile gray snapper, which use the fringing mangroves of the keys as shelter during the day, forage in adjacent sea grass beds at night. In the absence of salinity barriers, predatory fishes probably enter the fringes of these mangrove islands on the rising tide. Included in this group are sharks, tarpon, jacks, snook, bonefish and barracuda. 7.6 GRADIENT OF MANGROVE COMMUNITY INTERACTIONS Mangrove communities occur under a wide range of conditions from virtually freshwater at the headwaters of tidal streams to nearly oceanic conditions in the Florida Keys. Attempting to present a single list of fish characteristic of mangrove environments (Appendix B) can be misleading. For this reason we presented the concept of a continuum or complex gradient in Figure 11 and have followed that scheme throughout section 7. The gradient stretches from seasonally fresh to oceanic conditions, from highly varia- ble salinities to nearly constant salini- ty, from muddy and limestone substrates to sandy substrates, from dark-stained and sometimes turbid waters to clear waters, and from food webs that are predominantly mangrove detritus-based to food webs based primarily on other energy sources. Clear- ly, there are other gradients as one moves from north to south in the State of Florida. At the northern end of the State, temperatures are more variable and seasonally lower than in the south. Sedi- ments change from predominantly silicious in central and north Florida to predomi- nantly carbonate in extreme south Florida. Nevertheless, the complex gradient shown in Figure 11, while greatly simplified for graphic purposes, suggests that charac- teristic fish assemblages replace one another along a gradient of changing physical and biogeographic conditions. Such a concept is useful in understanding the factors controlling the composition of fish assemblages associated with mangroves of the four major community types in south Florida. 57 CHAPTER 8. COMMUNITY COMPONENTS - AMPHIBIANS AND REPTILES Food habits and status of 24 species of turtles, snakes, lizards, and frogs of the Florida mangrove region are given in Appendix C. Any of three criteria had to be met before a species was included in this table: (1) a direct reference in the literature to mangrove use by the species, (2) reference to a species as being present at a particular geographical location within the mangrove zone of Florida, and (3) North American species recorded from mangroves in the West Indies or South America, but not from Florida. This last criterion assumes that a species which can utilize mangroves outside of Florida will be able to use them in Florida. Ten turtles are listed of which four (striped mud turtle, chicken turtle, Florida red-bellied turtle, and softshell turtle) are typical of freshwater. Two (mud turtle and the ornate diamondback terrapin) are found in brackish water and the remainder (hawksbill, green, logger- head, and Atlantic ridley) are found in marine waters. Freshwater species usually occur in the headwater regions of mangrove-lined river systems. All four freshwater species are found in habitats other than mangrove swamps including streams, ponds, and freshwater marshes. The brackish water species are found in salt marshes in addition to mangrove swamps. Mangroves, however, are the principal habitat for the ornate diamondback terrapin (Ernst and Barbour 1972). Cam and Goi n (1955) listed two subspecies of the diamondback: Malaclemys terrapin macrospilota and M. t. rhizophorarum. Malaclemys terrapin macro- spilota inhabits the southwest and south- ern coasts, and M. ;t. rhizophorarum is found in the Florida Keys. The two sub- species intergrade in the region of north- ern Florida Bay. All four of the marine turtles are associated with mangrove vegetation at some stage of their lives. Loggerhead and green turtles are apparently much less dependent on mangroves than the remaining two, although we strongly suspect that recently hatched loggerheads may use man- grove estuaries as nursery areas. Green turtles are generally believed to feed on a variety of submerged aquatic plants and sea grasses; recent evidence has shown that they also feed on mangrove roots and leaves (Ernst and Barbour 1972). The Atlantic ridley's preferred habitat is "shallow coastal waters, especially the mangrove-bordered bays of the southern half of the peninsula of Florida" (Carr and Goin 1955). Hawksbill turtles feed on a variety of plant materials including mangrove (especially red mangrove), fruits, leaves, wood, and bark (Ernst and Barbour 1972). Three species in the genus Anol i s have been reported from Florida mangroves: the green anole, the cuban brown anole, and the Bahaman bank anole. All are arboreal lizards that feed on insects. The green anole is widespread throughout the Southeastern United States and is not at all dependent on mangrove swamps. The other two species have much more restricted distributions in the United States and are found only in south Florida. They also are not restricted to mangrove ecosystems. Of the six species of snakes listed, the mangrove water snake (Figure 13) is most dependent upon mangrove habitats. Two important species of reptiles found in mangrove swamps are the American alligator and the American crocodile. The alligator is widespread throughout the Southeastern United States and is only incidentally found in low salinity sec- tions of Florida mangrove areas (Kushlan 1980). The American crocodile is rare; historically its distribution was centered in the mangrove-dominated areas of the upper and lower Florida Keys (particularly Key Largo) and the mangrove-lined shore- lines and mud flats along the northern edge of Florida and Whitewater Bays (Kushlan 1980). Mangroves appear to be critical habitat for this species. Its range has shrunk considerably in south Florida since the 1930's, even though Florida Bay was added to Everglades National Park in 1950 (Moore 1953; Ogden 1978). Much of the decrease in range is due to increased human activity in the Florida Keys. The remaining population centers of the American crocodile are in Figure 13. The mangrove water snake, Nerodja fasciata compressicauda, curled on a red mangrove prop root. Photograph by David Scott. 59 northern Florida Bay and adjacent coastal swamps and the northern end of Key Largo (Ogden 1978; Kushlan 1980). The species uses a variety of habitats for nesting in the Florida Bay region including open hardwood thickets along creek banks, hardwood-shrub thickets at the heads of sand-shell beaches, and thickets of black mangroves behind marl banks (Ogden 1978). On Key Largo the crocodile locates its nests on creek and canal banks in red and black mangrove swamps (Ogden 1978). Man- grove areas thus appear to be important in the breeding biology of this endangered species. Interestingly, only three species of amphibians, to our knowledge, have been recorded in Florida mangrove swamps (Ap- pendix C). This is due to two factors: (1) lack of detailed surveys in low sa- linity swamps and (2) the inability of most amphibians to osmoregulate in salt water. No doubt, several additional species occur in the freshwater -dominated hammock and basin mangrove communities inland from the coast. Possible addi- tional species include: the eastern narrow-mouthed toad, Gastrophryne caro- 1 in en si s, the eastern spadefoot toad, Scaphiopus holbrooki, the cricket frog, Acris gryllus, the green tree frog, Hyl a cinerea, and the southern leopard frog, Rana utricularia. 60 CHAPTER 9. COMMUNITY COMPONENTS - BIRDS 9.1 ECOLOGICAL RELATIONSHIPS Because mangroves present a more diverse structural habitat than most coastal ecosystems, they should harbor a greater variety of birdlife than areas such as salt marshes, mud flats, and beaches (MacArthur and MacArthur 1961). The shallow water and exposed sediments below mangroves are available for probing shorebirds. Longer-legged wading birds utilize these shallow areas as well as deeper waters along mangrove-lined pools and waterways. Surface-feeding and diving birds would be expected in similar areas as the wading birds. The major difference between mangrove swamps and other coastal ecosystems is the availability of the trunks, limbs, and foliage comprising the tree canopy. This enables a variety of passerine and non-passerine birds, which are not found commonly in other wetland areas, to use mangrove swamps. It also allows extensive breeding activity by a number of tree-nesting birds. The composition of the avifauna com- munity in mangrove ecosystems is, in fact, highly diverse. Cawkell (1964) recorded 45 species from the mangroves of Gambia (Africa). Haverschmidt (1965) reported 87 species of birds which utilized mangroves in Surinam (S. America). Ffrench (1966) listed 94 species from the Caroni mangrove swamp in Trinidad while Bacon (1970) found 137 in the same swamp. In Malaya, Nisbet (1968) reported 121 species in mangrove swamps and Field (1968) observed 76 from the mangroves of Sierra Leone (Africa). Use of mangrove ecosystems by birds in Florida has not been recorded in de- tail. Ninety -two species have been ob- served in the mangrove habitat of Sanibel Island, Florida (L. Narcisse, J.N. "Ding" Darling Natl. Wildlife Refuge, Sanibel Is., Fla.; personal communication 1981). Robertson (1955) and Robertson and Kushlan (1974) reported on the entire breeding bird fauna of peninsular south Florida, including mangrove regions. Based on limited surveys, these authors reported only 17 species as utilizing mangroves for breeding purposes. Because their studies did not consider migrants or non-breeding residents, a significant fraction of the avifauna community was omitted. Based on information gleaned from the literature, we have compiled a list of 181 species of birds that use Florida mangrove areas for feeding, nesting, roosting, or other activities (Appendix D). Criteria for listing these species is the same as that used for listing reptiles and amphi- bians (see Chapter 8 of this volume). Often references were found stating that a given species in Florida occurred in "wet coastal hammocks", "coastal wet forests" or the like, without a specific reference to mangroves. These species were not included in Appendix D. Thus, this list is a conservative estimate of the avifauna associated with Florida man- grove swamps. Sources for each listing are provided even though many are redun- dant. Food habit data are based on Howell (1932) and Martin et al. (1951). Esti- mates of abundance were derived from bird lists published by the U.S. Fish and Wildlife Service for the J.N. "Ding" Darling National Wildlife Refuge at Sanibel Island, Florida, and by the Ever- glades Natural History Association for Everglades National Park. Frequently, species were recorded from mangrove swamps at one location, but not the other. We have divided the mangrove avifauna into six groups based on similarities in methods of procuring food. These groups (guilds) are the wading birds, probing shorebirds, floating and diving water- birds, aerially-searching birds, birds of prey, and arboreal birds. This last group is something of a catch-all group, but is composed mainly of birds that feed and/or nest in the mangrove canopy. 9.2 WADING BIRDS Herons, egrets, ibises, bitterns, and spoonbills are the most conspicuous group of birds found in mangroves (Figure 14) and are by far the most studied and best understood. Eighteen species (and one important subspecies) are reported from south Florida mangroves. 61 Figure 14. A variety of wading birds feeding in a mangrove-lined pool near Flamingo, Florida. Photograph by David Scott. 62 Mangrove swamps provide two functions for wading birds. First, they function as feeding grounds. Two-thirds of these species feed almost exclusively on fishes. Although much of their diet is provided by freshwater and non-mangrove marine areas, all of them feed frequently in mangrove swamps. White ibis feed predominantly on crabs of the genus Uca when feeding in mangroves (Kushlan and Kushlan 1975; Kushlan 1979). Mollusks and invertebrates of the sediments are principal foods of the roseate spoonbill although some fish are eaten (Allen 1942). Yellow-crowned night herons and American bitterns eat crabs, crayfish, frogs, and mice in addi- tion to fishes. Snails of the genus Pomacea are fed upon almost exclusively by the limpkin. The sandhill crane is an anomaly in this group since a majority of its food is vegetable matter, especially roots and rhizomes of Cyperus and Sagittaria. Its use of mangroves is probably minimal, occurring where inland coastal marshes adjoin mangroves (Kushlan, unpubl . data). The remaining 12 species are essentially piscivorous although they differ somewhat in the species and sizes of fishes that they consume. Mangrove swamps also serve as breeding habitat for wading birds. With the exception of the limpkin, sandhill crane, and the two bitterns, all wading bird species in Appendix D build their nests in all three species of mangrove trees (Maxwell and Kale 1977; Girard and Taylor 1979). The species often aggregate in large breeding colonies with several thousand nesting pairs (Kushlan and White 1977a). The Louisiana heron, snowy egret, and cattle egret are the most numerous breeders in south Florida mangroves (based on data in Kushlan and White 1977a). In wet years over 90% of the south Florida population of white ibis breed in the interior, freshwater wetlands of the Everglades; during these times the man- groves are apparently unimportant, sup- porting less than 10% of the population (Kushlan 1976, 1977a, b). During drought years, however, production is sustained solely by breeding colonies located in mangroves near the coast (Kushlan 1977a, b). Mangroves are critically important for the survival of the white ibis popula- tion even though they appear to be utilized to a lesser extent than fresh- water habitats. This pattern of larger but less stable breeding colonies using inland