For Online Publication Only s1

FOR ONLINE PUBLICATION ONLY

Appendix A: Tables summarizing the porewater salinity, forest structure, growth responses, spatial characteristics, leaf traits, and nutrients for the three study locations used in this study. These data are from Feller and Chamberlain (2007), Feller and others (2003a, b, 2009), Lovelock and others (2005). Distances from the fertilized trees to the mainland and upland vegetation were measured with Google Earth Pro.

Table A1. Summary of the Explanatory Variables Related to the Canopy Characteristics of Rhizophora mangle at Indian River Lagoon (IRL), Florida; Twin Cays, Belize; and Bocas del Toro, Panamá in Response to Nutrient Treatment (Control, +N, +P) in Fringe and Scrub Zones

Location / Zone / Nutrient
Treatment / Salinity
(‰) / Tree Height
(m) / LAI / Growth
(cm∙yr−1∙kg−1) / Distance to mainland (km) / Distance to upland vegetation (m)
IRL / Fringe / Con / 33.0 (2.4) / 5.1 (1.5) / 2.3 (0.8) / 9.6 (2.2) / 2.67 (0.02) / 122.3 (4.5)
+N / 35.5 (1.0) / 4.2 (1.0) / 2.5 (0.5) / 41.2 (16.0) / 2.67 (0.02) / 114.8 (3.2)
+P / 31.5 (2.9) / 3.9 (0.4) / 2.2 (0.9) / 6.4 (1.6) / 2.68 (0.02) / 110.0 (2.6)
Scrub / Con / 32.4 (1.3) / 1.6 (0.3) / 1.6 (0.7) / 3.7 (1.2) / 0.26 (0.01) / 298.9 (7.9)
+N / 32.8 (1.6) / 2.7 (0.7) / 2.0 (1.0) / 51.1 (10.4) / 0.26 (0.01) / 298.8 (8.3)
+P / 32.6 (1.3) / 1.7 (0.2) / 1.8 (0.4) / 5.9 (4.7) / 0.26 (0.01) / 302.6 (6.2)
Twin Cays / Fringe / Con / 36.9 (4.0) / 3.5 (0.8) / 2.6 (0.7) / 9.3 (2.3) / 15.99 (0.08) / 18986.3 (91.4)
+N / 36.8 (2.2) / 4.1 (1.4) / 3.5 (1.2) / 47.0 (18.7) / 16.07 (0.35) / 19070.0 (82.8)
+P / 36.9 (2.4) / 4.3 (1.1) / 3.7 (0.9) / 10.0 (1.7) / 15.97 (0.08) / 18965.6 (83.0)
Scrub / Con / 39.4 (3.6) / 1.2 (0.3) / 1.2 (0.9) / 4.7 (1.0) / 15.96 (0.09) / 18963.3 (91.4)
+N / 39.9 (3.3) / 0.9 (0.2) / 1.4 (0.7) / 12.3 (4.7) / 15.96 (0.1) / 18961.1 (98.5)
+P / 40.9 (3.5) / 2.2 (0.7) / 4.6 (1.6) / 122.7 (27.7) / 15.96 (0.08) / 18960.0 (82.8)
Bocas del Toro / Fringe / Con / 33.7 (0.9) / 4.6 (0.8) / 2.2 (0.9) / 19.4 (4.7) / 8.81 (3.4) / 270.0 (232.0)
+N / 30.9 (4.5) / 4.5 (1.2) / 2.9 (0.7) / 77.9 (21.4) / 8.81 (3.4) / 275.0 (247.6)
+P / 33.9 (0.9) / 4.2 (1.4) / 2.6 (0.7) / 31.3 (6.9) / 8.8 (3.4) / 253.7 (225.3)
Scrub / Con / 32.8 (2.8) / 1.1 (0.2) / 1.5 (1.4) / 11.3 (3.2) / 8.83 (3.0) / 278.4 (256.7)
+N / 33.2 (3.0) / 1.3 (0.4) / 1.0 (0.4) / 19.2 (6.9) / 9.35 (3.21) / 257.3 (258.5)
+P / 32.8 (1.9) / 1.5 (0.2) / 1.90 (0.6) / 25.0 (4.0) / 9.34 (3.21) / 250.1 (239.7)
Nested ANOVA / F / P / F / P / F / P / F / P / F / P / F / P
Location / 74.249 / 0.000 / 7.899 / 0.001 / 9.595 / 0.000 / 17.384 / 0.000 / 1044.809 / 0.000 / 1215.064 / 0.000
Zone/Location / 6.694 / 0.000 / 205.330 / 0.000 / 18.152 / 0.000 / 6.857 / 0.000 / 42.734 / 0.000 / 10.480 / 0.000
Treatment/Zone/Location / 1.289 / 0.233 / 6.803 / 0.000 / 7.412 / 0.000 / 12.176 / 0.000 / 0.031 / 1.000 / 0.039 / 1.000

Values are means ± 1 SD with the F- and P-values for the nested ANOVAs below.

