<%BANNER%>

Phosphorous Storage Dynamics in Wetland Vegetation and Forage Grass Species: Facilitating Wetland Hydrologic Restoration...

xml version 1.0 encoding UTF-8
REPORT xmlns http:www.fcla.edudlsmddaitss xmlns:xsi http:www.w3.org2001XMLSchema-instance xsi:schemaLocation http:www.fcla.edudlsmddaitssdaitssReport.xsd
INGEST IEID E20110217_AAAACG INGEST_TIME 2011-02-17T20:23:04Z PACKAGE UFE0015200_00001
AGREEMENT_INFO ACCOUNT UF PROJECT UFDC
FILES
FILE SIZE 8423998 DFID F20110217_AABXNV ORIGIN DEPOSITOR PATH smith_j_Page_075.tif GLOBAL false PRESERVATION BIT MESSAGE_DIGEST ALGORITHM MD5
7c128f2a2da7b6a79d4c2cf58ee2ff0f
SHA-1
c597ce621fbffc6c70070a34e61145227f33e150
F20110217_AABXOJ smith_j_Page_104.tif
210c7d8dc67926a4f4bf9f241edab3f8
f941a5df16ed5c28c16e4ad95b1e36e1ead134fa
F20110217_AABXNW smith_j_Page_076.tif
a0a61bbf2546af468d00ce09603cd8dd
09ef755b43d9010ae78659ec3be084e5b78d39c9
F20110217_AABXOK smith_j_Page_106.tif
4fa0bb572d672bfef5b03f5b80851bb1
73795736b7c516f123952e270f85187e55068cd0
F20110217_AABXNX smith_j_Page_078.tif
2269161c913c10f0139c1cc7b1149306
4cd870ee52132f0a9779d3c5d1e4d16d63cb4235
F20110217_AABXPA smith_j_Page_135.tif
c2aa3057346245ae555543bf7bc5017c
1dcc648572561474860163785eaabd901ec37b4b
F20110217_AABXOL smith_j_Page_107.tif
4eaa23b2483711149a238d7f113e9534
66d4be50bd66a1e0658f6194904886e30b36cb0a
F20110217_AABXNY smith_j_Page_079.tif
138f8f0a17c9998d59debd403349bf0f
dcef9dc72bb3e5d0e7d2623b1d6564b1e8910191
F20110217_AABXPB smith_j_Page_136.tif
36da7ceebc2ca9ecce1953d536000fd6
15d7536d2cbf2694aa0f6ede641b2d413b74c6d2
F20110217_AABXOM smith_j_Page_110.tif
5c17e8801dbf04dc5e2b617803187196
08467f3035244b96ecb667a272615d8bd0354f1c
F20110217_AABXNZ smith_j_Page_082.tif
db3183f97b7963daeacbbec98fe7f459
a32175116036548805521137db654bd59c081db8
F20110217_AABXPC smith_j_Page_138.tif
50dc5876be065bf74b76db339fdf6a1c
ea267baf55301c335c3be449598a590e7cbf6398
F20110217_AABXON smith_j_Page_111.tif
fe6a7d4d27d57511aaead14f157e74bb
308e8099bf0cdc51d6614471614e151af9e05e08
F20110217_AABXPD smith_j_Page_139.tif
a71d0ca5e96993cbbea1415307c2e59d
a566d02b579fd147dd1d14c719cfee47c511e295
F20110217_AABXOO smith_j_Page_113.tif
581594a7982a08fe68553a3303c79f06
1b6bf0684a67c63f4b1b1bc2afcf5e277250b616
F20110217_AABXPE smith_j_Page_144.tif
17193adc7c915f0ac878237e7626ce08
7672762ffd147fc1e9b5ca3796ca53f06162cd7d
F20110217_AABXOP smith_j_Page_114.tif
e4443eefe3acd854df3eeea6350a89eb
a48c27b41b6f25fc2b163f8a3283d34f6a25135b
F20110217_AABXPF smith_j_Page_147.tif
f04317cc51a0445245749bfa22f2f066
204f5c9587ac78bb5de0021ec80014dab1162594
F20110217_AABXOQ smith_j_Page_115.tif
dbd476a1859e41caf0c1c4731f21e07c
f1d49597604c66f228bc176d3aee1f477b210601
F20110217_AABXOR smith_j_Page_116.tif
55bc97321b7402331ce0a4c0f2e01988
da0a3e076e1582fe4684629b61fe578c919323f6
F20110217_AABXPG smith_j_Page_149.tif
86633fe2c985dc380ba802f2887ac99f
e7f56cc2ef8bccd96ab773a1664f9d485abe308d
F20110217_AABXOS smith_j_Page_118.tif
fdfa4a9cbec0d625bd235300dd2a8bc1
629810bd68f3a67f47b4da24d5b645eddf05ec86
F20110217_AABXPH smith_j_Page_151.tif
117e6b43187afcdadce1165980abd76b
3a4332e496c17d3c11d73461024956e265cbf863
F20110217_AABXOT smith_j_Page_119.tif
1b4ef16e4ba6f0f2b0988a3d36171802
551077ae684da5b145e5984c7aa5a1b26fb8e7cd
F20110217_AABXPI smith_j_Page_152.tif
52e3467738257115223ffd33c6244f97
b02fc0ee2b45c159d4c82f0d1ff8617a31386e92
F20110217_AABXOU smith_j_Page_120.tif
e8b761caa5b18709278ae5b3571e00ce
407c441639b0ccea00a50047ad61c2a2076c86dd
F20110217_AABXPJ smith_j_Page_154.tif
4bb86504d1e5d5642d5e27f695f2a090
20eede9d72287f75fb0a911da9fce21765218927
F20110217_AABXOV smith_j_Page_121.tif
4c9f4a0af4bd08a824ed77d12a8ac88b
85feed7d67390ddf4f9754d9dd03b1a0ecf0e400
F20110217_AABXPK smith_j_Page_155.tif
b4ce49d081115083c616a3dd35a3b0bf
50abdd7def8078794b98e16e57d3723b4494da25
F20110217_AABXOW smith_j_Page_123.tif
fedb5d62ad36a8d9a642430bb101865e
726127b849fc501c14e2e944cddcad592475fea5
1947 F20110217_AABXQA smith_j_Page_028.txt
182fc7ab399160b5ba7a5fd8f045a652
b250e557ecc884647cbec28a06725d2c98824c6d
F20110217_AABXPL smith_j_Page_156.tif
40c63d6b825415db5eed8a0b5f52585b
f748d953189356d80caee527948c2ef65fc1da1b
F20110217_AABXOX smith_j_Page_125.tif
30ccf4e15c3b9d6d5d8bfe44e33e7484
cc99343dc762f591ba03c57fda347ff622322d8c
1884 F20110217_AABXQB smith_j_Page_029.txt
57d1db07b71fdab0bd3e97bd29e507d8
8d9a025fe653ffea6f1e926d3a2a03336776af98
F20110217_AABXPM smith_j_Page_158.tif
e80207dc40f4bf86ce1329ebd60a7611
5f3773c6b5fdecd276bed09d4863ae43f53e2aeb
F20110217_AABXOY smith_j_Page_131.tif
648e0dda0e88bd4d4d32d2e2da1d46ff
53abd1f2df9b5f50ede7a8cc728325adf0346947
2039 F20110217_AABXQC smith_j_Page_033.txt
c55d233ecb11b688c745ff12b27b0371
bd2d29d5bfea6826ba5438165be2a1d41bbd86e7
F20110217_AABXPN smith_j_Page_159.tif
37981321a59225af4db1407d9bf0a939
1bbe1dabfd8945183e6f68d7aeb160b2df0ed4ef
F20110217_AABXOZ smith_j_Page_133.tif
1b3011c339d61db1854b73f5362da3f8
853f273676a2c1a6db2c351cf865e7ed2d43fc88
1989 F20110217_AABXQD smith_j_Page_034.txt
60379e10fac139a54aca022188a23554
c1c91221934a1187c1fee09cb14bc06d23824966
F20110217_AABXPO smith_j_Page_160.tif
95d8274a102a6bf0e92dbcf9a58ab00f
d9f5206faa10a80b846c8b42522caa4cf699a367
1711 F20110217_AABXQE smith_j_Page_035.txt
0ad303edc1f950458df36c3162ba9724
8d4779c0633dcf8a2eea08087e2052a3cabc3fb5
515 F20110217_AABXPP smith_j_Page_001.txt
941196c7aad451c28cc2ab477345c450
566bcf0d94960bed56da15a7ede9487f707bef01
679 F20110217_AABXQF smith_j_Page_036.txt
711ceb1d28d5f8438da594fec1f57824
f73cdcd01af99d986fc6362d3298cc4a3057b87d
118 F20110217_AABXPQ smith_j_Page_002.txt
4a4bfe2d61468edcb985f801ac42eadf
97d8dc80b9ed4e4e2cff921114d6dee0353e9acc
1960 F20110217_AABXQG smith_j_Page_037.txt
f3f126ed61c2e38d4b92ca296349ff5f
f66025b4b614c899b606c41cb6d4943b430ef829
924 F20110217_AABXPR smith_j_Page_004.txt
d60a95d63df56e1e34d98033bf6b6b3b
e0543ad22063c87a38d86808fe14add67f2b73b0
3640 F20110217_AABXPS smith_j_Page_005.txt
cc7494f30aafbee40fff503c083026cc
3fc11d844dab4b32e58021736e7d785b600c290d
1752 F20110217_AABXQH smith_j_Page_038.txt
10b5ce9be9cc053299c430d6668e72c1
54c5b22e0706d5c455f10f075ddd1f1ff27b45a7
1498 F20110217_AABXPT smith_j_Page_007.txt
1f5cb1692bf51b89ecbdb9f9e140c347
c01fc7f7f4784c3b0d2d7b4b430feaf90f5bcc71
995 F20110217_AABXQI smith_j_Page_040.txt
0852616d9f354f5ace3e46e85e026a10
fffdc21fa0d9e7bd514ddaea35d8550dbbf45b45
2317 F20110217_AABXPU smith_j_Page_008.txt
e6a9de2ac341e5206c1f0574c02100cd
8b08bed4a02d8f921f74439c2fce6b4e1e7063e1
2300 F20110217_AABXQJ smith_j_Page_045.txt
2bbd10881a37ac0bd4e87abbc01b98ac
7c71801c6da362be0967049f429d11272abc3213
2591 F20110217_AABXPV smith_j_Page_009.txt
8e39afacf12221fa18322c06de7c7680
8320aae3ed18e1dc13eb8d3d07ffdafcf6289b47
2009 F20110217_AABXQK smith_j_Page_046.txt
dc646507484c0d85a3f082fd0bfad26b
8230605213533144c76695d1e59f460d732ac5a2
2273 F20110217_AABXPW smith_j_Page_011.txt
3d8a26ad0a3ce2196325aa6b9e7f4579
da23c0e3eea8c4c2f160bce3e16ab7c3b9b94420
978 F20110217_AABXQL smith_j_Page_049.txt
c3d39d37be8ea37394d81490860bd555
a83f63b429a767f448409e1821761f0e490c178d
883 F20110217_AABXPX smith_j_Page_013.txt
190a61f96849137dc7ae00b4353c7875
f0514028eb9ad8acb6f05692e2b602400d38c685
1250 F20110217_AABXRA smith_j_Page_072.txt
65965e1b6b546ace16cf28305758b8e8
b85e346d6efb344899bdc8b05fb63b2fada552b9
1863 F20110217_AABXQM smith_j_Page_050.txt
8138bda1c4d75a3ded4c33c402bde262
b220956df0b3f086fc16f9eb3159d612e3eb6ff8
1735 F20110217_AABXPY smith_j_Page_022.txt
08132696d942b2830112026113adacc8
241b50e413732d00b0eaff4124643ca6008828ce
2214 F20110217_AABXRB smith_j_Page_073.txt
c5dc81bdf6d9176e4384f68882d41d88
3c016fba86e1e4ed2ef0574f0f640ce26a6a87f4
1575 F20110217_AABXQN smith_j_Page_051.txt
85115d404facb95d37cf45b9366463a2
3a9182fc5cc1315a5a50cfbfaee905745b0aefde
1995 F20110217_AABXPZ smith_j_Page_027.txt
9b14df39c208f97db51d0d2098ba83c5
3143da88f23fed4e0bd4c967d3e401c919f7cc8d
1697 F20110217_AABXRC smith_j_Page_075.txt
3d96bd37e6df4258a2521706e7197c2d
f73b3f443f321e4cc1108244b0fbf1e606f041eb
991 F20110217_AABXQO smith_j_Page_052.txt
460a5bd246315d35c72f4fb23862a28a
faa0d9f1027f9714e7452010d93883cc02ec656c
1416 F20110217_AABXRD smith_j_Page_076.txt
a84b20fc067d22c1ae9dc008700d77d9
6cb64f23504f07cf05a1c76da1e6b142c15b02e9
1895 F20110217_AABXQP smith_j_Page_054.txt
40fa31724a9ff288af46564c2b0b9755
2e264e0f656f44372ce69b8b188b0e9f3668b666
1290 F20110217_AABXRE smith_j_Page_077.txt
da97a41e675a4e39406f9aaed4034f67
1f17ab1435b4818ce0270a2f449ab073c8e0f5bb
2147 F20110217_AABXQQ smith_j_Page_055.txt
435071355932da735c90745929bc9af4
26e423f18374c375aeb403d3c9f8e2f36bb47222
1084 F20110217_AABXRF smith_j_Page_081.txt
d503a649b24ccbeb39e547278d2ff71d
0ec689fede00a47c739f8330725738d82036c34c
1242 F20110217_AABXQR smith_j_Page_057.txt
73620172062a010dd180baa56e197cc8
3f0d0465e877dd0d877f224f5ec4b6c5b9642a4f
1913 F20110217_AABXRG smith_j_Page_084.txt
05355382a3dbef076987c70ebcd1ab0d
9330a2cfdbf47268b97fba8baa7fd2aef6cb293e
1454 F20110217_AABXRH smith_j_Page_085.txt
6ef9bf6a71c5033cb937a1dcdb1a1206
41b498a499cc65aea5dfdcbdfb85f23253e5543d
1780 F20110217_AABXQS smith_j_Page_058.txt
629fdc55842a4baaa9e294177ffb2468
fbe7a7025a0f6f1bcd776cce63b455f51d00746f
1957 F20110217_AABXQT smith_j_Page_062.txt
5cd3dd957c491b8b8cfcd98a696a0117
ee564594c18377a218e25f626bf503606c8c3653
2022 F20110217_AABXRI smith_j_Page_089.txt
b1275aa305c9d3117f822b94869771fa
520c542f609e680bead5d39b764b733d55e22cc9
1934 F20110217_AABXQU smith_j_Page_064.txt
49eb614b6238bff850e1595eae9e987f
624963ecf5aac6f00cc0c150f4cd3da8989c23ef
1902 F20110217_AABXRJ smith_j_Page_090.txt
dae850f93a084cdea068877b69c33837
f91b79fc813bfc3139948d8230fd97f986777bd0
1803 F20110217_AABXQV smith_j_Page_066.txt
5b74e2194bdd44ed79856708cc6428f6
ca90b844b6fb20d04489db2d32ea06739439637b
1940 F20110217_AABXRK smith_j_Page_091.txt
5daaf6067c37cc77db8e045eaca69dcb
74a6bc2c9187bc1e185d879b1d20a89016288613
1776 F20110217_AABXQW smith_j_Page_068.txt
d3def105543d102d4857408507f528ea
0a148b9f9cc72bb18c3112fcc8a355186a8b4109
1991 F20110217_AABXSA smith_j_Page_115.txt
adc50a7904898880ce64bf19047136d4
5619482eeb96e050abd60dc4d0b5d37ede391d1c
1833 F20110217_AABXRL smith_j_Page_092.txt
7b6e11d21579551ff14a254977c6c2f9
191aa0e83e5248452932425065e1aed520a00d66
1840 F20110217_AABXQX smith_j_Page_069.txt
014fa8291b7ecc510bf637acc82d7703
4599b3cd2e85e27d2f2d646c0690b5e9d02c442d
1974 F20110217_AABXSB smith_j_Page_117.txt
bac35935fbf52c09a17d4ec2f838e94a
926a71a206cfd3a72e0ab95c8b89eedc2d3c3cc4
2064 F20110217_AABXRM smith_j_Page_094.txt
759f2e37138abfd8c0dd25725596df92
5acd01abe08f12069747dac42573e8d7a6390210
1809 F20110217_AABXQY smith_j_Page_070.txt
0333e7ae643697d435014863cf82d348
4f2fc831c23163ad32a9d5f56b0bd3cfcdc560f2
2526 F20110217_AABXSC smith_j_Page_119.txt
77aa13fbf6068af3a9442efc2bd3adaf
7e9824df72a1eabf6c6263bf301200413b79ed74
1862 F20110217_AABXRN smith_j_Page_095.txt
2361dc3b9065a9b79e7104eb7a5dd1c7
1f4d2668e68f6441d161f215415c60e4d883e427
1463 F20110217_AABXQZ smith_j_Page_071.txt
1c9969c6fd2ca8ec87336320c04fed82
d8668695d8e1f39a6b3e37af69102dd16049f676
2494 F20110217_AABXSD smith_j_Page_120.txt
b5e424e2ea38f8979e1a8c58e1475b21
7184969ca541a8122e54cfadf916faa77f6ad3d0
1018 F20110217_AABXRO smith_j_Page_097.txt
6a50212b0f3d4c2b95ac48477812a050
9e7e3c2ad9f137e11dcda6ac44f23e5d647a956b
2895 F20110217_AABXSE smith_j_Page_121.txt
d5a49ec3972a012cac497d4d6ad94668
cd1efbecbada2562eda5657d91dac3022f864c0a
1643 F20110217_AABXRP smith_j_Page_098.txt
a2313d0e80b41132570c09e8ae40ae8f
74a68a7959ff267135287e69d4579a5248fd6d12
448 F20110217_AABXSF smith_j_Page_123.txt
b8a4ada24a6d719a86319a09d5e1d27a
402f184ffd301123ecf5f074e9797dddef6609ed
F20110217_AABXRQ smith_j_Page_099.txt
2af76f2879e48b67783fb08118071ab8
733f7366374874fc5ce24a17a1f4ad6fd9c7f5ee
1692 F20110217_AABXSG smith_j_Page_128.txt
a10361e0e712460ac7eb958ba57a48cc
6bc09e5d8a324ceefe51d5cacbe88436799d9267
2223 F20110217_AABXRR smith_j_Page_102.txt
b9f1e97d0774ff89691a6f83b2eed168
272c9ebaf5da6050a1dfe15e11ea3fab296baafd
2374 F20110217_AABXSH smith_j_Page_129.txt
d810404559988740af00360db4de3a64
be4f1110bb86558635f52aca283d76f59de20ce1
2016 F20110217_AABXRS smith_j_Page_103.txt
9c2164135af783cb1691ee429b26b9f1
8780f6f9e770990d587eeadc6aeaf0e93e591db0
1844 F20110217_AABXSI smith_j_Page_130.txt
8a7054ed01c4ed210a69cbec5b3ae79d
21b0c23c0e8aa06fc963e61ae61f9e9a24bcbe06
1637 F20110217_AABXRT smith_j_Page_105.txt
a985be05d4a94083c033dec12857c05a
04d25fe4973a34357485765bb27eb1080d32c920
1701 F20110217_AABXRU smith_j_Page_106.txt
7382350742c282388b034275e4277df6
9259f630b09394788c8b04ecb8f5d0724403a0ea
1629 F20110217_AABXSJ smith_j_Page_132.txt
140fe195bc366a5ac09bd1aac24512f2
c43d4f300aac102da96efd5b6c6878e735e9529c
1453 F20110217_AABXRV smith_j_Page_107.txt
8c4e5db377d66fc5c4e8ad8919dc98d2
108d0de2dc435c31c438d3df3126f83607d1fa64
2472 F20110217_AABXSK smith_j_Page_134.txt
66a66dc95800cc0ee2afe818d1e67156
d330e86238eef38fe3eacbd5f5376d66b4b1b2c0
968 F20110217_AABXRW smith_j_Page_108.txt
6875dd0d5a0d06b79083b77ba44cf01f
80b8fcd9781d57dedc1d888e13d9fddb652fc80c
2497 F20110217_AABXSL smith_j_Page_135.txt
d5969d6f16656b72a2fd905263f35657
2a53c0d31256d4350412a158765fb18f92f1e078
873 F20110217_AABXRX smith_j_Page_111.txt
884466261492184310004b5be5560be6
0f912766e76fbb2c1c366412c459a3e1c0b65686
888 F20110217_AABXTA smith_j_Page_161.txt
817d02d6eb563e70c756b32f343fbbd3
4736c0e83eb792156d612c806563b63d3d381e97
2115 F20110217_AABXSM smith_j_Page_139.txt
ee52d40747ef274fe119e6754a097d33
2a338fec4ea853e9e2b598dd0b716e7c6f489e1b
F20110217_AABXRY smith_j_Page_112.txt
0a92050b9e19450266957daf336e3868
4890b0123c245684a1a85692c9ccf6a7c0c438fb
9768 F20110217_AABXTB smith_j_Page_001.pro
3a3f0b0a7a12763e6e315d3945af275d
9727b12d40220188aa02727f3e0ec3b78ccfcc66
1006 F20110217_AABXSN smith_j_Page_141.txt
7bd2c623ca26e390720677b6e1518b9c
53f7922354df17135bc67b9af2f3bddcf73b5a72
501 F20110217_AABXRZ smith_j_Page_113.txt
e83d456c6195cebfee17fcd2af6424f3
068d8d1fa0b5f6eb994ac77e8ccb49e9c383d7ae
1278 F20110217_AABXTC smith_j_Page_002.pro
323822f00a60b1b61d02263fd35732bc
f858a23d102b9f53c993c2729e0d6b322815a0bb
2379 F20110217_AABXSO smith_j_Page_142.txt
9757802b963594c05763a73dcb592fa8
82a7136f365de4849eea672851d0ae367a444fea
12544 F20110217_AABXTD smith_j_Page_003.pro
49847c3fa407899ae1f44e46b50f13dc
5bbd7c2e85ffc4320e10f9ebd29234c3d5879e20
2976 F20110217_AABXSP smith_j_Page_145.txt
bae3c9412e6c1ff5015fcdcd684e54c5
a6e871f45588303fe4fcc80959bfd53bf9532148
87951 F20110217_AABXTE smith_j_Page_005.pro
3a12ba96e01db00a6bd7b615cbd5d819
93fa524b9f57854c605a0b9d98de70437de99fc6
3339 F20110217_AABXSQ smith_j_Page_147.txt
264b0e5987d6475e1107e7926e1ff92e
1018b38aabc829badf4b325a989f02e166392a70
37307 F20110217_AABXTF smith_j_Page_007.pro
201ec598b8f175a9d13633141db532a9
3d73a4941542b4cd2faac4ca9ba49765a5d9bb3b
3028 F20110217_AABXSR smith_j_Page_149.txt
38265ec82fc3008bac46550f8517f177
08a07301409cfd5a17dc61400c215f7363154c7c
57856 F20110217_AABXTG smith_j_Page_008.pro
0f57e1249bad65ad2a61c63bc9f36876
d16c19e8c893f71e21a9fea55d53492c58420d8f
3225 F20110217_AABXSS smith_j_Page_151.txt
6547c6fbc8121e1cf159914ab406e98c
6d15db3a03e641d0166c2ba6b9c257686dcc72ab
33545 F20110217_AABXTH smith_j_Page_010.pro
feb55899461e884dca76314743266c07
8151739844afb59edc5cf8f2c9bb14c993f7608b
1600 F20110217_AABXST smith_j_Page_152.txt
c722ff7f083f36d027fdf46a9a5623a4
d20ffefbb1662d73b39af09b8cb1d740529d01e6
57238 F20110217_AABXTI smith_j_Page_011.pro
380639dac08b99c8b7837d284edecff8
2c521dfbb93c8d1561583180219d42a8ed1ea6c8
2228 F20110217_AABXSU smith_j_Page_154.txt
642302c74ee7b2c8bc93877f39bccc9b
8337a961793c9ee81f1b9b2e18d3b04f9dd1b413
67633 F20110217_AABXTJ smith_j_Page_012.pro
d4e8953391aa7e03d0f8b16f1bbc2110
2120689cd415918b95f173443ded9ffce23837fd
2322 F20110217_AABXSV smith_j_Page_156.txt
072800919f8fb29179703a9557c86d1b
d07134ffaeb483ac8c90ee2816dcac04db15758b
2444 F20110217_AABXSW smith_j_Page_157.txt
588623c2f35fdb9dcc7049570db4cf8c
1caf17994197d52ca6bb393aa258f50ab68797f4
45018 F20110217_AABXTK smith_j_Page_014.pro
d53a1aa30f269bf86bcb065ab62184e5
6fe92b9ae85227ce3a7eabfaacfa403e776fdedd
2336 F20110217_AABXSX smith_j_Page_158.txt
977119e3b52284139227315a4054256a
cb4836515faaf624c4749b561b3d9201de480a90
42994 F20110217_AABXUA smith_j_Page_038.pro
9608399fae556a431cc4209b7cd70a15
8ab23d694b8c5fdcd247d99efb671561d6bd77ae
50665 F20110217_AABXTL smith_j_Page_015.pro
00a5c4fb070fa9bed2eae032694a46cd
7d18eb62e3c7bfe6f600cb7252c64601dba53814
2396 F20110217_AABXSY smith_j_Page_159.txt
ccc1aced01079651858c49cbbb8ca5b5
fa4ab9469c47a18f13fd482401cc0e6b04d733fb
23644 F20110217_AABXUB smith_j_Page_040.pro
05d523659595f4402cbd258a72db8fba
042db59a4ce4777ecee3272cd829e46348018546
44734 F20110217_AABXTM smith_j_Page_017.pro
1effeb6cdaf11767a8a7dc79dda54d95
1a42c2bd5f6be531bc41fa24a9c663945f4ac74e
853 F20110217_AABXSZ smith_j_Page_160.txt
11983ac5e7ac34c3646e950108d4628d