marshes and smaller but more stable colonies using mangroves is also charac- teristic of heron populations (Kushlan and Frohring, in prep.). Table 5 gives the number of active nests observed in mangrove regions during the 1974-75 nesting season and the percen- tage this represents of the entire south Florida breeding population for the nine most abundant species of waders and three associated species. The dependence of roseate spoonbills, great blue herons, Louisiana herons, brown pelicans, and double-crested cormorants on mangrove regions is evident. Nesting by the red- dish egret was not quantified during this study although Kushlan and White (1977a) indicated that the only nests of this species which they saw were, in fact, in mangroves. Further observations indicate that this species nests in mangroves ex- clusively (Kushlan, pers. comm.). Similar- ly, the great white heron is highly depen- dent upon mangroves for nesting; they use the tiny mangrove islets which abound along the Florida Keys and in Florida Bay (Howell 1932). During many years the Everglades population of wood storks is known to nest almost solely in mangroves (Ogden et al. 1976); this population comprises approxi- mately one-third of the total south Florida population. Successful breeding of all these mangrove nesters is un- doubtedly correlated with the abundant supply of fishes associated with man- groves. Meeting the energetic demands of growing young is somewhat easier in habi- tats with abundant prey. This is especially important for the wood stork which requires that its prey be concen- trated into small pools by falling water levels during the dry season before it can nest successfully (Kahl 1964; Kushlan et al. 1975; Odgen et al. 1978). Breeding activity by wading birds in mangroves along the southwest and southern Florida 63 Table 5. Nesting statistics of wading birds and associated species in south Florida, 1974-1975 (based on data in Kushlan and White 1977a). Species % of total active Active nests in nests in south mangroves Florida 1914 7 500 100 1335 31 458 92 1812 39 2377 46 71 15 3410 70 2180 13 741 100 White ibis Roseate spoonbill Wood stork Great blue heron Great egret Snowy egret Little blue heron Louisiana heron Cattle egret Brown pelican Double-crested cormorant 1744 83 64 coasts takes place throughout the year (Table 6); at least one species of wader breeds during every month. Colonies on the mangrove islands in Florida Bay were noted to be active nesting sites during all months of the year except September and October (Kushlan and White 1977a). The seasonal movements of wood storks and white ibises between the various south Florida ecosystems were described by Ogden et al. (1978) and Kushlan (1979). Mangrove ecosystems appear to be most heavily used for feeding in summer (white ibis) and early winter (white ibis and wood stork). The remaining species of wading birds appear to use mangrove areas most heavily in the winter months, reflec- ting the influx of migrants from farther north. Wading birds play an important role in nutrient cycling in the coastal man- grove zone. Mclvor (pers. observ.) has noted increased turbidity, greater algal biomass, and decreased fish abundance around red mangrove islets with nesting frigate birds and cormorants. Onuf et al. (1977) reported results from a small (100 bird) rookery on a mangrove islet on the east coast of Florida. Additions of ammonium-nitrogen from the bird's droppings exceeded 1 g/m /day. Water beneath the mangroves contained five times more ammonium and phosphate than water beneath mangroves without rookeries. Although the wading birds were shown to be a vector for concentrating nutrients, it must be noted that this is a localized phenomenon restricted to the areas around rookeries in the mangrove zone. The effect would be larger around larger rookeries. Onuf et al. (1977) also reported that mangroves in the area of the rookery had increased levels of primary production, higher stem and foliar nitro- gen levels, and higher herbivore grazing impact than mangroves without rookeries. Lewis and Lewis (1978) stated that man- groves in large rookeries may eventually be killed due to stripping of leaves and branches for nesting material and by poisoning due to large volumes of urea and ammonia that are deposited in bird guano. This latter effect would be more pronounced in rookeries within mangrove regions subject to infrequent tidal flush- ing. 9.3 PROBING SHOREBIRDS Birds in this group are commonly found associated with intertidal and shal- low water habitats. Wolff (1969) and Schneider (1978) have shown that plovers and sandpipers are opportunistic feeders, taking the most abundant, proper-sized invertebrates present in whatever habitat the birds happen to occupy. Of the 25 species included in this guild (Appendix D), two are year-round residents (clapper rail and willet), two breed in mangrove areas (clapper rail and black-necked stilt), and the remainder are transients or winter residents. Baker and Baker (1973) indicated that winter was the most crucial time for shorebirds, in terms of survival. Coi ncidental ly, winter is the time when most shorebirds use mangrove areas. The invertebrate fauna (mollusks, crustaceans, and aquatic insects) which occur on the sediments under intertidal mangroves forms the principal diet of these species. Willets and greater yellowlegs eat a large amount of fishes, especially Fundulus, in addition to inver- tebrates. Many of the species listed in this guild obtain a significant portion of their energy requirements from other habi- tats, particularly sandy beaches, marshes, and freshwater prairies. Of the species in this guild, the clapper rail is prob- ably most dependent on mangroves for survival in south Florida (Robertson 1955), although in other geographical locations they frequent salt and brackish marshes. 9.4 FLOATING AND DIVING WATER BIRDS Twenty-nine species of ducks, grebes, loons, cormorants, and gallinules were identified as populating mangrove areas in south Florida (Appendix D). Eight species are year-round residents while the remainder are present only during migra- tion or as winter visitors. 65 Table 6. Timing of nesting by wading birds and associated species in south Florida. Adapted from data in Kushlan and White (1977a), Kushlan and McEwan (in press). White ibis Wood stork Roseate spoonbill Great blue/white heron Great egret Little blue heron Cattle egret Double-crested cormorant Brown pelican Months Species SONDJFMAMJJA 66 From the standpoint of feeding, mem- bers of this guild are highly hetero- geneous. Piscivorous species include the cormorant, anhinga, pelicans, and mergan- sers. Herbivorous species include the pintail, mallard, wigeon, mottled duck, and teals. A third group feeds primarily on benthic mollusks and invertebrates. Scaup, canvasback, redhead, and gallinules belong to this group. The ducks in this last group also consume a significant fraction of plant material. Species of this guild are permanent residents and usually breed in mangrove swamps. As shown in Table 5, the brown pelican and double-crested cormorant are highly dependent upon mangroves for nesting in south Florida even though both will build nests in any available tree in other geographical regions. It seems that when mangroves are available, they are the preferred nesting site. The anhinga breeds in mangrove regions but is more commonly found inland near freshwater (J. A. Kushlan, So. Fla. Res. Ctr., Everglades Natl. Park, Homestead, Fla.; personal communication 1981). For the other species listed in this guild, mangrove swamps provide a common but not a required habi- tat; all of these species utilize a variety of aquatic environments. Kushlan et al. (in prep.) provide recent data on the abundance and distribu- tion of 22 species of waterfowl and the American coot in south Florida estuaries. The American coot is by far the most abun- dant species, accounting for just over 50% of the total population. Six species of ducks were responsible for more than 99% of the individuals seen: blue-winged teal (41%), lesser scaup (24%), pintail (18%), American wigeon (9%), ring-necked duck (5%), and shoveler (3%). The major habi- tats included in these authors' surveys were coastal prairie and marshes, mangrove forests, and mangrove-lined bays and waterways of the Everglades National Park. From these data it appears that waterfowl and coots are most abundant in regions where mangrove, wet coastal prairies, marshes, and open water are interspersed. Overall, the Everglades estuaries support from 5% to 10% of the total wi nteri ng waterfowl population in Florida (Goodwin 1979; Kushlan et al. in prep.). As Kushlan et al. point out, however, the Everglades are not managed for single species or groups of species as are areas of Florida supporting larger waterfowl populations. Although the importance of south Florida's mangrove estuaries to continental waterfowl popula- tions may be small, the effect of 70,000 ducks and coots on these estuaries probably is not (Kushlan et al. in prep.). Kushlan (personal communication) thinks that the estuaries of the Ever- glades have an important survival value for some segments of the American white pelican population. In winter, approxi- mately 25% of the white pelicans are found in Florida Bay and 75% in the Cape Sable region. They feed primarily in freshwater regions of coastal marshes and prairies and use mangroves where they adjoin this type of habitat. 9.5 AERIALLY-SEARCHING BIRDS Gulls, terns, the kingfisher, the black skimmer, and the fish crow comprise this guild of omnivorous and piscivorous species (Appendix D). These birds hunt in ponds, creeks, and waterways adjacent to mangrove stands. Many fishes and inverte- brates upon which they feed come from mangrove-based food webs. Only six of the 14 species are year-round residents of south Florida. The least tern is an abun- dant summer resident and the remainder are winter residents or transients. Only the fish crow actually nests in mangroves. Gulls and terns prefer open sandy areas for nesting (Kushlan and White 1977b) and use mangrove ecosystems only for feeding. All of the species in this guild are recorded from a variety of coastal and inland wetland habitats. 9.6 BIRDS OF PREY This guild is composed of 20 species of hawks, falcons, vultures, and owls 67 which utilize mangrove swamps in south are common inhabitants of mangrove areas. Florida (Appendix D). The magnificant This could also be true for the merlin, frigatebird has been included in this which like the peregrine falcon, feeds on group because of its habit of robbing many waterfowl and shorebirds. The remaining of these birds of their prey. Prey con- species in this guild are probably not so sumed by this guild includes snakes, dependent on mangroves; although they may lizards, frogs (red-shouldered hawk, be common in mangrove ecosystems, they swallow-tailed kite), small birds (short- utilize other habitats as well, tailed hawk), waterfowl (peregrine falcon, great-horned owl), fishes (osprey, bald eagle), and carrion (black and turkey 9.7 ARBOREAL BIRDS vultures). This guild is the largest (71 Eleven of these species are permanent species) and most diverse group inhabiting residents, one a summer resident, and the mangrove forests. Included are pigeons, remainder are winter residents. Their use cuckoos, woodpeckers, flycatchers, of mangrove areas varies greatly. The thrushes, vireos, warblers, blackbirds, magnificent frigatebird, which occurs and sparrows. We have lumped this diverse principally in extreme southern Florida group together because they utilize man- and the Florida Keys, utilizes small over- grove ecosystems in remarkably similar wash mangrove islands for both roosts and ways. Invertebrates, particularly nesting colonies. Both species of vul- insects, make up a significant portion of tures are widely distributed in south most of these birds' diets, although the Florida mangrove regions; large colonial white-crowned pigeon, mourning dove, and roosts can be found in mangrove swamps many of the fringilids (cardinal, townee) near the coast. Swallow-tailed kites are eat a variety of seeds, berries, and common over the entire Florida mangrove fruits, region (Robertson 1955; Snyder 1974). Snyder (1974) reports extensively on the As the name given this guild implies, breeding biology of the swallow-tailed these birds use the habitat provided by kites in south Florida. The nests he the mangrove canopy. Many birds also use observed were all located in black man- the trunk, branches, and aerial roots for groves although they do nest in other feeding. Several different types of habitats. searching patterns are used. Hawking of insects is the primary mode of feeding by The bald eagle, osprey (Figure 15), the cuckoos, chuck-wi 1 1 s-wi dows, the and peregrine falcon are dependent upon kingbirds, and the flycatchers. Gleaning mangrove ecosystems for their continued is employed by most of the warblers, existence in south Florida. Both the bald Woodpeckers and the prothonotary warbler eagle and osprey feed extensively on the are classic probers, wealth of fishes found associated with mangrove ecosystems. Additionally, man- Several of the birds in this guild groves are used as roosts and support are heavily dependent upon mangrove areas, structures for nests. Nisbet (1968) indi- The prairie warbler and the yellow warbler cated that in Malaysia the most important are subspecies of more widespread North role of mangroves for birds may be as American species (see Appendix D for wintering habitat for palaearctic mi- scientific names). They are found largely grants, of which the peregrine falcon is within mangrove areas (Robertson and one. Kushlan (pers. comm.) stated that Kushlan 1974). The white-crowned pigeon, recent surveys have shown falcons to mangrove cuckoo, gray kingbird, and black- winter in mangroves, particularly along whiskered vireo are of recent West Indian the shore of Florida Bay where they estab- origin. They first moved into the lish feeding territories. They forage on mangrove-covered regions of south Florida concentrations of shorebirds and water- from source areas in the islands of the fowl. These prey species of the peregrine Caribbean. Confined at first to mangrove 68 Figure 15. Osprey returning to its nest in a red mangrove tree near Whitewater Bay. Photograph by David Scott. 