Table A2. Summary of the Explanatory Variables Related to Leaf Traits of Rhizophora mangle at Indian River Lagoon (IRL), Florida; Twin Cays (TC), Belize; Bocas del Toro (BdT), Panamá in Response to Nutrient Treatment (NE) as (Control (Con), +N, +P), in Fringe (F) and Scrub (S) Zones (Z) at Each Location (Loc)

* / LMA / %C / %N / %P / %K / C:N / C:P / N:P
Loc / Z / NE / Mean (SD) / Mean (SD) / Mean (SD) / Mean (SD) / Mean (SD) / Mean (SD) / Mean (SD) / Mean (SD)
IRL / F / Con / 0.016 (0.001) / 45.1 (0.9) / 1.26 (0.19) / 0.10 (0.01) / 0.74 (0.10) / 36.5 (6.0) / 473.8 (58.8) / 13.1 (1.0)
+N / 0.015 (0.001) / 46.3 (1.0) / 1.53 (0.13) / 0.11 (0.00) / 0.81 (0.09) / 30.5 (2.5) / 441.0 (23.5) / 14.5 (1.5)
+P / 0.017 (0.002) / 46.3 (0.6) / 1.23 (0.28) / 0.09 (0.02) / 0.67 (0.06) / 39.4 (9.9) / 521.5 (102.7) / 13.4 (4.5)
S / Con / 0.021 (0.002) / 44.3 (1.1) / 1.15 (0.07) / 0.09 (0.01) / 0.78 (0.11) / 38.5 (2.8) / 474.0 (41.0) / 12.3 (1.0)
+N / 0.021 (0.001) / 44.5 (0.6) / 1.04 (0.06) / 0.10 (0.01) / 0.80 (0.14) / 42.9 (2.2) / 463.9 (70.2) / 10.9 (2.1)
+P / 0.020 (0.002) / 43.9 (0.9) / 1.10 (0.04) / 0.10 (0.01) / 0.72 (0.07) / 40.0 (1.7) / 452.3 (29.5) / 11.6 (1.1)
TC / F / Con / 0.022 (0.003) / 43.4 (0.6) / 0.90 (0.19) / 0.06 (0.01) / 0.71 (0.05) / 50.5 (10.9) / 749.8 (122.1) / 15.3 (4.3)
+N / 0.020 (0.002) / 44.0 (1.2) / 1.05 (0.16) / 0.06 (0.01) / 0.79 (0.17) / 42.8 (6.5) / 762.1 (74.4) / 18.0 (2.7)
+P / 0.021 (0.001) / 44.0 (1.3) / 0.91 (0.25) / 0.06 (0.01) / 0.75 (0.19) / 50.4 (10.1) / 689.6 (77.9) / 14.4 (4.8)
S / Con / 0.027 (0.001) / 43.6 (1.2) / 0.96 (0.16) / 0.04 (0.00) / 1.10 (0.22) / 46.4 (7.5) / 1087.8 (90.3) / 24.0 (4.6)
+N / 0.027 (0.001) / 44.7 (1.7) / 1.25 (0.20) / 0.04 (0.00) / 1.16 (0.20) / 36.7 (6.2) / 1182.6 (143.4) / 32.9 (5.2)
+P / 0.021 (0.001) / 44.6 (1.6) / 1.02 (0.15) / 0.08 (0.01) / 0.52 (0.05) / 44.4 (5.5) / 583.8 (48.4) / 13.3 (1.6)
BdT / F / Con / 0.022 (0.001) / 47.0 (0.9) / 1.12 (0.08) / 0.07 (0.01) / 0.60 (0.07) / 42.1 (2.5) / 695.1 (74.0) / 16.5 (1.5)
+N / 0.023 (0.001) / 46.4 (0.7) / 1.15 (0.09) / 0.06 (0.01) / 0.65 (0.09) / 40.6 (3.1) / 752.8 (76.7) / 18.7 (2.3)
+P / 0.023 (0.001) / 47.4 (0.8) / 1.09 (0.06) / 0.07 (0.00) / 0.62 (0.06) / 43.6 (2.4) / 702.3 (52.8) / 16.1 (1.3)
S / Con / 0.025 (0.002) / 46.5 (0.9) / 1.14 (0.04) / 0.05 (0.01) / 0.72 (0.08) / 40.8 (1.9) / 899.4 (101.9) / 21.7 (2.4)
+N / 0.026 (0.002) / 47.2 (0.6) / 1.41 (0.18) / 0.05 (0.00) / 0.75 (0.11) / 33.9 (4.3) / 984.3 (75.5) / 29.5 (4.4)
+P / 0.024 (0.001) / 47.1 (0.7) / 1.02 (0.09) / 0.07 (0.01) / 0.60 (0.05) / 46.7 (3.8) / 639.9 (74.1) / 13.8 (1.4)
Nested ANOVA / F / P / F / P / F / P / F / P / F / P / F / P / F / P / F / P
Loc / 153.559 / 0.000 / 108.391 / 0.000 / 20.649 / 0.000 / 361.014 / 0.000 / 26.178 / 0.000 / 16.661 / 0.000 / 270.156 / 0.000 / 91.913 / 0.000
Z/Loc / 80.112 / 0.000 / 9.798 / 0.000 / 12.869 / 0.000 / 16.088 / 0.000 / 6.879 / 0.000 / 6.725 / 0.000 / 42.239 / 0.000 / 34.677 / 0.000
T/Z/Loc / 11.045 / 0.000 / 1.634 / 0.090 / 6.175 / 0.000 / 22.793 / 0.000 / 14.659 / 0.000 / 4.823 / 0.000 / 31.876 / 0.000 / 25.476 / 0.000