a0f79aa4783e6c63326f37159d04b8d49e7daafc
50131 F20110217_AABXUC smith_j_Page_041.pro
1fb721f50061c3041e54b4f3898346f5
d5956894f7a869c5cb773857633b8b27e8d9e9f1
48424 F20110217_AABXTN smith_j_Page_020.pro
676f0dbad9403fa14d85a80350b3dfa7
b98c5f91e7ed53ed5c1778e9c451a8db7b7a7812
42283 F20110217_AABXUD smith_j_Page_043.pro
741d358b5baa169c7e8388fdd90859f2
f991b54cf689700ef99322735274e7de120c80a9
52668 F20110217_AABXTO smith_j_Page_021.pro
b936421547097de22e83c066a4327a10
5d8fe95fd122cee208b21d7de920da114ac51a12
58553 F20110217_AABXUE smith_j_Page_044.pro
9f18b606d565147bc4b535b87bb6e131
25fc17142274c6cf2d748c18668b96623dd55727
43561 F20110217_AABXTP smith_j_Page_022.pro
9af9ea9567335a5e204a574d28d94e2c
85f8c7764b35dbcdb270f74785f31d026140f743
46927 F20110217_AABXUF smith_j_Page_045.pro
71084917698ef123e1e47b39b72fa7cb
0be56aabdbbe3abc56ffcfc20edbf66b17277467
6223 F20110217_AABXTQ smith_j_Page_023.pro
c9edbf5fd5ca1cbb44382b4cbb191e95
eeba2599d180b7e18a6659ea78d40a2aeed89e45
60030 F20110217_AABYAA smith_j_Page_126.jpg
bbff9625f833dd741e1809087b5c2a18
9a186e5bc48cef99d5d289b02f49cfe97c37482c
41691 F20110217_AABXUG smith_j_Page_046.pro
e78d11277576c8a51db43bce60f7d6ec
d97f78da45e9619f7e46204bf9d59c9f2d4bb55b
48489 F20110217_AABXTR smith_j_Page_025.pro
8ea77650b9528f5c597252a64444d74d
00a9a1fb499cc1c63509be46dcae73f127a90ca0
57212 F20110217_AABYAB smith_j_Page_127.jpg
a809550eca1bf878e12dc243b57b1d90
5c080c90986d50429bbeb482450c47b47cf33967
25023 F20110217_AABXUH smith_j_Page_047.pro
c4eb2d48dd6e65c3ae71e9c03385570e
676beba2e42366030a8f75de570105ed75965e55
51270 F20110217_AABXTS smith_j_Page_026.pro
dec637bf4729b18bb20f1684a8dc4ac7
5fb2180dd78ebd89aa296445518ff8d22a2b3ea8
53649 F20110217_AABYAC smith_j_Page_132.jpg
c8023da6a230c815942e3be482bbc7e0
fdb378e45df99361180d3274343b72b54c21567c
35015 F20110217_AABXUI smith_j_Page_048.pro
23209c0a4168bdf26f8964dca09f4983
1f0c12f1d196fb6a59d7d3a8afb6fa5c78b83599
49320 F20110217_AABXTT smith_j_Page_028.pro
cac0c2b483273beb2cd3e1f134cc6562
5faaed90ea31a54d2715617ab3cfe915ea133eec
75779 F20110217_AABYAD smith_j_Page_135.jpg
85cb77996f89939c828672675af645a1
f1a663688066c5190cd6a36fc0a7ae09ad173f9d
17330 F20110217_AABXUJ smith_j_Page_049.pro
9365044b242264c195ae2a9ffb6f067b
b5780a7f7b52fa94ef1f124a5e87c00fdbaada8b
47687 F20110217_AABXTU smith_j_Page_029.pro
dbadebed6f443529311fed227cf2168d
64536a9fe95fb875398dd9387723ed9cb9dfe68f
53623 F20110217_AABYAE smith_j_Page_136.jpg
993875ed75cf7119397dddd1f1f99388
a3958ecc62048b53a59868363ca34240d004e83a
33602 F20110217_AABXUK smith_j_Page_051.pro
b26903a5ec8b90786d25062fb939796e
8274b39eb59f75b7dcbda3495855c417808e850d
51834 F20110217_AABXTV smith_j_Page_033.pro
85459df5c569f766267a233391c8f128
f71cfc90ca34c0746951a2b453bca84e3945c2d1
51969 F20110217_AABYAF smith_j_Page_137.jpg
9c552ad3343da62c5efb44faf0941c48
827cea501595b77a7433f0940e78482dd3a242d5
50473 F20110217_AABXTW smith_j_Page_034.pro
d533fc51f9a10d77fda789946146d102
99321df36a167a41de83fad7fd343e250cf3ea00
54192 F20110217_AABYAG smith_j_Page_138.jpg
5534b0b7f10d2b6d8e396149e4a4f005
7656ba6ae1f3e3199c0e4fe832fb4067dc4ec0f2
33713 F20110217_AABXVA smith_j_Page_092.pro
cd08fdd9e37c9d23dcff5fec2ec6c9eb
9ee04880d37fad9af5d0e1f15d998f52b211b6a3
20144 F20110217_AABXUL smith_j_Page_052.pro
3ecef50e020cc07cd36487e9a337fd45
4eb70125cccb0525aa48751c15d3ac8ca0196bc9
43071 F20110217_AABXTX smith_j_Page_035.pro
210addae89ac7bf6cb9ce3030ef5474f
fcb2314834160b1f1f9493d666428429ac744408
52398 F20110217_AABYAH smith_j_Page_139.jpg
a653a7886b1b0f37e887935a61085874
4d945f0d008031f63ca798d86ae01e33e5c24849
49088 F20110217_AABXVB smith_j_Page_093.pro
d9c73c31bf2545355227b563c9c089e2
3d8b230d92b63eabc728a29b91bfe0516f495b1c
26076 F20110217_AABXUM smith_j_Page_057.pro
5e9407f695c98743f06001c26ff45ca9
5efb1a8d6a49ef2f75a13c1eb3ba8cdd852da560
15858 F20110217_AABXTY smith_j_Page_036.pro
ddd1f88893cbaf96c7e2b65905dff178
d23e25ea641c917ae4be9f429585bea8326027bb
62469 F20110217_AABYAI smith_j_Page_140.jpg
f93cad077ac0795ffef3531c0368fd2e
515d9458e6d4712c04df50154eaece2025fe324d
52672 F20110217_AABXVC smith_j_Page_094.pro
be4ac60bd733c0085f22831ae8cd1057
7fcb09b01e510db2b84a5bb342f70f2c3a470598
51821 F20110217_AABXUN smith_j_Page_059.pro
f72fe76855137c25409d33a806fa6aa0
ee0374e3c9ecac5afb7e719f738e73d825d4074c
46689 F20110217_AABXTZ smith_j_Page_037.pro
40f30a965103172e1829534b58f07e1d
aaea867e02525a71b547945e386a392f30f4b81c
57985 F20110217_AABYAJ smith_j_Page_141.jpg
e1f1a7d35fdfe9bfdb138ea13f9d8dc4
f6a9988e3c6cf05daa6ff616c8017165fabd9ffe
47169 F20110217_AABXVD smith_j_Page_095.pro
76689a0ccf2d65ab6b1722f91b8b0239
b023cead534a6296feb1fb924e0d3f2f7a45c324
49241 F20110217_AABXUO smith_j_Page_060.pro
d66a462b5d6ba88d6cf15f13262feb80
833eecb8e9e08083fd72283095ded0633c5170f5
70246 F20110217_AABYAK smith_j_Page_142.jpg
aacc21879f09ef153c361b572908e96f
80b83a15459d61b11b861751447afb3935fcae0e
47543 F20110217_AABXVE smith_j_Page_096.pro
671b5ca8aa3f0c8a3919ee67b8945d7d
b5f133db2279ec3e7976dda00a5d72cd28d85e37
43118 F20110217_AABXUP smith_j_Page_066.pro
161a3ab8274bbdeab85ab526f38c4d98
ee4fa0ebb2601a77f7229c858b781e67e877c592
128395 F20110217_AABYAL smith_j_Page_145.jpg
9dc1b84d5005f475a0d79f210327cc46
2be15129a5a03358c988a7bbdbd60ff379b04d36
25245 F20110217_AABXVF smith_j_Page_097.pro
991ca70d1ac8bf96c78b49c20d3c79da
1273e9b4a4151d6bd35d340e4bff68484907512e
42017 F20110217_AABXUQ smith_j_Page_070.pro
d4cacb7f709dbbb043c81cd50642863f
edc9a7033bad8168263537adc7c4fa705d1175a9
498822 F20110217_AABYBA smith_j_Page_010.jp2
960348960510d759e0642f31ff71df85
23b1010c825d2613b3f0ff7eb1c72932d22ef76f
131736 F20110217_AABYAM smith_j_Page_147.jpg
3f63d23f3a889a5675ca4c4c59809fa6
6c04464b7e8678dc3127ad9d87dbae0ab824e31c
38260 F20110217_AABXVG smith_j_Page_098.pro
3030fef27702dbed70035d7d287726b7
333e431976da1b56f6683fc4abbb6200b02c2598
36018 F20110217_AABXUR smith_j_Page_071.pro
1b833e45fbbad122e66cf5d9a114b583
650a7a4aec2bed03b175c665ca770149877b2a68
933426 F20110217_AABYBB smith_j_Page_012.jp2
1f753db6cb84fc42b4afab7a6eb6539b
9a3133e6837f0accb067da01180e8c0655c770d7
115619 F20110217_AABYAN smith_j_Page_148.jpg
598986ed38103ad54e723941d30043ac
4113d4e6a619ebd0b649864b4bb1997fbb5b5ed3
49240 F20110217_AABXVH smith_j_Page_100.pro
23617f2ce733a594eb271516fd4a694d
6de5ba727f46b4b42bb06c42010346bdd468292e
40536 F20110217_AABXUS smith_j_Page_079.pro
e4cb3aba7b91e1b839d27cde2c0f473a
f0702bf7d8a9b5d614e5396ac0007888e31c9368
285286 F20110217_AABYBC smith_j_Page_013.jp2
6ecaebcaab980a2712a34eb09900da39
1f1ea7a607150c142b228da2a89a7d0cf09d3bf5
132292 F20110217_AABYAO smith_j_Page_150.jpg
587d4897651f7f473be4eadbbe74d078
f023fe7802b3a1ac419e98abe6646e5e015a016f
50031 F20110217_AABXVI smith_j_Page_103.pro
ef061dd1364852edd88b1f07036550cb
6c72fe9d2d68b59f1e4332b4bef6a123a411e98c
20824 F20110217_AABXUT smith_j_Page_082.pro
9f2a7646da834568c07d332f7d7b0933
2ba943bfddac9623b34d61f3920ae911fe19730a
1051971 F20110217_AABYBD smith_j_Page_014.jp2
602c9efa6c67d9e0b962976fe60b1784
2309b9481628f475559d691218a6fe326edf776d
129619 F20110217_AABYAP smith_j_Page_151.jpg
e72176ca7ed567aecc91aee4d89086e1
d4e122a00f5433a2f719cdf454b824f1b4ef9e61
38101 F20110217_AABXVJ smith_j_Page_104.pro
2c4b4f3c10b91d1bf6891c15bf0a6dfa
13277db7db0cd2ebaf540c64b131e2eb107e6e3f
48518 F20110217_AABXUU smith_j_Page_083.pro
256ea5f071c75e3b1c0d72e39debc77f
4bca816f945cc31ef45479f1a0054f4301426027
1051977 F20110217_AABYBE smith_j_Page_015.jp2
e3d5845d2a6f42d1b3dee17f5029c8f1
14df65b5aa5370152063c58e1f18d308fa8268b0
67584 F20110217_AABYAQ smith_j_Page_152.jpg
fbd34a50f600ac7c308e18ccace3b13a
131204e04d445dfc7d73f0984acec4c8ee716bc5
38517 F20110217_AABXVK smith_j_Page_105.pro
89ca008baf6021bf0b40210b4978b8d1
cfd6ef92ca044b4e80f84bccb45308dc5fe127f6
41092 F20110217_AABXUV smith_j_Page_084.pro
95618035fda983b1a24b030b46795b93
a3a6a6aedde835c4f6cc4cf0a73a12487a8067de
898741 F20110217_AABYBF smith_j_Page_016.jp2
1ec165f8ad23ad4972acaafecbc2841e
aa7d38db7e14802840faab9d9651d8b8eee7c350
112877 F20110217_AABYAR smith_j_Page_154.jpg
543e6abee830b30cdb4ecb2f3362e69e
4b8d3c169985b00c4ef733cd098baf44895a013f
28042 F20110217_AABXVL smith_j_Page_107.pro
8a70bb45629a2fb143372aa57014ad05
f940e569ff2467755d64ac31b9ffb8d970a81b6e
32737 F20110217_AABXUW smith_j_Page_085.pro
13f44fac6eda38a7412d923f97b28e3f
8632789fd13a80ea8ae95df893ce088d2209a6f4
1012211 F20110217_AABYBG smith_j_Page_017.jp2
9e1d3e89b6cc9edd5c7c86d9f7e0e969
adb6068f2e39f078810a5db95c987d87d12ce084
112232 F20110217_AABYAS smith_j_Page_155.jpg
c77c1171f195c6e214bf69e871a89aa7
3ddcebf88c3d3d0f5a9fe8798158378edd607ba5
41609 F20110217_AABXUX smith_j_Page_086.pro
85f6da630de3310bea760369ac5b5482
c7b0955fc45cf53aa97221f7a722191637b17c28
29144 F20110217_AABXWA smith_j_Page_138.pro
1abb7ba049d09d4c9187a914151bdd88
a853c2ce59884a17160d5476639db74bec22aad7
1047650 F20110217_AABYBH smith_j_Page_019.jp2
6dcd41f19dec5c5c456f0b4e5eac0a31
314efd7d2e4d8a3cafc648d06450a8fe822b6b83
119537 F20110217_AABYAT smith_j_Page_158.jpg
fbbea1e63d158df69085242d5a4bd7da
89528d538bf2cf9223ac896da068886db982ca63
20872 F20110217_AABXVM smith_j_Page_108.pro
ed8cd27ee2a24436536f9e24ffa8e8d3
493c89e3655a4534daec024641da739726f700f0
23424 F20110217_AABXUY smith_j_Page_087.pro
d4c8b8ad76231bb534d2313f87529df4
61b01b5e3b5da67dae6092b77c51cb91580292d6
29214 F20110217_AABXWB smith_j_Page_139.pro
c8d71491c6f2d0b2277a27b381d4b735
021524eafeb5acce8aca44c940862c599bdc93af
897279 F20110217_AABYBI smith_j_Page_021.jp2
a8be100d4a3fe13b130f9cc066a635ad
fd21cfd84ca6e0c363d071143d6ae5731627e184
120342 F20110217_AABYAU smith_j_Page_159.jpg
43e446e5075435f551d45662d39724ae
2100fdfe6d6c45d8e4d3021c9022a70149ad7990
31257 F20110217_AABXVN smith_j_Page_110.pro
9f97d6e1ca8c0091b34d67efa2660750
e5b3f9af06a679cf29025a1036b128e2644d7e91
49118 F20110217_AABXUZ smith_j_Page_091.pro
0e4f1b50c31d6dce98e8aec4bbd0d8be
cfe83ed9e0e040e0e56c0964f74f28c9f87582e5
15048 F20110217_AABXWC smith_j_Page_140.pro
612d3c729f467645fc68323e146feca1
4617225d149665ff63c2a1070cda0eb58a2684ea
1051982 F20110217_AABYBJ smith_j_Page_027.jp2
8f6e00a783be0fb11ea6a199b7879fca
d993733a82d6f63a7be58bc6df5c4fef3b75c2d0
47259 F20110217_AABYAV smith_j_Page_161.jpg
4e4c99f4199271bbf5da9011f3268429
78a0c464d351604d1f1c9e61cb50fb4e5a41225c
33881 F20110217_AABXVO smith_j_Page_117.pro
87d4e16714ffe925c221f1ef8b7b72eb
97bb5988956e44919816e8d06a62a51cad3fde1b
38282 F20110217_AABXWD smith_j_Page_142.pro
afaf3d7d6d3d77630a839eeccc7e4efb
47d6947157d98a74ef40d7d115035566b72ec962
1051932 F20110217_AABYBK smith_j_Page_028.jp2
360e2868b12be2ab6a08c09eb5a39a6a
d4b3c08fdd5cacc355746d1f6607ac47e8a4875f
11266 F20110217_AABXWE smith_j_Page_143.pro
4465a5c9d667bbfe7b37463a96010ed2
0f789361cda1a38fcd825ccc7f4433020712be18
30787 F20110217_AABXVP smith_j_Page_118.pro
512fdeefc03e600a1cda760cc8b0849c
90d54b3ade78d5ed19ed15355c529810f896c112
1051976 F20110217_AABYBL smith_j_Page_029.jp2
48f99281aa00fb20639bbb80d06b00b2
a95e06c5bb8ebd014617af9c968cf0a02e28be15
296354 F20110217_AABYAW smith_j_Page_001.jp2
99e5814ab01efcc56099223944e2c2ce
053b63e3e2756997bb32b74d285e5ada88ab2302
74907 F20110217_AABXWF smith_j_Page_144.pro
8cf030682baf89b158b2700e81397ffc
e932c03d3c33d8b8e06824022262b5bd9d1f7392
41637 F20110217_AABXVQ smith_j_Page_119.pro
60d7a232c5c08fc10b5498cd60a69be9
3e66e4bdb3bb02e862997c106813c81d07f633a9
834867 F20110217_AABYCA smith_j_Page_053.jp2
95265ee5f235d14920fbc8601e4d8e8c
be7a0733c295e7642c2cc89d23c756e2b68a80c7
1051948 F20110217_AABYBM smith_j_Page_030.jp2
24a9d2aa95214792e4c03cd2da045b30
7e3496558ab5f4f6f47d6a9a2558eecf0c7ef731
460113 F20110217_AABYAX smith_j_Page_007.jp2
3d18533a0c340b1a5ff61301319560b6
cdc929df7f51d9f9382e82051c517bc85010be73
74184 F20110217_AABXWG smith_j_Page_145.pro
20d4df9e31e6ff9c3bea04c16e8fef30
ef8c8ee73a5075402b65c015131fa904d34cc081
14943 F20110217_AABXVR smith_j_Page_122.pro
a9d7de83cab834d9a737a7cc58d0eca6
1c50dd7f1c06a9a86de70b53997d365002e5fdda
723647 F20110217_AABYCB smith_j_Page_054.jp2
de21b58e02224805630541cdb57d6d10
c05378d25f648900bde332de0ad7e86dc955529b
374807 F20110217_AABYBN smith_j_Page_031.jp2
11057a47b203a81d5f845f97e485ec51
db6021a7f042b7f040b6441ccedc0a3633524527
728629 F20110217_AABYAY smith_j_Page_008.jp2
05f2b517512f59ed94cbca48803345f1
e2fac5d33437409b22a66fa6b56dd4f2639483fe
73827 F20110217_AABXWH smith_j_Page_149.pro
20c45801a2d9e6e60692cc9cd4d82f36
14d3529970557d4490ede8e3f5196ebe89c14167
7222 F20110217_AABXVS smith_j_Page_123.pro
22e4fd36155896cf9cae33df87d4874f
05936b5697686e35dc9ee509b39d746bd3275d90
706616 F20110217_AABYCC smith_j_Page_057.jp2
7fda55d673e06922135695ba1090b212
b6f091268ac9ce879ef8ee4e29d5bb6d0bf8a4d3
972185 F20110217_AABYBO smith_j_Page_032.jp2
10cad2103e04a5bed292c097bcb48a4b
ef386f093bb95c3af6008dc7eead33f67e06c171
968896 F20110217_AABYAZ smith_j_Page_009.jp2
fb7a03a02a05afcbb124e70b795a1320
3a4ab041fc291d90faeafe6b3377759d2608c5c5
82255 F20110217_AABXWI smith_j_Page_150.pro
50907d1399edd37c30bf55f543b4b0f4
6e54306c17d45c33711323c6285ea1ac37d56912
32445 F20110217_AABXVT smith_j_Page_124.pro
f209754971bb9a8ab45efe4d12f4ce88
afbc0a31e5000b6ad1b3bc517ad0e3787aaffc70
1004746 F20110217_AABYCD smith_j_Page_058.jp2
a692e795a627e42fd5c2068f682c04c2
2b0b2b2b15b857435e753fd9ed19b02ea9dd0e2e
1051983 F20110217_AABYBP smith_j_Page_034.jp2
6c52ba7f37ebd6a21ca72250d4607dcc
20f8ffd642f8089aeba50eb22bb8b630ede8b0e2
31742 F20110217_AABXWJ smith_j_Page_153.pro
42b46302981395f6eabbdc5a36ddef96
9b50dfc1ed5b2342e9660d28c25b8c1b19778682
32830 F20110217_AABXVU smith_j_Page_126.pro
f07a866b7d055c90e49282a4ec5502e9
576620f3b270524b591a8eff74b3c749c0264f24
F20110217_AABYCE smith_j_Page_061.jp2
b8495bd2c66feb4a5cc270ce48394283
ba689c07f10e3f5f89e49036c0474f53b4de819e
962204 F20110217_AABYBQ smith_j_Page_035.jp2
b90cc29bef8aeaeffc3697a5562301af
c5b7ea2d22b9af228a449286a064142ed8200054
53869 F20110217_AABXWK smith_j_Page_154.pro
6cb06bfa6ba1484dac3a0406cce5799d
1d8a81c4beb86792c2b7c089df1405af235c54b8
30853 F20110217_AABXVV smith_j_Page_130.pro
973f7de7df266681db4e49a06136f5d9
e52e013d6fe8bc803f46b36d02d6c0eae2f4b914
1051946 F20110217_AABYCF smith_j_Page_064.jp2
3204ccddd6f1ab447505113ca3ca1783
70f50384750a6914dc5067be8ee89cfbaea42b6d
962222 F20110217_AABYBR smith_j_Page_036.jp2
f12aec57219d6c628bc1f960896c2cb8
1867d4eef896ffa076aca1b068d43f50580b257c
56327 F20110217_AABXWL smith_j_Page_156.pro
fff3ee346dd65d631711b712b2a3251d
b44505ff2904dc54868c54d91ab07542dbaba1de
13268 F20110217_AABXVW smith_j_Page_133.pro
cba1075f5b8010e77799ac7a6aa27d17
eabe0efc7beff34f48b97c438fdbcc7b4a9b54c1
399879 F20110217_AABYCG smith_j_Page_065.jp2
53269b4e8369288c7429ecb54ee5c837
757a03522f2fc6ce113012b503c7e30a1e8e1295
1000941 F20110217_AABYBS smith_j_Page_038.jp2
2929932484ad67a9aab919662801c0e4
60d174604f4ec519503aa24a36574ce93f1f6224
73133 F20110217_AABXXA smith_j_Page_011.jpg
1ca1afb9fc7edfb18309cb992349cb24
c7cdd077949ced698d60cd4aa68f772fc1d5e460
56270 F20110217_AABXWM smith_j_Page_158.pro
cedfec1112aa0739867e113cbe62f170
b806d3193faf57e2e182d88028ba0d9fe90670c1
41015 F20110217_AABXVX smith_j_Page_134.pro
2650ef911dfa207ac455fecffbdfea72
d8057db58b435764ae5ba6bfa62a7f141f9c4042
1028640 F20110217_AABYCH smith_j_Page_066.jp2
68d2643119d8468fbe8868b11cd32381
a46bb640a7ceb25c0fb6d190ba8fe54ad9993e9f
785555 F20110217_AABYBT smith_j_Page_040.jp2
d997bfcb27da91b23d5364a70ebd48a5
77ab592800eb04d4125681aa8db49d11307d3519
105058 F20110217_AABXXB smith_j_Page_015.jpg
65e1995b529d100d94cb9654540521e3
470dd2eadafa216e9f406f960724f083f54490b4
41555 F20110217_AABXVY smith_j_Page_135.pro
ff488d6a2e59ddfb665aa837f45e9425
4aba1a52d073dbe36b1a23efcda07ffcbb30af83
1051931 F20110217_AABYCI smith_j_Page_071.jp2
ef6438a27a85c35c8ff896cdfdf4ac7e
b61e9a559f5b501322949271223304d49c73a78d
993172 F20110217_AABYBU smith_j_Page_042.jp2
8cd6b8f360936663e8e0077bb8c9f355
fc6523e99ed488711526cefab4c0011b15c4858f
79707 F20110217_AABXXC smith_j_Page_018.jpg
4b93c8c9f2352c023d90bbe9cf5c6e45
9c4e2538c5ec33f229d57fdbf0f87f0235954ba2
57964 F20110217_AABXWN smith_j_Page_159.pro
bd9e3a829f7dbaed4b79be1b8c1a81c7
80ad8b0ea0934eccaf6e9245f6d3047bae8ecbad
28552 F20110217_AABXVZ smith_j_Page_136.pro
6ca4d361f4c21035d93ebe6fe4ed59e0
e9a58260d24f65edab5f6cc1d43ae05cd225bbe0
797885 F20110217_AABYCJ smith_j_Page_072.jp2
27f2c6e166de44b632444bfa45467057
c1c0272b0eade5c27658e7f253ffbb367e77ea07
958627 F20110217_AABYBV smith_j_Page_043.jp2
edfb56a38f509cd97e13e8a97fb13927
cf7ea6367b6a6136850358b59190904fdaceb24b
96607 F20110217_AABXXD smith_j_Page_019.jpg
abef1eba3f188a5f02e886e09bdc298c
0c19d4f31a363f86b5949805939530b3bb73513e
19735 F20110217_AABXWO smith_j_Page_160.pro
fadac7d754b14f2c738380028fa4a990
5a5144090ea53c89c521a48b4f404767263e62b8
1051960 F20110217_AABYCK smith_j_Page_073.jp2
2c2dff9ae08a70ef860f6f1c1fec27e9
ac692039d62ac99c9590c5066c6546aa902898c1
933519 F20110217_AABYBW smith_j_Page_044.jp2
2b21fdd5b2033a39cd390494de5dbb91
09c2cd185158aa1dc41f2128d8703e64fd83aefd