69 swamps, all but the mangrove cuckoo have expanded their range in peninsular Florida by using non-mangrove habitat. In this vein it is interesting to note that many species of rare and/or irregular occur- rence in south Florida are of West Indian origin and use mangroves to a considerable extent. These include the Bahama pintail, masked duck, Caribbean coot, loggerhead kingbird, thick-billed vireo, and stripe- headed tanager (Robertson and Kushlan 1974). Twenty-four of the species in this guild are permanent residents, 27 are win- ter, and 6 are summer residents. Fourteen species are seen only during migrations. 9.8 ASSOCIATIONS BETWEEN COMMUNITY TYPES AND BIRDS MANGROVE Estimating the degree of use of mangrove swamps by birds as we have done (Appendix D) is open to criticism because of the paucity of information upon which to base judgements. Estimating which mangrove community types (see section 1, Figure 4) are used by which birds is open to even more severe criticism. For this reason the following comments should be regarded as general and preliminary. In terms of utilization by avifauna, the scrub mangrove swamps are probably the least utilized mangrove community type. Because the canopy is poorly developed, most of the arboreal species are absent, although Emlen (1977) recorded the red- winged blackbird, hairy woodpecker, north- ern waterthrush, yellow-rumped warbler, common yel 1 owth roat , orange-crowned warbler, palm warbler, yellow warbler, mourning dove, and gray kingbird in scrub mangroves on Grand Bahama Island. Of 25 different habitats surveyed by Emlen (1977), the yellow warbler and gray kingbird were found in the scrub mangroves only. Aerially-searching and wading birds might use scrub mangroves if fishes are present. Overwash mangrove islands are utilized in a variety of ways by all of the bird guilds. Most of the wading birds plus the magnificent f ri gatebi rd, the anhinga, the cormorant, and the brown pelican use overwash islands for nesting (Kushlan and White 1977a). Wading and aerially-searching birds commonly feed in close proximity to overwash islands. A variety of migrating arboreal and probing species use the islands for feeding and roosting. Yellow and palm warblers are common around mangrove islands in Florida Bay as are the black-bellied plover, ruddy turnstone, willet, dunlin, and short- billed dowitcher. Rafts of ducks are common near the inshore islands and birds of prey such as the osprey, the bald eagle, and both vultures use mangrove islands for roosting and nesting. Fringe and riverine mangrove com- munities are important feeding areas for wading and probing birds. Floating and diving and aerially-searching birds use the lakes and waterways adjacent to these mangrove communities for feeding. Many of the wading birds nest in fringe and riverine forests. For example, when the wood ibis nests in coastal areas, it uses these mangrove communities almost exclu- sively (Kushlan, personal communication). Most of the arboreal birds and birds of prey associated with mangroves are found in these two types of communities. This is not surprising since the tree canopy is extremely wel 1 -devel oped and offers roosting, feeding and nesting opportuni- ties. Hammock and basin mangrove communi- ties are so diverse in size, location, and proximity to other communities that it is difficult to make many general statements about their avifauna. Since there often is little standing water in hammock forests, wading and diving birds probably are not common. Proximity to terrestrial communities in some cases may increase the diversity of arboreal species in both hammock and basin forests; proximity to open areas may increase the likelihood of bi rds of prey. It seems safe to conclude that each of the six mangrove community types has some value to the avifauna. This value differs according to community type and 70 kind of bird group under consideration. Certainly, more information is needed, particularly concerning the dependence of rare or endangered species on specific community types. 9.9 MANGROVES AS WINTER HABITAT FOR NORTH AMERICAN MIGRANT LAND BIRDS An interesting observation based on the data in this chapter is the seemingly important role that mangrove ecosystems play in providing wintering habitat for migrants of North American origin. Lack and Lack (1972) studied the wintering warbler community in Jamaica. In four natural habitats including mangrove forest, lowland dry limestone forest, mid- level wet limestone forest, and montane cloud forest, a total of 174, 131, 61, and 49 warblers (individuals) were seen, respectively. When computed on a per hour of observation basis, the difference is more striking with 22 warblers per hour seen in mangroves and only 1, 2, and 1 seen in the other forest habitats, respec- tively. For all passerines considered together, 26 passerines/hour were seen in mangroves with 5, 13, and 3 respectively in the other forest habitats. On a species basis only 9 were recorded from mangroves whereas 19, 13, and 16 species, respectively, were seen in the other habi- tats. This large number of species from the other habitats appears to result from the sighting of rare species after many hours of observation. Only 9 hours were spent by Lack and Lack (1972) in the man- groves whereas between 30 and 86 hours were spent in other habitats. More time in the mangrove zone would have undoubted- ly resulted in more species (and in- dividuals) observed (Preston 1979). Hutto (1980) presented extensive data concerning the composition of migratory land bird communities in Mexico in winter for 13 habitat types. Mangrove areas tended to have more migrant species than most natural habitats (except gallery forests) and also had a greater density of individuals than other habitats (again except for gallery forests). In both Lack and Lack's and Hutto's studies, disturbed and edge habitats had the highest number of species and greatest density of individuals. The percentage of the avifauna community composed of migrants was highest in mangrove habitats, however. From this we can infer the importance of mangroves in the maintenance of North American migrant land birds. 71 CHAPTER 10. COMMUNITY COMPONENTS - MAMMALS Thirty-six native and nine introduced (Layne 1974). Hamilton and Whittaker species of land mammals occur in the south (1979) state that it is the coastal ham- Florida region (Layne 1974; Hamilton and mocks of south Florida, including mangrove Whittaker 1979). Of these, almost 50% (18 areas, which serve to preserve this species) are found in the mangrove zone species in the Eastern United States. (Layne 1974). In addition, two species of Shemnitz (1974) reported that most of the marine mammals are known from mangrove remaining panthers were found in the areas. Data on the abundance and food southwest portion of Florida along the habits of these 20 species are summarized coast and in the interior Everglades in Appendix E. All are permanent resi- regions, dents. The criteria for inclusion in this table are similar to those used for the The extent to which other carnivores avifauna. Sight records in mangroves or use mangrove areas varies widely among locality data from known mangrove areas species. Schwartz (1949) states that were required before a species was in- mink, although rare, prefer mangroves to eluded. This has produced a conservative other coastal habitats in Florida. Layne estimate of the mammal species that uti- (1974, see his figure 1) gives a disjunct lize mangrove areas. distribution for this species in Florida, with the major geographical range being Several mammals do not appear in the southwest coast. River otters also Appendix E because they have not been utilize mangrove habitat heavily. Otters recorded from mangrove swamps in south have been found even far from shore on Florida; however, they occur so widely small mangrove overwash islands in Florida that we suspect they will be found in this Bay (Layne 1974). Gray fox are not depen- habitat in the future. This group dent upon mangroves, although they occa- includes the cotton mouse, Peromyscus sionally use this habitat. Less than 20% gossypinus, the hispid cotton rat, Si g- of all sightings of this species in Ever- modon hispidus, the round-tailed muskrat, glades National Park were from mangroves Neof iber a! leni , the house mouse, Mus (Layne 1974). Bobcat are found in almost musculus, the least shrew, Cryptoti s all habitats in south Florida from pine- parva, and the short -tailed shrew, Blarina lands to dense mangrove forests. The brevicauda. preponderance of recent sightings, how- ever, has been made from the mangrove Few rodents and no bats are included zone, particularly on offshore mangrove in Appendix E. Compared to the rest of overwash islands (Layne 1974). Black bear the State, the south Florida region is are apparently most abundant in the Big deficient in these two groups (Layne Cypress Swamp of Collier County (Shemnitz 1974). Although we have no confirmative 1974) and are rare in the remainder of field data, we suspect that mangrove south Florida, swamps along the central and north Florida coasts contain more mammal species, par- ticularly rodents and bats. The small mammal fauna of the man- grove zone of south Florida are predomi- A number of medium-sized and large nately arboreal and terrestrial species carnivores, including panther, gray fox, which are adapted to periodic flooding, bobcat, striped skunk, raccoon, mink, Opossum, marsh rabbits, cotton rats, and river otter, and black bear, appear to rice rats are commonly found in mangrove utilize south Florida mangrove areas. swamps. The Cudjoe Key rice rat is a Only three of these species (striped newly described species found only on skunk, raccoon, and bobcat) are common in Cudjoe Key in the Florida Keys. This mangroves, but several of the rarer species appears to be closely associated species seem to be highly dependent on with stands of white mangroves (Hamilton mangrove swamps. Of 18 recent sightings and Whittaker 1979). of the panther in Everglades National Park, 15 were from mangrove ecosystems White-tailed deer are common in 72 Florida mangrove swamps, although they utilize many other habitats. The key deer, a rare and endangered subspecies, is restricted to the Big Pine Key group in the Florida Keys, although it ranged onto the mainland in historical times. Al- though this little deer makes use of pine uplands and oak hammocks, it extensively exploits mangrove swamps for food and cover. Two marine mammals, the bottlenose porpoise and the manatee, frequent mangrove-lined waterways. The bottlenose porpoise feeds on mangrove-associated fishes such as the striped mullet, Mugi 1 cephal us. Although the manatee feeds primarily upon sea grasses and other submerged aquatic plants, it is commonly found in canals, coastal rivers, and embayments close to mangrove swamps. Except for the Cudjoe Key rice rat, none of the mammals found in Florida man- groves are solely dependent upon mangrove ecosystems; all of these species can utilize other habitats. The destruction of extensive mangrove swamps would, how- ever, have deleterious effects on almost all of these species. Populations of panther, key deer, and the river otter would probably be the most seriously affected, because they use mangrove habi- tat extensively. 73 CHAPTER 11 VALUE OF MANGROVE ECOSYSTEMS TO MAN Mangrove swamps are often hot, fetid, mosquito-ridden, and almost impenetrable. As a consequence, they are frequently held in low regard. It is possible that more acres of mangrove, worldwide, have been obliterated by man in the name of "recla- mation" than any other type of coastal environment. Reclamation, according to Webster's, means "to claim back, as of wasteland". Mangrove swamps are anything but wasteland, however, and it is impor- tant to establish this fact before a valuable resource is lost. We can think of six major categories of mangrove values to man; no doubt, there are more. 11.1 SHORELINE STABILIZATION AND STORM PROTECTION The ability of all three Florida mangroves to trap, hold and, to some extent, stabilize intertidal sediments has been demonstrated repeatedly (reviewed by Scoffin 1970; Carlton 1974). The contem- porary view of mangroves is that they function not as "land builders" as hypo- thesized by Davis (1940) and others, but as "stabilizers" of sediments that have been deposited largely by geomorphological processes (see section 3.2). Gill (1970), Savage (1972), Teas (1977), and others have emphasized that land stabilization by mangroves is pos- sible only where conditions are relatively quiescent and strong wave action and/or currents do not occur. Unfortunately, no one has devised a method to predict the threshold of physical conditions above which mangroves are unable to survive and stabilize the sediments. Certainly, this depends to some extent on substrate type; mangroves appear to withstand wave energy best on solid rock substrates with many cracks and crevices for root penetration. From our own experience, we suspect that mangroves on sandy and muddy substrates cannot tolerate any but the lowest wave energies, tidal currents much above 25 cm/s, or heavy, regular boat wakes. The concept that the red mangrove is the best land stabilizer has been ques- tioned by Savage (1972), Carlton (1974), and Teas (1977). These authors argue that the black mangrove (1) is easier to transplant as a seedling, (2) establishes its pneumatophore system more rapidly than the red mangrove develops prop roots, (3) has an underground root system that is better adapted to holding sediments (Teas 1977), (4) is more cold-hardy, and (5) can better tolerate "artificial" substrates such as dredge-spoil, finger fills, and causeways. Generally, the white mangrove is regarded as the poorest land stabilizer of the Florida mangroves (Hanlon et al. 1975). Although mangroves are susceptible to hurricane damage (see section 12.1), they provide considerable protection to areas on their landward side. They cannot prevent all flooding damage, but they do mitigate the effects of waves and breakers. The degree of this protection is roughly proportional to the width of the mangrove zone. Very narrow fringing forests offer minimal protection while extensive stands of mangroves not only prevent wave damage, but reduce much of the flooding damage by damping and holding flood waters. Fosberg (1971) suggested that the November 1970 typhoon and accom- panying storm surge that claimed between 300,000 and 500,000 human lives in Bangladesh might not have been so destruc- tive if thousands of hectares of mangrove swamps had not been replaced with rice paddies. 