food-web structure in big cypress swamp wetlands based on

1
FOOD-WEB STRUCTURE IN BIG CYPRESS SWAMP WETLANDS BASED ON STABLE-ISOTOPE RESULTS William F. Loftus 1 and David P.J. Green 2,3 1 USGS FISC, Everglades NP, Homestead, FL, USA; 2 Audubon of Florida, Tavernier Science Center, Tavernier, FL, USA; 3 FL Gulf Coast University, Ft. Myers, FL, USA RESULTS: We analyzed 777 individual fishes of 24 taxa; 7 were non-native. We also analyzed 751 invertebrate samples of 24 taxa and 733 vegetation samples of 24 taxa. Fish species names are in Table 1. Stoichiometric data for C:N showed that as δ 15 N values increased, C:N ratios decreased (Fig. 4). Most vegetation taxa had δ 13 C values within + 2 0 / 00 of -30 0 / 00, except algae and moss. Values from BCNP biota were remarkably similar to those from Everglades marshes (Fig. 5). FUTURE WORK We will explore the dataset with analytical tools such as the IsoSource (Phillips and Gregg 2003) and circular statistical models (Schmidt et al. 2007) to examine mixing of primary producers in the cypress. We will investigate effects of lipids on values, and will also compare data quantitatively to food webs from other systems. APPLICATIONS Hydrological restoration should restore aquatic food webs supporting populations of higher vertebrates. Our data can help test that premise by defining spatio-temporal patterns in cypress-wetland food webs as a baseline for post-restoration comparisons. Hydrologic restoration will affect aquatic plant communities at the food-web base. We hypothesize that changes at the base will affect key invertebrate groups, with consequences resonating through the web. Stable-isotope analyses will permit tracing of food-web changes at both local and landscape scales. Our data will complement other south Florida food-web studies to provide a more comprehensive understanding of the entire ecosystem. ACKNOWLEDGMENTS Funding provided by USGS South Florida PES Program, administered by G.R. Best, and by Army Corps of Engineers CERP-MAP. Special thanks to S. Liston for project assistance, to B. Dunker, N. Katin and N. Silverman for field assistance, and J. Lorenz, J. Trexler, and C. McIvor for help in the study. We also thank K. Dunlop, L. McCarthy, W. Anderson, M. Kershaw, and C. Rebenack for sample prep and analysis at the FIU-SERC mass-spectrometer facility. METHODS AND STUDY LOCATIONS Primary producer, invertebrate, and fish samples of common species (Table 1) were taken three times per year: wet season, transitional season, and dry season. Sampling was at three sites within BCNP cypress habitats: L-28, Bear Island (BI), and Raccoon Point (RP) (Figure 2). Site descriptions, sampling techniques, and physico-chemical data were reported in Liston et al. (2008) (Fig. 3). Three to five individuals or sub-samples were collected for each taxon from each sub-habitat (marsh & dome). Samples were field-frozen, then measured in the lab prior to drying appropriate tissues at 50 C. Plant material was acid-treated to remove carbonate. Dried tissue was pulverized, weighed, and prepared for analysis by mass spectrometer at FIU (see Williams & Trexler, 2006). Error bars not shown on figures for clarity of presentation. Bear Island HQ I-75 Raccoon Point US 41 L-28 BIG CYPRESS NATIONAL PRESERVE N N 5 0 5 10 Km Bear Island HQ I-75 Raccoon Point US 41 L-28 BIG CYPRESS NATIONAL PRESERVE Figure 2. Maps of study locations Distinct food webs are evident in south Florida studies; data may serve to relate spatial data to key indicator species. For example, Roseate Spoonbills (pink circle; mean + 1 SE; N=8) receive energetic inputs from Taylor Slough mangroves, near nesting locations and flight paths to foraging grounds (Lorenz data). CROSS-LANDSCAPE COMPARISONS 0 2 4 6 8 10 12 14 -33 -28 -23 -18 -13 Species used in comparison: Gar / Mangrove snapper Gambusia / Rivulus Grass Shrimp Snails Amphipods δ 13 C (‰) δ 15 N (‰) Big Cypress Freshwater ‘Glades TS Mangrove Zone SR Mangrove Zone Spoonbill chicks LITERATURE CITED Liston, S. et al. 2008. Development and testing of protocols for sampling fishes in forested wetlands in southern Florida. Draft report to USACOE and USGS-PES. Loftus, W.F. 2000. Mercury bioaccumulation through an Everglades aquatic food web. Ph.D. Dissert., Florida International University, Miami. Phillips, D.L., and J.W. Gregg. 