98715 F20110217_AABXXE smith_j_Page_020.jpg
fcbf4d8afcad2a07f40f002ba87db4ad
edfc279773cbbef3bf85d6c7d1942274aad3df82
21159 F20110217_AABXWP smith_j_Page_161.pro
0edc738833211fa4aa7212e73f24674c
ac3dc83ec7e0689b183b696e35dd67b82838f447
1051940 F20110217_AABYCL smith_j_Page_074.jp2
8ecde6869e813f1aa23596e982da1d65
70209effcf667ff263b3015fadcafe655b0ae1b3
81166 F20110217_AABXXF smith_j_Page_021.jpg
e70a339f541accf1b8b858ed0fad5c12
75be9331c57bc52ed30688695400b440eaa37a71
31325 F20110217_AABXWQ smith_j_Page_001.jpg
5f0f7e0c2581af417d5fd4c7d654a268
95db9e498b83776cf6ebefa688f8021c3e2c43af
1051933 F20110217_AABYDA smith_j_Page_096.jp2
7d6b65bdad28b7569bb923fd61a8919d
c78b73c3c1f832e091f0646f1f52c695b36e0755
944761 F20110217_AABYCM smith_j_Page_075.jp2
385ad18f318e198685e034695b7cafd1
1f6048f7752275c2c824c5a01559f66e5c356b63
1051985 F20110217_AABYBX smith_j_Page_045.jp2
f1706c551b474ddce394f214bad9e7ce
6d96e0998fdcceb997350e56a418f814a12b85af
89859 F20110217_AABXXG smith_j_Page_022.jpg
1703bb5a951c44533f7b8d56d7735acc
60122043e23bcebcc855bd4e807a21313afd96c8
4897 F20110217_AABXWR smith_j_Page_002.jpg
17a7a54d177a5cbca7af2d603d6eb48f
4e2df2739560435f141bab5dd19d1b8d99975524
576363 F20110217_AABYDB smith_j_Page_097.jp2
87c2338a88bbaace4c5bad5fdad3cb6f
bf13aed4f7f41597557a05528ac60e35513399c0
805687 F20110217_AABYCN smith_j_Page_076.jp2
f5207a0772b9e0b8179151bb39d3f454
3d957badf18f1f90c5fd00af5609b90feb02104d
737890 F20110217_AABYBY smith_j_Page_048.jp2
bbbbb7413cfe111ea9a12579d61bc06a
f41f06c44d0862cfadcb56b84c75827a8183af3b
76868 F20110217_AABXXH smith_j_Page_023.jpg
43ce8dacf6f1983940dcdaf29defe2e2
b888f0980fb1cc7f7061f06ff1a3f0f58ed90ae9
27975 F20110217_AABXWS smith_j_Page_003.jpg
64a3d63ed0efb0f3e5975e6b29fe65f1
06d9134c3704fdab5ae71aab70f1869bc3a4e787
870461 F20110217_AABYDC smith_j_Page_098.jp2
80f7508155f56a5963be57231db79c9e
76fc6988dbda8c384653ec4c9a2699ea55f7893f
514811 F20110217_AABYCO smith_j_Page_077.jp2
43e6dcf688396551a6fe9528735ffece
8fd786950afab56cca8e0fcdbe2a1bee46f05469
675114 F20110217_AABYBZ smith_j_Page_052.jp2
7db2887686d1e573718f750aeb606ac8
ff8a4aa1ab1b91157275a0bd3325b944763a6b3d
78702 F20110217_AABXXI smith_j_Page_024.jpg
07267c060ab37ffed1718cde5e2305f7
136a28f6599afb114b475239f200699967556214
48273 F20110217_AABXWT smith_j_Page_004.jpg
b16ff4222708038768f93e0593804d02
92122b3c9ddafb30b221401d7c5d7f80b3325600
22855 F20110217_AABXAA smith_j_Page_077.pro
a158964a763127be0ff3908109d7fd82
23eba0fb2f2b5d0d767df8124a8a82c52cf9270c
1017169 F20110217_AABYDD smith_j_Page_099.jp2
cdb7f70e1afe2b4e8d04f9416e32ce73
f5e078ffb0c78ec2246ebb204e4ac8e3d0d02db7
899159 F20110217_AABYCP smith_j_Page_079.jp2
fbcdc62dc4c0478c945e78e0001ba5bb
218ccf9525b9059096d462edae7040e33c3d088b
101840 F20110217_AABXXJ smith_j_Page_025.jpg
1c57b103f386bd3bfb65376af4de1dca
2e6008a9f57700f1ed723382a752d9c8aff02a93
89996 F20110217_AABXWU smith_j_Page_005.jpg
01be8e808f7b9ed9fd4579246560b9e4
2340bb4c57576bde4926b115b3316de3a69ea7b9
41401 F20110217_AABXAB smith_j_Page_120.pro
e771653ae1d7bab272c0e0727eebd5c6
45f6fc8bdf1837c64a1e8542b51d56fa7d5e7bc1
1048093 F20110217_AABYDE smith_j_Page_102.jp2
67e3b834e2a3ee49b5d4337dc1fb79ba
cdc3d8e329237a91db911d8217086a6fbe6213ef
961881 F20110217_AABYCQ smith_j_Page_080.jp2
d32476d099dff4d48227408d5fcc28ca
071bcf12401828491b79b3c895282efa6ee3e18b
106565 F20110217_AABXXK smith_j_Page_026.jpg
86d1968244c4b36441eff427f437a1e5
f789c3afc476a77a10b8f31c221b75a09d3a0401
109174 F20110217_AABXWV smith_j_Page_006.jpg
c99684d2c30ed0fb98d497e2aa35c38b
8c7fbbae59ce092ee3807d5ec43e3b45e230d338
269791 F20110217_AABXAC smith_j_Page_003.jp2
7ad06039cfb0ed8caf22942537bb91b7
565edc76fbab5c10db306ca650a00fb920114040
1051958 F20110217_AABYDF smith_j_Page_103.jp2
85590b786b286d1c33a6ebc07719fa0c
2c7f54d6ca864bb8acf84ccdd19baacd4725b692
564263 F20110217_AABYCR smith_j_Page_081.jp2
8a2df39de8b1fdb4bc5c023d26eb81ca
a201218fe74c7343435bb04217e4d1568db562a7
104585 F20110217_AABXXL smith_j_Page_027.jpg
e10e201f622d72a4fe9ca613de5ba0bb
d8618696ea331f16455ab4cddda10bf7abe4d023
48355 F20110217_AABXWW smith_j_Page_007.jpg
cff332e08ea4ca33a76dc271228cd6d2
40489a4aa74d1364345cb3773e68c8f060bbc5ea
21175 F20110217_AABXAD smith_j_Page_112.QC.jpg
bd1ce1fcdde41dcc258c07bf3fd91fca
91f5c5619ecd3e6a329a174b6cb2a487d7bd38d2
852092 F20110217_AABYDG smith_j_Page_104.jp2
14c6a242ebdff0249dad35af79038311
faeb4c8eda73f6fde45da7e139c238a0be8cce93
970895 F20110217_AABYCS smith_j_Page_083.jp2
48fa0d868fef60a88789279cc4a55a9f
f991d764479d0d237be3272db7b830a8099fe21f
101771 F20110217_AABXXM smith_j_Page_028.jpg
a55551e9ba0a187b0cef7e9fbb94eb7c
26d4b9992ca61c629eeab71c34e88c66edaa6d08
70073 F20110217_AABXWX smith_j_Page_008.jpg
fd1e32e09326356f7d0249dd587f1ac9
0fe59c033a72bc59e867c6d0a1f11fed75f81386
8212 F20110217_AABXAE smith_j_Page_025thm.jpg
5de563331f97d5cc63ebbddf9a61f137
539c7fe78da3747cd70336ab14f7fac79538c770
95904 F20110217_AABXYA smith_j_Page_044.jpg
8154c01d51ce4717d6f1733218b8757d
8462e285f0d34a459dbf672ac8dd8468e1394f4c
709256 F20110217_AABYDH smith_j_Page_105.jp2
6603d60393ad5e826dac2712467989ca
791d9ebee51c48ab7b5bf44fee444c490319d7ee
960480 F20110217_AABYCT smith_j_Page_084.jp2
084b19e366a7a29998a28da99ff14a6c
0417bb3abf9f50d3ed7d894565acf780db229baf
95661 F20110217_AABXXN smith_j_Page_029.jpg
11401348078b9123bfb5313f1d0beba6
1ae12a2bf05b8dcdaab03f35968c78123a3677a6
92430 F20110217_AABXWY smith_j_Page_009.jpg
71005d9d3bd352eebaff68c9b505402d
06cf5493523c05bfa0d4f8a9ef60e636a8cb3529
25948 F20110217_AABXAF smith_j_Page_062.QC.jpg
07e0cde4cd73b1c90e3c3118950f3dc4
988b08ca175f14d5ff682dd1eb12a17b68219f11
104255 F20110217_AABXYB smith_j_Page_045.jpg
4740b737daf96e7aea927165a6d94ebd
4d308420f4e4febba38e3d8ec61bbb3329e46aea
410448 F20110217_AABYDI smith_j_Page_106.jp2
f1bcb6d140a451de801bedf3d6baf42d
d34576b69b61dd4c27dba65979ea09d8530eb614
768302 F20110217_AABYCU smith_j_Page_085.jp2
ac4fef658f7bae4c9aef1b1d18679ed0
8d12cd7d8adc338f661fc8849dcff6ccb165691d
48521 F20110217_AABXWZ smith_j_Page_010.jpg
edcb4356c9bd6d85d693f18e6a30ade4
ed52b5850eff1f86229d3b386b433070223b4695
81728 F20110217_AABXAG smith_j_Page_016.jpg
bfde33db2ef1645f1733fb11ce42ffb5
10d900be3565832e285d088e5f372a3b201fc462
34560 F20110217_AABWVA smith_j_Page_059.QC.jpg
b6d8035edc3f4d0242711ee8c6e718df
51754630acfb7d1e33c2ba7ff89c1e8de90f6d7f
67659 F20110217_AABXYC smith_j_Page_047.jpg
2336fb092a3f88bfc627762ee05b4056
394ca5a86d8bcb33364dbb14c9092ffc65c8b0b3
540296 F20110217_AABYDJ smith_j_Page_107.jp2
919c3a2a85fe203661f54487a9bf9980
bdd0ed701528e27ccfbcec15db74e105e62a58f5
618632 F20110217_AABYCV smith_j_Page_087.jp2
ef1991c7145be4bff485112ba94bb091
f6685485fb11eb0f21a8e8f2f4be3af78f0a9293
1095 F20110217_AABWUM smith_j_Page_109.txt
149cc9249da32ae9aa8e10c4d0d1880d
3dc63c9c7a246f06550ea3055e99d3ac98a6159f
95789 F20110217_AABXXO smith_j_Page_030.jpg
dbe2732ec239ddbf7b68b419d4824143
2c185c32c907ea77cfb8b38c115654f5c1069e5d
4528 F20110217_AABXAH smith_j_Page_153thm.jpg
bae6ae20db6ee47733c3ff57a9b79a6e
e737d1f30be18881675678cfad09bb061de16729
21813 F20110217_AABWVB smith_j_Page_129.QC.jpg
8596868cee0bc18dada9bf12de5e5f1a
d0531fd114ab476197063845a90de1049cf9ba26
72651 F20110217_AABXYD smith_j_Page_048.jpg
603fa7346cabcb515a0ae073c34dc937
12408bc9a0282a322a975571812e25f3d32d80ec
425548 F20110217_AABYDK smith_j_Page_108.jp2
da82116dda6a39fc33b4952398547997
c9cc4a016ac28befe6fef9963ec2b1b6ab863809
820322 F20110217_AABYCW smith_j_Page_088.jp2
95622c8ec07076ef3830a2e528a73555
b898b631f6307255adc8d9c658b2a89ceacc314f
F20110217_AABWUN smith_j_Page_014.tif
a7dbbd253050d70bc80f0809773da15d
5a1e9da497d5333a1a644de5a1611af2a2be8dd8
90656 F20110217_AABXXP smith_j_Page_032.jpg
dbae0443458eff2084b65b59c38b0ad0
5cee03686021796d52382cb57de28cb00c79d0cd
55979 F20110217_AABXAI smith_j_Page_130.jpg
66e9d341afe0eadf6dd59d87db5672cf
40451fafba3a8637a033ce870455f4dbf48d4f68
F20110217_AABWVC smith_j_Page_032.tif
d862aace0709ecdef4fdd58d86df1b3e
09872b078453c8f282193eda2df0a1678e82f9d8
53694 F20110217_AABXYE smith_j_Page_049.jpg
4f89f9b2d08110b18e1fe027ef27759a
be3ed27e2a0043573e6485083e52f944bed9b471
718607 F20110217_AABYDL smith_j_Page_109.jp2
20dee7c4012b37d13bfa14160bb80fff
5a5ebfdf1b47f42acc5500e9e57e2d0b6c577803
990955 F20110217_AABYCX smith_j_Page_090.jp2
7ca7687562558db55107260b9da735b3
170e55cc75bf946a06e3f65cee5eeb557b7c0990
26212 F20110217_AABWUO smith_j_Page_128.pro
1c5a36fd0511fd321ccb02da1197fb4a
d634c4aae306a27bfd88c02715f1c327bc883774
106063 F20110217_AABXXQ smith_j_Page_033.jpg
80313dce243649cdd1758af6ae450042
6ffd8482b1350ec80a779fd9d936bd61c4805641
2204 F20110217_AABXAJ smith_j_Page_124.txt
bd8d21f35d70ab28e08be3bcdceec383
b807ef8e2f1099eeb66912e04a013d5382d666f6
8007 F20110217_AABWVD smith_j_Page_019thm.jpg
5859744aef2cd27a65ed7c14ad408b5d
20f5f76610fb3fe974a9c518c057c5904c56b931
89799 F20110217_AABXYF smith_j_Page_050.jpg
b969053b3c36114660abe4a16aea523a
0c373c4ffc94d60e4cf411d2b187a1bd3cf9d665
473418 F20110217_AABYEA smith_j_Page_139.jp2
f8cdb27b164441886c020796bb933031
f6f00435ae15d0ff6ec4393db54c7f37ec12518c
582425 F20110217_AABYDM smith_j_Page_110.jp2
e3979a49d21651b61823e410cd898e15
d70d47328ac31e37a6fed4a32e87f627df83b7c9
5432 F20110217_AABWUP smith_j_Page_134thm.jpg
aa21f88eb7118a9ac3be3f4de415a1b3
77771c250d57f388c2e23cc6db5dab2dc0e7397d
103155 F20110217_AABXXR smith_j_Page_034.jpg
1dbc48939160c3507f74b6a83aa18dd7
340089b074556233ab9e568fae0f4a3c67d1b385
84813 F20110217_AABXAK smith_j_Page_079.jpg
2a3b658750066857aa3bd20d53c5957a
391ca58a8b4c8b622420f0fd75b3e030434317d1
1929 F20110217_AABWVE smith_j_Page_096.txt
efc4e9ce14258ee9d3c2258b8695bce3
32c448e881e353a3ed342697d8fa976f19985640
72179 F20110217_AABXYG smith_j_Page_052.jpg
2f3db8d1b246b7e89dcc399706b15904
aa704b680eff927846e7da42f197c6bd71c759d5
648897 F20110217_AABYEB smith_j_Page_142.jp2
aefb423becc01b2f3959a509aa2622df
b0e33da13b9ab7454b5c783a57bc97120ffae620
650864 F20110217_AABYDN smith_j_Page_120.jp2
e37b640fc41ebd697bd02cb16266aa0b
1d903c866ef7fa5bab8cc9810e19eda6c5551a5e
1051980 F20110217_AABYCY smith_j_Page_093.jp2
41bfc4abaffbd8cd9a79fd29d1bdb2c9
6962188d34de6c0cba6b1495acdff92c58d84f27
F20110217_AABWUQ smith_j_Page_013.tif
da33be6b5e28f48687f17c9ae7139588
48ac6fff756ab77657cfc3a2a383a674bbc775e0
89136 F20110217_AABXXS smith_j_Page_035.jpg
8a02fd7f0ab8421f021a6e083a713f3b
29cfd792c02b259a4115d38c52664989274e6efc
430581 F20110217_AABXAL smith_j_Page_143.jp2
bd71cf5fbb730deddb8dedafff4f858b
d07388cb37b7d3c2a8632715152d1849cdcca2cf
999719 F20110217_AABWVF smith_j_Page_068.jp2
fabce68e9b5a6575da4e2f9549a40d53
836442c200d30d04f2e630d9764fdeeb553c96e4
74078 F20110217_AABXYH smith_j_Page_054.jpg
aa583b5526369555d2996ed8e0298b22
c5a82765823daa8c21c8fd587043a87886da467e
1051969 F20110217_AABYEC smith_j_Page_145.jp2
3375f986b8074a6cf75a8f6eb03981fb
fafe865c483e5c0cf03c43e225b521ff2bb78cb5
816046 F20110217_AABYDO smith_j_Page_121.jp2
8132d1df7d11154a23ae4d4c246a82da
d3be4f1745ec13628226d0a54ef641d3f04443eb
1042400 F20110217_AABYCZ smith_j_Page_095.jp2
ccc40ca785143540b160b11f3fedb01f
ba661daca40837055c500d58e620e97408f707d8
7498 F20110217_AABWUR smith_j_Page_032thm.jpg
db5c84d5010cecf43ddb31389fcfb601
bec738aa3e1b0a93ee332898e3fa7952a0978787
96959 F20110217_AABXXT smith_j_Page_037.jpg
efc9c023e5e3a89649170e758b5da1be
38b603b0e7c9e516ac7ed3837459e8b31bbb9661
6400 F20110217_AABXBA smith_j_Page_009thm.jpg
aac6c897cdeac4d2a7447d9c98a46e0f
42a62a9fdf019812f8c648f89d5f8a2997b291df
917309 F20110217_AABXAM smith_j_Page_005.jp2
17fe96757573f46e072b3b87c5c41693
562b03d6acff8adfba3ec1a465a60512e4e6427e
F20110217_AABWVG smith_j_Page_108.tif
167124d953909b7f85d75558f1284450
0665dc7e8445094e57d3ecfda740928a1e5b56ac
83701 F20110217_AABXYI smith_j_Page_055.jpg
34b4c1f823f65fb165911732c60f791b
869aaf0f06985c94cbca7d4a8022b141bd522a33
1051949 F20110217_AABYED smith_j_Page_146.jp2
29006be697402cfd2eb93e7cc910941b
ef2f49e6fa5a30215ef0e5acb9d5cfba232d8f8d
392292 F20110217_AABYDP smith_j_Page_122.jp2
283c48a3a1c7dbbb613e08db84623e54
b01f82d612dbe0fd4c43f0a08a0ff067205fd8f7
45467 F20110217_AABWUS smith_j_Page_099.pro
4db3e4183e422df22a5d3c48b5a5d460
5f37f011fa1e017118d30ab6b30e6a0ce6965bf4
91410 F20110217_AABXXU smith_j_Page_038.jpg
d27732fad442800bec15e86dbb97bd98
e0305bcd2fff3748db110f1af541bb3f862395b4
41471 F20110217_AABXBB smith_j_Page_075.pro
22db73c7dbe227e89c623ba5b30adef0
3ab575dbf033e94438957985347f10a10ed03fbf
1856 F20110217_AABXAN smith_j_Page_018.txt
19636aa26530c60ab2858e6b50b39f99
298f8a70ad04930ffc9e13d1fee1dd2bd0299fa9
36775 F20110217_AABWVH smith_j_Page_116.pro
31a3ae47b63b94330c6c292f4792e5d9
fd4c451874a9ba41a2fbaf11a1043bb70d84b7f4
74356 F20110217_AABXYJ smith_j_Page_057.jpg
258d5af925d4f2b94a61c0222d5048ba
be0db3a86ae6b92ee193583911b0fcfb918f31a9
1051930 F20110217_AABYEE smith_j_Page_148.jp2
e451c89caec7d614dd9925a6190fb6c0
d0008d5143e4f970b6885c60595adcffbbe420d3
202008 F20110217_AABYDQ smith_j_Page_123.jp2
a207e40e570ba21283c14fce9fb494d3
90a3e2993b09f126e54d6cdf69db0b2e5668333e
43432 F20110217_AABWUT smith_j_Page_042.pro
11792aebe79302e1ad51adac1a17682b
11daafdf7c0291fe14b111e555d0f38345105672
82459 F20110217_AABXXV smith_j_Page_039.jpg
33d716770a9381da44355a1ee0778ffc
ff0de8f23e9f235092047afae793939dd7a6972c
852520 F20110217_AABXBC smith_j_Page_086.jp2
49e3b7385e53a9195bf4a8c20f082385
8fcb8c838e4ee167ed2d0fe8070d2b497447922b
1051952 F20110217_AABXAO smith_j_Page_006.jp2
e0f869d1dbd656dcd9c79ec3c23a0680
4edf45108420bcea42c66dc53a6799f5badf3982
30124 F20110217_AABWVI smith_j_Page_058.QC.jpg
e6d90f0f6d3006f0d41239f68ec7d940
9f554a168cffee8717c686b913e40b756a43811b
90351 F20110217_AABXYK smith_j_Page_058.jpg
80c161bd0dd4bdf24b5639868901cb3d
c91aab9ffab0d86dc29a8f0d389d8d4876b31a9e
F20110217_AABYEF smith_j_Page_150.jp2
4e2ac4a096b1e199da1c28c2d20067c3
a448f4799adfee156aa4c7d310de3098f8b5a0b1
518087 F20110217_AABYDR smith_j_Page_124.jp2
03bd3cd2a088348da5a2b68b10b46623
a00847a364331bc0717578fe2b45e6eb7568cba3
39955 F20110217_AABWUU smith_j_Page_053.pro
92b5b1b51326264acb7f916dabced486
69f475c8c9f4f0fd1d7895916696bbe001458a9a
72949 F20110217_AABXXW smith_j_Page_040.jpg
f49c52360d98a6c3b49d930458d5ee47
9c3d6197dd79232a31513d633d3128657e7daa94
1051950 F20110217_AABXBD smith_j_Page_041.jp2
7d251aaae06ccbcf1eda6a2402353996
d72a7c4a2afb8b28b7a3d3816d308f93b6fb5797
6757 F20110217_AABXAP smith_j_Page_016thm.jpg
bac7950b81a33e57efd7b4a74f5610c3
389b20c42c2f48ec065aadf8b00831845dec3e18
52983 F20110217_AABWVJ smith_j_Page_067.pro
14b8510521ed55e84acae18acc36965d
2f2afbee702a34b31650b7f9a97fb3a40fc2d04c
97558 F20110217_AABXYL smith_j_Page_061.jpg
4e665fa0610cc2a1c41d75ec79d696b7
af0c28efb994d4c98086af3cdaef7a1ae3e38bab
1051968 F20110217_AABYEG smith_j_Page_151.jp2
e8cbad4363b13d6b78e54671bf22b62b
9309916cef139f09143b89e2108d32b00fea17bb
514591 F20110217_AABYDS smith_j_Page_125.jp2
6ebfab9017b45a20c1b026c25020c032
4a8c407a6c841d98c90a8bd28ef38ef02dd1712f
87308 F20110217_AABXZA smith_j_Page_080.jpg
110d7899541371b501b3cef3bd51e3b5
ff77d37f057534c213ea79f31113a79cbb304b42
977485 F20110217_AABWUV smith_j_Page_022.jp2
9ac8bf06aa2702c73354abc4e544a3f4
5da1afd81383d053adf83bfb74d58b9cdc01e9bb
102270 F20110217_AABXXX smith_j_Page_041.jpg
fc0c0dccba85a2cf346da78c90ee8c82
43fca7499fca9b1ace29fd3ad845b81e0b3389d1
31338 F20110217_AABXBE smith_j_Page_046.QC.jpg
abe661201689a902258565bce103d29b
63240f45be9a30a3d3f6ac41065283f464cea9ca
F20110217_AABXAQ smith_j_Page_109.tif
282af07be13afee5b4b0f53ad71aadad
f342114f0df1cf68520aaab0c957c0242a789ae0
94668 F20110217_AABWVK smith_j_Page_017.jpg
f0dc209c8679685e16dc1f8924401e20
dbc047fe6e126a183bc73106fce7131c4470e200
84173 F20110217_AABXYM smith_j_Page_062.jpg
164ef5e22e5166b3be31323c7371546b
2f46b0cb502f909bfbad92cb1e183c0a136121f1
711957 F20110217_AABYEH smith_j_Page_152.jp2
e027342334f4480519c55b046122cf32
1b991f5445ced3c10cb45cbdd5c32e7cb2224d9b
518747 F20110217_AABYDT smith_j_Page_126.jp2
54a434294701d1c39b7604beea18e6ca
7d58812ce5be26dd45e9cd0c74e2f7a2c49ccc5d
52682 F20110217_AABXZB smith_j_Page_081.jpg
0f0b8f2e7ebe1255c78eaa580e88fdd8
28e5b954aecbb2808d8ddb340d069f18b110fd55
F20110217_AABWUW smith_j_Page_132.tif
bee411807a925272f818ba6fa9be2599
a6da0651e29bcbdeacba3acf967cf5b53b9e950e
90339 F20110217_AABXXY smith_j_Page_042.jpg
c28d88d3eedb2e0442991ea465fe1e5a
fb8a3d5c7409241e1f743f389919fb7947e55c55
7331 F20110217_AABXBF smith_j_Page_080thm.jpg
e11278b8c58e42288fec4afe7bfc0311
77788e5c98f24c00d4a3a681e334af9a660027c8
7619 F20110217_AABXAR smith_j_Page_099thm.jpg
a8ae1db08c31c1db24a93855283c09f6
963a8d3c501d4f2db95d5759ebe8d7afc61e209d
F20110217_AABWVL smith_j_Page_101.tif
2b5fba8a9b69a6a9d03af80af2047b63
04ee2b54c7088cc546c5f2a6955ad09ab4c97b60
100799 F20110217_AABXYN smith_j_Page_063.jpg
4eb8ef1f6ddeab1953bc6c166b39b8c2
8c839efe9e83d0f3b3a54dfeef16c7615ba5663c
1051986 F20110217_AABYEI smith_j_Page_154.jp2
96a814f0aea2efc114a79e73ecee4ef6
ab4e9a12343408bbf19d4b8c0a9c657c1887aca9