11.2 HABITAT VALUE TO WILDLIFE Florida mangrove ecosystems are important habitat for a wide variety of reptiles, amphibians, birds, and mammals (see sections 8, 9, and 10). Some of these animals are of commercial and sport importance (e.g., white-tailed deer, sea turtles, pink shrimp, spiny lobster, snook, grey snapper). Many of these are important to the south Florida tourist industry including the wading birds (e.g., egrets, wood stork, white ibis, herons) which nest in the mangrove zone. 74 11.3 IMPORTANCE TO THREATENED AND ENDAN- GERED SPECIES The mangrove forests of south Florida are important habitat for at least seven endangered species, five endangered sub- species, and three threatened species (Federal Register 1980). The endangered species include the American crocodile, the hawksbill sea turtle, the Atlantic ridley sea turtle, the Florida manatee, the bald eagle, the American peregrine falcon, and the brown pelican. The endan- gered subspecies are the key deer (Odocoi leus vi rginianus cl avi urn), the Florida panther (Felis concolor coryi ), the Barbados yellow warbler (Dendroica petechia petechia), the Atlantic saltmarsh snake (Nerodia fasciata taeniata) and the eastern indigo snake (Drymarchon corais couperi). Threatened species include the American alligator, the green sea turtle and the loggerhead sea turtle. Although all of these animals utilize mangrove habitat at times in their life histories, species that would be most adversely affected by widespread mangrove destruc- tion are the American crocodile, the Florida panther, the American peregrine falcon, the brown pelican, and the Atlantic ridley sea turtle. The so-called mangrove fox squirrel (Sciurus ni ger avicennia) is widely believed to be a mangrove-dependent endangered species. This is not the case since it is currently regarded as "rare", not endangered, and, further, there is some question whether or not this is a legitimate sub-species (Hall 1981). As a final note, we should point out that the red wolf (Cam's rufus), which is believed to be extinct in Florida, at one time used mangrove habitat in addition to other areas in south Florida. 11.4 VALUE TO SPORT AND COMMERCIAL FISHERIES The fish and invertebrate fauna of mangrove waterways are closely linked to mangrove trees through (a) the habitat value of the aerial root structure and (b) the mangrove leaf detritus-based food web (see sections 6 and 7). The implications of these connections were discussed by Heald (1969), Odum (1970), Heald and Odum (1970), and Odum and Heald (1975b) in terms of support for commercial and sport fisheries. A minimal list of mangrove-associated organisms of commercial or sport value includes oysters, blue crabs, spiny lobsters, pink shrimp, snook, mullet, menhaden, red drum, spotted sea trout, gray and other snapper, tarpon, sheepshead, ladyfish, jacks, gafftopsail catfish, and the jewfish. Heald and Odum (1970) pointed out that the commercial fisheries catch, excluding shrimp, in the area from Naples to Florida Bay was 2.7 million pounds in 1965. Almost all of the fish and shellfish which make up this catch utilize the mangrove habitat at some point during their life cycles. In addi- tion, the Tortugas pink shrimp fishery, which produces in excess of 11 million pounds of shrimp a year (Idyll 1965a), is closely associated with the Everglades estuary and its mangrove-lined bays and rivers. 11.5 AESTHETICS, INTANGIBLES TOURISM AND THE One value of the mangrove ecosystem, which is difficult to document in dollars or pounds of meat, is the aesthetic value to man. Admittedly, not all individuals find visits to mangrove swamps a pleasant experience. There are many others, how- ever, who place a great deal of value on the extensive vistas of mangrove canopies, waterways, and associated wildlife and fishes of south Florida. In a sense, this mangrove belt along with the remaining sections of the freshwater Everglades and Big Cypress Swamp are the only remaining wilderness areas in this part of the United States. Hundreds of thousands of visitors each year visit the Everglades National Park; part of the reason for many of these visits includes hopes of catching snook or gray snappers in the mangrove-lined water- ways, seeing exotic wading birds, croco- diles, or panthers, or simply discovering 75 what a tropical mangrove forest looks like. The National Park Service, in an attempt to accommodate this last wish, maintains extensive boardwalks and canoe trails through the mangrove forests near Flamingo, Florida. In other, more developed parts of the State, small stands of mangroves or mangrove islands provide a feeling of wilderness in proximity to the rapidly burgeoning urban areas. A variety of tourist attractions including Fairchild Tropical Gardens near Miami and Tiki Gardens near St. Petersburg utilizes the exotic appearance of mangroves as a key ingredient in an attractive landscape. Clearly, mangroves contribute intangibly by diversifying the appearance of south Florida. 11.6 ECONOMIC PRODUCTS Elsewhere in the world, mangrove forests serve as a renewable resource for many valuable products. For a full dis- cussion of the potential uses of mangrove products, see de la Cruz (in press a), Morton (1965) for red mangrove products, and Moldenke (1967) for black mangrove products. In many countries the bark of man- groves is used as a source of tannins and dyes. Since the bark is 20% to 30% tannin on a dry weight basis, it is an excellent source (Hanlon et al. 1975). Silviculture (forestry) of mangrove forests has been practiced extensively in Africa, Puerto Rico, and many parts of Southeast Asia (Holdridge 1940; Noakes 1955; Macnae 1968; Walsh 1974; Teas 1977). Mangrove wood makes a durable and water resistant timber which has been used successfully for resi- dential buildings, boats, pilings, hogsheads, fence posts, and furniture (Kuenzler 1974; Hanlon et al. 1975). In Southeast Asia mangrove wood is widely used for high quality charcoal. Morton (1965) mentions that red man- grove fruits are somtimes eaten by humans in Central America, but only by popula- tions under duress and subject to starva- tion. Mangrove leaves have variously been used for teas, medicinal purposes, and livestock feeds. Mangrove teas must be drunk in small quantities and mixed with milk because of the high tannin content (Morton 1962); the milk binds the tannins and makes the beverage more palatable. As a final note, we should point out that mangrove trees are responsible for contributing directly to one commercial product in Florida. The flowers of black mangroves are of considerable importance to the three million dollar (1965 figures) Florida honey industry (Morton 1964). Other than the honey industry, most of these economic uses are somewhat destructive. There are many cases in which clear-cut mangrove forests have failed to regenerate successfully for many years because of lack of propagule dispersal or increased soil salinities (Teas 1979). We believe that the best use of Florida mangrove swamps will continue to be as preserved areas to support wildlife, fishing, shoreline stabiliza- tion, endangered species, and aesthetic values. 76 CHAPTER 12. MANAGEMENT IMPLICATIONS 12.1 INHERENT VULNERABILITY Mangroves have evolved remarkable physiological and anatomical adaptations enabling them to flourish under conditions of high temperatures, widely fluctuating salinities, high concentrations of heavy metals (Walsh et al. 1979), and anaerobic soils. Unfortunately, one of these adap- tations, the aerial root system, is also one of the plant's most vulnerable compo- nents. Odum and Johannes (1975) have referred to the aerial roots as the man- grove's Achi 1 1 es' heel because of their susceptibility to clogging, prolonged flooding, and boring damage from isopods and other invertebrates (see section 6 for a discussion of the latter). This means that any process, natural or man-induced, which coats the aerial roots with fine sediments or covers them with water for extended periods has the potential for mangrove destruction. Bacon (1970) men- tions a case in Trinidad where the Caroni River inundated the adjacent Caroni Mangrove Swamp during a flood and deposited a layer of fine red marl in a large stand of black mangroves which sub- sequently died. Many examples of damage to mangrove swamps from human activities have been documented (see section 12.2). One of the few natural processes that causes periodic and extensive damage to mangrove ecosystems is large hurricanes (Figure 16). Craighead and Gilbert (1962) and Tabb and Jones (1962) have documented the impact of Hurricane Donna in 1960 on parts of the mangrove zone of south Florida. Craighead and Gilbert (1962) found extensive damage over an area of 100,000 acres (40,000 ha). Loss of trees ranged from 25% to 100%. Damage occurred in three ways: (1) wind shearing of the trunk 6 to 10 ft (2 to 3 m) above ground, (2) overwash mangrove islands being swept clean, and (3) trees dying months after the storm, apparently in response to damage to the prop roots from coatings by marl and fine organic matter. The latter type of damage was most widespread, but rarely occurred in intertidal forests, presumably because the aerial roots were flushed and cleaned by tidal action. Fish and invertebrates were adversely affected by oxygen depletion due to accumulations of decomposing organic matter (Tabb and Jones 1962). Hurricane Betsy in 1965 did little damage to mangroves in south Florida; there was also little deposition of silt and marl within mangrove stands from this minimal storm (Alexander 1 967). Lugo et al. (1976) have hypothesized that severe hurricanes occur in south Florida and Puerto Rico on a time interval of 25 to 30 years and that mangrove ecosystems are adapted to reach maximum biomass and pro- ductivity on the same time cycle. 12.2 MAN-INDUCED DESTRUCTION Destruction of mangrove forests in Florida has occurred in various ways including outright destruction and land filling, diking and flooding (Figure 17), through introduction of fine particulate material, and pollution damage, par- ticularly oil spills. To our knowledge there are no complete, published docu- mented estimates of the amount of mangrove forests in Florida which have been destroyed by man in this century. Our conclusion is that total loss statewide is not too great, probably in the range of 3 to 5% of the original area covered by mangroves in the 19th century, but that losses in specific areas, particularly urban areas, are appreciable. This con- clusion is based on four pieces of infor- mation. (1) Lindall and Saloman (1977) have estimated that the total loss of vegetated intertidal marshes and mangrove swamps in Florida due to dredge and fill is 23,521 acres (9,522 ha); remember that there are between 430,000 and 500,000 acres (174,000 to 202,000 ha) of mangroves in Florida (see section 1.3). (2) Birnhak and Crowder (1974) estimate a loss of approximately 11,000 acres (4,453 ha) of mangroves between 1943 and 1970 in three counties (Collier, Monroe, and Dade). (3) An obvious loss of mangrove forests has occurred in Tampa Bay, around Marco Island, in the Florida Keys, and along the lower east coast of Florida. For example, Lewis et al. (1979) estimated that 44% of the intertidal vegetation 77 Figure 16. Damaged stand of red and black mangroves near Flamingo, Florida, as it appeared 7 years after Hurricane Donna. 78 Figure 17. Mangrove forest near Key West as it appeared in 1981 after being destroyed by diking and impounding. 