2003. Source partitioning with stable isotopes: coping with too many sources. Oecologia 136:261-269. Post et al. 2007. Getting to the fat of the matter: models, methods, and assumptions for dealing with lipids in stable isotope analysis. Oecologia 152: 179-189. Schmidt, S.N. et al. 2007. Quantitative approaches to the analysis of stable isotope food web data. Ecology 88:2793-2802. Williams, A.J. and J.C. Trexler. 2006. A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades. Hydrobiologia 569:494-503. INTRODUCTION Southern Florida wetlands have been modified by drainage. A major restoration plan, CERP, is attempting to reestablish natural hydrology. CERP success implies aquatic food webs will be restored to support important predator populations. That premise is difficult to measure without data on those webs. From 2005-07, we collected biotic samples to map pathways of energy flow and trophic status of biota in freshwater wetlands of Big Cypress National Preserve (BCNP) (Fig. 1 ). Food webs in cypress wetlands have been relatively unstudied throughout their range. Food webs provide maps of their biotic constituents, their roles, and interactions. We used stable-isotope analysis to trace the movement of carbon (δ 13 C) from primary producers to fishes, and to determine the trophic positions of animals by using nitrogen (δ 15 N). Basic questions included: What are the major primary producers? How long are food chains? Is there spatio-temporal variability in the web? What roles do non-native fishes play? Figure 1. Cypress Forest. BI Jun 05 Sep 05 Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 -20 0 20 40 60 80 100 Estimated water depth (cm) Date BI Jun 05 Sep 05 Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 -20 0 20 40 60 80 100 Estimated water depth (cm) Date Date Estimated water depth (cm) -60 -40 -20 0 20 40 60 80 100 120 L28 Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 Date Estimated water depth (cm) -100 -80 -60 -40 -20 0 20 40 60 80 100 120 L28 Jun 05 Sep 05 Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 Estimated water depth (cm) -20 0 20 40 60 80 100 RP Date Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 Estimated water depth (cm) -40 -20 0 20 40 60 80 100 RP Date Jun 05 Sep 05 Dec 05 Mar 06 Jun 06 Sep 06 Dec 06 Mar 07 Jun 07 Figure 3. BCNP water depths showing strong seasonal patterns at the three study sites: Bear Island (BI), L-28, and Raccoon Point (RP). Table 1. Relative abundance (%RA) and % Incidence in samples of a) common fishes and b) common invertebrates collected at the study sites (Liston et al., 2008). All taxa were analyzed for stable-isotope values. Sailfin molly (Poecilia latipinna) Least killifish (Heterandria formosa) Ea. mosquitofish (Gambusia holbrooki) a) Fishes Scientific Name Common Name % RA % I Gambusia holbrooki Eastern mosquitofish 56.7 84.5 Jordanella floridae Flagfish 20.8 40.8 Heterandria formosa Least killifish 10.2 43.7 Lepomis marginatus Dollar sunfish 2.8 29.6 Lucania goodei Bluefin killifish 2.6 46.5 Lepomis gulosus Warmouth 2.1 30.3 Labidesthes sicculus Brook silverside 1.6 15.5 Ameiurus natalis Yellow bullhead 1.3 4.2 Poecilia latipinna Sailfin molly 0.8 11.3 Cichlasoma bimaculatum Black acara 0.7 22.5 Elassoma evergladei Everglades pygmy sunfish 0.6 16.9 Fundulus chrysotus Golden topminnow 0.4 15.5 Clarias batrachus Walking catfish 0.3 1.4 Hoplosternum littorale Brown hoplo catfish 0.2 7.7 Enneacanthus gloriosus Bluespotted sunfish 0.2 9.2 Cichlasoma urophthalmus Mayan cichlid 0.2 8.5 Notemigonus crysoleucas Golden shiner 0.1 4.2 Tilapia mariae Spotted tilapia 0.1 4.2 Lepomis punctatus Spotted sunfish 0.1 2.1 Lepisosteus platyrhincus Florida gar 0.01 1.4 Belonesox belizanus Pike killifish 0.01 0.7 b) Invertebrates Scientific Name Common Name % RA % I Palaemonetes paludosus Grass shrimp 46.2 62 Procambarus alleni Everglades crayfish 12.4 62.7 Procambarus fallax Slough crayfish 5.6 54.9 Gyrinus spp. Whirligig water beetle 4.4 13.4 Anisoptera Dragonfly 3.6 63.4 Coleoptera