630114 F20110217_AABYDU smith_j_Page_129.jp2
7d629fe54bf044f68c32d452a1b7fd9b
9bccee9d8f690729e1192629a320253f8a832688
89012 F20110217_AABXZC smith_j_Page_084.jpg
e956bb780cd9467395242d105dba4a6d
6512602e438505a4399a85cd5b3b8ecfa6d921f7
F20110217_AABWUX smith_j_Page_137.tif
aac23cbb1cd2889a0f940b8b3afb0bba
99b6002391c7103b761bda36672a3d2cc8187063
87707 F20110217_AABXXZ smith_j_Page_043.jpg
4c8253e64bc2f16d8c6f0cceff06895a
3f9044398fa85503fc7e72b320857f90f60c333b
1638 F20110217_AABXBG smith_j_Page_016.txt
f852a8b095135fcd91a5a7665572411c
c7dca5f948ed3f39eff0ca9bd5d2090c99ef3eae
38044 F20110217_AABWWA smith_j_Page_151.QC.jpg
4084a41d681d3d03c2f79c0ffd3db707
b1aa2f576e9804934db80391da81715e4f335e9e
74501 F20110217_AABXAS smith_j_Page_072.jpg
af815f08be6a2a7657bb16b35ee9cacd
4adec6c4364fb48dfde1ee6e2f3da031cc48ee15
7993 F20110217_AABWVM smith_j_Page_044thm.jpg
30466aec155478a3a12a8bb5ca794f0c
9c9625a768076392c38056030f037bff31db1843
98872 F20110217_AABXYO smith_j_Page_064.jpg
05d9745a8e57b8b3f91f84d2614e4077
079f151698970a7fc8b5b991ddcb464b577f9642
F20110217_AABYEJ smith_j_Page_156.jp2
653409755ae3e99958b0c22d3215f0dd
1634a6ba33c148005eaea84fd8907928a7944c47
486287 F20110217_AABYDV smith_j_Page_130.jp2
5565c5850d220aab453caf31c746e9e1
f75603385bb3edddf54e1b9de38636355cd878a6
78235 F20110217_AABXZD smith_j_Page_085.jpg
c87bbd71dfe6f6a35858d34f376967bf
1b5e6f2e2e835eacf4f8fbf193722674cd4c2cbc
27129 F20110217_AABWUY smith_j_Page_078.QC.jpg
744ce87eadaff45685707ed54376946f
971c3eeca12ea44b2d0167d28b869afe2df28133
11964 F20110217_AABXBH smith_j_Page_007.QC.jpg
6c9c4544119f418afac157c9c1047422
7b18cbd63d8c5eb72133bb6ddcc42b18f90afafc
9222 F20110217_AABWWB smith_j_Page_001.QC.jpg
d3d8bc855a056bdeb12148c663d9253e
b93081f021ab17c634161befd9cb08d7a00e1d45
38316 F20110217_AABXAT smith_j_Page_074.pro
b0a62afb26e41c77b68ecbfc002e2b1c
0f9d3dc9f89eddac819900359a73c93b3af8e308
F20110217_AABYEK smith_j_Page_158.jp2
9763bcc819ae0a179c0cad2e8ef2b5bb
062e76c2fa529c8aa11ca7ba58d8e8b8af10655e
223466 F20110217_AABYDW smith_j_Page_133.jp2
16af02e8d8ef8deaf3fb385b4f1de46e
d48bd71f85e7d5f8da5b44a23fcdd218f6edd69c
84568 F20110217_AABXZE smith_j_Page_086.jpg
5b94569462bdf4fbceba3f9567c6bdaa
06230e2d15a5e7810b472e64abf991ffc981a08a
1544 F20110217_AABWUZ smith_j_Page_104.txt
969e679fd2862b82e5156648d7544236
7f5804b134c36a048621d187cfa9f0d6586bf7e9
2086 F20110217_AABXBI smith_j_Page_067.txt
afabfeb7b21b1f03c1a977b4e57fc06b
6d5982c97146f9ed6997ddb4d5770421d650d55a
321 F20110217_AABWWC smith_j_Page_023.txt
f21b5606c2bad2dbb5128e69ccdbd75f
5572639cbe74d82e4c044b0f868ab0587651cfc1
3098 F20110217_AABXAU smith_j_Page_144.txt
ff9750981b2ef505b2cf945034f98227
f605c9e21cfd5ae014adfee6a3816c780b864bde
F20110217_AABWVN smith_j_Page_089.tif
e51fb50855856098260af3cd0cddb395
2bf2c5309ce39d169c4371f45375b3214f811d90
38413 F20110217_AABXYP smith_j_Page_065.jpg
a00995c22ba742adccf7bf6012b8e7ed
0dbf5209524fd520ca2bb495707098a700b9c199
F20110217_AABYEL smith_j_Page_159.jp2
ef2645a502f7dda1e2befec3925c1ab4
2310e99ceb87584d56bb81cb019252a94a2ab369
656686 F20110217_AABYDX smith_j_Page_134.jp2
36e6941732a1e69514c16b1c33984894
f06396063b0eadb56cefacfe905932f0e052e8fa
60103 F20110217_AABXZF smith_j_Page_087.jpg
2cc0ac8305e2cb15cf2e252ee6f0d59a
cf5825fdb2c7d860f1fab897859473b021c6fc36
41941 F20110217_AABXBJ smith_j_Page_032.pro
19f8efbc1522896a8f6eb97f4dec5f75
59cd030bee847f687438e605c3d7b358a376e3c9
71701 F20110217_AABWWD smith_j_Page_148.pro
f588ba3769d81ebb31bf8f650093f091
091be7300a7f14e31221a5ceb594244e0d012003
F20110217_AABXAV smith_j_Page_003thm.jpg
2ac5a31618a0b134e44dd61bc061d547
0c4acf6888c9ca7916730ca10dcd0f4003d4dab3
51924 F20110217_AABWVO smith_j_Page_128.jpg
8b5b3fa49ae3d8229a72a09677333c2b
916157d78732ae78f6970378c402327aab564280
91369 F20110217_AABXYQ smith_j_Page_068.jpg
a1c6b2488bd6891ca59b1d1db2091c08
45ead56e5c15593b24bf4b5f08793d392b012241
14263 F20110217_AABYFA smith_j_Page_106.QC.jpg
3cdada1d65887a16237dc2582dc866bb
b82a3476e25fe51060e8cd48a7eb63b0c2dba170
464397 F20110217_AABYEM smith_j_Page_160.jp2
7a252854b8799b9fed23adc2d8aaf756
b83fb4f3a66cbef0b15939f80aca9e62243f196f
656281 F20110217_AABYDY smith_j_Page_135.jp2
6f94204bbe7c2e9888ba3b05263d8cb7
c313a4f297eac51cd4971f61f0fc66f63a2ea1e0
79006 F20110217_AABXZG smith_j_Page_088.jpg
d5f6aa71ec38a82523b68e9072c95e59
11325dd1d83e4927cfcd7ad64d4e68646a2e3eb8
F20110217_AABXBK smith_j_Page_155.jp2
eceb44407d0ba29f2220a7e719fc0c74
f49f40ed5055bf9b4eeba48d9ac3381ff963a59b
6422 F20110217_AABWWE smith_j_Page_024thm.jpg
14d451c62d8c1d90224196734d76e5e0
40933b15545315d8289d764d293a4a0cacab64b1
2045 F20110217_AABXAW smith_j_Page_136.txt
d044d8c3e57fe57b079e6d1fb662d0bf
e62bae1927340d9abf6befa36b9b59ac11c9ce87
50751 F20110217_AABWVP smith_j_Page_027.pro
5112a387eb35207d1d8caf90bf1e7455
02cb479c48789e3a8fdfe339b664d671e2b486e5
87295 F20110217_AABXYR smith_j_Page_069.jpg
2593efcd24bb68d4661e2e302788e05f
4946c8cb801f45f2b08ee686eb1ded4b813da181
17771 F20110217_AABYFB smith_j_Page_131.QC.jpg
555b0e41d2a03f56011ad34eb9db878e
487bc34a7834aff6b9fc8e0479f7947fd7e6f086
7939 F20110217_AABYEN smith_j_Page_055thm.jpg
70e71424315c6a80621771f8c6c6def4
2a59f5d5a0b129402cf5228b5cfb940416f992fa
82164 F20110217_AABXZH smith_j_Page_089.jpg
b16434e051dc5ffd399759f1f2d8db14
34585f4b5a339e4472b816cd7c280332a0ae2f18
107731 F20110217_AABXYS smith_j_Page_070.jpg
a8e54f2c605e8a0c4775b49de2d98c67
c6bb24114eb1d8c815916477e5ca7de053d6aaff
29031 F20110217_AABXBL smith_j_Page_032.QC.jpg
a79eb990a5391804ce6bd1b8cf9e431c
36555dc37b9db84ea2ab24682b1a78969b105d6c
696 F20110217_AABWWF smith_j_Page_143.txt
12c01cb75be2ae487e755d7cbafa3561
9e4a8aab51bb9b9e573dbfe26a0c7947ffbe6492
8039 F20110217_AABWVQ smith_j_Page_046thm.jpg
72b9a34a6ca5493ac1e39ce3d4703030
e84ae9440c53fccf5b5d4ad57f9cb23e34145db1
8394 F20110217_AABYFC smith_j_Page_034thm.jpg
366f41ad418192415430c3d4afb30299
fdb8b0dd4d71f97c872563edbddd62ca4c3e9db3
8006 F20110217_AABYEO smith_j_Page_100thm.jpg
dc26fbaedacb1f4aae87b69010a01282
79e3a54a0318f7b94cf636aa71ee9c96b5b81af4
471571 F20110217_AABYDZ smith_j_Page_136.jp2
c78e30dd317cac1fa17931d4a49dce4e
a36438006e3e329a107534ad7dcb4013e21a73d0
98662 F20110217_AABXZI smith_j_Page_091.jpg
cab5fa4316ddd1991585574ce0474d16
0716b57e50e0b65f741d01aa83d0c25cec4c06f9
96565 F20110217_AABXYT smith_j_Page_071.jpg
d6d8ad7e50ce498bcfb741fbfe757b7d
55e4b2e2e95c8fb2e8a5a086138471adc10398bb
8248 F20110217_AABXBM smith_j_Page_063thm.jpg
8c48bff0f6894185bd9da1dc2e42e2a0
a73da1a008ef5e770c16e3892bc3bc4babb2797a
86974 F20110217_AABWWG smith_j_Page_121.jpg
bc2318c25341fccdd8c55709e8648285
94eb835e26e9dce1645505ac3a29a02442b50caa
481744 F20110217_AABXAX smith_j_Page_111.jp2
389250e0a98e940f32b38ff5d0088592
d81acca4fe53f534bf1d4f13e7e647fec3d646ab
2630 F20110217_AABWVR smith_j_Page_012.txt
d7fe1f510442c84cd62dcdfe9448bc22
a2e8aef078e6aad5b28ec68934e2442eb71454a8
461007 F20110217_AABXCA smith_j_Page_128.jp2
35b4c919c9129e7d4304ec5b9df9ebb5
fca39d04f4082ee91847d8c09baf103652b9069f
6675 F20110217_AABYFD smith_j_Page_021thm.jpg
400d4688262c8bde7e7cd30d75684a95
37f4490bbe8f228bf6b9e89b80c2a28eca8409fa
2640 F20110217_AABYEP smith_j_Page_133thm.jpg
3bacc127458b3c277afd92ae856b92de
25c8faef311606434b55d2f04b24e0fb4384d2b8
106474 F20110217_AABXZJ smith_j_Page_094.jpg
1168558c886d6fe544507a7313972d06
5ab525f19ef441856714c5e1021ff1333e99e615
95679 F20110217_AABXYU smith_j_Page_073.jpg
6143cf53b8832b50520ae53e8d696c25
6154aec53eb1802d12254496ccf8cabb6d42a87d
119182 F20110217_AABXBN smith_j_Page_149.jpg
de510641c0edcea5293a14863e8cf99a
cba6e58a155c644111d5fef15a7ba296344a3219
8922 F20110217_AABWWH smith_j_Page_013.QC.jpg
e0a02d888cbf326f88af94740359deff
5568e7b1ebd0ab7c97f1d9332926716be39668d4
100289 F20110217_AABXAY smith_j_Page_100.jpg
6886cb6e2db28e05ea860f7d9b1dfa71
8924fdc280bb95c677fc023acc0dfb6744a747d8
8344 F20110217_AABWVS smith_j_Page_045thm.jpg
e1bbe35f1de8ed50eb57209adf4b8c84
80765763174c1670d3f7b7a1226613168722a035
F20110217_AABXCB smith_j_Page_038.tif
88c979596bdd822985d9f29dc706f8c0
ca752f4d71d5850189cbc5cdcf00e04c1610a5ca
5431 F20110217_AABYFE smith_j_Page_127thm.jpg
e22b42eb49c4cbbb2bf222ae4952c207
596604f842fdd681569281040be424cf16803470
21054 F20110217_AABYEQ smith_j_Page_143.QC.jpg
79d03ddb9e3d1c060e02e32cd28cf7e3
b6ea5a6d13d235a7b1a93bd87ca66747d90adfdc
94728 F20110217_AABXZK smith_j_Page_095.jpg
c07f74e6b28e4893bc0d50651e533cb0
d16fdd034d4e1fa01ff8e92150cf3c7f4c09c231
102608 F20110217_AABXYV smith_j_Page_074.jpg
352c39673c69a1c85f2ad2a7d550ff42
765401a258d3cb764c9cf39f31464794a9e91ff1
22407 F20110217_AABXBO smith_j_Page_013.pro
541b636715790294e7343629bca2ebf8
d8801b3513d8f728d3033fba81f9d83f37902850
6116 F20110217_AABWWI smith_j_Page_105thm.jpg
82a8abe6fab9d7a5153feae0e809312d
79e7408f18585682a1b619fb1acd881f9590d62d
4614 F20110217_AABXAZ smith_j_Page_132thm.jpg
a62a79ec923e0b889d7379afdebb316e
a6bf83062c610b0da21908f671322d462f4b86f8
2001 F20110217_AABWVT smith_j_Page_015.txt
3e1759eaa1a4c446e23fe414ca1ba7ee
14cd55197035aad432e489cb1035783f1428ea49
37831 F20110217_AABXCC smith_j_Page_129.pro
3134c054c13be5a75a9cec594de37d0d
8ff04a6bbb1ee544dd733e3e7d16e6c9b0f6b09a
7137 F20110217_AABYFF smith_j_Page_035thm.jpg
c60055b253d14d2913dc3b3c7f77fdda
4d1499650b1358755768251d7678e1be24ab3c2e
4162 F20110217_AABYER smith_j_Page_004thm.jpg
1bc81100d16858ef33e1f9b2e17fe418
026777e8ccfe12fdd6f607ed912ef56656cdd4ef
97117 F20110217_AABXZL smith_j_Page_096.jpg
536af07f0f414799ddbde1ed2be32b72
578aeb0fd9a9bac8507ee43f8a6bd506ab0ab749
86242 F20110217_AABXYW smith_j_Page_075.jpg
c684bc730fd53ffce82904656cb53481
7e23d6e6a96ac10b3074a5c0d6f2c47cc3ca3247
1169 F20110217_AABXBP smith_j_Page_082.txt
309fb18f1b2dbc8dffab528e705110cb
c8e5caef3d0215b94744358c7ff8aa6b259319ac
F20110217_AABWWJ smith_j_Page_142.tif
06d05023a3abc6424ec7f85b3b2623eb
7c0e1a2792eefeb02a870eedc92cec2884e6f3e7
F20110217_AABWVU smith_j_Page_077.tif
ee35ec1f8fa7befb65143174ed291b4e
ff4275837a2922fe0bc9c4b912cae8e613126259
34611 F20110217_AABXCD smith_j_Page_106.pro
06b8012fb25e36b5135cc4c984bffa64
4e8f023488a6fd436a707b0ff718c09140ca8d27
12218 F20110217_AABYFG smith_j_Page_160.QC.jpg
25ff20cb6daa3b38f803db1fe915dbdc
96ea5c8c24322d83c18ffb88f0104a731a583ea7
34762 F20110217_AABYES smith_j_Page_033.QC.jpg
7dcc1931fe525bce23d6c5e8833322e5
19865f07ce3e2700bf939d4bc6046517a6493fdf
80004 F20110217_AABXZM smith_j_Page_098.jpg
69dfc90d104f451680eb3dd0cffa1b14
ae23cf91b6d3aee43fc110b94a24090b5ec17ab2
75246 F20110217_AABXYX smith_j_Page_076.jpg
1fc30ad5def9feb29032c34c83c4d7dc
b1e42c597d3660e042c197d3cdede3b193fab8ff
592 F20110217_AABXBQ smith_j_Page_003.txt
2f03e05672ecc6c678fbf4b0b0c43332
d6aa46ac00bb6d27800866bd65a113dd0a2960cb
1038 F20110217_AABWWK smith_j_Page_087.txt
3f4b3a08326205364b0ac6620fe953ba
24c4ff94bb917646ba8904887c5358af3abad94b
632588 F20110217_AABWVV smith_j_Page_153.jp2
4a0e00cfb418e663fe39893fd2d94e69
718753366f6c6160e32028c045a99e643cf1a0c7
37821 F20110217_AABXCE smith_j_Page_062.pro
96d1beccb25ce3aa1affa89be1f09614
7fe689bece2e98d87aa18e8165aa112318869464
17298 F20110217_AABYFH smith_j_Page_153.QC.jpg
b5420a979c107be81a68d30a63ac6533
65201fc581db36d084f3c2f14742f74e6be8bf4b
7554 F20110217_AABYET smith_j_Page_056thm.jpg
4a4a2cab70823897faab8821d4e4b06f
e9290b90eedc175988804d0d92ce49cd0c8a59b2
93073 F20110217_AABXZN smith_j_Page_099.jpg
456163ede69c6e7fff43c353fe3bd52a
49bf459300bf39579fb65f515010d7d1f2ca79d2
55099 F20110217_AABXYY smith_j_Page_077.jpg
390a8a4548cc7212821a4c6ebdc4a175
eb5cc040eb93ed79e22331c5e4fcc3f47e087642
F20110217_AABXBR smith_j_Page_068.tif
f8b7c373a840327c64e2565af086f9d7
328afbdddc248f5dc13de546fbecbafba012b44a
5728 F20110217_AABWWL smith_j_Page_120thm.jpg
e481b34958ceb7e1f9f4741753c6bef1
215aa1f7e02f7b28dad7ee0235a3f0e29d5cb9bf
7624 F20110217_AABWVW smith_j_Page_083thm.jpg
d091d8a88a2e37678c1747d4abfc7aac
766ad3662cacbb82a66245ac3e67476a708b6c61
42712 F20110217_AABXCF smith_j_Page_069.pro
6f680dd8d188b1adf3d6e6bce5997e48
a0a245f388d9c6c9db9d292f84b7357b9c539775
7844 F20110217_AABYFI smith_j_Page_102thm.jpg
2d2e4fb5b520e74025d9fbef5d49f08d
23089fc66ea906b7a89e4392abfbe892ba5184d0
34298 F20110217_AABYEU smith_j_Page_070.QC.jpg
02e54c8e79fa0bb487589f66f2799eb6
f7026eeb53ba81a3b1033ad7bd0f0c135030e8a9
71422 F20110217_AABXZO smith_j_Page_105.jpg
f0712092ca3281a1ddda48224169a5e9
dacb7cee67eb70795e5de7182358cf614116f6a7
85386 F20110217_AABXYZ smith_j_Page_078.jpg
13243dbb84214a2b793f34f73c08d94a
34f620ea03354879f811bb3325b10a326e534bd6
5877 F20110217_AABWXA smith_j_Page_142thm.jpg
140cb522fb9ab0d8c80e2832293e195b
acfc0cf280d3fc9c68edfc77b7c8a3d92ff4dd0b
1951 F20110217_AABXBS smith_j_Page_086.txt
bf396b12ff4f887d3d1be1438a2e8806
dd733a9f46bceb14e77bd468e828984fd7dae18f
31543 F20110217_AABWWM smith_j_Page_045.QC.jpg
cbf16b598b3e40225b90d302913b8054
163ef549a1172dc7b8eb12cd572b91ff694e29ae
1051955 F20110217_AABWVX smith_j_Page_063.jp2
750fb8cd4cf1645cae71aec3f3e55e6d
784b55b2c08b895ae8710d1d04e5f9ac3fca81eb
40884 F20110217_AABXCG smith_j_Page_050.pro
adb0cae035411bb70a37e35bc057f2b8
bb143642f84303c0b1fafa9b8e93736b8b6c29f4
28563 F20110217_AABYFJ smith_j_Page_056.QC.jpg
27c40538cc66a2f8073b63367c2a5629
c128633f4265c0cfdbeeaba907d1c344698eff37
16092 F20110217_AABYEV smith_j_Page_004.QC.jpg
96f20cf1a8331a97cafe7976514d4e43
0a29a925d96720cb618021139f214d22bd317897
58379 F20110217_AABXZP smith_j_Page_107.jpg
50eb92c46a58f5786bfc1a3162a4b6d4
e338444f088ce0d4fcb7c8de163b16aacc13421c
500335 F20110217_AABWXB smith_j_Page_004.jp2
473f67a9324a742e08b0a9d4cce732f6
085787510b10d60547e322f7edd0c0dd584e70f6
F20110217_AABXBT smith_j_Page_026.tif
d78a68520c13dabede2bb4d3994910a1
abbe8191292fe5304ef31abf80b8f62845c18e47
F20110217_AABWWN smith_j_Page_140.tif
79eb0527fc3e27e15cf491f33c72bbef
3de5587d997cb261a1187226d6eefe92190d2aad
5186 F20110217_AABWVY smith_j_Page_116thm.jpg
c75b8b894d4e4e15038bf92854b395a3
1bac4528edbe67e697c835d88e1cd075cccbb1c3
961686 F20110217_AABXCH smith_j_Page_069.jp2
5f231b81cde8e2cdc1d468ec2ba331c9
eb19f04fd5001df8665a3dc665573c4b2c9cec7d
33780 F20110217_AABYFK smith_j_Page_060.QC.jpg
8c5f8de7a32b4850858fc2b7312816c2
3cc74ad306c77252113f0a72909d0524b943940c
22374 F20110217_AABYEW smith_j_Page_105.QC.jpg
ad8771e404336514917b41439ae0835c
658ab8f0e93449dfe425c1861c6409f4681fc7a6
7129 F20110217_AABXBU smith_j_Page_054thm.jpg
312a4b556f267a9f4f95c410018cbd5a
18e1545f673a70afcedd5fe8323aae1415499f5d
1949 F20110217_AABWVZ smith_j_Page_063.txt
afa645ade542bb9f298aa5c8ae9945db
f23380e0d53a138ff48f0b648ec3affb311a0caf
1051961 F20110217_AABXCI smith_j_Page_101.jp2
b2e10926f476d8a0197f8601f1dc543b
a7fd73bcdf765cfb8bebf6ccf1262f37e4a3e7c0
59287 F20110217_AABWXC smith_j_Page_124.jpg
116b416587cc505ba5b5abce5b74f6c4
3719a908f54786a3b5facdb8fe8a4f3f2317505f
5128 F20110217_AABYFL smith_j_Page_130thm.jpg
1729d6c5ba4a63f71813dc7c551922cc
51fe14805b795ca2b35fed49421d86a0c1a25fe7
7482 F20110217_AABYEX smith_j_Page_085thm.jpg
00faa4456ca4b81e7e0e98e7129fc827
4f2ef7acdccc1592a004ddbdcd4ce9df39f6b4fe
47704 F20110217_AABXZQ smith_j_Page_108.jpg
e78b8943576207aa9a7604a21bac534f
ca62da311937b476fa5f166e82b4f34144f75ec5
1051945 F20110217_AABXBV smith_j_Page_147.jp2
33559f9e1e08c5df17a7090df7634e38
333b8078de0dc9b3064f5928f0136c025ad27d04
622828 F20110217_AABWWO smith_j_Page_112.jp2
a80e6959f0abc47be236a863d7ffb554
fee992d4dbfd68bc0d208f8a3ab8fc079cec4bc1
488559 F20110217_AABXCJ smith_j_Page_132.jp2
847901a0eea83b125856b8fb5d2cb9c6
615cf4c4cacb8e110b2682bd256ec4a52ecf87ff
106782 F20110217_AABWXD smith_j_Page_067.jpg
1675064c4a8dc0ea0ff062cfcd93121c
c8e1b3eec27d903fc8594404b4946a803b6b28f4
3107 F20110217_AABYGA smith_j_Page_123thm.jpg
2e9aa2e5689ef5114c61594fd0b8c371
1763251a878e76b18850385766c33f667034a859
7727 F20110217_AABYFM smith_j_Page_017thm.jpg
98e26c89ca0d9c2ee1e6b7dea30bf909
21852789127ba1ba528f33345d6142664b5a97c9
6954 F20110217_AABYEY smith_j_Page_088thm.jpg
82422c9f383a3af7bd4c77b83e27cc0a
f7ee922c0c987a6e8d74f74a70c6e393eacf66f9
69374 F20110217_AABXZR smith_j_Page_109.jpg
dd6365967628d43b4ace9e99e7ece155
963729985a05a1c7d957f42abc0c9f00b362f1a6
1051972 F20110217_AABXBW smith_j_Page_067.jp2
dd52ee1e0b4ab050121d2ecf9504b98c
dc5e6b8f41a9f25133ec232ac3f3eee5e7f68786
701732 F20110217_AABWWP smith_j_Page_092.jp2
b34db0b2a3732d2bd16df7a5a7c9a8f0
4cfc7e8b0bbba3d4f8724e42f89138eb741060f9
39429 F20110217_AABXCK smith_j_Page_078.pro
6cc6c9a2e1a683aac8253f3e6edf9da2
b316ad2097d7ce5be6ab2cd7c752486c9aed0dd3
F20110217_AABWXE smith_j_Page_143.tif
03a9131405778aac2663c0f4c4f4c569
b36358da5d4ca9d8383da040d26085995329db68
32778 F20110217_AABYGB smith_j_Page_156.QC.jpg
00b209bd54b8415dd273f71dd65ac626
34d151cff4354bd5dc2ceb68f003a0a301aff315
7902 F20110217_AABYFN smith_j_Page_053thm.jpg
52d153fc770293a518c4c7a6c99b9410
21f0b937df0bcd655ceb9fd0a6966aed76fd61c7
3930 F20110217_AABYEZ smith_j_Page_108thm.jpg
b3790719dacee0e6b3eb717e5a17ac58
7327fd1d2727357306bce5535c9855d0951d0d4e
63501 F20110217_AABXZS smith_j_Page_110.jpg