*LMA = leaf mass per unit area; C = carbon; N = nitrogen; P = phosphorus; K = potassium

Values are means (± 1SD). F- and P-values for the nested ANOVAs are listed below.

Literature Cited for Appendix A

Feller, I.C., and A.H. Chamberlain. 2007. Herbivore responses to nutrient enrichment and landscape heterogeneity in a mangrove ecosystem. Oecologia 153: 607-616.

Feller, I.C., C.E. Lovelock, and C. Piou. 2009. Growth and nutrient conservation in Rhizophora mangle in response to fertilization

Feller, I.C., K.L. McKee, D.F. Whigham, and J.P. O'Neill. 2003a. Nitrogen vs. phosphorus limitation across and ecotonal gradient in a mangrove forest. Biogeochemistry 62: 145-175.

Feller, I.C., D.F. Whigham, K.L. McKee, and C.E. Lovelock. 2003b. Nitrogen limitation of growth and nutrient dynamics in a disturbed mangrove forest, Indian River Lagoon, Florida. Oecologia 134: 405-414.

Lovelock, C.E., I.C. Feller, and K.L. McKee, and R. Thompson. 2005. Forest structure of the extensive Caribbean mangrove forests of Bocas del Toro, Panama. Caribbean Journal of Science 41: 456-464.

Appendix B: Descriptions of the three sites sampled for the effects of nutrient enrichment on herbivory in mangrove forests along a latitudinal gradient from a warm temperate climate in central Florida to tropical in Belize and Panamá.

This study was conducted at three locations along the Atlantic and Caribbean coasts from Florida to Panama (Figure B1). These locations represented a climatic gradient from a warm temperate climate in central Florida to the tropics in Belize and Panama over more than 2050 km and 18.4º of latitude at: 1) Indian River Lagoon, Florida; 2) Twin Cays, Belize; 3) Bocas del Toro, Panama. The three locations were located at sea level and had similar forest structure, which was characterized by complex gradients in tree height that included a narrow seaward fringe of pure stands of uniformly tall (5–6 m) R. mangle, varying in width from 5–20 m wide (Figure B2). Tree height decreased rapidly to landward with interior areas dominated by old-growth stands of low stature, or ‘scrub’, (≤1.5 m tall) trees. The black mangrove (Avicennia germinans L.) and the white mangrove (Laguncularia racemosa (L.) Gaertn. f.) were also present in each of these locations, primarily near the landward ecotone. The hydrogeomorphic settings were variable among the three locations (Table B1). IRL and Bocas del Toro were continental in contrast with Twin Cays, which was an offshore archipelago of small mangrove cays. However, the mineralogies of Twin Cays and Bocas del Toro were more similar (Phillips and others 1997; Macintyre and others 2004; Coates and others 2005), with mangrove forests developing atop carbonate platforms and forming deep peat deposits. All sites were microtidal with mixed semidiurnal tides (Kjerfve 1982; Kaufman and Thompson 2005). The fringe zones at the three locations were similarly well-flushed, but the hydrologies of the scrub zones varied. At IRL, the scrub R. mangle stands drained completely at low tide during the summer, but remained inundated for days during the winter (Feller and others 2003b). At Twin Cays, many of the scrub R. mangle stands in the interior portions of the island were waterlogged and perennially flooded except at unusually low tides (McKee and others 2007). In contrast, the scrub zone at Bocas del Toro regularly drained completely at low tide (Lovelock and others 2005).