Aquatic beetle 2.4 20.4 Coenagrionidae Damselfly 0.7 14.8 Planorbella spp. Planorbid snail 0.7 21.8 Corixidae Water boatman 0.4 9.2 Pelocoris femoratus Alligator flea 0.4 16.2 Dytiscidae Predaceous diving beetle 0.3 5.6 Lethocerus spp. Toe biter 0.3 11.3 Physella spp. Physid snail 0.3 8.5 Belostoma spp. Giant water bug 0.1 7.7 Ephemeroptera Mayfly 0.01 1.4 Ranatra spp. Water scorpion 0.01 3.5 CONCLUSIONS Isotope values varied temporally and spatially, and were similar to those from the Everglades. Cypress dome δ 13 C values tended to be more depleted compared with other south Florida systems. Both detritus and algae were food base end-members. Snails, crayfishes, and amphipods were major 1 o consumers, while fishes, shrimp, and most insects were mainly carnivorous. As prairies dried in fall, animals moved into cypress domes but food-web plots showed little evidence for movement. Piscivorous Florida gar had the highest δ 15 N values and highest relative trophic positions. Non-native fishes functioned at similar trophic levels to native species and used a similar range of primary producers. Fig. 8. Introduced fish values overlapped those of native species. Piscivorous Pike killifish had highest trophic position, as expected. 0.00 10.00 20.00 30.00 40.00 50.00 60.00 0.00 2.00 4.00 6.00 8.00 10.00 del 15 N C:N Ratio V I F Py M A 0.00 10.00 20.00 30.00 40.00 50.00 60.00 -40.00 -35.00 -30.00 -25.00 -20.00 del 13 C C:N Ratio V I F Py A M Fig. 4. Bi-plots of δ 13 C and δ 15 N with C:N ratios for groups of biota. V=Vascular Plants, I=Insects, F=Fish, Py=Python, A=Algae, and M=Moss. -4.0 -2.0 0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 -35.0 -30.0 -25.0 -20.0 -15.0 -10.0 -5.0 0.0 Del 15 N del 13 C Algae Amphipod Crayfish Fish Insect Shrimp Vascular Pl. Everglades -4.00 -2.00 0.00 2.00 4.00 6.00 8.00 10.00 12.00 14.00 -40.00 -35.00 -30.00 -25.00 -20.00 -15.00 -10.00 -5.00 0.00 del 15 N del 13 C Algae Amphipod Crayfish Insect Moss Python Snail Shrimp Vascular Pl. Fish Big Cypress Fig. 5. Comparison bi-plots for taxa with all data combined from the Everglades (Loftus 2000) and BCNP (this study). General Patterns δ 15 N values from Everglades animals appear enriched compared to BCNP values. Myriad factors influence δ 15 N values. For example, enrichment may indicate Everglades organisms function at higher trophic levels than conspecifics in BCNP, or it may imply differences in source, fractionation, or assimilation processes in primary producers at the base of the food web. 4.00 5.00 6.00 7.00 8.00 9.00 10.00 -34.00 -33.00 -32.00 -31.00 -30.00 -29.00 -28.00 -27.00 del 15 N del 13 C Oscar Pike Acara Mayan Clarias Hoplo SpotTilap Native Non-native Fishes Spatial/ Temporal Comparisons By Year 2005-06 δ 15 N 2006-07 δ 13 C Fig. 6. Stable-isotope values for taxa consistent among locations & seasons. The three locations had very similar isotope values, though fish/shrimp seemed more enriched at L-28. Most fish and insects were omnivores/carnivores. The web seems to have but three levels: producers, 1 o consumers, and 2 o consumers, with both algal and detrital bases important. By the dry season, domes had few species surviving. L-28 and RP were sites with contiguous marshes. Amphipod Fish Insect Shrimp Vascular P. V - Algae V - Moss Seasonal Comparisons: As animals move from marsh to dome from wet to transition (Liston Et al, 2008), do isotope values reflect those movements? 0 1 2 3 4 5 6 7 8 -33.50 -31.50 -29.50 -27.50 -25.50 -23.50 -21.50 AMPHIPOD CRAYFISH FISH INSECT SNAIL Algae PERIPHYTON UTRICULARIA 0 1 2 3 4 5 6 7 8 -33.50 -31.50 -29.50 -27.50 -25.50 -23.50 -21.50 AMPHIPOD CRAYFISH FISH INSECT SHRIMP SNAIL DETRITUS UTRICULARIA 0 1 2 3 4 5 6 7 8 -33.50 -31.50 -29.50 -27.50 -25.50 -23.50 -21.50 AMPHIPOD CRAYFISH FISH INSECT SHRIMP SNAIL ALGAE DETRITUS UTRICULARIA RP Wet, Marsh RP Transition, Dome RP Wet, Dome Figure 7. Values in wet-season marshes are enriched compared to domes. We see little evidence of enrichment signal in the transition-season domes, 1.5 months later, that we would expect as animals enter the domes for dry-season shelter. We will continue to examine these data. δ 15 N δ 13 C δ 13 C δ 15 N