9864367dc4d9ec15d363c76a33369c31
296ccb3c17c8a419f8f3baa3be1ee166014fe18c
F20110217_AABXBX smith_j_Page_071.tif
341158d59abd71678cb2d1ecfd671db1
46b9a02b1769298956e5defd66615a15ab8bebc1
1051942 F20110217_AABWWQ smith_j_Page_060.jp2
7587eabe2fd195d48ca7b5c767f00ad8
1aa52f9457afb48e147c5964471d3800c68f8643
4002 F20110217_AABXCL smith_j_Page_161thm.jpg
7f19872eb57aa25857b85ad5c1eedeee
f2a123ddec42503b0c7d29f822eff90b9a2fef68
23423 F20110217_AABWXF smith_j_Page_134.QC.jpg
9d64689a901d5e308833c6c6f16d1f47
1165129e9c1d53f0e884ad9d3b20e02661ee5cdb
8151 F20110217_AABYGC smith_j_Page_156thm.jpg
920d6bf4bd0ba9e815071e046565f320
0c6542b1e7629f1cdf7ff43b4c14fd4d04ded2c0
28526 F20110217_AABYFO smith_j_Page_089.QC.jpg
2c29143d36edc7b0cc39baa7135a15d7
e616eb20023087b12bb6fa19b3ad27d8185f2578
69174 F20110217_AABXZT smith_j_Page_112.jpg
ecd95b5d815c5e80d9ec556697d562b1
8a7f94d83b2d24eaacf483dfe802341a90b973d0
1234 F20110217_AABWWR smith_j_Page_047.txt
70cb61a692321807942a83a3fae524ef
52c7b9bb0a868e7e701ae2c7dec78b3969afc230
5847 F20110217_AABXDA smith_j_Page_087thm.jpg
68d57a65ca11c13e6cbcb7c2276403da
d5e84907167a2f7912ce0e23068dd5bb5d638343
F20110217_AABXCM smith_j_Page_146.tif
454396762da052ac2a0f7156d4360bdc
0c9bd5b7c972aa97d4278eb742df035bf4a23db2
F20110217_AABWXG smith_j_Page_102.tif
1148c058f88f338ee75712a425bf8206
b53d6d595a3293b85e12ad9f18e76e62ca1b1d16
29422 F20110217_AABYGD smith_j_Page_035.QC.jpg
4a09f53ecfa4f9bf52f73209720ff03c
38641909ba2b8d31638f4c5c5aef57117ef313f5
5961 F20110217_AABYFP smith_j_Page_112thm.jpg
5841ed7af551ebbaba00c745a05bfa5e
4c28ec1260c3251a22817d93456b9d8bc4cefa93
52789 F20110217_AABXZU smith_j_Page_113.jpg
46be57667a120cdebab2fd6ef3bc51a5
4664576895ab64fb69293c783704c65397d5e69b
815629 F20110217_AABXBY smith_j_Page_018.jp2
74e2344f2113cce5b8380a90fb5a8231
1ee04b3b62209832b82ea575cb719a10999bfad8
63750 F20110217_AABWWS smith_j_Page_153.jpg
f5055c0b2195db4e97a12547050b2344
c456c686d5ecc864bffcea2a1a775213890b9a01
2053 F20110217_AABXDB smith_j_Page_137.txt
e480e740e50186f1bb71fd1f6791e045
f09cb56cf6b0f0a06b14b9120b8961e821ce7b26
1939 F20110217_AABXCN smith_j_Page_093.txt
cd8afbf78707bfa1aaa75706bb19d840
183d1002ee8259130afe2f7f6b6b2f7e8480f3f6
F20110217_AABWXH smith_j_Page_098.tif
832ff6bc863cfacce2b8f759b9f5b521
48c14ac2bf87d9397652d5682856cd3b48068c74
20113 F20110217_AABYGE smith_j_Page_141.QC.jpg
bac0a8203d5c5fca3ee6d60b2917618c
24746e2fdddb7a48509b3b7712dae2f8b545d698
6808 F20110217_AABYFQ smith_j_Page_039thm.jpg
4c492a1d3e54b6100a6b6b3ca856f75a
12413a40e3d5086b61af96238c85dd92b6e9d5be
49072 F20110217_AABXZV smith_j_Page_114.jpg
e02621c9b813c2b47541596e00ef5866
5821efa9a4d4432451cb97031b3414d1dd629c07
6981 F20110217_AABXBZ smith_j_Page_048thm.jpg
99b4567802e95b97277eb5c491af196e
cdbaab2bf47180b6e9f085e5a57e957fa867f7f6
F20110217_AABWWT smith_j_Page_117.tif
a73557a29cf898a672e9d522796a6fe3
c31d82926d3c742a473c672e5f18381bfc178645
32867 F20110217_AABXDC smith_j_Page_159.QC.jpg
5636e3ae19040b1ea0033ded785ee00e
13fb935551d304c900b73b75984a80fe3c67b3bb
F20110217_AABXCO smith_j_Page_067.tif
170e934b32ac685d01894ac4346c0c4a
8e5676ff937238969781cbc67bc5b0b9c6e005ba
F20110217_AABWXI smith_j_Page_127.tif
ea0bd66f85b39e101d5dd3ca323669fc
d4174e98f0d1b9e789351a50bd6c6ae88e1dad58
31006 F20110217_AABYGF smith_j_Page_037.QC.jpg
6c0e1cf24d653e3f8e8241eeeaeac541
69c77ddf2e138a40b263825d1773acff5bd4effd
8345 F20110217_AABYFR smith_j_Page_074thm.jpg
10777c89b068adacc9d5e37359e9b26c
1d92422a3a6b7f85e2c684f9dbba50443a076c33
60530 F20110217_AABXZW smith_j_Page_115.jpg
04fa4fc68c7445df0162b54dcbb7e2b9
4bb71549fd9de02090a2b9852213684a26f9b9ec
16680 F20110217_AABWWU smith_j_Page_039.pro
768af38d7a72337abb6661dd9cae6603
f1c73f3dc4608c8c09be522254973242a5686844
4971 F20110217_AABXDD smith_j_Page_117thm.jpg
60878e725395943b15fc0a6838523ae9
7ab2afdcc2ed1b4b20906754b7c56a099c7e0a54
51822 F20110217_AABXCP smith_j_Page_102.pro
b6b72ba5145aaa800bd571136fa109d8
96ba6b201055ff395f1daf7ab74348fa11acd6f9
3084 F20110217_AABWXJ smith_j_Page_044.txt
6e4c0fa0c84b8ea59cb85e0ef9641cdf
e010c9a4fa79207919c535efdb0b5309af93920c
5999 F20110217_AABYGG smith_j_Page_040thm.jpg
d74493f80be252a83d33d04ab8930e97
041d4b9f6b1aeffe3d29599bb9c402921ea6f111
3202 F20110217_AABYFS smith_j_Page_160thm.jpg
1664719cfc678a63f17adb2132dc0aaf
d862113d594da926c945c755815ee35786585f97
55626 F20110217_AABXZX smith_j_Page_116.jpg
60bdb1219bd053ce9e933de34b9e22cd
baec2cf366863e754799b4cee6fa8460806b86b1
34020 F20110217_AABWWV smith_j_Page_148.QC.jpg
44606bcc5215023920d7df500cd0362f
4de62417b53d222eab9ebc032dd2f80f1d46f645
F20110217_AABXDE smith_j_Page_134.tif
c8e766f3957a1b6b1910291dbd721a78
8897a7f5e4f8c884edbf6366a78cc6a1ec61e8d7
96387 F20110217_AABXCQ smith_j_Page_102.jpg
1664a1c0588ae4e3c15a3ac37aedbc07
48e664b430a8dce99b69309ee50bc887b1f44dd5
644109 F20110217_AABWXK smith_j_Page_119.jp2
1c0ac9a959f9c50966035ffa8d6f2aab
d65843394e1e06df4ad1d88e62cc74fae689bada
5553 F20110217_AABYGH smith_j_Page_135thm.jpg
512fde7734a3b00485218100aa84abb3
6e8e6b08afdfd0d71e2f41e9c49e5a8e49cb30d4
6952 F20110217_AABYFT smith_j_Page_023thm.jpg
1656ef8a90861dfb9439d27fd5937dcb
a30aafb2d62ab9ab016dc4b8f1a06e6b7bd642a8
48439 F20110217_AABXZY smith_j_Page_122.jpg
aa03ec0dbe8e6c4fc25f6a1f69511fd6
43cb406344dd6cec4e5b47cf7094e21c7956b0b4
2913 F20110217_AABWWW smith_j_Page_021.txt
0e6c88946736346c7006c87c39e5928a
c1b3272da85af65f6c6eb6d294034ee90e75c0bc
20206 F20110217_AABXDF smith_j_Page_005.QC.jpg
9773a10aeadae6e803f0ca6fd41902e7
50fa87650cfdb7b9c8b4421e9ec7911a1b3e2bc9
907727 F20110217_AABXCR smith_j_Page_039.jp2
be6dc0671cdd1c5e40db9a8de1642538
de33a3b8013c4989d92e802940368ae1658dba0d
25486 F20110217_AABWXL smith_j_Page_104.QC.jpg
50feef68e42bc12901d8d11def53078f
3f0343100e5a3d99b1f30e64237ee81df2228b13
8398 F20110217_AABYGI smith_j_Page_157thm.jpg
622c74bcd9ebc97f9db99ef28119b340
d9f81136eeaa2f5996efd304171697f978bac16a
18322 F20110217_AABYFU smith_j_Page_111.QC.jpg
c2ad5753a4bc1460166809478004920f
33cc3e747c505247590be275da2acfa806df9be4
26413 F20110217_AABXZZ smith_j_Page_123.jpg
ae4f8d39360e5145b205a0e88a4a859b
42970c7359ce7e1801c8fadac56cbce5aa9a0576
31993 F20110217_AABWWX smith_j_Page_061.QC.jpg
e9dabe04e4d096fbbcc718761ba6e114
cc59ef89626c842ef4e18fde79af705a761161f3
1695 F20110217_AABXDG smith_j_Page_110.txt
416ee7e2e88577ae8e857d3bd5d2c14a
28af3ec55ef62afda0b5948fc6ed871cfd06d2f2
3977 F20110217_AABWYA smith_j_Page_146.txt
d590fe2da35cae0843421506140421c4
d452632e36887c590acf51f4ee31d9fc93fabd7f
26274 F20110217_AABXCS smith_j_Page_133.jpg
7b7e56c8154bf04d26325ab267e820c8
9c2231d0acaa1913f9e9eb44e0a6b545e078276c
F20110217_AABWXM smith_j_Page_012.tif
893bb2c83a314618da67178db7482c63
1cec3fc1f4126cde692eaa784d8d289a45cfa2d8
6719 F20110217_AABYGJ smith_j_Page_098thm.jpg
719ec1072bbfb223f7ef04354a42a782
b7c127f18a8c07c2496c9cdd4a9c7d06e30ea5a9
17878 F20110217_AABYFV smith_j_Page_136.QC.jpg
d5fd80d396f1604a25a6e386392afbbe
ddc578b96e906c36b89f5e8dfa38970369943910
4542 F20110217_AABWWY smith_j_Page_118thm.jpg
b618adcf334be61f77ba923ca039eb46
4d9e212c314677b03c893bdee07023e8b4635dd0
37399 F20110217_AABXDH smith_j_Page_115.pro
f08cad226365f8e62b82e2ba338d836a
5e6aee8881d7774c4af39d97f24e5f25453b91ea
48039 F20110217_AABWYB smith_j_Page_061.pro
ba1910c3cb6708ea6015a706bbc737c6
3346b39bbde5fde8b6401725c3500bef8cf4b521
28351 F20110217_AABXCT smith_j_Page_132.pro
0212ffb3c3029539b922f7d053ee1f5a
d330c6f36cedf01e04b6276dbc797078df44b3c4
17686 F20110217_AABWXN smith_j_Page_137.QC.jpg
a0b0cb20fb6824a5d58d4ee43c0e2866
dfc0d9b9f88017a3d6edf5df35ca604ce07ae286
23217 F20110217_AABYGK smith_j_Page_119.QC.jpg
139136647e88362089e9b3a3cfc9a097
a7e51da44128a589a93e2ed0fc628c5e76d2c0e9
30841 F20110217_AABYFW smith_j_Page_073.QC.jpg
c38889b3da2f380bfb078b14d831a410
b8f386f128a75e5598f4a004c7a9d31d39def0cb
F20110217_AABWWZ smith_j_Page_025.tif
c412223bc7b4ee712191c58ccd2acd80
68e9b5e2d085942213a26d39ef639d21da3dd7c5
4569 F20110217_AABXDI smith_j_Page_097thm.jpg
0ecbaa889d851db4ba32a368a246d966
1fcf03e2c8f3a8d25221c04a32e78bc45c370992
72413 F20110217_AABWYC smith_j_Page_119.jpg
5d67030761ab7cfdc2dec8aa672f1a05
d5f8fa1c92ce6e91dd2fe1d6e83865050ed3ee7e
2013 F20110217_AABXCU smith_j_Page_026.txt
e3f54c02720fd21819e944beba0f749f
b8c21c8adbd9a8b2a10fb3c1fa1610790d36edea
82846 F20110217_AABWXO smith_j_Page_147.pro
4161e01209d2eac6a0942a6acf38d7c6
d55e49648a730d00c3bbe5305b6e463c10b49217
30352 F20110217_AABYGL smith_j_Page_102.QC.jpg
e1e8fc00b8d25e8cb09dd8e295b5dd78
4717de363abd8859acdbc9a43be97771de845870
4886 F20110217_AABYFX smith_j_Page_138thm.jpg
8a99705d8cf573263a4affb0b48e41c3
d995a6e252fa420672709f28c0acb804ddc65d9d
32029 F20110217_AABXDJ smith_j_Page_074.QC.jpg
0f75b99bbbd9050bbcf99cd3766447ef
456099a0c8e6617db30e13c74335516de16603e6
1051947 F20110217_AABWYD smith_j_Page_026.jp2
92c07038874ec9ca3ff535bce63ddda9
7e04b29ad58a7cc127e9cd1de00e21c53e256c2e
38689 F20110217_AABXCV smith_j_Page_146.QC.jpg
622710d22d63112a8222d6172a6f09a5
f664b6c74b3efff5474960523a679733fd8eacb5
8028 F20110217_AABYHA smith_j_Page_149thm.jpg
ccdaf9e68cc65e50abfa810414b3e9b2
ecef7b43adf2c2d36587061c0d6f270a871c67b2
17762 F20110217_AABYGM smith_j_Page_077.QC.jpg
0edf4231c8103eaed08c21e24155ac12
e6639c0decf2a5da313d16100f525796bb48c50f
30183 F20110217_AABYFY smith_j_Page_017.QC.jpg
9ca799c6a2674221db2468a02beda4db
72f1da94a43d9baa37c6571b3962b9c524db03d8
6615 F20110217_AABXDK smith_j_Page_140thm.jpg
b17a1ad4a74f985ac0ba98abd880fe19
84890bcae661ae95e4a9e5e7967d04eb5a25bba7
49253 F20110217_AABWYE smith_j_Page_121.pro
249d93b06145648107df9e1072270aa4
92242e921198a387ca0bed73c9741ff56de68a33
7834 F20110217_AABXCW smith_j_Page_029thm.jpg
58ce8d2dfa25b643218aa975c19ee987
9bf1f1295acfbeafdb6920b21f2c43515499e648
24277 F20110217_AABWXP smith_j_Page_072.pro
d30f63b723caf5b08ec9a5436a6c5237
2f33bfb04c43b359fed1e5ba607cc725e6bbb3ef
7857 F20110217_AABYHB smith_j_Page_096thm.jpg
f8f26c21e154e66c1fb8a675019c3749
9ff22ff82746a4446db4a4738f28bfcbfb780575
18018 F20110217_AABYGN smith_j_Page_115.QC.jpg
1cf9e7a4a64f16040eb5c79384e8ccfd
d40974218d48939e8d528026b8c66def0afd4e59
6638 F20110217_AABYFZ smith_j_Page_143thm.jpg
94e61c19d9ef944571791af33e60d8f2
20e5aba1e115a5baf6bd6ede4181400da4740a58
30233 F20110217_AABXDL smith_j_Page_013.jpg
5d2399dbce2f8feea94ce09c5a9347f1
cdccfd79f83ebea2888088fd1e699f53e841e911
2215 F20110217_AABWYF smith_j_Page_155.txt
d06214d6f684cd0f1685f13638b1914b
ea96737c09c53bf2c0a7a3ed29281ddf237ada77
16111 F20110217_AABXCX smith_j_Page_081.pro
2abaa7784ae006aaaee346a20dbb18c6
d460e43f33cb6415a2963214cd610df2fd9d6eb3
56381 F20110217_AABWXQ smith_j_Page_125.jpg
465ce7c79358e74f65ed7dacfc302fa5
314d243dfed2d84bc5d4dcfb7430f4c8645795e6
30056 F20110217_AABYHC smith_j_Page_083.QC.jpg
2f1c6651eb7712f51265c2fa2893bdc5
a06a9a6219cc1bd2fee87e305c028824bb9959bb
3229 F20110217_AABYGO smith_j_Page_007thm.jpg
25b030ac14750f81b5317df686d46a83
f5673621887f7ef23d27b3aa78a46897b8cda62f
3239 F20110217_AABXDM smith_j_Page_150.txt
4ca7a673d047fa371dd2db39a254afdb
a7128fa24b84f149c08467ceecc59fe945e5673c
45217 F20110217_AABWYG smith_j_Page_106.jpg
e7ca28df7a552ef8cb6c68cacbf858a8
cb31f7581191b7ddf588efc38304dbb89148ce00
1980 F20110217_AABXCY smith_j_Page_100.txt
3f2ea0151f20b3302eddfbfa0ef80fd5
9ba843269407d46483a6a6cb133670cd918db069
5665 F20110217_AABWXR smith_j_Page_110thm.jpg
ffb6da9903ca54e7a62f4b69a38cced3
a87f3d3414bf048b79697ece125863396622db94
F20110217_AABXEA smith_j_Page_001.tif
8bb779fefc4061fd9e4b228068522ca2
93a4dac12bc5140301c87a9a03feeb5602dcf220
7453 F20110217_AABYHD smith_j_Page_038thm.jpg
2b3623155d8502ef2631a7180bb792cd
62c105e0be1901600d5c525fc02fd737d5454caa
8019 F20110217_AABYGP smith_j_Page_041thm.jpg
85e859a03492bb6271bb55b0b02e853e
1a52958d28680506eddfbf46a649ab0289f84527
477887 F20110217_AABXDN smith_j_Page_131.jp2
87f7a9dce4fd3a5dffb064c5932b5581
c08106023d2fb864ccb2085564c41b1a7fe14a24
38445 F20110217_AABWYH smith_j_Page_018.pro
23e374165e9aad30fe96a9fe1daa2b7c
830c4e90edad0aa6a6059e4a53569a3cabe3999e
1324 F20110217_AABWXS smith_j_Page_019.txt
7a7a7689a69402d38b906b9d47a70c4e
cc0221c6d1192f4e0afa7c8f7b7df0f7108af10c
7679 F20110217_AABXEB smith_j_Page_154thm.jpg
66cceecdbe0af5bda290a45008e7c774
3924682d944905af500376ae0b5958b68caf1df0
4952 F20110217_AABYHE smith_j_Page_139thm.jpg
64281750bb4acc93e1ddb3ff9b52f456
70511848845b70bbcacc0d4f28d6f69acb950be7
5289 F20110217_AABYGQ smith_j_Page_124thm.jpg
8c20606dfad02ec19d49d70c524f0e1b
a9ee8cfe12d691239e518c0f7795d6dd495b3b84
33234 F20110217_AABXDO smith_j_Page_020.QC.jpg
df381d564f490b234253be5aed55dc38
f98b1260e515879e4679519923573128d142520c
F20110217_AABWYI smith_j_Page_057.tif
2a8ee4ca4643d3fef9a74c8bf0c0ec14
ddb6c966b18f35a3e1bc38fc6bc30f921413b67a
F20110217_AABXCZ smith_j_Page_025.jp2
ed6c51c4ef604525ca0d776453d164c1
24ea4961d0fbdd9e385ebfa3657487b79d96df3d
70563 F20110217_AABWXT smith_j_Page_051.jpg
c9711481bb8ecb6436ed4b10c6a19503
f826ef55e6ecd2dd98c5ad77f372ceff5a3cc41a
93536 F20110217_AABXEC smith_j_Page_066.jpg
98aaf6fe559de91fce83f74f2c2357d8
eab012e55b9d4032cfe65cf922252a04d0e98f70
26315 F20110217_AABYHF smith_j_Page_088.QC.jpg
4259a053131e68f269e3ccb262e6f504
d454097b5831726c9adab8de0a1ec2a7b925301a
30477 F20110217_AABYGR smith_j_Page_022.QC.jpg
c37dbdec506576194c3af57159152183
451a5d7269296ab8d8a89cc17e3a494d3088897a
F20110217_AABXDP smith_j_Page_046.tif
3e78d2ab9cf96bbc5cba9348f28b43b0
119c31616d8e1d58203aed172ecee6b2fc3fcaf7
6372 F20110217_AABWYJ smith_j_Page_051thm.jpg
9424c29cc1b615222abda0baf658328a
ac783c325c7d6eaefc0d08416f11f121e331ce20
100040 F20110217_AABWXU smith_j_Page_060.jpg
c35b0de74297cf130c27a52d1a496177
ccc660c7ccca6c8a4293ffcdd15d4c489fd55114
1976 F20110217_AABXED smith_j_Page_041.txt
2c0d18c50ad8a47b44eac807b7d455e8
75a3c45a97ec8fea19fc7747811e69624a830f5e
37717 F20110217_AABYHG smith_j_Page_145.QC.jpg
fb6a898e052c06636876e3ca6c4a7ab5
a338c8f0d61f113269ba72b9d122ba424aa71adc
33814 F20110217_AABYGS smith_j_Page_041.QC.jpg
809733d4069898699de18bb9e5fbbd37
28867bd8f68f5cc53ee8cb2042963c304ace845f
66651 F20110217_AABXDQ smith_j_Page_117.jpg
8a55acd4b2fae472aa1f676147787823
d92299fecb218bebaeedce43789197e72fec7a31
805205 F20110217_AABWYK smith_j_Page_055.jp2
389576507a1f2c7b89e7c18fe25a5ae5
efebfa163363fe8eb54de3f87b4ec2979b7ba012
5278 F20110217_AABWXV smith_j_Page_131thm.jpg
1d36899e55f384d54d616dae983328cc
db365fc5e1241bb7224ed108b10a121ae8c2ff26
21039 F20110217_AABXEE smith_j_Page_087.QC.jpg
6da9627d1f3a02430646f99b99406f73
b49f7a70c9aa271839e67f986c662ccb507f839e
25915 F20110217_AABYHH smith_j_Page_092.QC.jpg
ca80f6c5b133ca83cfa3491f7d21e16a
1889f6adf1d5631e07faed32f0a895549787063e
17944 F20110217_AABYGT smith_j_Page_130.QC.jpg
0eaade31c69d2813a3fe4e78bb0417da
980cbbbedc8f6f0eff90c98e17fcd3d068dca7eb
54908 F20110217_AABXDR smith_j_Page_143.jpg
3cda79eb24270ac03551f3dd090de6ce
77644a2f768f779093d40b7e9eb9705c8adea74c
7802 F20110217_AABWYL smith_j_Page_061thm.jpg
c4f7fdaa75e6cc1d5885739b8345ac86
1fd1a68a5aadb608819ff0e07be036baadd223c9
1878 F20110217_AABWXW smith_j_Page_131.txt
7aafb123d7125e8bfdf48f8cc4d86022
2d303ade90ea37989abf74eac1479ccfbc2f3529
491282 F20110217_AABXEF smith_j_Page_161.jp2
d601a6dff5168d4f41f121d1ec61e265
0da56b2c828047d7b2cdd763c5c9873ba76b1a70
7984 F20110217_AABYHI smith_j_Page_103thm.jpg
0319ad1e7f88d9bf80b17602b5873e73
77de48b6b59fdce7a9573a02de0976b93947b884
7801 F20110217_AABYGU smith_j_Page_090thm.jpg
2d5109712fc4bdc8b9ff1a1eae4edd1d
3fd4f9b9b992472c4cf473af0af43c2fd273e5ec
32443 F20110217_AABWZA smith_j_Page_019.pro
725b9285a63d700d0403052bba0fb79a
50d94a83490b86df87509924376715633916a8a6
8419 F20110217_AABXDS smith_j_Page_059thm.jpg
8be76360ce30714c35a07c58139984fd
0c530b9096df6417b2432d25306cc5b891fbf5c8
F20110217_AABWYM smith_j_Page_009.tif
cb0f14e6a5882fb1a8b62e3ccf5d4280
59bbe00243ec859807ad46a5e21c49ea4a9bd4f3
44135 F20110217_AABWXX smith_j_Page_068.pro
0784e5dac57fb74003cda36170982d34
663cd73e7582431b8aa6606ad743bc059fc1a8f4
6152 F20110217_AABXEG smith_j_Page_113thm.jpg
8109876b7d678f6b2c63b1b8cf777dc1
2b3fc536a5fbfecfa16ef0d6ea355e1d7d3eb706
6893 F20110217_AABYHJ smith_j_Page_078thm.jpg
067ef28e0ac97063eec87f831d6a774d
80a9f6b61a6a0d6a184dc97dbebaf0bbbfe16bc9
34345 F20110217_AABYGV smith_j_Page_028.QC.jpg
95b53a09906f6d1649f1d87176d80727
29ac411a1470230050247c128d338d2114a03fd4
32259 F20110217_AABWZB smith_j_Page_103.QC.jpg
14c8256c7f5890c75173a9195612da51
60a2ddac255175ccf1778f1bfa49be023a3209a2
35241 F20110217_AABXDT smith_j_Page_076.pro
cbdcfc69e481ce2a6bf52742974e1e85
cc94aab009033a607d83f25314133dd4b0a68194
F20110217_AABWYN smith_j_Page_086.tif
e3b0a32e15ebf339408e813cd447d1fa
0f2709f0dfb95236b5667b5a843ae9ec80466517
6460 F20110217_AABWXY smith_j_Page_049thm.jpg
d56be7343c8daf9e316cf4332f7ea335
fe3fc66bc430e4fa4c0c0c496373c21fdbc9324b
F20110217_AABXEH smith_j_Page_084.tif
d844556fa573b58d81ef8f6acb6a5204
c090b8bc599a3d136ffacf6e396360f676ff4038
5142 F20110217_AABYHK smith_j_Page_081thm.jpg
3efed584535ee60ea6a076f83d1e8d37