79 including mangroves in the Tampa Bay estuary has been destroyed during the past 100 years. (4) Heald (unpublished MS.) has estimated a loss of 2,000 acres (810 ha) of mangroves within the Florida Keys (not considered by Birnhak and Crowder 1974). So while loss of mangrove ecosys- tems throughout Florida is not over- whelming, losses at specific locations have been substantial. Diking, impounding, and long-term flooding of mangroves with standing water can cause mass mortality, especially when prop roots and pneumatophores are covered (Breen and Hill 1969; Odum and Johannes 1975; Patterson-Zucca 1978; Lugo 1981). In south Florida, E. Heald (pers. comm.) has observed that permanent impoundment by diking which prevents any tidal exchange and raises water levels significantly during the wet season will kill all adult red and black mangrove trees. If condi- tions behind the dike remain relatively dry, the mangroves may survive for many years until replaced by terrestrial vege- tation. Mangroves are unusually susceptible to herbicides (Walsh et al. 1973). At least 250,000 acres (100,000 ha) of man- grove forests were defoliated and killed in South Viet Nam by the U.S. military. This widespread destruction has been docu- mented by Tschirley (1969), Orians and Pfeiffer (1970), Westing (1971), and a committee of the U.S. Academy of Sciences (Odum et al. 1974). In many cases these forests were slow to regenerate; observa- tions by de Sylva and Michel (1974) indi- cated higher rates of siltation, greater water turbidity, and possibly lower dis- solved oxygen concentrations in swamps which sustained the most damage. Teas and Kelly (1975) reported that in Florida the black mangrove is somewhat resistant to most herbicides but the red mangrove is extremely sensitive to herbicide damage. He hypothesized that the vulnerability of the red mangrove is related to the small reserves of viable leaf buds in this tree. Following his reasoning, the stress of a single defoliation is sufficient to kill the entire tree. Although mangroves commonly occur in areas of rapid sedimentation, they cannot survive heavy loads of fine, floculent materials which coat the prop roots. The instances of mangrove death from these substances have been briefly reviewed by Odum and Johannes (1975). Mangrove deaths from fine muds and marl, ground bauxite and other ore wastes, sugar cane wastes, pulp mill effluent, sodium hydroxide wastes from bauxite processing, and from intrusion of large quantities of beach sand have been documented from various areas of the world. 12.3 EFFECTS OF OIL SPILLS ON MANGROVES There is little doubt that petroleum and petroleum byproducts can be extremely harmful to mangroves. Damage from oil spills has been reviewed by Odum and Johannes (1975), Carlberg (1980), Ray (in press), and de la Cruz (in press, b). Over 100 references detailing the effects of oil spills on mangroves and mangrove- associated biota are included in these reviews. Petroleum and its byproducts injure and kill mangroves in a variety of ways. Crude oil coats roots, rhizomes, and pneu- matophores and disrupts oxygen transport to underground roots (Baker 1971). Various reports suggest that the critical concentration for crude oil spills which may cause extensive damage is between 100 and 200 ml/m of swamp surface (Odum and Johannes 1975). Petroleum is readily absorbed by lipophylic substances on sur- faces of mangroves. This leads to severe metabolic alterations such as displacement of fatty molecules by oil hydrocarbons leading to destruction of cellular permea- bility and/or dissolution of hydrocarbons in lipid components of chloroplasts (Baker 1971). As with other intertidal communities, many of the invertebrates, fishes, and plants associated with the mangrove com- munity are highly susceptible to petroleum products. Widespread destruction of organisms such as attached algae, oysters, tunicates, crabs, and gobies have been reported in the literature (reviewed by de 80 la Cruz in press, b; Ray in press). 12.4 MAN-INDUCED MODIFICATIONS Damage from oil spills follows a predictable pattern (Table 7) which may require years to complete. It is impor- tant to recognize that many of the most severe responses, including tree death, may not appear for months or even years after the spill. In Florida, Chan (1977) reported that red mangrove seedlings and black mangrove pneumatophores were particularly sensitive to an oil spill which occurred in the Florida Keys. Lewis (1979a, 1980b) has followed the long-term effects of a spill of 150,000 liters (39,000 gal) of bunker C and diesel oil in Tampa Bay. He observed short-term (72-hour) mortality of inverte- brates such as the gastropod Mel ongena corona and the polychaete Laeonerei s cul veri. Mortality of all three species of mangroves began after three weeks and continued for more than a year. Sub- lethal damage included partial defoliation of all species and necrosis of black mangrove pneumatophores; death depended upon the percentage of pneumatophores affected. In addition to the damage from oil spills, there are many adverse impacts on mangrove forests from the process of oil exploration and drilling (Table 8). This type of damage can often be reduced through careful management and monitoring of drilling sites. Although little is known concerning ways to prevent damage to mangroves once a spill has occurred, protection of aerial roots seems essential. Prop roots and pneumatophores must be cleaned with com- pounds which will not damage the plant tissues. Dispersants commonly used to combat oil spills are, in general, toxic to vascular plants (Baker 1971). If pos- sible, oil laden spray should not be allowed to reach leaf surfaces. Damage during clean-up (e.g., trampling, compac- tion, bulldozing) may be more destructive than the untreated effects of the oil spill (de la Cruz in press, b). In south Florida, man has been re- sponsible for modifications which, while not killing mangroves outright, have al- tered components of the mangrove ecosys- tem. One of the most widespread changes involves the alteration of freshwater runoff. Much of the freshwater runoff of the Florida Everglades has been diverted elsewhere with the result that salinities in the Everglades estuary are generally higher than at the turn of the century. Teas (1977) points out that drainage in the Miami area has lowered the water table as much as 2 m (6 ft). Interference with freshwater inflow has extensive effects on estuaries (Odum 1970). Florida estuaries are no excep- tion; the effects on fish and invertebrate species along the edge of Biscayne and Florida Bays have been striking. The mismanagement of freshwater and its effects on aquatic organisms have been discussed by Tabb (1963); Idyll (1965a,b); Tabb and Yokel (1968) and Idyll et al. (1968). In addition, Estevez and Simon (1975) have hypothesized that the impact of the boring isopod, Sphaeroma terebrans, may be more severe when freshwater flows from the Everglades are altered. One generally unrecognized side effect of lowered freshwater flow and salt water intrusion has been the inland expan- sion of mangrove forests in many areas of south Florida. There is documented evi- dence that the mangrove borders of Biscayne Bay and much of the Everglades estuary have expanded inland during the past 30 to 40 years (Reark 1975; Teas 1979; Ball 1980). Sections of many mangrove forests in south Florida have been replaced by filled residential lots and navigation canals. Although these canal systems have not been studied extensively, there is some evi- dence, mostly unpublished, that canals are not as productive in terms of fishes and invertebrates as the natural mangrove- lined waterways which they replaced. 81 Table 7. General response of mangrove ecosystems to severe oil spills (from Lewis 1980b) Stage Observed impact Acute 0 to 15 days 15 to 30 days Deaths of birds, turtles, fishes, and invertebrates Defoliation and death of small mangroves, loss of aerial root community Chronic 30 days to 1 year 1 year to 5 years 1 year to 1 0 years ( ? ) 10 to 50 years (?) Defoliation and death of medium-sized mangroves (1-3 m), tissue damage to aerial roots Death of large mangroves (greater than 3 m), loss of oiled aerial roots, and regrowth of new roots (often deformed) Recol oni zati on of oil-damaged areas by new seedlings Reduction in litter fall, reduced re- production, and reduced survival of seedl 1 ngs Death or reduced growth of young trees colonizing spill site (?) Increased insect damage (?) 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Q •r- +-> o 3 "0 o t- •r— CL LO •r— O 83 Weinstein et al. (1977) found that artifi- cial canals had lower species diversity of benthic infauna and trawl -captured fishes and generally finer sediments than the natural communities. Courtney (1975) reported a number of mangrove-associated invertebrates which did not occur in the artificial channels. Mosquito production is a serious problem in black mangrove-dominated swamps in Florida (Provost 1969). The salt marsh mosquitos, Aedes taeniorhynchus and A. sol 1 icitans, do not reproduce below the mean high tide mark and for this reason are not a serious problem in the inter- tidal red mangrove swamps. Mosquitos lay their eggs on the damp soil of the irregu- larly flooded black mangrove zone; these eggs hatch and develop when flooded by spring tides, storm tides or heavy rains. As with the "high marsh" of temperate latitudes, there have been some attempts to ditch the black mangrove zone so that it drains rapidly after flooding. Although properly designed ditching does not appear to be particularly harmful to mangrove swamps (other than the area destroyed to dig the ditch and receive the spoil), it is an expensive practice and for this reason is not widely practiced. Properly managed diking can be an effec- tive mosquito control approach with mini- mal side effects to black mangroves (Provost 1969). Generally, ditching or diking of the intertidal red mangrove zone is a waste of money. Mangrove swamps have been proposed as possible tertiary treatment areas for sewage (see discussion by Odum and Johannes 1975). To our knowledge, this alternate use is not currently practiced in south Florida. Until more experimental results are available on the assimilative capacities and long-term changes to be expected in mangrove forests receiving heavy loads of secondary treated sewage, it would be an environmental risk to use mangrove forests for this purpose. In many areas of the world mangrove swamps have been converted to other uses such as aquaculture and agriculture (see de la Cruz, in press, a). Although some of the most productive aquaculture ponds in Indonesia and the Philippines are located in former mangrove swamps, there is some question whether the original natural system was not equally productive in terms of fisheries products at no cost to man (Odum 1974). Conversion to aquaculture and agriculture is cursed with a variety of problems including subsequent land subsidence and the "cat clay" problem. The latter refers to the drastically lowered soil pH which often occurs after drainage and has been traced to oxidation of reduced sulfur compounds (Dent 1947; Tomlinson 1957; Hesse 1961; Hart 1962, 1963; Moorman and Pons 1975). Experience in Africa, Puerto Rico, and Southeast Asia confirms that mangrove forests in their natural state are more valuable than the "reclaimed" land. 12.5 PROTECTIVE MEASURES INCLUDING TRANSPLANTING Protection of mangroves includes (1) prevention of outright destruction from dredging and filling; (2) prevention of drainage, diking and flooding (except for carefully managed mosquito control); (3) prevention of any alteration of hydrologi- cal circulation patterns, particularly i nvol ving tidal exchange; (4) prevention of introduction of fine-grained materials which might clog the aerial roots, such as clay, and sugar cane wastes; (5) preven- tion of oil spills and herbicide spray driftage; and (6) prevention of increased wave action or current velocities from boat wakes, and sea walls. Where mangroves have been destroyed, they can be replanted or suitable alter- nate areas can be planted, acre for acre, through mitigation procedures (see Lewis et al. 1979). An extensive body of literature exists concerning mangrove planting techniques in Florida (Savage 1972; Carlton 1974; Pulver 1976; Teas 1977; Goforth and Thomas 1979; Lewis 1979b). Mangroves were initially planted in Florida at least as early as 1917 to protect the overseas railway in the Florida Keys (Teas 1977). Both red and black mangroves have 84 been used in transplanting. As we men- tioned in section 11, black mangroves seem to have certain advantages over red man- groves. Properly designed plantings are usually 75% to 90% successful, although the larger the transplanted tree, the lower its survival rate (Teas 1977). Pruning probably enhances survival of trees other than seedlings (Carlton 1974). Important considerations (Lewis 1979b; Teas 1977) in transplanting mangroves are: (1) to plant in the intertidal zone and avoid planting at too high or too low an elevation, (2) to avoid planting where the shoreline energy is too great, (3) to avoid human vandalism, and (4) to avoid accumulations of dead sea grass and other wrack. Costs of transplanting have been variously estimated. Teas (1977) suggests $462 an acre ($l,140/ha) for unrooted propagules planted 3 ft (0.9 m) apart, $1,017 an acre ($2,500/ha) for established seedlings planted 3 ft (0.9 m) apart and $87,500 ($21 6,130/ha) for 3 year-old nur- sery trees planted 4 ft (1.2 m) apart. Lewis (1979b) criticized Teas' costs as unrealistically low and reported a project in Puerto Rico which used established seedlings at a cost of $5,060 an acre ($12,500/ha); he did suggest that this cost could be cut in half for larger projects. 12.6 ECOLOGICAL VALUE OF BLACK VS. RED MANGROVES One unanswered question of current interest in Florida concerns the ecologi- cal value of black mangrove forests com- pared to intertidal red mangrove forests. In many respects, this is identical to the "high marsh" versus "low marsh" debate in temperate wetlands. One hypothetical argument which has been presented fre- quently in court cases during the past decade suggests that black mangrove forests have less ecological value than red mangrove forests to both man and coastal ecosystems. This argument is based on an apparent lack of substantial particulate detritus export from black mangrove forests above mean high tide and the generally perceived lack of organisms, particularly gamefishes, which use black mangrove forests as habitat. The counter argument states that black mangrove forests are important for the support of wildlife and the export of substantial quantities of dissolved organic matter (DOM). Lugo et al. (1980) provide evidence that black mangrove forests do, in fact, export large quanti- ties of DOM. They point out that (1 ) black mangrove leaves decompose more rapidly than red mangrove leaves and thus produce relatively more DOM and (2) abso- lute export of carbon from these forests, on a statewide scale, is equal or greater than from red mangrove forests. 