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Page 1: FOOD-WEB STRUCTURE IN BIG CYPRESS SWAMP WETLANDS BASED ON

FOOD-WEB STRUCTURE IN BIG CYPRESS SWAMP WETLANDS BASED ON STABLE-ISOTOPE RESULTS

William F. Loftus1 and David P.J. Green2,3

1 USGS – FISC, Everglades NP, Homestead, FL, USA; 2 Audubon of Florida, Tavernier Science Center, Tavernier, FL, USA; 3 FL Gulf Coast University, Ft. Myers, FL, USA

RESULTS: We analyzed 777 individual fishes of 24 taxa; 7 were non-native. We also analyzed 751 invertebrate samples of 24 taxa and 733 vegetation samples of 24 taxa. Fish species names are in Table 1. Stoichiometric data for C:N showed that as δ15N values increased, C:N ratios decreased (Fig. 4). Most vegetation taxa had δ13C values within + 2 0/00 of -30 0/00, except algae and moss. Values from BCNP biota were remarkably similar to those from Everglades marshes (Fig. 5).

FUTURE WORK

We will explore the dataset with analytical tools such as the IsoSource (Phillips and Gregg 2003) and circular statistical models (Schmidt et al. 2007) to examine mixing of primary producers in the cypress. We will investigate effects of lipids on values, and will also compare data quantitatively to food webs from other systems.

APPLICATIONS

Hydrological restoration should restore aquatic food webs supporting populations of higher vertebrates. Our data can help test that premise by defining spatio-temporal patterns in cypress-wetland food webs as a baseline for post-restoration comparisons.

Hydrologic restoration will affect aquatic plant communities at the food-web base. We hypothesize that changes at the base will affect key invertebrate groups, with consequences resonating through the web. Stable-isotope analyses will permit tracing of food-web changes at both local and landscape scales. Our data will complement other south Florida food-web studies to provide a more comprehensive understanding of the entire ecosystem.

ACKNOWLEDGMENTS

Funding provided by USGS South Florida PES Program, administered by G.R. Best, and by Army Corps of Engineers CERP-MAP. Special thanks to S. Liston for project assistance, to B. Dunker, N. Katin and N. Silverman for field assistance, and J. Lorenz, J. Trexler, and C. McIvor for help in the study. We also thank K. Dunlop, L. McCarthy, W. Anderson, M. Kershaw, and C. Rebenack for sampleprep and analysis at the FIU-SERC mass-spectrometer facility.

METHODS AND STUDY LOCATIONS

Primary producer, invertebrate, and fish samples of common species (Table 1) were taken three times per year: wet season, transitional season, and dry season. Sampling was at three sites within BCNP cypress habitats: L-28, Bear Island (BI), and Raccoon Point (RP) (Figure 2). Site descriptions, sampling techniques, and physico-chemical data were reported in Liston et al. (2008) (Fig. 3). Three to five individuals or sub-samples were collected for each taxon from each sub-habitat (marsh & dome).

Samples were field-frozen, then measured in the lab prior to dryingappropriate tissues at 50 C. Plant material was acid-treated to remove carbonate. Dried tissue was pulverized, weighed, and prepared for analysis by mass spectrometer at FIU (see Williams & Trexler, 2006).