82889f497b9b5d81fa2ff82806bb2469117c202a
35148 F20110217_AABYGW smith_j_Page_149.QC.jpg
d3c746055c56a7fdf34a141888815a18
d09ab9ec8d3bd06ddeb5ca8deb89423620f8bfc7
54202 F20110217_AABWZC smith_j_Page_097.jpg
1c7ae19f624708c9e88d74f12d13c1ac
cc4c1bb8589e3237150ac4d6c6910e9d9fd9977c
16510 F20110217_AABXDU smith_j_Page_031.pro
c6565ecf0085cd5ffdc5ac35c9a54474
8940b782d3a1d29ca3a3d064c035782d67a8b1b0
8829 F20110217_AABWYO smith_j_Page_146thm.jpg
39000d44fd334da42434fdf5cda61a9d
fe134f4384a16407a7923bded9a2dcf414b3b7ab
F20110217_AABWXZ smith_j_Page_157.tif
ceca3637e39f009caff7d7e31bb3ace4
b592ef7a9203fa95b65b8e4c7173049785cf1f0e
F20110217_AABXEI smith_j_Page_016.tif
6e9cab19d5604b9aecc02ac59cb8c06a
d8de127fed2419d66bc71ad76d29268ec99517a5
31521 F20110217_AABYHL smith_j_Page_071.QC.jpg
e87e8d8069c8b18cbe18bf703a58c3e9
24313f8d17e630365a770bbc564a77dddbd4be49
18861 F20110217_AABYGX smith_j_Page_049.QC.jpg
a64e544403db1a0f15cc6d6853cafab3
fe3b7116914a962b7cdb0b8050af173c8d0bdc18
2028 F20110217_AABWZD smith_j_Page_056.txt
a7cb65da654f0a071de88d49ff7e0b94
ab86543e674586634b71b230c41df7f5eb516ad6
472587 F20110217_AABXDV smith_j_Page_137.jp2
1104a97648899cd9064ab674e5a7b0f1
4b2c10b229d2431b77769a46264d06d5cd943f8f
480442 F20110217_AABWYP smith_j_Page_049.jp2
f02427be17bd2fa85b50b107171a3de9
59951613a6db94f53c2bb34e255ed436005307e2
115412 F20110217_AABXEJ smith_j_Page_156.jpg
015e2c70ac6d641f80a2574fc7252a5c
3b3a375db82e2f15b490481df32eefcfada792b4
25959 F20110217_AABYIA smith_j_Page_018.QC.jpg
da73f8e749d505938c44908dfed14730
cf9ed707f6fdd10f4bc788eaf01b4a44adbdb469
17983 F20110217_AABYHM smith_j_Page_139.QC.jpg
06ac9f87a9eeba497e5e24b61751e6f6
a9a37a0593fc5461430d54d617c663f87002fd57
31462 F20110217_AABYGY smith_j_Page_064.QC.jpg
49c6491edc55756b5ff52cb59665ef0e
2e9e1d98018823ca53a03099f7701a3f44565703
5249 F20110217_AABWZE smith_j_Page_077thm.jpg
7e9c3534accf8465d4d302c1d13467c8
b2013a9039c2f4137e1c13ebebc78c9f710afdfe
2975 F20110217_AABXDW smith_j_Page_148.txt
fb35327957d90b5f7a7e8f75a39bb55f
92eefe3919edf3d1d2ce4529aa8ad593efc89a44
949534 F20110217_AABXEK smith_j_Page_024.jp2
182dee9a292b80ef1d66ef3f9f0bc7ae
9f96d945b5d418f678d4f4cbd014a6e7516df6ab
33223 F20110217_AABYIB smith_j_Page_015.QC.jpg
0859ae794b9d8c7e3355597d375baa0a
ea280d7437eb9e33a15d98b5ae1c930b9906a115
26823 F20110217_AABYHN smith_j_Page_086.QC.jpg
a7805bc345d7139d1a27f4cd7959ff65
1bb286b275460b5bebf22ed1651dbbdc282d73cf
23071 F20110217_AABYGZ smith_j_Page_006.QC.jpg
e088470555e7160e2541564772152381
d732b35ab334a79d60edf4f0076cb7b152ab73ee
8289 F20110217_AABWZF smith_j_Page_101thm.jpg
5494a5941a552f24906cd3024da2b457
c2949559d1e8b2d67a4698fbdb5da13212b0d563
F20110217_AABXDX smith_j_Page_080.tif
dfc9cf1e773f8fc9cbd28ff6e95e69e1
3d86a4c4be9be98caa5efdb1cafe449eaee58acf
1665 F20110217_AABWYQ smith_j_Page_088.txt
84e89e3b510cf0522bc7bca9bd229a82
103544bfc64d394858167de85be47335caea2047
55745 F20110217_AABXEL smith_j_Page_131.jpg
12f9feb0cc04f235a31fde54dc1fdfaf
9c4e13e0067da9a56c1a72abc520728f9c67cc2e
25754 F20110217_AABYIC smith_j_Page_098.QC.jpg
29e250099576e7217993386ea1390a50
5c88e6894d8dac7daa8ba0765b7eddd0ffd1f0ec
5411 F20110217_AABYHO smith_j_Page_122thm.jpg
93f6154051eb0846e7b6921d5e1ecd93
d7202c3fa608504764be98576b5c8dacaba75d2f
449410 F20110217_AABWZG smith_j_Page_114.jp2
91f2556a37082d87e28989702e6d18c3
c8c365e5bbcf3ade833585ac0736e9a2de12ee70
F20110217_AABXDY smith_j_Page_096.tif
100860f9ba0a0609f081c93258d113b8
0ed6b906ffcd4cdd52d40afc4df15dfad68dbeff
F20110217_AABWYR smith_j_Page_090.tif
921650fdd0466b9cd90d575b764dc0af
bebe1a49e54437e3800dcee43dceb02e3634c637
F20110217_AABXFA smith_j_Page_145.tif
633136b72703885c0473e3e097637fa3
6403e2d9a5f24f3892ce1444e7d6ff55b68e92b2
98030 F20110217_AABXEM smith_j_Page_014.jpg
a363c5e0859891ae11b7d0d59e2bb8db
71b585eb8022050a40595b0c7b5cd969fbc3cb9d
19369 F20110217_AABYID smith_j_Page_127.QC.jpg
9edbca7f621ed6c19f04d850b6acdc7c
03782672931e417d02f65d1252d3b89255c6572a
5593 F20110217_AABYHP smith_j_Page_111thm.jpg
741463cce2071d217e74b877f12723e7
cdea43167cb3945506367af912770d6ab8d7a5b5
1838 F20110217_AABWZH smith_j_Page_078.txt
9ee49d6b80bb3664f2953a242687f76e
578324b9f84839e1909ca2b128f713d527d4d866
119498 F20110217_AABXDZ smith_j_Page_157.jpg
ef51894e4172f98be71fcb489b109bb7
f1c1388a3b7ffe107bbf523be0f9fcbf9802f6e3
38230 F20110217_AABWYS smith_j_Page_056.pro
02fab90653e398fc970342fb79461261
8da847aa32f64bb70610eb63eadcc433fad9bf78
80682 F20110217_AABXFB smith_j_Page_053.jpg
3b61befc20862c098d68c392f03d8b4d
9a9409e07b2e886f7524c9836a53ff8229d89cab
7573 F20110217_AABXEN smith_j_Page_030thm.jpg
883a4f8dbb54b9b0ef77784212d22563
053a041ac2eeb692b02bf64ca9d63fd1fad96d0e
5380 F20110217_AABYIE smith_j_Page_126thm.jpg
43b374ab533ca536a461e8e41a8eee44
f62afb2c2d4f71ffeaf689ede0cad21654813fb5
5344 F20110217_AABYHQ smith_j_Page_125thm.jpg
9de00d50b9a5d457824a40f6b5e5ee53
91275c162362e52da0c8a6c24891826db638ff84
7520 F20110217_AABWZI smith_j_Page_042thm.jpg
70dea8a08d3dc21854e0596b0654f226
ac250c6635aa87e4ac99f9691da782e3da9f1d7a
F20110217_AABWYT smith_j_Page_072.tif
f601915cf81493bc847ee52322e96e1c
6977e48686426245d962691ffcf1f0185d440f88
1914 F20110217_AABXFC smith_j_Page_020.txt
53022accb09f527df8724cdb128cf1e9
d5b1d6900dd491da4446319d796af45077192a51
51892 F20110217_AABXEO smith_j_Page_111.jpg
55c2c80c241e7e611855c82ad7a3ee3f
e7c6407fed0b037ea9427c3dd22cee84ab30e494
12889 F20110217_AABYIF smith_j_Page_065.QC.jpg
e54c7546dc06dcfea90f6efad891ba6e
3e4b5c2241f0ea67878298d0f9b3f96ce9929a15
23994 F20110217_AABYHR smith_j_Page_120.QC.jpg
2cd9b1c9babb8aefacd997ae05d37f43
89bb041e8f05cdb11ec802a7d2ae4463864a3fa9
1903 F20110217_AABWZJ smith_j_Page_080.txt
0f34b5e5a1ee2618378d0b838cbacf4d
104160848cc74ef53c075170f716b713a86be901
F20110217_AABWYU smith_j_Page_091.jp2
71e6b7bdd4fe27c38e764b47672adfc5
4adeae87b7b2a0c9b52f947eb2b0be1462844f2b
609074 F20110217_AABXFD smith_j_Page_047.jp2
1f451ed07374defec576d933bf33a52f
373cba15467bc135139bd89454da1cedca7f071b
F20110217_AABXEP smith_j_Page_105.tif
cf21925f65e63ad891fddfa2fde0710a
d5665538f8d13cd6826316c4e603db95a456cc69
29731 F20110217_AABYIG smith_j_Page_099.QC.jpg
1f3fcda7aea94ea7d7ac4d3c5382ea45
1c79e83b6cf51044f99e695417d3b985ca84f93a
5714 F20110217_AABYHS smith_j_Page_119thm.jpg
c2cf35d1942d43239a469c5d0b022a07
b5af64112e55b47287e788bdd2d72d321c2804f1
F20110217_AABWZK smith_j_Page_124.tif
2a3e7b528d5953a5923c7c4793d5e733
e81e8763715b1f617b9a381e6bd556aa66def4cf
8760 F20110217_AABWYV smith_j_Page_147thm.jpg
17aeb15ef274c79e6f5eabad760e1acb
9c9b89576761ead6ac63e228c1defc4f9fe585c0
606317 F20110217_AABXFE smith_j_Page_117.jp2
0600d27f79844e8621b3d5aadb49a901
ba0b1c245c2253af960fb338842dfa24c0b1b68f
1815 F20110217_AABXEQ smith_j_Page_017.txt
c56a54593064e4450da231e45a2c4bbc
b2147e1bbb6d5724c3a54a739e50ef125a396237
19687 F20110217_AABYIH smith_j_Page_126.QC.jpg
a38f0b31b066c9489b11f877ec2b1b21
772a3980b8a8ae097dc9c84d46af1df366ae4f9d
6841 F20110217_AABYHT smith_j_Page_062thm.jpg
f69789029531b7c87c4e5710c3dc97c6
bef1115b06a0d9ccdaabec96a1fd3f4a50b8f811
1893 F20110217_AABWZL smith_j_Page_061.txt
4caf0d4e5aa6e8541ee264402518d840
c5e20dfa8792fd4cfaa8e890e58965c9c20b1a36
17810 F20110217_AABWYW smith_j_Page_111.pro
c8d13819955f40bf8d3dcb43d42b30ae
6b897ece3127a253d16a02cef538a486a4da30ca
22518 F20110217_AABXFF smith_j_Page_040.QC.jpg
dab58a0b914a837909d96787259cc7b4
237bd96cf169afb3785ab0d754e2c136646dfd9d
26265 F20110217_AABXER smith_j_Page_085.QC.jpg
7eddd6eecf02022ad9c598c6ddebe3f2
4bcaf0060dfaafced26ab853217b3fc3527daa6a
18320 F20110217_AABYII smith_j_Page_097.QC.jpg
52382a609b5b28d77059a15a7ad8b522
1bfda33cc7c2abd9cd3e8be439d2fca529fe3e6f
6924 F20110217_AABYHU smith_j_Page_018thm.jpg
904de39691c7ff519f6e13063a80a90c
36d40fb94e58f30799a81765305f5ed734bb5846
26191 F20110217_AABWZM smith_j_Page_072.QC.jpg
f0c4102b8035e2b30474308c2def9388
956861ddfd6a0426243507c66939708af5305d82
8603 F20110217_AABWYX smith_j_Page_033thm.jpg
e3cdd8175d2aaa4259e815e989f737d6
b6fafc089e6950a874f1c83fd3f69dc8aeaf67de
107097 F20110217_AABXFG smith_j_Page_006.pro
f87fc580a5d6c5c5bfef705181660ae3
0256c3131cb93a90dee19093b9f75d93d2d015b2
19493 F20110217_AABXES smith_j_Page_124.QC.jpg
c484f766a43dcb472a7a25fedf5d88e8
3c291907f27f9e74df148701f45e332a28a25baa
31841 F20110217_AABYIJ smith_j_Page_030.QC.jpg
4c4495053d6a5eac7ae4c1dcf3931e24
d29e9262bf7a8c4ea12004aea8a3f4f5695cd452
26256 F20110217_AABYHV smith_j_Page_016.QC.jpg
35a8528c49f4a74008bad770b1b89086
a306f378eb18758d7e02d9e91d0bb6d689d57aa2
1943 F20110217_AABWZN smith_j_Page_053.txt
414bc7b1874dd05d1dda96f2d183995e
4e180fa52b435545cb040a837831b133d7a4ecbb
8167 F20110217_AABWYY smith_j_Page_028thm.jpg
8a2af89833679e7573deca18bcd3cb18
5bb0f0ba2f33d81e7cf23c444c250378fc60d39d
89406 F20110217_AABXFH smith_j_Page_146.pro
e484625321ca333ecbc5b31a9c4153b7
8930798ba028129f7b9e7dd8f819613a089727d7
79836 F20110217_AABXET smith_j_Page_104.jpg
02d0bb535181ccdbf9f1b3e8ae9db05a
03616bb19f564679b3b41cf07502a5343fbd4994
6427 F20110217_AABYIK smith_j_Page_052thm.jpg
81fad39530d45b48990bb71d4b575445
0247dbb0180c2e49ebd7472d646aea87baf330b0
3058 F20110217_AABYHW smith_j_Page_031thm.jpg
5c103a57de164e24b10154f381266372
50fb0edbfefb7d103d0bd91796c0c710d98ee7c3
42312 F20110217_AABWZO smith_j_Page_160.jpg
a75bc6889492e5a79c3c93eb40cf317f
3b3bfd91424502b8d6e144ee6b9976ed45ee62b2
1734 F20110217_AABWYZ smith_j_Page_043.txt
02ef876d9213b2d5f8463b059b96a1bd
d3744665c0ac51cec3fc501db13b6918de8a45d3
8945 F20110217_AABXFI smith_j_Page_133.QC.jpg
313bdd9490d615859f9d3ae9fa975df6
3c5261d80a3e405673dabc1c5a1e76d7f5c56a47
22062 F20110217_AABXEU smith_j_Page_024.pro
a111f3ed5d371eec0db43a16064fd3d3
29e2aed6580972105258076be91539dbe34c6001
23980 F20110217_AABYIL smith_j_Page_052.QC.jpg
0aebfea441b9f54e0103df08cc252ea0
a7fb9a1a4352a74ea57d9f1e37c803e6978e9a85
7350 F20110217_AABYHX smith_j_Page_072thm.jpg
5f49444ba59286a1dda3d2893fb2c212
df095749152c68a568203de4bcb57f02ed19ad3e
1779 F20110217_AABWZP smith_j_Page_118.txt
6486afddb20a6a73b1077b792ada4cc2
6d509443e6901362879380dca6c892c2fc0ba1c0
29733 F20110217_AABXFJ smith_j_Page_084.QC.jpg
bdffbde37ccb9885f40f50fdf605be07
3a4da346f565a4b90c5d9e1a1d83631be1601e76
927497 F20110217_AABXEV smith_j_Page_050.jp2
eebc82d19e4d3dc353294edc9ff04c80
4721d2861b63b57ac668a7db5ff1dd98c0ed7b24
32775 F20110217_AABYJA smith_j_Page_101.QC.jpg
26ba154c6e7cae0e6ff2f54f05c2260d
3dda85273e0b8672347353e83a8ae24a5c0e1299
5123 F20110217_AABYIM smith_j_Page_107thm.jpg
d26ac88156815471ee53038bb9896010
187a9cb909c182615402eeb932be0fa5e37ab35a
29659 F20110217_AABYHY smith_j_Page_090.QC.jpg
9b259a2cca4d1f6267c04ecebfd36804
f21e4ba4bbb15c87065aa7702bd36a2314ef204d
32318 F20110217_AABWZQ smith_j_Page_100.QC.jpg
2c71074c68324375f48428f0c366fba2
4f29acdaeb763efc1aad4264fbb0130f4f0e3c03
33571 F20110217_AABXFK smith_j_Page_025.QC.jpg
9bcc5970480fd0bd844473ff2a99072e
4a2d78b73fb46decb31129de2dbacf6267efa048
7831 F20110217_AABXEW smith_j_Page_095thm.jpg
221595fdeb221db80b6d19958fc58843
67278b9763e15f6effffd0a645110460271ae8b5
31608 F20110217_AABYJB smith_j_Page_095.QC.jpg
b32e732bf47ca84c23cf1fc61eb88aa1
3d1315f94b800d1a0320f9e8d56d3d6dfc55628d
3901 F20110217_AABYIN smith_j_Page_106thm.jpg
2e4a672c00319729e91c23681bb62203
123501bc4ca36d61c39c92eb4153b898184039fc
6779 F20110217_AABYHZ smith_j_Page_086thm.jpg
8ffcebed152e5c14b42c917e9062833a
83ce8aabc10d98fc025565b5f04c2b2a4cd7bd4f
15148 F20110217_AABXFL smith_j_Page_161.QC.jpg
8105fbe00063625a2cf9432b9c431f5c
c369be324af09e38b95f4d90a7a62b8a9c4f26e1
1566 F20110217_AABXEX smith_j_Page_074.txt
be9d7090ddedbc0322be145047a56074
f305bf449803be0d024a9f957384451119f35e70
29797 F20110217_AABYJC smith_j_Page_042.QC.jpg
8fa2c20230e449720dc76dbdcc3c7bd9
3438f8abd92519c03f74b8e04bda5c17772fd984
4922 F20110217_AABYIO smith_j_Page_152thm.jpg
b7c58e4e94471c535ac092ceaff37268
2522eecb53d27c055cda754be05389d00c1f61f7
517165 F20110217_AABWZR smith_j_Page_113.jp2
86e3e7d02bc0ef2d8ce8d04ceaadb1cf
e899aa38a64cae4e578b66c820155959ac898f25
F20110217_AABXGA smith_j_Page_011.tif
fc57a81eba5ac2728394934604a90031
ac16decb27dc7ca5de9eea89cbe3f7ebdf17841c
13195 F20110217_AABXFM smith_j_Page_114.pro
689e1adf064dd8f9e07da7276f3d7e72
0b8f7c741fc4cc2d8ff7daed947821e8663fa1dd
1267 F20110217_AABXEY smith_j_Page_153.txt
6b7f57077a856d0869640622cbfbbad4
5232dc6043083cd1ef0f47698fa89e10cd30dfb8
5555 F20110217_AABYJD smith_j_Page_011thm.jpg
c16cb8ebe02afe016607f3d1f1ea2eda
ac6421b09ece31693005165ecde47e2ddf09b6a6
29790 F20110217_AABYIP smith_j_Page_038.QC.jpg
a40e6cea065b6c3aeb8afe3589fe1707
830ca2b81a2fc41950a3967980d2fafb3b9b08b9
F20110217_AABWZS smith_j_Page_161.tif
4f41bbcdf267b854da74e3c67aa4dc35
3fddd76608648e3bab9292c5d2014350a1a7cad2
20481 F20110217_AABXGB smith_j_Page_152.QC.jpg
04246e40ac205e9e31879089e69decf9
cec9bc0698a016d1a7660ce9692f972a98498cf7
34520 F20110217_AABXFN smith_j_Page_094.QC.jpg
295de48ce8378d4983d8fc0162c4f0a5
b574778e9bc804212d604ab3740987f7a875eb0c
6416 F20110217_AABXEZ smith_j_Page_104thm.jpg
d8ade5abbb739b872e7ec3bf9fe781ff
9a3bb653a475ce334d368363dbc3ba5f009ad660
38997 F20110217_AABYJE smith_j_Page_150.QC.jpg
f1a7b3e296df740f26638e1f035ddf18
f19d15033eecc1284bce61a9b7fbeeb7f59a7c36
2457 F20110217_AABYIQ smith_j_Page_001thm.jpg
a90c40af1f70a2ff9c3b5f5c86bf5a6f
2e62aeebf69410147001086ad9a11445dc2dc662
581170 F20110217_AABWZT smith_j_Page_118.jp2
9ac92fdc57ffd16e2750c2b9df77f555
3b5fc9dc3c2cd80df0c9ce6785fdd5caca4f562e
1911 F20110217_AABXGC smith_j_Page_025.txt
cb2c97a7cba4ceedd998ea7052f0932c
a83bd528e00859177a4a89a606bb2e763e6ffcf0
47933 F20110217_AABXFO smith_j_Page_064.pro
2198563b8b616283e75045734946193b
805df74948914530883443bbe121c3c3baa7c2d0
6442 F20110217_AABYJF smith_j_Page_012thm.jpg
d8f20e6b6f875c2270dc5caf58f71e44
85e90333a691003e2272957ad1a63baa208dbc4d
8044 F20110217_AABYIR smith_j_Page_158thm.jpg
9f05375bce04281b98d6bd12a4307956
eda891be6b6a6f589c9a98d233af40555028c176
1026 F20110217_AABWZU smith_j_Page_024.txt
28ece469544eef362f24c40e0a0706f0
aea63aec840fc5b00a8b111e46808581406793f1
23470 F20110217_AABXGD smith_j_Page_047.QC.jpg
e70b1cc2eafa6c45ce145da45d8c223c
431550bb2ccc34d6f416a87c7ba70b13f0ed0fed
F20110217_AABXFP smith_j_Page_002.tif
406391a6fbe1c56f7bb1d1c4355ca839
c46e0e211d0249ad80e7cca9d2b497a5240d0bc9
7903 F20110217_AABYJG smith_j_Page_091thm.jpg
575b7f3eca6ce253f9b60c6513f6e034
2508955ae9f37ce9d93da3af85267f1d7f611c7d
7142 F20110217_AABYIS smith_j_Page_003.QC.jpg
24d6b88a54acdeb7368b33483ad672bf
e8ca8d522f28b71a09f9e7a6331b2a08b1862400
49367 F20110217_AABWZV smith_j_Page_063.pro
8aa280accfc3d4b21b4a34cf8d5375c8
504a425fe2aaddee4c5a6a658085457cf3830cbc
1946 F20110217_AABXGE smith_j_Page_060.txt
016b9a090dbd93d62e3b90fcfce8fedc
e9ef46b8d28bb712688c3fff4a255ef0c64722ba
4392 F20110217_AABXFQ smith_j_Page_006.txt
f441c1c1bb0b34fecdb75f9739063dc0
15425e784c8e8a0c51d90eef826ae2b554704156
7556 F20110217_AABYJH smith_j_Page_014thm.jpg
23a77cea938f18f7f9c6b08bec5f4f79
308ba87629da778cdd717bc8c81d663264ec46d0
27893 F20110217_AABYIT smith_j_Page_075.QC.jpg
4986cfbc0e8bd32ce659c2709fcc473b
9767ce723c4a8f1fea6ef2d10398a850df1e96ea
882 F20110217_AABWZW smith_j_Page_140.txt
bbb6ab35f3c6a469962f9f9a927e9cda
b282c5da05326560cb7a27eeb19d42c4c8af47ad
602921 F20110217_AABXGF smith_j_Page_082.jp2
cc4c457fa4cb0a35f5b27c12f0116aa8
97fb2dfb59702b6925afdcbace88ab7a0507a8a8
42561 F20110217_AABXFR smith_j_Page_080.pro
16f2484ed842016f3adfcc8fcb99a117
fbd741bb5d04a71433539c99d0118341681cc8d2
6022 F20110217_AABYJI smith_j_Page_114thm.jpg
0cace3e300c203a6decd0fed6e191013
d3f6b8ba59f8331d0363d72367be913053db3000
23539 F20110217_AABYIU smith_j_Page_024.QC.jpg
beccfda47f78e55c10fc7727381ff6e4
0cb6d4c72351fd4dc426760b7dbacd92c75d15b2
6775 F20110217_AABWZX smith_j_Page_047thm.jpg
ef9bcc70acca574ed0894f873a949e96
28474d6ad3627c47a5b174d571af58177e822dfd
39114 F20110217_AABXGG smith_j_Page_152.pro
d89b70c7d57344d01346901a76d16547
047460ea49648e52cbcd3dfef35e45a1db89e017
F20110217_AABXFS smith_j_Page_148.tif
1faced7124e040a15bcffacd69c5a10a
b483ac35e85b174dd5b3c3b96e2f049fedcff059
31719 F20110217_AABYJJ smith_j_Page_019.QC.jpg
d9a0517984391c1610694776617071b0
f38fc7651d573dfb62fb1baabc22d04e24e307ed
24563 F20110217_AABYIV smith_j_Page_048.QC.jpg
f4c03ee8ed2f8e861c908e85a92048fd
0a435546628aa6e2d399c23011a5d7d8148280ca
7983 F20110217_AABWZY smith_j_Page_058thm.jpg
03020c330212bd14eb15b933270043f3
4f2e63135d60b04bbe15dc84c05d38495019384e
24585 F20110217_AABXGH smith_j_Page_039.QC.jpg
4e268db3ca94706cd72e6c3decaaced0
16dea5ae90501279705077e43d00dc1bba414e7d
F20110217_AABXFT smith_j_Page_122.tif
f77e4cade269f79d7925daf321095096
25f2657a44cf485ecb3a7b2dda41c23d45ebd609
7172 F20110217_AABYJK smith_j_Page_022thm.jpg
ca055888489cc0d35d4aef61de79d3f4
deaf79971c3d16254a9b2a329f1f2c30a00d3f0f
25030 F20110217_AABYIW smith_j_Page_036.QC.jpg
da8f2def37c0e0e51783a13e3b2142d9
ee4aca3db58a7be475fad7d210bda1084117c545
F20110217_AABWZZ smith_j_Page_149.jp2