12.7 THE IMPORTANCE OF INTER-COMMUNITY EXCHANGE From previous discussions (sections 6 and 7.5 and Appendices B, C, D and E) it is clear that many species of fishes, invertebrates, birds, and mammals move between mangrove forest communities and other habitats including sea grass beds, coral reefs, terrestrial forests, and the freshwater Everglades. For example, the gray snapper, Lut janus gri seus, spends part of its juvenile life in sea grass beds, moves to mangrove-lined bays and rivers, and then migrates to deeper water and coral reefs as an adult (Croaker 1962; Starck and Schroeder 1971). The pink shrimp, Penaeus duorarum, spends its juve- nile life in mangrove-lined bays and rivers before moving offshore to the Tortugas grounds as an adult. During its juvenile period it appears to move back and forth from mangrove-dominated areas to sea grass beds. The spiny lobster, Panulirus argus, as a juvenile frequently uses mangrove prop root communities as a refuge; when nearing maturity this species moves to deeper water in sea grass and coral reef communities (see discussion section 6.1). Many of the mammals (sec- tion 10) and birds (section 9) move back and forth between mangrove communities and a variety of other environments. These are only a few of many 85 examples. Clearly, mangrove ecosystems are linked functionally to other south Florida ecosystems through physical pro- cesses such as water flow and organic carbon flux. As a result, the successful management and/or preservation of many fishes, mammals, birds, reptiles, and amphibians depends on proper understanding and management of a variety of ecosystems and the processes that link them. Saving mangrove stands may do the gray snapper little good if sea grass beds are destroyed. Pink shrimp populations will be enhanced by the preservation of sea grass beds and mangrove-lined waters, but shrimp catches on the Tortugas grounds will decline if freshwater flow from the Everglades is not managed carefully (Idyll et al. 1968). Successful management of south Florida mangrove ecosystems, including their valuable resources, will depend on knowledgeable management of a number of other ecosystems and the processes which link them. 12.8 MANAGEMENT PRACTICES: PRESERVATION Based on years Florida and based of research in south on the information reviewed for this publication, we have concluded that the best management prac- tice for all types of Florida mangrove ecosystems is preservation. Central to this concept is the preservation of adjacent ecosystems that are linked signi- ficantly by functional processes. The continued successful functioning of the mangrove belt of southwest Florida is highly dependent on the continual exis- tence of the Everglades and Big Cypress Swamp in an ecologically healthy condi- tion. At no cost to man, mangrove forests provide habitat for valuable birds, mam- mals, amphibians, reptiles, fishes, and invertebrates and protect endangered species, at least partially support exten- sive coastal food webs, provide shoreline stability and storm protection, and generate aesthetically pleasing experi- ences (Figure 18). In situations where overwhelming economic pressures dictate mangrove destruction, every effort should be made to ameliorate any losses either through mitigation or through modified development as described by Voss (1969) and Tabb and Heald (1973) in which canals and seawalls are placed as far to the rear of the swamp as possible. 86 Figure 18. Mangrove islands in Florida Bay near Upper Matecumbe Key. Note the extensive stands of seedling red mangroves which have become established (1981) after a long period without major hurricanes. Mangrove islands in the Florida Keys tend to expand during storm- free intervals. 87 REFERENCES Abele, L.G. 1974. 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Zuberer, D.A., and W.S. Silver. 1975. Mangrove-associated nitrogen fixa- tion. Pages 643-653 2D. S. Walsh, S. Snedaker and H. Teas, eds. Proceed- ings of the international symposium on the biology and management of man- groves. Univ. of Florida, Gaines- ville. Zuberer, D.A., and W.S. Silver. 1978. Biological nitrogen fixation (acety- lene reduction) associated with Florida mangroves. Appl. Environ. Microbiol. 35: 567-575. 105 APPENDIX A. Summary of the site characteristics and sampling methodology for fishes in: A-l - mangrove- fringed tidal streams and rivers, A-2 - mangrove-lined estuarine bays and lagoons, and A-3 - mangrove- lined oceanic bays and lagoons . 106 V) 0> s- o 4- >> a Ol CO o s- t— 01 o > ■o •r— o t- JZ +J-o 0) C B *> O) 10 c E •I— <0 ^- 0) Q. S. 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C 00 id QJ -r- O 3 0J QJ r0 IC s. o S (T3 S- ^l 2 QJ QJ 00 •o JZ (^ 4-> ^ +J c 00 4-> fO 00 C fO c c a> OO aj oo aj O oO E 00 E -Q cn S- TD cn S- -p « ■o *D C 03 fO 3 V- 3 o LD c X CL<-H 1 c o It) 03 QJ • 1 iT3 • O O s- S-OCM CC r-^ o o u- CD CO C CM OJ 3 enrs i Q.cn < «i -* o 00 i- 0J o OJ c > 3N ■D CO i-( "O O f-v 00 (T3 s_ C A3 CU C n3CM +J j: 1 r- S--DET3 or _i O «i- O x: fl3 S- 00 OJ -i- ^^ -r- E as. wo +j U OJ oO t— i 4J S- E TD >>^ 1- OT3N 3 oo -a cu r^ w OiO£ r- ai o Or- 3Cft O ■1- i at >, Reference Diet Diet Reference Comments Orectolobidae - carpet sharks Ginglymostoma cirratum nurse shark 5, 7 Fish, cephalopods, molluscs, shrimp, sea urchins Randall 1967 Clark & von Schmidt 1965 Bohlke & Chaplin 1968 Carcharhinidae - requiem sharks Carcharhinus leucas bull shark Carcharhinus limbatus blacktip shark Negaprion brevirostris lemon shark 11 4, 5, 7 Juveniles: fish (Arius felis, Lophoqobius , Mugil cephalus , Brevoortia patronus, Micropogon undulatus) , crus- taceans including penaeid shrimp, blue crabs Fish (Caranx sp., Centropomus undecimalis, Chilomycterus schoepfi, Arius felis, Lacto- phrys trigonnus Lagodon rhomboides) , crabs Young: crustaceans, fish Adults: fish, crustaceans Odum 1971 Clark & von Schmidt 1965 Randall 1967 Clark & von Schmidt 1965 Sphyrnidae - hammerhead sharks Sphyrna tiburo - bonnethead + + 2, 5 Mantis shrimp, shrimp, isopods, Bohlke s barnacles, bivalve molluscs, Chaplin 1968 cephalopods, fish Pristidae - sawfishes Pristis pectinata smalltooth sawfish + 5, 15 Fish, benthic crustaceans Bohlke s Chaplin 1968 Rhinobatidae - guitarfishes Rhinobatos lenti- guitarfish Torpedinidae - electric rays Narcine brasiliensis • + 1 + + 1, 17, 18 lesser electric ray Rajidae - skates Raja texana roundel skate Crustacea, fish, annelids Reid 1954 Dasyatidae - stingrays Dasyatis americana - southern stingray Dasyatis sarjina Atlantic stingray + + 2, 4, 5, 7 2, 8, 13, 17 Fishes, sipunculid and poly- chaete worms , crabs , bivalves shrimp , mantis shrimp Benthic invertebrates inclu- ding bivalves , xanthid and portunid crabs , shrimps , amphipods, annelids, chirono- mid larvae Randall 1967 Darnell 1958 Gymnura micrura - smooth butterfly ray Urolophus jamaicensis ■ yellow stingray 17 Fish, molluscs, annelids, shrimp, other small crustaceans 1 Probably small burrowing invertebrates Peterson s Peterson 1979 Bohlke s Chaplin 1968 This and all subsequent Odum 1971 citations refer to W.E. Odum 1971. Ill Family and Species Habitat Type *-> > . Reference Diet Reference Comments Myliobatidae - eagle rays Aetobatus narinari - Spotted eagle ray Lepisosteidae - gars Lepisosteus platyrhincus - + Florida gar 2, 1, 15 13, Clams, oysters Boh Ike & Chaplin 1968 Fish (poeciliids, cyprinodonts , Odum 1971 small centrarchids) , crustaceans (caridean shrimp), insect larvae Elopidae - tarpons El ops saurus - ladyfish 2, 3, 7, 8, 13, 15 < 45 mm: zooplankton, chaeto- gnaths, polychaete worms > 45 mm: caridean & penaeid shrimp, various small fish Odum 1971 Austin & Austin 1971 Mega lops atlantica - tarpon , 8, 13, < 45 mm: plankton (cyclopoid 15 copepods) juveniles: fish (Gambusia, Fundulus heteroclitus , Mugil cephalus) , crustaceans (ostra- cods, caridean shrimp) adults: wide variety of fish, crabs , shrimp , ctenophores , insects Odum 1971 Austin & Austin 1971 Obligate air breathe rs . Juv- eniles inhabit shallow brackish pools low in oxygen, often containing H,S (Wade 1962) Albulidae - bone fishes Albula vulpes - bonefish Anguillidae - eels Anguilla rostrata - American eel Ophichthidae - snake eels Myrophis punctatus speckled worm eel 4, 5 8, 13 2, 3, 17, 18 Clams, snails, shrimp, small fish 50-200 mm: amphipods, isopods 180-472 mm: xanthid crabs,, caridean shrimp, fish ( Lophogobi us cyprinoides) Polychaetes, Branch iostoma caribaeum, sand crabs Bohlke & Chaplin 1968 Odum 1971 Springer & Woodburn 1960, Reid 1954 Members of this family burrow in mud or sand, undersampled by most methods (Bohlke & Chaplin 1968) Bascanichthys scuti- caris - whip eels Ophichthus gome si - shrimp eel Clupeidae - herrings Brevoortia smithi - yellowfin sardine 3, 17 2, 5, 17 Brevoortia patronus Gulf menhaden 38-48 mm: phy toplankton , zoo- Darnell plankton, plant fragments , 1958 detritus 85-103 mm: organic matter, silt, diatoms, foraminiferans, copepods 112 Family and Species Habitat Type a Reference Diet Diet Reference Harengula pensacolae scaled sardine 2, 3, 8, 30 mm: planktonic copepods, 13 zoea, nauplii, larval fish 64-96 mm: amphipods, harpactlcoid copepods, isopods mysids, chironomid larvae Odum 1971 Opisthonema oglinum - Atlantic thread herring 2, 3, 5, 13, 17 Copepods, polychaetes, shrimp, fishes, crab larvae, mysids Odum 1971 Sardinella anchovia Spanish sardine 17 Engraulidae - anchovies Anchoa cub ana - Cuban anchovy 2, 16 Ostracods, copepods Springer & Woodburn 1960 Anchoa hepsetus striped anchovy 2, 3, 13, 32-114 mm: copepods, isopods, Springer & 16, 17 mysids, caridean shrimp, small Woodburn bivalves 1960 Anchoa lampro taenia bigeye anchovy Anchoa mitchilli - bay anchovy 1, 2, 3, <25 mm: microzooplankton Odum 1971 5, 7, 8, 31-62 mm: amphipods, zooplank- 13, 16-18 ton, mysids, ostracods, plant detritus, copepods, small molluscs, chironomid larvae Synodontidae - lizardfishes Synodus foetens - inshore lizardfish 1-3, 5, 8, Small fish, crabs, shrimp, 17, 18 polychaete worms Odum 1971 Catostomidae - suckers Erimyzson sucetta " lake chubsucker A freshwater stray Ictaluridae - freshwater catfish Ictalurus natal is - yellow bullhead 14 A freshwater stray Noturus gyrinus tadpole mad torn 14 A freshwater stray Arriidae - sea catfishes Arius felis - sea catfish + 2, 3, 5, 100 mm: copepods, zooplankton 7, 8, 13, amphipods, mysids, chironomid 17 larvae, isopods, small crabs 100-200 mm: benthic inverte- brates 200-330 mm: crabs, amphipods, mysids, fishes, bark, crayfish, caridean and penaeid shrimp Odum 1971 Bagre marinus - gafftopsail catfish 2, 8, 17 262-445 mm: blue crabs, small fishes Odum 1971 Batrachoididae - toadfishes Opsanus beta - Gulf toadfish 1-3, 5, 18-60 mm: amphipods, chironomid 7, 12, 13, larvae, mysids, isopods, few fish 15, 17, 18 >60 mm: caridean shrimp, xanthid crabs, snapping shrimp, mussels, fish, mangrove bark Odum 1971 Salinities 10 o/oo > (Odum 1971) 113 Family and Species at Type Htn ma o Manning 1961) Syngnathus pelagicus sargassum pipefish Centropomidae - snooks C e n t r opojmus parallelus fat snook Family as a whole shows preference for estuarine man- grove habitat (Rivas 1962) Centropomus pectinatus tarpon snook 7, 8, 13 116 Habitat Type Family and Species u « Reference Diet Diet Reference Centropomus undecimalis + snook 2, 5, 7 , 8, Juveniles: caridean shrimp, Odum 1971 11, 13, 14 small cyprinodont fishes, Austin & gobies, mojarras Austin Adults: fish, crabs, penaeid 1971 shrimp, crayfish, snapping shrimp By far most abundant of three species (Rivas 1962) Serranidae - sea basses Centropristis striata black seabass Diplectrum formosum sand perch Epinephelus itajara jewfish Epinephelus morio red grouper 2, 3, 11, 16-18 2, 5, 7, 8, 11, 13, 15 Family in general carnivorous on fish, crustaceans Caridean & penaeid shrimp, copepods, crabs, fish Juveniles: penaeid shrimp, xanthid crabs 228-340 mm: crustaceans, crabs, fishes Randall 1967 Re id 1954 Odum 1971 The most abundant of the seabasses in mangrove habitats Randall 1967 Epinephelus striatus Nassau grouper 11 170-686 mm: fish, crabs, stomatopods , cephalopods , shrimp, spiny lobsters, gastropods, bivalves, isopods Randall 1967 Hypoplectrus puella barred hamlet 11 54-98 mm: snapping shrimp, Randall crabs, fish, mysids, stomato- 1967 pods , isopods Mycteroperca microlepis gag Centrarchidae - sunfishes Elassoma evergladei Everglades pygmy sunfish + 1, 2, 5, 11, 71-100 mm: 17, 18 fish penaeid shrimp. Reid 1954 Family is primarily freshwater , fish occasionally enter headwater area of mangrove- fringed stream Lepomis auritus redbreast sunfish Lepomis gulosus warmouth Shrimp (Palaemonetes), fish ( Gob io soma bosci , Lepomis macrochirus) , detritus, Vallisneria, amphipods , xan- thid crabs, blue crabs Desselle et al. 1978 Diet from Lake Pontchartrain salinities 1.6- 4 . 1 o/oo Lepomis macrochirus bluegill 2, 15 Amphipods, blue crab (Cal- linectes sapidus) , xanthid crabs, detritus, Vallisneria, clams (Rangia cuneata) , sponge (Ephydatia fluviatilis) , barnacles, insect larvae Desselle et al. 1978 Diet from Lake Pontchartrain salinities 1.6- 4.1 o/oo Lepomis microlophus redear sunfish 2, 13-15 Chironomid larvae, amphipods, xanthid crabs, clam (Rangia cuneata) , sponge (Ephydatia fluviatilis) , detritus Desselle et al. 1978 Diet from Lake Pontchartrain salinities 1.6- 4 . 1 o/oo Lepomis punctatus spotted sunfish 8, 14, 15 Cladocerans, small crabs, odum 1971 mysids , chironomids , amphipods , insects, molluscs, isopods, fish, algae Salinities < 15 o/oo (Odum 1971) 117 Family and Species Habitat Type ■32 3 3 4 to o « Reference Diet Diet Reference Mlcropterus salmoldes largemouth bass Apogonldae - cardlnalflshes Astrapogon alutus bronze cardinalf ish 13-15 1. 