Error bars not shown on figures for clarity of presentation.

N

5 0 5 10 Km

Bear

Island

HQ

I-75

Raccoon

Point

US 41

L-28

BIG CYPRESS NATIONAL

PRESERVE

NN

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Bear

Island

HQ

I-75

Raccoon

Point

US 41

L-28

BIG CYPRESS NATIONAL

PRESERVE

Figure 2. Maps of study locations

Distinct food webs are evident in south Florida studies; data may serve to relate spatial data to key indicator species. For example, Roseate Spoonbills (pink circle; mean + 1 SE; N=8) receive energetic inputs from Taylor Slough mangroves, near nesting locations and flight paths to foraging grounds (Lorenz data).

CROSS-LANDSCAPE COMPARISONS

0

2

4

6

8

10

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14

-33 -28 -23 -18 -13

Species used in comparison:

• Gar / Mangrove snapper

• Gambusia / Rivulus

• Grass Shrimp

• Snails

• Amphipods

δ13C (‰)

δ1

5N

(‰

)

Big Cypress

Freshwater ‘Glades

TS Mangrove Zone

SR Mangrove Zone

Spoonbill chicks

LITERATURE CITED

Liston, S. et al. 2008. Development and testing of protocols for sampling fishes in forested wetlands in southern Florida. Draft report to USACOE and USGS-PES.

Loftus, W.F. 2000. Mercury bioaccumulation through an Everglades aquatic food web. Ph.D. Dissert., Florida International University, Miami.

Phillips, D.L., and J.W. Gregg. 2003. Source partitioning with stable isotopes: coping with too many sources. Oecologia 136:261-269.

Post et al. 2007. Getting to the fat of the matter: models, methods, and assumptions for dealing with lipids in stable isotope analysis. Oecologia 152: 179-189.

Schmidt, S.N. et al. 2007. Quantitative approaches to the analysis of stable isotope food web data. Ecology 88:2793-2802.

Williams, A.J. and J.C. Trexler. 2006. A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades. Hydrobiologia 569:494-503.

INTRODUCTION

Southern Florida wetlands have been modified by drainage. A major restoration plan, CERP, is attempting to reestablish natural hydrology. CERP success implies aquatic food webs will be restored to support important predator populations. That premise is difficult to measure without data on those webs. From 2005-07, we collected biotic samples to map pathways of energy flow and trophic status of biota in freshwater wetlands of Big Cypress National Preserve (BCNP) (Fig. 1 ). Food webs in cypress wetlands have been relatively unstudied throughout their range.

Food webs provide maps of their biotic constituents, their roles, and interactions. We used stable-isotope analysis to trace the movement of carbon (δ13C) from primary producers to fishes, and to determine the trophic positions of animals by using nitrogen (δ15N).

Basic questions included:What are the major primary producers? How long are food chains?Is there spatio-temporal variability in the web?What roles do non-native fishes play?

Figure 1. Cypress Forest.

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Figure 3. BCNP water depths showing strong seasonal patterns at the three study sites: Bear Island (BI), L-28, and Raccoon Point (RP).

Table 1. Relative abundance (%RA) and % Incidence in samples of a) common fishes and b) common invertebrates collected at the study sites (Liston et al., 2008). All taxa were analyzed for stable-isotope values.

Sailfin molly

(Poecilia latipinna)

Least killifish

(Heterandria formosa)

Ea. mosquitofish

(Gambusia holbrooki)