5a2d67736f7fabac9edde694febcd8b9
8df8c0a12e33cf7bdcb892b99e93c5e7a553e965
8562 F20110217_AABXGI smith_j_Page_026thm.jpg
adc9205a06e4d9d4af2a5df0bd4eb004
b46223faa5ec6056b8aeeb9d48676622fc9d2257
5097 F20110217_AABXFU smith_j_Page_005thm.jpg
6cb14ed3f23a7f474465163e88a4639a
ca55fd7acb39fc96f5de2b808874d9aa795d6cb1
2404 F20110217_AABYJL smith_j_Page_013thm.jpg
2b174c1373ce1df8a283b435ed2deb5d
82699bc4a1b9ef92d588fb1c656e86b4dd970d27
7612 F20110217_AABYIX smith_j_Page_089thm.jpg
d5d9321fa593287a3e1217d150b19d0f
ce00ea9a6b79c4034ec1823ab09d7c8c68d9fa5d
2251 F20110217_AABXGJ smith_j_Page_083.txt
ead730eeff4e1d070297ae6788d85859
45b07968f25feefe29990de442bdbd68043a707e
947485 F20110217_AABXFV smith_j_Page_078.jp2
529b5cc0c004517b88d52d01b6722012
0d0093870479793a5c5361673e690e1814e9e6bc
7786 F20110217_AABYKA smith_j_Page_073thm.jpg
ff01e56830a21dfd42900d78901471c7
3140fbb6c96e6087f3caa09342efe9378606c9ea
19061 F20110217_AABYJM smith_j_Page_113.QC.jpg
d2de7cb9f76631dda2aaa69c203f3f34
e5c8cb0073a671ee30fc138b3ca27bd29caadaa0
7427 F20110217_AABYIY smith_j_Page_109thm.jpg
4789db5fbd9691b3de463b418bd5acf6
c11061629b7c4aa1cc1d9bf030ae60e56e4a292e
F20110217_AABXGK smith_j_Page_153.tif
c3af1756bb20c03c76cf8e4070825363
557f1fc23015f22ecb01a0fce076fdba172ba669
1723 F20110217_AABXFW smith_j_Page_042.txt
42f9f2e43020adb335a2db91761373c8
52ac54e3ec1efa25f9581b89529e9c0d44b870e0
30560 F20110217_AABYKB smith_j_Page_050.QC.jpg
4835bfaf73475a3724259b35f0c6db01
60cb718c6be4d37701bba57f397d6011533ebd92
38371 F20110217_AABYJN smith_j_Page_147.QC.jpg
6454edf9d8015be8b593c95966a26c6d
77a71234251ce230191dda5a130a62c55f5d6cda
1659 F20110217_AABYIZ smith_j_Page_002.QC.jpg
09b99b9efa13ee75f75900047bc78a1f
3d34d7fa5c9bef5267f35960cd768ee3de803e08
17411 F20110217_AABXGL smith_j_Page_081.QC.jpg
4f16c6a31be3553c4f42e1319b276185
4e7cccfbdb1dc2d0cba352c08d48dc41ae1459d5
17808 F20110217_AABXFX smith_j_Page_118.QC.jpg
ad3031700a0ceae589d487b7451a0e0e
4509da883354d4d678bfa22ab1a8a35bae1972fc
4794 F20110217_AABYKC smith_j_Page_136thm.jpg
5a759145fa5af814092b4be461cbd3bd
33018b8125b63c00592a9dd2aaf34fcdaf4e51cd
24232 F20110217_AABYJO smith_j_Page_076.QC.jpg
e0b8c71a28cfcb6c446ae2af9b9f41c5
56dc1f4d0b5320b12f0d19018e0390c586a6a030
8836 F20110217_AABXGM smith_j_Page_145thm.jpg
6800cf87593d80a5cf396484f45312df
07a815ad5567369171f45d48732d86cd4b022bd9
90465 F20110217_AABXFY smith_j_Page_090.jpg
48b8f085de5f2fa3df11b5bba5cc461e
4aed901ed005ffb0682c908e97788d70c72ebf70
F20110217_AABXHA smith_j_Page_150.tif
8061e22beca38039f89c789887093cea
f3c3eb3c8253ecd6f46229d5e949a10f8a25f4da
18302 F20110217_AABYKD smith_j_Page_107.QC.jpg
49cc77ff6b9272254608e5e3b0d67b3b
30ad3d95aec4b0a79e2bb28c5635fe5b61c1e8e1
3276 F20110217_AABYJP smith_j_Page_065thm.jpg
7739b27fb062c80874deb903ce7a4f9e
dee30f8474c005d2c9a5dd89d796a2d06653a4db
F20110217_AABXGN smith_j_Page_049.tif
dcf6898f25dd4119204f00181c65cf74
e11332e9eb26cff4ca4358d2f6378b582d7da837
69909 F20110217_AABXFZ smith_j_Page_129.jpg
fbdfd80cb3538ba9863e70cb456c1616
ff5c936985c84f7b4df49afd6ef5be8eae502c93
19523 F20110217_AABXHB smith_j_Page_117.QC.jpg
982293db0a55e6608e29877acf97785f
5877255464e50d76c91ead35a412b3a902d895a0
4752 F20110217_AABYKE smith_j_Page_128thm.jpg
9259640fa073225950c8f3e8f6c07bbb
ef8f13141ef3ff7ed38ec7cc9b24f3a99fc1bf62
8201 F20110217_AABYJQ smith_j_Page_050thm.jpg
42b9b53f1dd80206875bee166aa7a556
6a7af57a277a1657364ac2fd26df86a47e551d6b
8935 F20110217_AABXGO smith_j_Page_144thm.jpg
243737e0fe04108a5f73ee3fb0c952db
51a57826c00764d5d5d964de902f412fdeb5f7c8
2036 F20110217_AABXHC smith_j_Page_059.txt
2bf68882a3bb413c0e284934e7a25057
07aa6ad2d044554f094c8b14645e1998c01720ce
29387 F20110217_AABYKF smith_j_Page_014.QC.jpg
97892773d5a3de0c07714f5a4681f424
57515550d5d368ea2b09e8f507c854ea60694af9
3543 F20110217_AABYJR smith_j_Page_010thm.jpg
4f4a9d0ba5319e8c3f29c364f2dfd563
0e7635fe22feb9a4373829d4e02e808f2e6b090f
31261 F20110217_AABXGP smith_j_Page_131.pro
1f05e3698cedd29f2fe3bca795c35176
6a0ed51eb12f07bae6644d254cb7056ec9cd8f4f
F20110217_AABXHD smith_j_Page_126.tif
f8533e0faeb594eda0a387f6ed3db533
5e87601d25ed74920146454bad3e3b3b38a5e827
28809 F20110217_AABYKG smith_j_Page_069.QC.jpg
1ca3885c01f74d8ce15743f8539bd37d
5d6387c8ce30c4c2baf1fe496c2e0e38f3a6b777
25334 F20110217_AABYJS smith_j_Page_057.QC.jpg
dcacdd663159c88bd61d532926655079
ffc4aaafbb1a54023f94077dcdf663146f1b6b0e
794 F20110217_AABXGQ smith_j_Page_039.txt
849b14f59e95acd48dc88803151b9fee
055efeb6e6536f8450886c034a7a998d760d49b0
F20110217_AABXHE smith_j_Page_070.tif
26796d606f61db1ef7431341f4624804
cdac3dea19d77c5f1b94ebedbf5526919a4433b4
8203 F20110217_AABYKH smith_j_Page_027thm.jpg
4714d28f24bf3b840682ce64ebc3fec0
43df5797a4350e8f0a606db77525355542327c37
17215 F20110217_AABYJT smith_j_Page_132.QC.jpg
1a502e69f6aa2d5ca3f03e3f5e0e29e7
1066e0e176b2af8b3e6b52e31aa4bb41f2a9ec54
7488 F20110217_AABXGR smith_j_Page_075thm.jpg
d42c3fef2389947579d8d768cb6bd2e3
7e4b8f8b41e89e0d691735a864a3ecb2d4195fd2
19113 F20110217_AABXHF smith_j_Page_082.QC.jpg
d72e7614a32fe5f6db5cd18535e2b170
f95888119c9bbbf09bf96499c39d9385cfbab554
8165 F20110217_AABYKI smith_j_Page_155thm.jpg
c20a17359afff9fb46103646a5562bf8
6ee046fb58591b33547159e48997d1272cafa195
6340 F20110217_AABYJU smith_j_Page_036thm.jpg
e1e7ca72c0358be0f6af2c496fb22387
192cf1e0579855efe2f46382a128920c841357d1
17255 F20110217_AABXGS smith_j_Page_128.QC.jpg
fce199b555528bbb8f5834fbd0a88f21
531fd436d1f07534a090eaed70361142b848d05c
80149 F20110217_AABXHG smith_j_Page_036.jpg
61897d55e8112dd9b15100369520abc1
b0a4b182cf5d8697440930dfa11a542646743c53
7357 F20110217_AABYKJ smith_j_Page_121thm.jpg
7fad80453f689d75180e1df92eea83fa
ac365a116bc18e70c741b2d038443cd171342059
24082 F20110217_AABYJV smith_j_Page_023.QC.jpg
17ca1754ad24e42b7f98bb033ad4982b
79aee9fe447221c4147faef03b8fab5d2a79688b
5028 F20110217_AABXGT smith_j_Page_115thm.jpg
a8313f082580c8007e0adcca1d0dc138
483c9a854749328af6d758fd31466791d8de6b75
2191 F20110217_AABXHH smith_j_Page_125.txt
c2cb399ab560de6d0388db7c7c5649b1
76a5fa4434070f4f86234727c90a4f537983f113
29016 F20110217_AABYKK smith_j_Page_043.QC.jpg
a687b12513ddc82c76f743c0eddb6b5f
f57e57f9b8644f0afcec7588c37ba82572348c57
29226 F20110217_AABYJW smith_j_Page_066.QC.jpg
f5c5fb3b96c76670ddded56e80532d4e
f881956e0f00d0a00eb6f35a3817eee7996f3ac8
27521 F20110217_AABXGU smith_j_Page_079.QC.jpg
580abec5ba815d97788bc941ed2f9aee
ecff1aeeab9737e1bef4d2a664c2f51f550cf070
2217 F20110217_AABXHI smith_j_Page_126.txt
d1a79a8e6940867f222a1c00376c318d
edd68e3c4a0a5c51de52f349d3a807e0ff83b603
4851 F20110217_AABYLA smith_j_Page_137thm.jpg
3b2a146fa1244daf7c889f9d6c517274
c1337fb814b91b9bef13cf5b04c6c4f4214c7315
17451 F20110217_AABYKL smith_j_Page_122.QC.jpg
6717ba95e8275ba68550fbf426f84fbb
a482237faa42ca76e3f9c6608a587a9589aa3199
7740 F20110217_AABYJX smith_j_Page_068thm.jpg
22cb0101e8150d833125e475c8f60baa
8de2559198e0d38610d2529dec37b08cb4e0e7b7
81740 F20110217_AABXGV smith_j_Page_056.jpg
cbc29f5a38d34df5eaa1a6048fff90d7
b9a415e6232ab0c11521946249f3ff432a6cc0e7
861487 F20110217_AABXHJ smith_j_Page_062.jp2
82a66d4ef2727ded337a069bf5b6f956
1ac224fb4cb38b6d5b25a4e8cf67d1b8ccc1bd26
17636 F20110217_AABYKM smith_j_Page_114.QC.jpg
efa68fd91bb37522524fefe08046d288
2dd6023eabff972d3b2e1f0e6136cecf2e8f5874
6049 F20110217_AABYJY smith_j_Page_076thm.jpg
da27c144fad87fca2c98fa8fdbbb4cca
6ac1dfa5dda67a20957b0e8c8e72dc9a863760c8
30093 F20110217_AABXGW smith_j_Page_002.jp2
523104259e364da9101808fd8c9b7c48
122c50ddb3f22cdbd5a13843ba8c41661aa98d5c
8490 F20110217_AABXHK smith_j_Page_094thm.jpg
0b280f276f6b65ad21a91ccb22722592
00a5698a66ff7c8f9a70155b52810bcb56c0b4b6
634 F20110217_AABYLB smith_j_Page_002thm.jpg
e0c8a0b67a6d1809b4dcaa652a54bf2b
9263ea436b1d508aba334863ebf5ab982b156a93
18075 F20110217_AABYKN smith_j_Page_138.QC.jpg
b7f285a55dff685b0ffed7240929eb24
0ba134fcd385ef4be89caa7fb068bc8b32f13a73
7972 F20110217_AABYJZ smith_j_Page_071thm.jpg
7804260f2a9c0dbdf29afa80b4dcf32c
fc9cc89ebad816c681d1469744aebf55898745e4
56119 F20110217_AABXGX smith_j_Page_082.jpg
984611324b656b09bbd7242889d209e3
f06e8d5162e41ad7f4afdca7fd07c000f2aa3783
1621 F20110217_AABXHL smith_j_Page_079.txt
78ad2d555a3724c287e87b378bbed08e
f30e55e11b63d31fdc8bf72d272f7012d52916fe
14493 F20110217_AABYLC smith_j_Page_010.QC.jpg
0cb93506706721fa1d9c9c5e8bc1ecb1
d0d70e1de31aaf0459d997cd3f0e710041cd09fb
24493 F20110217_AABYKO smith_j_Page_135.QC.jpg
aa03c679b9a5f9adf8d63c49c2ea7dd9
0d208817992ce9ab3c3ce94de09658c507ed123f
707 F20110217_AABXIA smith_j_Page_031.txt
c324ee65c202ce82012ecb62a8c7ba97
788cd49f7927a1e1cfa141036b1b69f2a91e8701
742549 F20110217_AABXHM smith_j_Page_051.jp2
c8aeaca0a5aaaee54351de77248c72ee
eb8f9d2d7e5654854d4265b09ffb3228c56fbf16
93284 F20110217_AABXGY smith_j_Page_083.jpg
62373d744963cb7503c153f67e6f4bc0
8726e151ea6a60319bdae705d2be326a4ffe3936
8450 F20110217_AABYLD smith_j_Page_070thm.jpg
8d8e377dab34b07c371a9eb915c32c8f
37835d12f2b244522d3df41b0f47e973bdc298e5
21771 F20110217_AABYKP smith_j_Page_011.QC.jpg
532acd98a89a94eab61a6cd7064f43e6
5fea55280a5f7e3c5fbe0c17a547c43add72585a
15060 F20110217_AABXIB smith_j_Page_108.QC.jpg
567dbb8ab45bd279a851ce435d78d3e3
e043e5be49276b57cbeba638a575859fdaceb602
F20110217_AABXHN smith_j_Page_112.tif
d43063ff5d7feb5e9c88c5920566ef72
8458e8ee00cfe5e535bc94a11204e8b64603b197
73238 F20110217_AABXGZ smith_j_Page_134.jpg
108d8276e1cf8a53ea4739e2818de298
8c6e4be109882f2daca2c7091688a4893b2a2ed8
19121 F20110217_AABYLE smith_j_Page_125.QC.jpg
a179e8b5ca219b92a7999b539b7918a6
d0c74c6dd277e8fb00509496caa69cc8ab5f8bb4
37495 F20110217_AABYKQ smith_j_Page_144.QC.jpg
a3e92e41638f073f5e35b574a350345c
a650ef9dd3570dc3e3a010f72d3373fa5fcb2106
F20110217_AABXIC smith_j_Page_103.tif
18794706f555f1afebb922a6d415cac0
d7a6eecac10dc85cf17c099984a6e6089d2cc462
F20110217_AABXHO smith_j_Page_037.tif
b84a75be2dcdaca699a84bc5d02b5214
9970b8a2c4cf1fb6fbe7e90127ef57aa8f012588
6731 F20110217_AABYLF smith_j_Page_079thm.jpg
8c0a206c01fc324098670ef80e4c7308
60488cc2588ddc36ae0333ce12016134961b2c32
F20110217_AABYKR smith_j_Page_092thm.jpg
3e3827f9b6b85dad1ea9fc7192465f46
007a54873f681bcd494b17c3af6528dece94ba17
F20110217_AABXID smith_j_Page_097.tif
93300900c584a09c73c599bb104eb884
42715d66f0cbe1b3f8ae98647af67b1fab18468f
7514 F20110217_AABXHP smith_j_Page_113.pro
8bd26b20c6710c85e5c772dd3ecc8f7c
092439cd35fd494c71293283443be0d8c0aa7bbc
31461 F20110217_AABYLG smith_j_Page_154.QC.jpg
e45bebb58eb492ea1799dbc0df685b47
36578d90b2e48e2119d1c46d9a08d67d0058643b
25426 F20110217_AABYKS smith_j_Page_054.QC.jpg
fb59771f5e3a598b5f0eb10771bd047f
a7bc07f3bf27b8c876db85612ee442d9916a9911
F20110217_AABXIE smith_j_Page_129.tif
7e3e44bd8d24e2cc07757da02940814c
6498bdc9f03e74dafcf0e01895ff05a12be5e460
6837 F20110217_AABXHQ smith_j_Page_057thm.jpg
b9be511751db65034a776cfaac3cc0c7
d535854dec27f07f4099ecd061a5183e7a3370fc
34632 F20110217_AABYLH smith_j_Page_027.QC.jpg
7dbbfbf6b01cb3b4050494c6c2c5c1bb
6634832f1c6b75418c70b6a9215a53d6bc730921
7962 F20110217_AABYKT smith_j_Page_037thm.jpg
eb93ac49dbd1518c4a3fcf081f89a771
8e6b9c650d4c30a56deca99b77b86067d0651905
5668 F20110217_AABXIF smith_j_Page_082thm.jpg
8c56fd1188f1200d993175d9d32ef904
8d26a888e3d9eb19565ea2e1d936b9f969fac013
76854 F20110217_AABXHR smith_j_Page_092.jpg
953b668e0fdd62d5bb7d17048383e878
40514107240647f315744e7538c1a67f217e9c87
258721 F20110217_AABYLI UFE0015200_00001.xml FULL
55650fd11a23459f714718706664b69f
88b04ce5c65c05c54275a001d6610645a6979365
6133 F20110217_AABYKU smith_j_Page_141thm.jpg
91298da23b0ed2b7d3c53e36c5f342ba
e60d6b84028f87b8e372e1779b76ef029f392735
105206 F20110217_AABXIG smith_j_Page_059.jpg
7b6b45465ddb23759a0e125724e796ce
82667393d347941f0dc832a2aa0913ec9b82de46
941490 F20110217_AABXHS smith_j_Page_023.jp2
a36e9d666efe15c5dda88fb0ab5bc6f3
2dbd3feab7633b4244db27e4630e351c7cbac236
20358 F20110217_AABYLJ smith_j_Page_008.QC.jpg
ebdbe46613ca07f49553f19e5400c3fc
6d8e39215c986b3374f48c4c77d0a71a463829ee
22873 F20110217_AABYKV smith_j_Page_109.QC.jpg
0378f9882e5e01792a3ef08b5beacb0f
8b90ea82f9aadc9d8f2ef355a658b494d0484149
8227 F20110217_AABXIH smith_j_Page_015thm.jpg
a8da9bffda9b51e54ad401112578a9ce
24cf1fa3797b03f9ea66191330c5c57468a9d42f
27750 F20110217_AABXHT smith_j_Page_080.QC.jpg
a6ad97926ba46b1024548c5ec228335d
94a01590e4b1550dea9a72ce91f77db5e2ed543f
25378 F20110217_AABYLK smith_j_Page_012.QC.jpg
39427067be6fd0cbccb771df4e982b5f
c344edc596f75c84c867bba95f12415e832849a6
21992 F20110217_AABYKW smith_j_Page_140.QC.jpg
cfca959aa9da61918aa5fce53731ae26
9359c0872a7363edc511411ea954d5f1e9a40fe8
44905 F20110217_AABXII smith_j_Page_058.pro
821e860785c1837ddeb8d4c7902d0608
8ab37ec87e27e72df0f85cc2b92af098c093047d
500501 F20110217_AABXHU smith_j_Page_116.jp2
88702d08f428993d1c157200c2b0cbfb
a9de295d1ce1a58a6401f43a43a5b72aa677e060
35446 F20110217_AABYLL smith_j_Page_026.QC.jpg
8ca6d98a501d66017cc1e7b9955ab45a
b424244859d846938ff6d73b13190189aa02b179
7298 F20110217_AABYKX smith_j_Page_066thm.jpg
f0573bc1f550a77a38cf68a5a77898b9
9c1e1cd224c45c93df9c915297a8a519621976af
66505 F20110217_AABXIJ smith_j_Page_009.pro
94123d3f781af8e7ab2e875bb7d4c907
bd040b5f0ee0105f93f41a374cd21c2923c730b8
1051953 F20110217_AABXHV smith_j_Page_144.jp2
b517dfe6bb58020e9ac61619e4f8ab8d
51935861d74ab77d06352fcc0aab36a3594d5624
31212 F20110217_AABYLM smith_j_Page_029.QC.jpg
fadb514d4600352ee5d16d8e968ef61d
24eff12f62c337e97241a7dfe73c9c38b0f09e22
20257 F20110217_AABYKY smith_j_Page_110.QC.jpg
0aa4b1f4d99c5db79c0ee4af9c36cc21
73b9bb4f2f7902abade8e4617ae15b6f26d86085
32511 F20110217_AABXIK smith_j_Page_125.pro
54e8b24ed5e296e6842fba5575b756de
02429776d1492d16bb189bd5100b366c38c0927a
28024 F20110217_AABXHW smith_j_Page_121.QC.jpg
ba060e1741e8e6abe63a447fee5abae1
b3e5d6e36c0b53d4115806936b5c588e737da447
11244 F20110217_AABYLN smith_j_Page_031.QC.jpg
4fd5b8eaf4ae8db5d75aa777453d6652
53c14bafb4109162d5265608edcedc1c80a8f295
31924 F20110217_AABYKZ smith_j_Page_096.QC.jpg
acddd617001ddacb51042ae235d68e77
47d02523814ff324164729d700f00883d17f1ac0
32500 F20110217_AABXIL smith_j_Page_158.QC.jpg
48782a63318d7a03b5963b541a5035fb
430f607697ae6eeace26e856f2e0d5ce2f6176da
8700 F20110217_AABXHX smith_j_Page_151thm.jpg
492f05ddd51f8659da992dafcbf22976
665bef68494a42dafc88ca713e390f6989e3215f
30602 F20110217_AABYLO smith_j_Page_044.QC.jpg
5053648ab954f2c3ad06367f8ebed124
93ff798fee6a8a2d21afde1304bde3e8bc90b07b
5256 F20110217_AABXIM smith_j_Page_006thm.jpg
3d04034108d106ada52235c0f0f52830
4f35dd0ae46dc83790908b46a41789264cd1cfa9
8312 F20110217_AABXHY smith_j_Page_060thm.jpg
b088428d40ec1e01a530c57d109184ee
f2c9245c9e5c5af6f933234323e46070f79eca38
89315 F20110217_AABXJA smith_j_Page_012.jpg
b2bfa86e0962b13398c0df98641d182a
4329d10828ba09efd45d14177ad3b4397dd23ead
26780 F20110217_AABYLP smith_j_Page_053.QC.jpg
51fc63e226242320e9bc9c1a4b49cf4c
35a1954e7a990988d5937a421f2d6850bc97874e
41319 F20110217_AABXIN smith_j_Page_090.pro
5491472be3a53577c2dbea878641b982
8f6e3bb95479a64debd541c3e9f47e6c98b402ef
1873 F20110217_AABXHZ smith_j_Page_048.txt
d00d07ccc62600216224cba0229d4613
0a053b20e6d3cc5e31d1997beca0b0f1d3df568a
34318 F20110217_AABXJB smith_j_Page_034.QC.jpg
994ef6720eabfb6631cdf6124c72131d
b65d419ec6143e8d585331dac5b956e90e115ab2
28712 F20110217_AABYLQ smith_j_Page_055.QC.jpg
b36ac254727bb42eb73893e53c6ab2e4
fe836d02df93d5d69c6fe84bc800229d700d080e
104842 F20110217_AABXIO smith_j_Page_046.jpg
bf2feb32adac4cb954374b54ff74c482
bca37c396bda70b502d805fecbad3b90969bdd47
17842 F20110217_AABXJC smith_j_Page_141.pro
78e46d8ae6da740d03cd29f924c303db
85bdbeedcb1a5f85ec0603b61d8de0c470f0a9cb
34243 F20110217_AABYLR smith_j_Page_067.QC.jpg
354a9f5c330a9b3750330973b334d997
6bd1ca546cc552697fd3b25ca6199fa668a944dc
2032 F20110217_AABXIP smith_j_Page_116.txt
f40c684865e8b85c056fae526d3ac6de
1b977725d97c58afdd8d87c2eaa65f427e5152e1
8236 F20110217_AABXJD smith_j_Page_064thm.jpg
b8933db079e1b3c6f2302fbec303680a
64dedc823b977a5f922653c71442b0d2110d42e2
33043 F20110217_AABYLS smith_j_Page_093.QC.jpg
3113107bfb0bac6e1a8587ade3705a9d
0cf855079a6eca9f161b13eb076018839a0eaa17
F20110217_AABXIQ smith_j_Page_101.txt
36fbb506671ebfdc75b746f1179ffce3
69465b4674f2c45afa1720b2fe2a6779198e469c
39373 F20110217_AABXJE smith_j_Page_055.pro
c2aa1c7cb5b6a9112c3c1a946f68ca6d
518f4ebb353a62fe1f102c60b7c6b3c5434b86da
21858 F20110217_AABYLT smith_j_Page_142.QC.jpg
cb687611a9f03f352ff0328700731bdc
3e997229acc65cdf099a312a226074fa11e2d18b
F20110217_AABXIR smith_j_Page_073.tif
ad1fb41e3ca8749a159535ae4297f3d7
403981b9fb7a6008cd785126c2bf4fef1ca5ef38
2094 F20110217_AABXJF smith_j_Page_138.txt
13c0a2ac228a3f77854789c4d16aecf8
a7c458be2b1b95e16513f4a3218a2cbcfa342457
31754 F20110217_AABYLU smith_j_Page_155.QC.jpg
d4c07575f18cf44e97c8e82eb04c3a0d
f8490148428373fdcadce3bca666dcc2d77faa38
1050879 F20110217_AABXIS smith_j_Page_037.jp2
83c742c2db28f0fc393e640ac4d18cff
6bafa2b2a71be1a88d5fa8dfba31c6451d254b06
1051938 F20110217_AABXJG smith_j_Page_033.jp2
cf2d1b7c4eb539b141971bbfd4be13d7