3 Caridean shrimp, small blue crabs, crayfish, xanthld crabs, 25 species of fish, Vallisnerla, Cladophora Darnell 1958 Astrapogon stellatus conchfish Pomatomldae - blueflshes Pomatomus saltatrlx bluefish 11 Young: mainly fishes (anchovies, sllversldes, killifishes, men- haden, shad, spotted seatrout), shrimp, crabs, other small crustaceans, annelids, snails Peterson & Peterson 1979 Rachycentrldae - cobias Rachycentron canadum cobia + 5, 7, 11 Fish, crabs Randall 1967 Echeneldae - remoras Echenels neucratoides vhltefln sharksucker 2, 11 Fish, isopods, other Crustacea Randall 1967 Members of this family attach to sharks and large bony fishes (Randall 1967) Remora remora 58-175 mm: copepods, Isopods, vertebrate muscle tissue, crab larvae, fish remains, crusta- ceans, amphipods Randall 1967 Carangidae - jacks, pompanos Caranx crysos - blue + runner + 2, 4, 5, 7. 11 Family of swift- svlmmlng, carniv- orous fishes, often running in schools, wide- ranging (Randall 1967) Caranx hippos crevalle Jack Caranx ruber bar jack Chloroscombrus chrysurus Atlantic bumper Ollgoplltes 8aurua leatherjacket Trachlnotus carolinus Florida pompano Trachlnotus falcatus permit + 2, 5, 7, Fishes, crustaceans 8, 11, 13 + 4, 11 160-547 mm: fish, shrimp, myslds, stomatopods, gastropods + 2, 11, 17, 18 + 2, 3, 5, Snapping shrimp, penaeld shrimp, 8, 11, 13 larval anchovies, lady fish, harpactlcoid copepods + 11 sardines (Harengula sp.) , mole crabs (Hlppa sp . ) , bivalves (Donax sp.) + 7, 11 15-70 mm: myslds, shrimp, anchovies, sllversldes, crabs, snails Odum 1971 Randall 1967 Tabb & Manning 1961 Springer & Woodburn 1960 Common over mud bottoms (Randall 1967) Carr 6 Adams More apt to occur 1973 over sandy bottoms than T. carolinas (Randall 1967) Selene vomer lookdown + +2,3, 7, Young: shrimp and other 11 crustaceans, small molluscs Peterson 6 Peterson 1979 118 Habitat Type Reference Diet Diet Reference Family and Species ■o u ■H U 01 q M 3 U 03 u 30 mm: polychaetes, shrimp, amphipods 2, 3, 5, 7, <40 mm: in grassbeds - copepods , 8, 11-13 , amphipods , chironomid larvae , 17-18 mysids , algae, molluscs >40 mm: in mangrove creeks - mussels, false mussels, crabs, snapping shrimp, crayfish, hydrazoans , algae , plant detritus 32-85 mm: in Puerto Rico man- groves - 100% blue-green algae (Lyngbya mojuscula) 5, 11 105-220 mm: seagrasses Cymodocea Randall & Thalassia, algae, crabs, gas- 1967 tropods, invertebrate eggs, bivalves Randall 1967 Carr & Strong preference Adams for vegetated sub 1973 strate in bay areas (Weinstein et al. 1977) Odum 1971 Austin & Austin 1971 Usually seen in mangrove sloughs, rare on reefs (Randall 1967) Copepods , amphipods , mysids , Reid shrimp, bivalves, gastropods (Hitrella, Bittium) , polychaetes 1954 Associated with grassy flats (Tabb s. Manning 1961) 190-250 mm: polychaetes, brittle Randall stars, bivalves, hermit crabs, 1967 sea urchins, gastropods, chitons 120 Habitat Type Family and Species 01 a ■H « OJ 3 TT U i-( U on a H W Diet Diet Reference Lagodon rhomboides pinfish 1-3, 5, 7, B, In mangrove creek - scorched 11, 12, 16- mussel, mysids, amphipods, 18 false mussel In Whitewater Bay - 100% plant material Odum 1971 Strong preference Reid 1954 for vegetated sub- strate in bay areas (Weinstein et al. 1977) Sciaenidae - drums Balrdiella batabana blue croaker Balrdiella chrysura silver perch Cynoscion arenarlus sand seatrout Cynoscion nebulosus spotted seatrout 3, 11 Lelostomus xanthurus spot Menticirrhus americanus Southern kingfish Menticirrhus littoralis Gulf kingfish Micropogon undulatus Atlantic croaker Pogonlas cromis black drum Sciaenops ocellata red drum + 1-3, 8, Il- ls, 16-18 Larvae: copepods, larval fish (Henidia beryllina) Odum 1971 127-181 mm: fish (Anchoa mitchilli) , mysids 2, 12, 17, 18 Mostly fish, caridean shrimp, mysids, amphipods, crab zoea Springer & Woodburn 1960 + 1-3, 5, 7, 8, 11-13, 15, 17, 18 <50 mm: copepods, planktonic Crustacea 50-275 mm: fish (Mugil cephalus, Lagodon rhomboides, Eucino- stomus gula, E. argenteus, Cyprinodon varieqatus, Gobiosoma robustum, Anchoa mitchilli) Odum 1971 2, 7, 12, 17-18 <40 mm: planktonic organisms >40 mm: filamentous algae, desmids, forams, amphipods. Springer & Woodburn 1960 2, 11-12, 17-18 11, 12 + 2, 7, 11, 12, 15 2, 3j 5, 8, 11-13, 15, 17 Equetus acuminatus high-hat mysids, copepods, ostracods, isopods , chaetognaths , bi- valves, snails, polychaete worms Fish, benthic crustaceans Polychaetes , bivalves (Dona*) , sand crab Emerita), razor clams Juveniles: copepods, mysids, caridean shrimp, polychaete worms, insect larvae, iso- pods, small bivalves <100 mm: molluscs, xanthid crabs >100 mm: bivalves, amphipods, blue crabs, penaeid shrimp, caridean shrimp <10 mm: planktonic organisms (copepods, crab zoea, larval fish) 34-42 mm: mysids, amphipods, caridean shrimp >50 mm: xanthid & portunid crabs, penaeid shrimp, small fish 308-403 mm: xanthid crabs 68-152 mm: shrimp S shrimp larvae, isopods, stoma topod larvae , copepods , amphipods Springer & Woodburn 1960 Springer & Most common off sandy Woodburn beaches (Springer & 1960 woodburn 1960) Springer & Woodburn 1960 Darnell 1958 Odum 1971 Randall 1967 Characteristic of coral reefs (Randall 1967) 121 Habitat Type Family and Species a •S H a 3 a ■d u *-> CO « H ■s> u oa Diet Reference Ephippidae - spadefishes Chaetodipterus faber ■Atlantic spadef ish Pomacentridae - damsel fishes Abudefduf saxatilis sergeant major 2, 3, 5, 11, 16-18 Worms, crustaceans, debris Darnell 1961 101-135 mm: copepods, algae, Randall fish eggs, fish, shrimp larvae, 1967 polychaetes Juveniles (7-12 mm) inhabit very shallow nearshore sandy beaches. Bear a deceptive resemblance to infertile red mangrove seed pods (Breder 1946) Characteristic family of coral reefs (Ran- dall 1967) A habitat generalist: reefs , grassbeds , rock piles, wharfs (Bohlke & Chaplin 1968) Labridae - wrasses Halichoeres bivittatus slippery dick 67-153 mm: crabs, sea urchins, Randall polychaetes , gastropods , brittle 1967 stars, bivalves , shrimp, fish, hermit crabs Shallow water patch reefs, sand bottoms, gr a s sbeds ( Randa 1 1 1967) Scaridae - parrotfishes Nicholsina usta emerald parrotfish + 1, 2, 11, 18 Family herbivorous, feeding primarily on algae growing on hard substrates, on seagrasses secondarily Randal] Family characteris- 1967 tic of coral reefs, ranging into grass- beds Scarus coeruleus blue parrotfish Scarus croicensis striped parrotfish Spar i soma chrysopterum redtail parrotfish Sparisoma rubripinne redfin parrotfish Sparisoma viride stoplight parrotfish Mugilidae - mullets Mugil cephalus striped mullet Mugil curema white mullet 11 + 2, 3, 5, 7, 11-13, 15 + 2, 5, 7, 11-12 Inorganic sediments, fine detritus , micro-algae 25-73 mm: plant detritus, blue- green algae (Lyngbya majuscula) Odum 1971 Austin & Austin 1971 Requires near marine salinities {Tabb & Manning 1961) Mugil trichodon fantail mullet Sphyraenidae - barracudas Sphyraena barracuda great barracuda Opistognathidae - jawfishes Opistognathus maxillosus mottled jawfish + 2, 7, 11, 12 1-5, 7, 8, 11, 13 135-369 mm: fish (Eucinostomus quia, Menidia beryllina, Archo- sargus probatocephalus) 53-110 mm: shrimp, isopods, fishes, polychaetes, mysids, copepods Odum 1971 Randa 1 1 1967 Salinities >10 o/oo (Odum 1971) Family lives in burrows in sediment, often in vicinity of reefs (Randall 1967) 122 Family and Species Habitat Type 0} 3 1 0) 3 M *-> >> u 41 >» ■H 4J a> Q O Q H v. W 03 O 03 Diet Reference Comments Microgobius gulosus clown goby Microgobius microlepis banner goby Microgobius thalassinus green goby 2,5,8, 11- Amphipods, copepods, chironomid Odum 1971 13, 15, 17, larvae 18 5 3, 12 Planktonic organisms Bird song 1981 Small crustaceans including Peterson & amphipods, other invertebrates Peterson 1979 Scombridae - mackerels, tunas Scomberomorus maculatus Spanish mackerel 2, 11, 12, Adults feeding on penaeid Tahb & 15 shrimp migrating from tidal Manning stream 1961 Scomberomorus cavalla king mackerel 11 350-1022 fish Randall 1967 Scorpaenidae - scorpion- fishes Scorpaena brasiliensis barbfish Scorpaena grandicornis plumed scorpionfish 11 Shrimp, other crustaceans, fish 1 37-102 ram: shrimp, fish, unidentified crustaceans 1967 Randall Most often found 1967 in seagrass Triglidae - searobins Prionotus salmonicolor blackwing searobin Prionotus scitulus leopard searobin Prionotus tribulus blghead searobin + 1-3, 11, Small molluscs, shrimp, crabs 16-18 fish, small crustaceans (ostracods , cumaceans) + 1-3, 11-13, Shrimp, crabs, fishes, amphi- 17 i 18 pods, copepods, annelids, bivalves, sea urchins Peterson & Peterson 1979 Peterson & Peterson 1979 Bothidae - lefteye flounders Bo thus ocellatus eyed flounder Citharichthys macrops spotted whiff Citharichthys spllopterus bay whiff i, n 1, 17, 18 68-130 ram: fish, crabs, shrimp, amphipods Mainly myslds, also shrimp, crabs, copepods, amphipods, fishes, annelids Randall 1967 Peterson & Recorded from Peterson salinity range 1979 2.5-36.7 o/oo (Darnell 1961) Etropus crossotus fringed flounder Farallchthys albigutta Gulf flounder Parallchthys lethostlgma Southern flounder 3, 11, 16 1-3, 7, 11, 12, 17, 18 Calanoid copepods, cumaceans, Peterson & amphipods, myslds, shrimp, Peterson crabs, isopods, annelids, 1979 molluscs, fishes 45 mm: fish (pigflsh, plnfish, 1954 lizardflsh, bay anchovy, labrids), crustaceans Mainly fishes (mullet, menha- Peterson & den, shad, anchovies, plnfish, Peterson mojarras, croakers), crabs, 1979 mysids, molluscs, penaeid shrimp, amphipods 124 Family and Species Habitat Type Reference Diet Diet Reference Comments Syacium papillosum dusky flounder Soleidae - soles Achirus lineatus lined sole 1-3, 5, 8, 11-13, 17- 18 32-74 mm: chironomid larvae, polychaete worms, foraminiferans Odum 1971 Trinectes inscriptus scrawled sole Trinectes maculatus hogchoker 2, 3, 8, 11-13, 17, 18 14-110 mm: amphipods , mysids Odum 1971 Cynoglossidae - tongue- fishes Symphurus plagiusa blackcheek tonguefish Balistidae - triggerfishes & filefishes Aluterus schoepfi orange filefish Balistes vetula queen triggerfish Monacanthus ciliatus fringed filefish Monacanthus hispidus planehead filefish Balistes capriscus gray triggerfish 1, 3, 11, 12, 16-18 1, 11 35-102 mm: polychaete worms, ostracods, portunid crabs, Ruppia and Halodule plant tips Seagrasses, algae, hermit crabs, gastropods 11 130-480 mm: sea urchins, crabs, bivalves, brittle stars, poly- chaetes, hermit crabs, gastro- pods, algae 1, 11, 17 47-97 mm: Algae, organic detri- tus , seagrass , copepods, shrimp & shrimp larvae, amphipods, tanaids, polychaetes, molluscs 1-3, 11, Detritus, bryozoans, annelids, 16-18 harpacticoid copepods , amphi- pods, hermit crabs, molluscs, algae, sea urchins Austin Austin 1971 Randall 1967 Randall 1967 Randall, 1967 Springer & Woodburn 1960 Peterson & Peterson 1979 Associated with grassbeds , sponge/sea fan habitats (Ran- dall 1967, Voss et al. 1969) Solitary reef fish ranging into grass- beds Closely associated with vegetated areas (Tabb & Manning 1961) Associated with vegetated areas {Tabb & Manning 1961) Ostraciidae - boxfishes Lactophrys quadracornis scrawled cowfish Lactophrys trigonus trunkfish Lactophrys triqueter smooth trunkfish Tetraodontidae - puffers Sphoeroides nephelus southern puffer I, 2, 5, 7, II, 16-18 1, 4, 11 1-3, 5, 11, 16-18 Vegetation , algae , bivalves 109-395 mm: crabs, bivalves, polychaetes, sea urchins, algae, seagrass, gastropods, amphipods 93-250 mm: polychaetes, sipun- culid worms, crabs, shrimp, gastropods, hermit crabs, sea urchins, bivalves Juveniles : detritus , fecal pellets, zooplankton, poly- chaetes, gastropods, crabs, shrimp Adults : small crabs , bivalves Reid 1954 Young mimic sea- grass blades (Bohlke & Chaplin 1968) Randall Primarily a resident 1967 of seagrass (Randall 1967) Randall Primarily a reef 1967 species (Randall 1967) Carr & Adams 1973 125 Habitat Type Family and Species M 3, Reference Diet Diet Reference Comments Sphoeroides spengleri bandtall puffer Sphoeroides testudineus + checkered puffer Diodontidae - porcupine- fishes Chilomycterus antenna tus bridled burrfish Chilomycterus ant 11 la rum web burrfish 1» 7, 11 Crabs, bivalves, snails, polychaetes, amphipods, shrimp 1, 7 85-92 mm: portunid megalops larvae, gastropods 11 Gastropods, hermit crabs, isopods, crabs, shrimp Randall Inhabits sea- 1967 grass, reef, rubble, man- groves (Randall 1967; Voss et al 1969) Austin & Austin 1971 Randall Reefs and grass 1967 beds (Voss et al. 1969) Chilomycterus schoepf 1 striped burrfish 1-3, 5, Gastropods, barnacles, crabs, 11, 16-18 amphipods Springer & Uoodburn 1960 Associated with grassbeds (Voss et al. 1969) Salinities >25 o/oo (Springer & Woodburn 1960) Reference Numbers Key 1. Bader & Roessler 1971 2. Carter et al. 1973 3. Clark 1970 4. Holm 1977 5. Hudson et al. 1970 6. Rush Ian & Lodge 1974 7. Nugent 1970 8. Odum 1971 9. Rivas 1969 10. Seaman et al. 1973 11. Schmidt 1979 12. Springer 6, Woodburn 1960 13. Tabb 1966 14. Tabb, Dubrow & Manning 1962 15. Tabb & Manning 1961 16. Weinstein et al. 1977 17. Yokel 1975a 18. Yokel 1975b 126 APPENDIX C. Amphibians and reptiles recorded from south Florida mangrove swamps . 127 AMPHIBIANS AND REPTILES OF FLORIDA'S MANGROVES Species Status Food Habits Mud Turtle (Kinosternon subrubrum) Abundant Insects, crustaceans, mollusks Striped Mud Turtle (Kinosternon bauri ) Common Algae, snails, dead fish Ornate Diamondback Terrapin (Malaclemys terrapin macrospilota and M.t_. rhizophorarum) Florida Red-bellied Turtle (Chrysemys nelsoni) Chicken Turtle (Deirochelys reticularia) Green Turtle (Chelonia mydas) Hawksbill (Eretmochelys imbricata) Loggerhead (Caretta caretta) Atlantic Ridley (Lepidochelys kempii) Florida Softshell (Trionyx ferox) Green Anole (Anolis carolinensis) Cuban Brown Anole (Anolis sagrei) Bahaman Bank Anole (Anolis distichus) Green Water Snake (Nerodia cyclopion) Mangrove Water Snake (Nerodia fasciata compressicauda) Uncommon Littorina, Melampus, Uca . Anomalocardia Rare - Uncommon Sagittaria, Lemna, Naias Uncommon Uncommon Rare Common Uncommon Common Common Common Uncommon Common Common Crayfish, insects, Nuphar Mangrove roots and leaves , seagrasses Rhizophora: fruits, leaves wood , bark Crabs, jellyfish, tuni- cates Snails, crabs, clams Snails, crayfish, mussels, frogs, fish, waterfowl Insects Insects Insects Fish Fish, invertebrates 128 AMPHIBIANS AND REPTILES OF FLORIDA'S MANGROVES (concluded) Species Status Food Habits Striped Swamp Snake Uncommon (Liodytes alleni) Eastern Indigo Snake Uncommon (Drymarchon corais) Rat Snake Uncommon (Elaphe obsoleta) Eastern Cottonmouth Uncommon (Agkistrodon piscivorus) American Alligator Common (Alligator mississippiensis) American Crocodile Rare (Crocodylus acutus) Giant Toad Common (Bufo marlnus) Crayfish, sirens, frogs Small mammals, birds, frogs Small mammals , birds Fish, frogs, snakes, birds , small mammals Fish, waterbirds Fish, waterbirds Invertebrates Squirrel Treefrog (Hyla squirella) Cuban Treefrog (Hyla septentrionalis) Abundant Common Insects Insects, frogs, toads, lizards References : Carr and Goin 1955; Ernst & Barbour 1972; Mahmuud 1965; L. Narcisse, R.N. "Ding" Darling Fed. Wildlife Refuge, Sanibell Is., Fla. ; personal communication (1981) . 