a) Fishes

Scientific Name Common Name % RA % I

Gambusia holbrooki Eastern mosquitofish 56.7 84.5

Jordanella floridae Flagfish 20.8 40.8

Heterandria formosa Least killifish 10.2 43.7

Lepomis marginatus Dollar sunfish 2.8 29.6

Lucania goodei Bluefin killifish 2.6 46.5

Lepomis gulosus Warmouth 2.1 30.3

Labidesthes sicculus Brook silverside 1.6 15.5

Ameiurus natalis Yellow bullhead 1.3 4.2

Poecilia latipinna Sailfin molly 0.8 11.3

Cichlasoma bimaculatum Black acara 0.7 22.5

Elassoma evergladei Everglades pygmy sunfish 0.6 16.9

Fundulus chrysotus Golden topminnow 0.4 15.5

Clarias batrachus Walking catfish 0.3 1.4

Hoplosternum littorale Brown hoplo catfish 0.2 7.7

Enneacanthus gloriosus Bluespotted sunfish 0.2 9.2

Cichlasoma urophthalmus Mayan cichlid 0.2 8.5

Notemigonus crysoleucas Golden shiner 0.1 4.2

Tilapia mariae Spotted tilapia 0.1 4.2

Lepomis punctatus Spotted sunfish 0.1 2.1

Lepisosteus platyrhincus Florida gar 0.01 1.4

Belonesox belizanus Pike killifish 0.01 0.7

b) Invertebrates

Scientific Name Common Name % RA

% I

Palaemonetes paludosus Grass shrimp 46.2 62

Procambarus alleni Everglades crayfish 12.4 62.7

Procambarus fallax Slough crayfish 5.6 54.9

Gyrinus spp. Whirligig water beetle 4.4 13.4

Anisoptera Dragonfly 3.6 63.4

Coleoptera Aquatic beetle 2.4 20.4

Coenagrionidae Damselfly 0.7 14.8

Planorbella spp. Planorbid snail 0.7 21.8

Corixidae Water boatman 0.4 9.2

Pelocoris femoratus Alligator flea 0.4 16.2

DytiscidaePredaceous diving beetle

0.3 5.6

Lethocerus spp. Toe biter 0.3 11.3

Physella spp. Physid snail 0.3 8.5

Belostoma spp. Giant water bug 0.1 7.7

Ephemeroptera Mayfly 0.01 1.4

Ranatra spp. Water scorpion 0.01 3.5

CONCLUSIONSIsotope values varied temporally and spatially, and were similar to those from the

Everglades. Cypress dome δ13C values tended to be more depleted compared with other south Florida systems. Both detritus and algae were food base end-members. Snails, crayfishes, and amphipods were major 1o consumers, while fishes, shrimp, and most insects were mainly carnivorous. As prairies dried in fall, animals moved into cypress domes but food-web plots showed little evidence for movement. Piscivorous Florida gar had the highest δ15N values and highest relative trophic positions. Non-native fishes functioned at similar trophic levels to native species and used a similar range of primary producers.

Fig. 8. Introduced fish values overlapped those of native species. Piscivorous Pike killifish had highest trophic position, as expected.

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Fig. 4. Bi-plots of δ13C and δ15N with C:N ratios for

groups of biota. V=Vascular Plants, I=Insects,

F=Fish, Py=Python, A=Algae, and M=Moss.

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Fig. 5. Comparison bi-plots for taxa with all data combinedfrom the Everglades (Loftus 2000) and BCNP (this study).

General

Patterns

δ15N values from Everglades animals appear enriched compared to BCNP values. Myriad factors influence δ15N values. For example, enrichment may indicate Everglades organisms function at higher trophic levels than conspecifics in BCNP, or it may imply differences in source, fractionation, or assimilation processes in primary producers at the base of the food web.

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Non-native

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Spatial/Temporal

ComparisonsBy Year

2005-06

δ15N 2006-07

δ13C

Fig. 6. Stable-isotope values for taxa consistent among locations & seasons. The three locations had very similar isotope values, though fish/shrimp seemed more enriched at L-28. Most fish and insects were omnivores/carnivores. The web seems to have but three levels: producers, 1o consumers, and 2o consumers, with both algal and detrital bases important. By the dry season, domes had few species surviving. L-28 and RP were sites with contiguous marshes.

Amphipod

Fish

Insect

Shrimp

Vascular P.

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Seasonal Comparisons: As animals movefrom marsh to dome from wet to transition (ListonEt al, 2008), do isotope values reflect those movements?

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INSECT

SHRIMP

SNAIL

DETRITUS

UTRICULARIA

0

1

2

3

4

5

6

7

8

-33.50 -31.50 -29.50 -27.50 -25.50 -23.50 -21.50

AMPHIPOD

CRAYFISH

FISH

INSECT

SHRIMP

SNAIL

ALGAE

DETRITUS

UTRICULARIA

RP – Wet, Marsh

RP – Transition, Dome

RP – Wet, Dome

Figure 7. Values in wet-season marshes are enrichedcompared to domes. We see little evidence of enrichmentsignal in the transition-season domes, 1.5 months later, thatwe would expect as animals enter the domes for dry-seasonshelter. We will continue to examine these data.

δ15N

δ13Cδ13C

δ15N