At the IRL, our experimental sites were situated on the lagoon side of two barrier islands. The fringe site was in Avalon State Park on North Hutchinson Island, St. Lucie County; the scrub site was in the Hobe Sound National Wildlife Refuge on Jupiter Island, Martin County. These sites were previously impounded for mosquito control. In this area, soil was composed primarily of sand with mangrove forests adjacent to coastal strand vegetation and maritime hammocks. Descriptions of forest structure, hydro-edaphic conditions, growth, nutrient dynamics, and photosynthesis at the Avalon State Park site were previously reported (Feller and others 2003a; Lovelock and Feller 2003). At Twin Cays, our fringe and scrub sites were located on the two largest cays in this 92-ha mangrove archipelago. Descriptions of forest structure, biogeochemistry, ecophysiology, growth, and nutrient dynamics were previously reported (McKee and others 2002; Feller and others 2003b, 2007; Lovelock and others 2006a,b,c,d). These offshore islands were underlain by deep mangrove peat 8–12 m thick (Macintyre and others 2004; McKee and others 2007). At Bocas del Toro, fringe and scrub sites were located on three islands (Isla San Cristobal, Isla Solarte, Isla Popa) in Almirante Bay and the Chiriqui Lagoon in a vast network of mangrove islands and mainland peninsulas covering approximately 2885 km2 (D’Croz 1993; Guzman and others 2005; Lovelock and others 2004, 2005). Here, mangroves occurred adjacent to tropical rainforests and grew on peat about 5 m deep over ancient coral reef limestone (Phillips and Bustin 1996; Phillips and others 1994, 1997). Bocas del Toro was outside the hurricane belt, but flooding was common. Episodic earthquakes were the major non-anthropogenic disturbance regime influencing these forests.

Literature Cited for Appendix b

Coates, A.G., D.F. McNeill, M.P. Aubry, W.A. Berggren, and L.S. Collins. 2005. An introduction to the geology of the Bocas del Toro Archipelago, Panama. Caribbean Journal of Science 41: 374-391.

D’Croz, L. 1993. Status and uses of mangroves in the Republic of Panama. Pp. 115-127 in Technical report of the project conservation and sustainable utilization of mangrove forests in Latin America and Africa regions. Part 1. Latin America. L.D. de Lacerda (Ed.). International Society for Mangrove Ecosystems and International Tropical Timber Organization.

Feller, I.C., K.L. McKee, D.F. Whigham, and J.P. O'Neill. 2003a. Nitrogen vs. phosphorus limitation across and ecotonal gradient in a mangrove forest. Biogeochemistry 62: 145-175.

Feller I.C., C.E. Lovelock, and K.L. McKee. 2007 Nutrient addition differentially affectsecological processes of Avicennia germinansin nitrogen vs. phosphorus limited mangrove ecosystems. Ecosystems 10: 347-359.

Guzman, H.M., P.A.G. Barnes, C.E. Lovelock, and I.C. Feller. 2005. CARICOMP mangrove, seagrass and coral reef sites in Bocas del Toro, Panama. Caribbean Journal of Science 41: 430-440.

Kaufman, K.W. and R.C. Thompson. 2005. Water temperature variation and the meteorological and hydrographic environment of Bocas del Toro, Panama. Caribbean Journal of Science 41: 392-413.

Kjerfve, B. 1982. Water exchange across the reef crest at Carrie Bow Cay, Belize. Smithsonian Contributions to Marine Science 12: 59-62.

Lovelock, C.E. and I.C. Feller. 2003. Photosynthetic performance and resource utilization of two mangrove species coexisting in hypersaline scrub forest. Oecologia 134: 455-462.

Lovelock, C.E., I.C. Feller, K.L. McKee, B.M.J. Engelbrecht, and M.C. Ball. 2004. The effect of nutrient enrichment on growth, photosynthesis and hydraulic conductance of dwarf mangroves in Panama. Functional Ecology 18: 25-33.

Lovelock, C.E., I.C. Feller, and K.L. McKee, and R. Thompson. 2005. Forest structure of the extensive Caribbean mangrove forests of Bocas del Toro, Panama. Caribbean Journal of Science 41: 456-464.

Lovelock, C.E., M.C. Ball, B. Choat, B.M.J. Engelbrecht, N.M. Holbrook, and I.C. Feller. 2006a. Linking physiological processes with mangrove forest structure: phosphorus deficiency limits canopy development, hydraulic conductivity and photosynthetic carbon gain in dwarf Rhizophora mangle. Plant, Cell and Environment 29: 793-802.