129 APPENDIX D. Avifauna of south Florida mangrove swamps. 130 WADING BIRDS Common Name (Latin name) Season of Occurrence3 Nestinya Food Habits References Great Egret (Casmerodius albus) Howell, 1932 Kushlan & White 1977a Snowy Egret (Egretta thula) Fish Howell 1932 Kushlan 6 White 1977a Ffrench 1966 Cattle Egret (Bubulcus ibis) Fish Howell 1932 Kushlan & White 1977a Great White Heron (Ardea herodias occidentalis) Fish Howell 1932 Kushlan & White 1977a Great Blue Heron (Ardea herodias) Howell 1932 Kushlan & White 1977a Reddish Egret (Dichromanassa ruf escens) Howell 1932 Kushlan & White 1977a Louisiana Heron Common (Hydranassa tricolor) Kushlan s. White 1977a Maxwell & Kale 1977 Girard & Taylor 1979 Little Blue Heron (Florida caerulea) Kushlan & White 1977a Maxwell & Kale 1977 Girard & Taylor 1979 Green Heron Common (Butorides striatus) Fish Robertson & Kushlan 1974 Maxwell & Kale, 1977 Girard & Taylor 1979 Black-crowned Night Heron (Nycticorax nycticorax) Fish , crustaceans , frogs, mice Ffrench 1966 Maxwell & Kale 1977 Girard & Taylor 1979 Yellow-crowned Night Heron (Nyctanassa violacea) Fish, crayfish, crabs Ffrench 1966 Girard & Taylor 1979 Least Bittern (Ixobrychus exilis) Fish Ffrench 1966 American Bittern (Botaurus lentiginosus) W,T Crayfish, frogs, small fishes Narcisse , pers . comm . Wood Stork (Mycteria americana) Common (locally abundant) Kahl 1964 Ogden et al. 1976 Kushlan 1979 Glossy Ibis (Plegadis falci- nellus Uncommon Fish Bacon 1970 Howell 1932 White Ibis (Eudocimus albus) Fish, crabs (Uca) Kushlan 1979 Kushlan & Kushlan 1975 Girard & Taylor 1979 Roseate Spoonbill (Ajaia ajaja) Sandhill crane (Grus canadensis) Rare to Uncommon Shrimp, fish, aquatic vegetation Roots, rhizomes of Cyperus & Sagit- taria Kushlan & White 1977a Howell 1932 Ogden 1969 Howell 1932 Limpkin (Aramus guar a una) Snails (Pomacea) Howell 1932 Bacon 1970 131 PROBING SHORE BIRDS Common Name (Latin name) Abundance Season of Occurrence3 Nesting* Food Habits References King Rail (Rail us elgans) Clapper Rail (Rallus longiro- stris) Virginia Rail (Rallus limicola) Sora (Porzana Carolina) Black Rail (Laterallus jamaicensis) Semipalmated Plover (Charadrius semi - palmatus) Wilson's Plover (Charadrius wilsonia) Black-bellied Plover (Pluvialis squatarola) Ruddy Turnstone (Arenaria lnterpres) Common Snipe (Capella gallinago) Common Uncommon- common Rare Uncommon to locally abundant Rare Locally common Locally common Yr W,T Uncommon W,T Beetles, grass- hoppers, aquatic bugs Crabs, shrimp Beetles, snails, spiders Insects, seeds of emergent aquatic plants Beetles, snails Crustaceans, mollusks Crabs, shrimp, crayfish Crabs , mollusks Insects, crus- taceans , mollusks Mollusks, insects, worms Narcise, pers. comm. Martin et al. 1951 Howell 1932 Ffrench 1966 Bacon 1970 Marcisse, pers. coram. Martin et al. 1951 Howell 1932 Bacon 1970 Narcisse, pers- coram. Ffrench 1966 Bacon 1970 Baker & Baker 1973 Howell 1932 Bacon 1970 Howell 1932 Bacon 1970 Ffrench 1966 Ogden 1969 Howell 1932 Howell 1932 Bacon 1970 Long-billed Curlew (Numenlus americanus) Whimbrel (Numenius phaeopus) Spotted Sandpiper (Ac tit is macularia) Solitary Sandpiper (Tringa solitaria) Willet (Catoptrophorus semipalmatus) Greater Yellowlegs (Tringa melanoleucas) Lesser Yellowlegs (Tringa flavipes) Rare-uncommon WtT Abundant Common Common Red Knot (Calidrls canutus) Dunlin (Calidris alplna) White-rumped Sandpiper Rare (Calidris fusclcollls) Common Uncommon Common W,T W,T Yr W,T W,T W,T Crustaceans, insects Mollusks, crus- taceans , worms , insects Mollusks, crus- taceans Ogden 1969 Ogden 1968 Howell 1932 Ffrench 1966 Bacon 1970 Russel 1980 Crustaceans, aquatic Howell 1932 insects, small frogs Bacon 1970 Crabs, crayfishes, Howell 1932 killifishes Bacon 1970 Fishes, crabs , crustaceans Snails, mollusks, crabs Marine worms , crustaceans Marine worms, mollusks Howell 1932 Ffrench 1966 Bacon 1970 Ffrench 1966 Bacon 1970 Baker & Baker 1973 Howell 1932 Ogden 1964 Ogden 1964 Baker & Baker 1973 Chironomids, snails Howell 1932 Bacon 1970 132 PROBING SHOREBIRDS (concluded) Common Name (Latin name) Abundance Season of Occurrence3 Least Sandpiper (Calidris minutilla) Common W,T Short-billed Dowitcher (Limnodromus griseus) Common W,T Stilt Sandpiper (Micropalama himantopus) Rare-uncommon W,T Semipalmated Sandpiper (Calidris pusilla) Common- abundant W,T Western Sandpiper (Calidris mauri) Common- abundant W,T Marbled Godwit (Limosa fedoa) Rare-common W Nesting3 Food Habits References American Avocet (Recurvlrostra americana) Uncommon W,T Pupae of beetles and flies Mollusks, crustaceans Chironomids Mollusks, insects Chironomids Crustaceans, mollusks, seeds of emergent aquatic plants Marine worms, aquatic insects Bacon 1970 Baker & Baker 1973 Bacon 1970 Baker 6. Baker 1973 Howell 1932 Bacon 1970 Bacon 1970 Baker & Baker 1973 Howell 1932 Bacon 1970 Howell 1932 Ogden 1969 Black-necked Stilt Common (Himantopus mexicanus) Aquatic beetles Howell 1932 Bacon 1970 133 SURFACE AND DIVING BIRDS Common Name Season of (Latin name) Abundance Occurrence' Common Loon Occasional w (Gavia immer) Horned Grebe Uncommon w (Podiceps auritus) Pied-billed Grebe Uncommon- Yr (Podilymb'us common podiceps) White Pelican Rare S (Pelecanus Common w ery throrhy nchos ) Brown Pelican Common Yr (Pelecanus occidentalia) Double-crested Common Yr cormorant (Phalacrocorax auritus) Anhinga Common Yr (Anhinga anhinga) Fulvous Whistling Duck Uncommon w (Dendrocygna bicolor) Mallard Uncommon W,T (Anas platyrhynchos) Black Duck Rare W,T (Anas rubripes) Mottled Duck Uncommon Yr (Anas fulvigula) Gad wall Uncommon W,T (Anas strepera) Pintail Abundant W,T (Anas acuta) Green-winged Teal Abundant W,T (Anas crecca carollnensis) Blue-winged Teal Abundant Yr (Anas discors) Common American Wigeon W,T (Anas americana) Northern Shoveler Common W,T (Anas clypeata) Wood Duck Rare W (Aix sponsa) Redhead Rare W (Aythya americana) Nesting' Food Habits References Fish, crabs, mollusks Narcisse, pers. co Fish, aquatic insects, Ogden 1969 mollusks Crayfish, fish, mollusks Widgeon grass Mollusks, crusta- ceans, widgeon grass Polygonum, snails, Ruppia Ruppia , Zostera, mollusks Saggitaria, mollusks, Cyperus Ruppia , Zostera.- aquatic insects Cyperus , snails, insects, crustaceans Ruppia , Zostera, mollusks mollusks, aquatic insects, Ruppia, Zostera Nuts, seeds Narcisse, pare, coram. Narcisse, pers. conm. Ffrench 1966 Bacon 1970 Kushlan & White 1977a Ffrench 1966 Ogden 1969 Smith, pers. obs . Ogden 1969 Kushlan et al., in prep. Ogden 1969 LaHunt & Cornwell 1970 Kushlan et al., in prep. Ogden 1969 Narcisse, pers. comm. Kushlan et al.,in prep. Narcisse, pers. comm. Kushlan et al., in prep. Narcisse, pers. coram. Ffrench 1966 Narcisse, pers. comm. Kushlan et al. , in prep. Narcisse, pers. comm. Ogden 1959 Snails, clams, aquatic Ogden 1969 insects, Ruppia, Zos- tera Ring-necked Duck Abundant (Aythya collar is) Polygonum, Ruppia , crayfish, snails Ogden 1969 Kushlan et al., in prep. Canvasback Uncommon (Aythya valiaineria) Vallisneria, Ruppia , Zostera Ogden 1969 Kushlan et al., in prep. 134 SURFACE AND DIVING BIRDS (concluded) Common Name (Latin name) Abundance Season of Occurrence Nesting ' Food Habits References Lesser Scaup Common- (Aythya affinls) abundant Mollusks, Ruppia Narcisse, pers. coram. Ogden 1969 Kushlan et al., In prep. Bufflehead (Bucephala albeola) Rare Ruddy Duck Common W (Oxyura jamaicensis) Gastropods, crabs, Ogden 1969 crustaceans Kushlan et al., in prep. Potamogeton, Najas, Ogden 1969 Zostera, Ruppia, Kushlan et al. mollusks in prep. Hooded Merganser (Lophodytes cucullatus) Red-breasted Merganser (Mergus serrator ) Purple Gallinule (Porphyrula martlnica) Common Gallinule (Gallinula chloropus) American Coot (Fulica americana) Rare-uncommon Common Rare Common Abundant W.T Yr Yr W,T Fish Fish Ogden 1969 Narcisse, pers. coram. Aquatic insects, Narcisse, pers. coram, mollusks, Ffrench 1966 Eleocharis, Paspalum Seeds, aquatic insects Ruppia, Na j as , Potamogeton, aquatic insects Narcisse, pers. coram. Ffrench 1966 Narcisse, pers. coram. 135 AERIALLY SEARCHING Common Name (Latin name) Abundance Season of Occurrence Nesting" Food Habits References Herring Gull (Larus argentatus) Ring-billed Gull (Larus delawarensis) Common Fish, mollusks, crustaceans Fish, insects, mollusks Narcisse, pers. comm. Ogden 1969 Narcisse, pers. comm. Ogden 1969 Laughing Gull Common (Larus atricilla) Bonaparte's Gull Uncommon (Larus Philadelphia) Gull-billed Tern Uncommon (Gelochelidon nilotica) Forster's Tern Uncommon- ( Sterna fosteri) common Fish, shrimp, crabs Fish, insects Narcisse, pers. comm. Ogden 1969 Ogden 1969 Mayflies, dragonflies Ogden 1969 Fish Narcisse, pers. comm. Ogden 1969 Common Tern (Sterna hirundo) Fish Ogden 1969 Least Tern (Sterna albifrons) Fish Narcisse, pers. comm. Ogden 1969 Royal Tern (Thalasseus maxima) Common W,T Fish Ogden 1969 Sandwich Tern (Sterna sand- vicensis) Uncommon Yr Fish Narcisse, pers. comm. Ogden 1969 Caspian Tern (Sterna caspia) Uncommon Fish Ogden 1969 Black Skimmer Common (Rynchops nigra) Belted Kingfisher Common (Megaceryle alcyon) Fish Crow Common (Corvus ossifragus) Yr Fish Fish Fish Ogden 1969 Narcisse, pers. comm. Narcisse, pers. comm. 136 BIRDS OF PREY Common Name (Latin name) Season of Abundance Occurrence* Nesting3 Food Habits References Magnificent Frigate- Common S bird Uncommon W (Fregata magnificens) Turkey Vulture (Cathartes aura) Common Fish Narcisse, pera. com. Smith, pers. obs. Narcisse, pers. comm. Orians 1969 Black Vulture Common (Coragyps atratus) Swallow-tailed Kite Common (Elanoides forf ica- tus) Sharp-shinned Hawk Uncommon (Accipiter striatus) Cooper's Hawk Uncommon (Accipiter cooper ii) Red-tailed Hawk Uncommon (Buteo jamaicensis) Red-shouldered Hawk Common (Buteo lineatus) Broad-winged Hawk (Buteo platypterus) Swainson's Hawk (Buteo swainsoni) Short-tailed Hawk (Buteo brachyurus) Bald Eagle (Haliaeetus leucocephalus) Marsh Hawk (Circus cyaneus) Osprey (Pandion haliaetus) Peregrine Falcon (Falco peregrinus) Merlin (Falco columbarius) American Kestrel Common (Falco sparverius) Barn Owl Uncommo (Tyto alba) Great Horned Owl Uncommo (Bubo virginianus) Barred Owl Uncommo (Strix varla) Rare- locally common (Fla. Bay) Uncommon Very rare- locally common (Fla. Bay) Uncommon Carrion Snakes, lizards, frogs Robertson & Kushlan 1974 Orians 1969 Howell 1932 Snyder 1974 Smaller passerines Howell 1932 Larger passerines Howell 1932 Small mammals, birds Howell 1932 Snakes, frogs, lizards. Insects Insects, small mammals Howel] 1932 Robertson & Kushlan. 1974 Howell 1932 Small mammals, grass- Howell 1932 hoppers Small birds Fishes Howell 1932 Howell 1932 Small mammals, shore- Howell 1932 birds Fishes Howell 1932 Waterfowl, shorebirds Nisbet 1968 Ogden 1969 Howell 1932 Small birds, shore- Howell 1932 birds Insects Small mammals Waterfowl , small mammals Hovel) 1932 Howell 1932 Howell 1932 Y Small mammals, frogs, Howell 1932 snakes 137 ARBOREAL BIRDS Common Name (Latin name) Mourning Dove (Zenaidura macroura) White-crowned Pigeon (Columba leucocephala) Mangrove Cuckoo (Coccyzus minor) Abundance Season of Occurrence3 Nesting3 Food Habits Uncommon Yr Seeds Berries, seeds fruits Caterpillars, man t ids References Emlen 1977 Howell 1932 Robertson & Kushlan 1974 Howell 1932 Ffrench 1966 Robertson & Kushlan 1974 Martin et al. 1951 Yellow-billed Cuckoo Common (Coccyzus americanus) Caterpillars, beetles Howell 1932 Ffrench 1966 Martin et al. 1951 Smooth-billed Anl Rare Yr (Crotophaga anl) Chuck-will 's-widow Uncommon Yr (Caprimulgus carolinensis) Common Flicker Uncommon Yr (Colaptes auratus) Pileated Woodpecker Uncommon Yr (Dryocopus pileatus) Mosqultos, moths Ants, beetles, fruits in winter Beetles, berries, fruits Howell 1932 Ffrench 1966 Martin et al. 1951 Narcisse, pers. comm. Narcisse, pers. coram. Martin et al. 1951 Howell 1932 Robertson 1955 Robertson 4 Kushlan 1974 Red-bellied Woodpecker Common (Melanerpes carolinus) Beetles, ants, grasshoppers, crickets Narcisse, pers. coram. Martin et al. 1951 Red-headed Woodpecker (Melanerpes erythrocephalus) Yellow-bellied Sapsucker (Sphyraplcus varius) Hairy Woodpecker (Picoides vlllosus) Eastern Kingbird (Tyrannus tyrannus) Gray Kingbird (Tyrannus dominicensis) Western Kingbird (Tyrannus vertlcalus) Great Crested Flycatcher (Mylarchus crlnitus) Acadian Flycatcher (Empldonax virescens) Eastern Phoebe (Sayornis phoebe) Eastern Wood Pewee (Contopus vlrens) Uncommon Common Uncommon (common S) Rare Common Rare-uncommon Yr W,T S,T W,T Beetles, ants, grasshoppers, caterpillars Beetles, ants-, caterpillars Insects, beetle larvae Ants, wasps, grasshoppers Bees, wasps, beetles, dragon Bees, wasps, grasshoppers Insects, berries Narcisse, pers. comm. Martin et al. 1951 Narcisse, pers. coram. Martin et al. 1951 Emlen 1977 Narcisse, pers. comm. Martin et al. 1951 Howell 1932 Robertson & Kushlan 1974 Narcisse, pers. comm. Martin et al. 1951 Howell 1932 Robertson 1955 Small flying insects Morton 1980 Bees, wasps, ants Narcisse, pers. comm. Martin et al. 1951 Bees, wasps, ants, Narcisse, pers. comm. moths Howell 1932 138 ARBOREAL BIRDS (continued) Common *