a8305e6ba53b2b679b120fc7afaec1c682f43564
34302 F20110217_AABYLV smith_j_Page_157.QC.jpg
62eb061244889a38a9cba91c77ed3e6d
cac0ee3ae0865876b2c51ea257957a7f4d2b3dd1
F20110217_AABXIT smith_j_Page_094.jp2
bad2aecd308cd92024d0162e1d56a6b2
f333e27fddd9f2cb51c524a7add240e1645c6df3
5327 F20110217_AABXJH smith_j_Page_008thm.jpg
9062afe11c3f43b44038a6eb10e06a7b
6c7144ecc20efcf49e6174eebce7ae58b369d07d
8107 F20110217_AABYLW smith_j_Page_020thm.jpg
a1b5562446c7cfaff85ad408b7e15261
d19b22df77e5332a58b22a355582900093bb14f0
F20110217_AABXIU smith_j_Page_058.tif
7f75fe2fc7822e8ec43e0f80278066ab
ba6a77244ab792d9c148e88ccdb6266eabc435bd
514474 F20110217_AABXJI smith_j_Page_127.jp2
7023bf2f6fc6475c2787cea4485a5120
9c8ce21f57b457d1a102b2c95956784e7f9231f9
7375 F20110217_AABYLX smith_j_Page_069thm.jpg
c54320700f4ea4290ba14a6901b55b12
3332ada4fd8caacd93f94d37683d7c617ea83d80
F20110217_AABXIV smith_j_Page_128.tif
91c9d61ef16c874225a2db613ff861ae
2a242177d583a4b24ad0f3010e16299c0556aaea
F20110217_AABXJJ smith_j_Page_081.tif
c21f2673c8c680c63221be872b9e0e80
e7d77b9f50189bfa4e27ad6554c97eab3afe690b
8021 F20110217_AABYLY smith_j_Page_093thm.jpg
919aac395e7bdf81be688180d079ecc7
12fab8cc05877e570644a672bc186803212a018e
585833 F20110217_AABXIW smith_j_Page_141.jp2
c001b4a2820d748574e5b5d0164f2291
bc3c81af8d894a326a9a7637ab00c9f149b6bcc2
54095 F20110217_AABXJK smith_j_Page_155.pro
c7d55672afaf518a2416e14ffa5cd12c
3738ef42b0ca3e8842b556bbd5ec0b427ce5a359
7889 F20110217_AABYLZ smith_j_Page_148thm.jpg
1a567674f068bffaec8159dad00805fe
ef4a4b9f210ea7480e82d79381981fc8e026b025
F20110217_AABXIX smith_j_Page_130.tif
1e5e14285be8a52cd71586265678a66b
aca7b0548476782b5cffe89446430ac08de46d7a
F20110217_AABXKA smith_j_Page_083.tif
6d6d972e46cc08ab484312141e0d0483
1b3d2ed2455894714d15833ac9640f607fcdc8da
887 F20110217_AABXJL smith_j_Page_133.txt
70553bda1368f998764970ed43d56a3d
3450a18c169b4880bcd768486bca47d760b7dc38
7133 F20110217_AABXIY smith_j_Page_043thm.jpg
7df800aa6516367b5ba41393da682dcb
4444dfc2486af695225d70988caa98eecb0007bc
800026 F20110217_AABXJM smith_j_Page_089.jp2
2415191dcd9b9647f82bb992615f4cd6
2b5a4a850bb7120a544267eda1e0a8bc8045cffc
8311 F20110217_AABXIZ smith_j_Page_159thm.jpg
f9ecc1ac6b0174c0644837343ccba25e
b05543d3c71785e5cacab6473eff05354de30956
33717 F20110217_AABXKB smith_j_Page_088.pro
9125212228912ebbc46483a12a371182
fa0ceb5bfaee0e93451f572e374f818c42375fe6
32947 F20110217_AABXJN smith_j_Page_112.pro
3f92410ff8181e99a5f01d6c149a71c8
8b11e6144efac8088fbc04e3cbadd88c52a800a2
662104 F20110217_AABXKC smith_j_Page_140.jp2
3a5f658596a11cda3c8deb7e7b069e90
37f9b9a76336c22bb65b29defdc59854ece22a62
1882 F20110217_AABXJO smith_j_Page_030.txt
112bf68824f6b4758c0cad19cb5fff04
3cb5f8ad1d54829253c262c1dae685db07b42718
2235 F20110217_AABXKD smith_j_Page_127.txt
43a9aa4ab00a108198d1ef0ff0b56d74
4d4023401e6762dcd7678b2e0454a820ddda6a44
128754 F20110217_AABXJP smith_j_Page_144.jpg
ff3afeb4490f759dbf9e2da7703b4f0d
bdfe4e62a8342a27b477f718c689fd0293c59543
5513 F20110217_AABXKE smith_j_Page_129thm.jpg
13e01c31a95dd8d82befd9757e942c0a
de0974443e1f80b797bf1ddaf3cf9693bfff1951
787884 F20110217_AABXJQ smith_j_Page_056.jp2
ce0e6d155137807ebf098b40fc821469
6a20e1bef0e9c342df7e184e367c87e36dfdd844
39404 F20110217_AABXKF smith_j_Page_016.pro
ad6c19278a7a5700846aba15ca79f586
0faace043ecd955e924d5cf6d96e278aff48c3c3
1051956 F20110217_AABXJR smith_j_Page_100.jp2
5ae870eccd3ed7185c09bfb8d318d362
200864b4b7bf19e891e64b6ec104c4498b786a74
F20110217_AABXKG smith_j_Page_036.tif
348bb6c97258df4d0c49aee337e30e04
0e9d14e9d937ad2a4539a33ab7e708d25ba66677
1051877 F20110217_AABXJS smith_j_Page_059.jp2
48a5236cd503dc0a4f3c4e45580b9e04
98697662f83cccc8e29080e0d2c8efdb63ed9181
475035 F20110217_AABXKH smith_j_Page_138.jp2
f1f76ec8d7c8f40c108194c8f3dbb235
030cbc22715138ac0007604528e17b8f977d3f25
59602 F20110217_AABXJT smith_j_Page_157.pro
4525066dedbaeed19c393e2d9666d90e
59cd568a49313dfe461bb8ffc84c073ba4401348
24149 F20110217_AABXKI smith_j_Page_051.QC.jpg
b575204e2496cdccce5abdfdcd35b2e8
e0ee03da6d1c408e5db53d6fe9e13cb54cf5c0b0
16434 F20110217_AABXJU smith_j_Page_109.pro
3e2d3a1fb28b5ee1da7ed8e054a0adf6
0c12df2ae852376828e9ee7672d19b0a3c3edff4
1749 F20110217_AABXKJ smith_j_Page_032.txt
e5e9fc3aff76b94c911103b1a15c58e9
91af028ae7342e121cad26682e73ce367ab5c803
17525 F20110217_AABXJV smith_j_Page_065.pro
dff0004c52e4feae54761609cd26c050
98a4e1393032371496b2aa24fb2867e07f216bb9
F20110217_AABXKK smith_j_Page_122.txt
81a6c0235139c3ad90a06560cd96dc56
1b04772dec439e8323a104c5ec39fb6849ff34c5
1322 F20110217_AABXJW smith_j_Page_010.txt
494a1c1bc6c77659823fe880dfbe000b
1924c209f143dbd4120dec87a1d43c9ae21c20cb
100763 F20110217_AABXLA smith_j_Page_101.jpg
7ca92031e331a91b5e51dd7652626a7b
329228c0e8aea9c1ffbfa72c687580e1463f3942
28286 F20110217_AABXKL smith_j_Page_137.pro
d27b5af6742ed92b1f9defdb9877ae59
15e13502d516d6b85783411df29f8cd4f3cc0b6d
10039 F20110217_AABXJX smith_j_Page_123.QC.jpg
d952df603629ea6c240d7f166999cfcb
88ebb9acbcba666a193c2e9a7b47274116b2a089
1051965 F20110217_AABXLB smith_j_Page_020.jp2
3106fa2082d782cbe046b7466b6f970b
7d50f8086dccacd3369291c7973f79861708c323
35402 F20110217_AABXKM smith_j_Page_031.jpg
fbffbdee23ad99a5d52a4c871cc9e77b
09547b8d242fd0392853a64a57cd050c699cee49
50029 F20110217_AABXJY smith_j_Page_073.pro
a5233a2bc30d8cecbc637713b6d38ef2
a74c78b9faa5ce3052b643bc54d15a2467b48cd7
7658 F20110217_AABXKN smith_j_Page_084thm.jpg
a8921f3ffed3309ce6eef25087969b0c
7bcb272fd96fc11bf7b91b04cf262219da1b7705
F20110217_AABXJZ smith_j_Page_095.tif
44b64ebdd3b7a3052afd8d601f362b62
1631ab476985f8231d35f787fcdccab32059994a
F20110217_AABXLC smith_j_Page_046.jp2
320503eb3f88aeec4678dfea12672e92
b5b9925c797c0709cef2fe09eacc18b5a42b2263
133401 F20110217_AABXKO smith_j_Page_146.jpg
fe1c9a4f3691c548a725217b945fba72
f11dd8601a95d967bc449f8e049d4fbae80f68de
101630 F20110217_AABXLD smith_j_Page_103.jpg
41f8b9edcb606ce22c4eb02be754ae06
b79bcce6384b7582547f72c147d7dc93da2dd21f
24734 F20110217_AABXKP smith_j_Page_021.QC.jpg
8aa261e279b5ec12b05baddb20a985f2
b92f4a38f6947db82fbbe8dab0a1452b8b65b5a2
47518 F20110217_AABXLE smith_j_Page_030.pro
1658734b5db9f2af55a95bd78e290c70
58918bd462f70799a19e87ae80a0cbcbb1445276
F20110217_AABXKQ smith_j_Page_027.tif
d5e6f7434d32c317b4f09b235bab4501
31d9d3d72fa6a992116ee6e320aa65340b587a5b
703 F20110217_AABXLF smith_j_Page_065.txt
c2fd0c67348513c0632b19aaf0b7206d
1fefd0001aee5736a9b6e0c399809e757c4229b0
32948 F20110217_AABXKR smith_j_Page_127.pro
ed4a14e69416db05b6cf6d21f708952c
9ade01dabe9efa977ba9e026cd14f3075d2b5bfe
35372 F20110217_AABXLG smith_j_Page_054.pro
d28ba9449b375d88de6012319d51e294
e250b1a1946547f8addde2e5d0ae6d20b29f51dc
17703 F20110217_AABXKS smith_j_Page_116.QC.jpg
a30d3351a771dde4e7150a170b2a8155
03454a9e46d74fcf079aa4b331deb55708d0bf15
6367088 F20110217_AABXLH smith_j.pdf
662b46a7732c1e9221e04f23b51e0a6a
6aaee0c57aeb99626570d5f0b9052463b7b62024
791 F20110217_AABXKT smith_j_Page_114.txt
d4c60418d87db610e5b30abc738418aa
4a027c9b7ab560be36db014bb41740b648195c04
26931 F20110217_AABXLI smith_j_Page_009.QC.jpg
783bb122ac169140f0176d5cce55798f
6bd0a43cdd11bc7304a3abc7dbec0666c4767f03
8821 F20110217_AABXKU smith_j_Page_150thm.jpg
cd06ccb3f3b34ef5fa64f43afd2cfcbd
b3a0d7026b2f6ff65ea0eeb739ba4cfa1c0f9226
38868 F20110217_AABXLJ smith_j_Page_089.pro
0f78fee3524eb22f0a6bdac2e22f81d9
1fde8d7c739e45c83cf96ed62e0c45dcb57eea4a
30437 F20110217_AABXKV smith_j_Page_068.QC.jpg
6fcc325c8492b7bca37d387e6055c97e
9b5b6fbcbdf5d4452c04db55c89515e850c906de
1051970 F20110217_AABXLK smith_j_Page_157.jp2
895ecedb0b020142bc34a949b7c9fbd5
8d3ff59de70c03c3dd61a980ea3f4ca0d7b223ac
F20110217_AABXKW smith_j_Page_034.tif
68641c0b98338f6085c6bf1d6a462728
7711e31600089e481a2fd5d9c8a220692362b8bc
73299 F20110217_AABXLL smith_j_Page_120.jpg
2b7f04b647f98a5309869b13e8bc1ed1
a77d49fcd123efb64010789cd8469cdddb8c3be9
33540 F20110217_AABXKX smith_j_Page_063.QC.jpg
ce030dd6fb1200334f7c154a3eea7c98
4e6f97a64310050cf316bca99b37004d4536b7df
F20110217_AABXMA smith_j_Page_004.tif
518a5a9ef51b34aee401adaa1c33da72
3defcfaf504918c9c1e08f22781a5df958aa214f
99394 F20110217_AABXLM smith_j_Page_093.jpg
5bd0255119bce151e57edb9ffbf806f2
8bd2bacd58e757a54b58c53d6395416fb770021b
60068 F20110217_AABXKY smith_j_Page_118.jpg
95c633b6407252909f311febcb5c74e8
189383d19982765eb0ba1988f340eb2bf4a7b05a
F20110217_AABXMB smith_j_Page_005.tif
ac092bf7376ece62e97b93d0c03b257a
acb99caf0e0c6f20be19263bdb567a255ce4ea0a
32648 F20110217_AABXLN smith_j_Page_091.QC.jpg
d043a7d669d86d1a53f3108b36d1998a
4062320e091f0fabca6ce2735f5a3dc689be0368
50151 F20110217_AABXKZ smith_j_Page_101.pro
51ab5c89babeb5b067519b73b6d8442c
21759793d8cf214af55db2d535c8c83cc010fd39
F20110217_AABXMC smith_j_Page_006.tif
9ca52098e74a8b3cf29574852ba470c3
d7e45c16d25211675db55dd44a7bab9e9e34d42e
1945 F20110217_AABXLO smith_j_Page_014.txt
278f68b814a06b2d4e9e1241c861037a
05fe016c99b63f0f329bf95b5686cf48dc1fa1a9
F20110217_AABXLP smith_j_Page_141.tif
528e606d63e3d834631d6750a53a73e5
9770fd963bb91a114849935b8e050984e2c974b3
F20110217_AABXMD smith_j_Page_007.tif
ae4c5436b6a7912d14e3eb4af2c0a511
31b4010bdea12e8452e887a7b77882c51b8341f9
555140 F20110217_AABXLQ smith_j_Page_115.jp2
847b08f92a45a71990f048a81ffc1292
500e14d291b30ad094d2dc9bbfc58a292d32a50c
F20110217_AABXME smith_j_Page_008.tif
578bfe7abaa7889412a06b04554a031b
258c268746170caab42ce71f064e588a432dc956
1051917 F20110217_AABXLR smith_j_Page_070.jp2
39aed72e756e8fbfc297b73c34a9e125
2274fc6253c35dae8413e3f60825ced11238039a
F20110217_AABXMF smith_j_Page_010.tif
cb87a66c77af25e8b7e4e009b172bfa4
10d9dc754b90e179b34c443dfde207a62efc6036
767964 F20110217_AABXLS smith_j_Page_011.jp2
812f512a8ad0b3a5d8ab06b4348322ad
befb722ed0cdd09c213fc416fe32541ae4f1b6f3
F20110217_AABXMG smith_j_Page_015.tif
84ce96518a29dccaaec408c930baa706
463183ca943b4f7a3427926cd5f6aa6523917f9c
8564 F20110217_AABXLT smith_j_Page_067thm.jpg
fb215f22fdf5e63d4641807a7d0e2a23
c4474690fe6baf6d460080869d90973c661f9344
F20110217_AABXMH smith_j_Page_017.tif
9340b35d375d23863d3930666783e595
63fa89e15a3725100f164250528b73b5e9fcf6f4
22069 F20110217_AABXLU smith_j_Page_004.pro
06ecbe9dd0a839d486a0c8c9155828cb
dd11afda3e97049c214f68896ddae67b0af6ed91
F20110217_AABXMI smith_j_Page_018.tif
d206bf1663b458ebd39b20775f0834a7
c356a5e7a9d15a4e309205d5afb6fb7a4f4d8476
F20110217_AABXMJ smith_j_Page_019.tif
9e51c4086499ef5a22d618863148b058
c85cafb530fc7c2c908e0e30a8adc9e3bc491819
81039 F20110217_AABXLV smith_j_Page_151.pro
e3f41787cd902a943c9bef5a071d7fe8
991a7864a86989664c7761a5d2e6a1516b77c373
F20110217_AABXMK smith_j_Page_020.tif
cb2c69ec782cf3bce3d5f650f73f5e87
b8e3b88d10a3d5826e39f5e30b45dfa924f64267
188680 F20110217_AABXLW UFE0015200_00001.mets
4dffe1ca9e0af15e32e0dfadc556ab0b
ba1517d8719888590c60350347009ebd188aa9e5
F20110217_AABXNA smith_j_Page_044.tif
bbb335006256e2e04a994a43f639e861
d536927ae48b40af5d75138418979e6694be8fa6
F20110217_AABXML smith_j_Page_021.tif
7a5acde56f21037aa7b634cfb591b004
ef29eb0e045768b0e182d619ebef0c143aeaf6e6
F20110217_AABXNB smith_j_Page_045.tif
2bc45add914a3b6c7c4e069ebe7178f7
fdf69ee02dcdf7c0b2e714d300252c09afcefa97
F20110217_AABXMM smith_j_Page_022.tif
60e41110b4d0a1e21614e352cfa4d1b2
649c8ef67886ec9a4c9c18159c5050826fc7062a
F20110217_AABXNC smith_j_Page_047.tif
f5b4e052e92b170c588d474dfefeab4f
9201e907fc13c03cf5feb3b5186c19afdebcf48d
F20110217_AABXMN smith_j_Page_023.tif
640e0a235e565ad0e6a9dbb7f9f48572
e2e9c20f70d15b4b5e1446e0360afe3368a684fc
F20110217_AABXLZ smith_j_Page_003.tif
6e70689f59c0b80ee13e6b8a7cb9e2a7
e2517f5ac9c6cd20f18470d9614d6bdd65718f0b
F20110217_AABXND smith_j_Page_048.tif
b68bc829115bbc284a11457cc8231b5b
3b395e5adfc9ffa96f1dc518bccc717a7068bb8d
F20110217_AABXMO smith_j_Page_024.tif
055735e558d226875bcee18cb266d643
219df19329868eecc31b07dc2ae4af03daa7057c
F20110217_AABXMP smith_j_Page_028.tif
c1ce26c77784dddccb71fa0f979a4137
9f2302faa151b9810f9c61a7c86aadb65eb8c1e4
F20110217_AABXNE smith_j_Page_050.tif
fb7ab6dd596f2890c4da7a2b79afdaae
759d2f76ffc1567dadffc4576138c32908315129
F20110217_AABXMQ smith_j_Page_029.tif
c56afba8489b37b47bda0b76f4dee9a4
c9efb0c8ab051c89012f3cf9f37049b9835dffc8
F20110217_AABXNF smith_j_Page_051.tif
a6f8efbf1a583428e4f983160d683f32
dbd93b31fe5cd7b9be7b3f1519e52fe11161a66a
F20110217_AABXMR smith_j_Page_030.tif
5fd7fd89c85dc0e5e01ece8f9bf6c54c
b419b0baf7ae214507a7384fcd39ae075fa94f8a
F20110217_AABXNG smith_j_Page_052.tif
65fff76c37c8097500c35ca38de021d0
b4e454e56428a730eef9120dce405ca523868367
F20110217_AABXMS smith_j_Page_031.tif
2bf846628d8b7746587b6ed47e32fe9e
eb59a83bf9d390d9195c8933d0425943ed22f526
F20110217_AABXNH smith_j_Page_053.tif
2068d56a4addf6e6d0b5b0f3c7cd44a8
075802544e4debe16be48d5b2165f5a2d1d8def3
F20110217_AABXMT smith_j_Page_033.tif
afc0469355b2e20a9635f6eab005da3a
491081f1111865d0d460ffe7221089590e91ffed
F20110217_AABXNI smith_j_Page_054.tif
23b57cd55e3dff3af0f5058069d9a60a
521ec6d358828e11660fce3203037478c3005c70
F20110217_AABXMU smith_j_Page_035.tif
9d8b7531452bc61deeb7f3f55cb21acb
bb637a38ff459d247f8500599dabbcd20faf03f0
F20110217_AABXNJ smith_j_Page_055.tif
e64adf0aaaab963634058a81df19f5a7
d3dcfb60e08efbbc3a097e458769fb359bead321
F20110217_AABXMV smith_j_Page_039.tif
157e1a74fea09eccb087362add48c04e
c28daf622aa82d7a7f4225d96de81e060cda1cf3
F20110217_AABXNK smith_j_Page_056.tif
72d5d8598229331c5353283cafc05e60
b9f5cefe757ac88b4b843f30090d66883f9fdba7
F20110217_AABXMW smith_j_Page_040.tif
324b0cbc924cdd00270f9e46d1abd4fc
d50c116f9f27d3b2a2efd6069e372388af9b1947
F20110217_AABXNL smith_j_Page_059.tif
9b13e3568984feae4733d9d61e27183b
61bc5c0221007646849cf85baaafe496e8c6e5c7
F20110217_AABXMX smith_j_Page_041.tif
c2eecf666c9465cc4efd616db5ba4702
fffbb2de3b751812f7935773f3cb26b211bbd543
F20110217_AABXOA smith_j_Page_085.tif
cac0e92bc313d6de36a396b9110d111a
00946aa00b14aebbf9fd4502c08b1eabaef94447
F20110217_AABXNM smith_j_Page_060.tif
cbfbd2f7e958e002fef9d8108d10cc01
17a50ee72e47dbc48a0ed0d315a2ac136f326460
F20110217_AABXMY smith_j_Page_042.tif
10f428b8ab9cdab2eec23fd6bdc8ae23
1e6946abbdfb9dd47070e081c3152fcbaed8075b
F20110217_AABXOB smith_j_Page_087.tif
2d50ca97aeccd042eabccb784d0453c6
7be0a094e39cb0a23c5deb8951330955b1d024f2
F20110217_AABXNN smith_j_Page_061.tif
c28bcdda60ecac65b63a8278109b7448
79123a4ab7f984015d8673222262262b381fe53a
F20110217_AABXMZ smith_j_Page_043.tif
f4d889507ba982ad8fb68249054ade1a
f699053e30ae82e87bee0c030818e6ea27197363
F20110217_AABXOC smith_j_Page_088.tif
8190c2972bf7eadd57492db334063ae4
93e4b58a5901e4570559781b1bc3540fc33981af
F20110217_AABXNO smith_j_Page_062.tif
73ac326d7522c096b2910ed10b56a088
311546bbc8c5c56a89fa729be8dca449f0816a86
F20110217_AABXOD smith_j_Page_091.tif
03ae9003340c2f068fe2a6b72ecc266d
cd42da627b2980e3ab26496c9b01a819fa284df5
F20110217_AABXNP smith_j_Page_063.tif
76da168e526ffe113a180f5e61ab1e2e
0f8d6ecfe750c286c7d5f75e9d04bb96bfab1dec
F20110217_AABXOE smith_j_Page_092.tif
2aa7758f790542e049d18fb53e2cdc4f
31f717bad3b482b00921202d54f703dd23b4d3ea
F20110217_AABXNQ smith_j_Page_064.tif
4365107d775d1e26ba91fa1842ce8f3e
2777fc982444d75dfcd92a2400f53ad26552b742
F20110217_AABXNR smith_j_Page_065.tif
d88076991e71a6e2734042594e8d380a
2e1f1c2028f9ef308c49dc1e51c825c411494235
F20110217_AABXOF smith_j_Page_093.tif
26d4f120579082936430e5bec46bd769
dcaf4dd585e644c9a2533e8b78d1280361001a9b
F20110217_AABXNS smith_j_Page_066.tif
8464c8f51baf65cf44e95281d819166c
d8324f13c25c48bd6b85f63950963d29005098fb
F20110217_AABXOG smith_j_Page_094.tif
62e5b4e5b4b2cc20363d63e90e97fef5
e9f4d9bc343281bf4cba0b74f93b7874377135d4
F20110217_AABXNT smith_j_Page_069.tif
4b28a5dfc106c11a8a1369b3d6dea161
3091940c7ebba23440dcb59062fd441f34f881c9
F20110217_AABXOH smith_j_Page_099.tif
91f5a3f103662f8665df7abb00f4ec0a
ef82e9829302b21ea4e13f82d8a1c3e8eb0a5596
F20110217_AABXNU smith_j_Page_074.tif
7148c35717ad2f623e762a4ab1976eb1
351c0a64bce96fa0566cfac72e01ad1d4bc37283
F20110217_AABXOI smith_j_Page_100.tif
555629d72aa52b08ca57fac2c8f42de6
d4ac23b886b7df80063b9e735c87cc9aa719ceb9



PAGE 1

PHOSPHORUS STORAGE DYNAMICS IN WETLAND VEGETATION AND FORAGE GRASS SPECIES: FACI LITATING WETLAND HYDROLOGIC RESTORATION IN THE LAKE OKEECHOBEE WATERSHED By JEFFREY D. SMITH A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

PAGE 2

Copyright 2006 by Jeffrey David Smith

PAGE 3

This thesis is dedicated to all who strive to educate themselves, and those who support education. “The question is, does the educated ci tizen know he is only a cog in an ecological mechanism? That if he will work with that mechanism his mental wealth and his material wealth can expand indefinitely? But that if he refuses to work with it, it will ultimately grind him to dust? If education does not teach us these things, then what is education for?”(Leopold, 1966).

PAGE 4

iv ACKNOWLEDGMENTS I would like to thank Dr. Mark W. Clark fo r his contagious enthusiasm for science, and life in general, and for his guidance a nd support throughout th e course of this research. I would also like to give speci al thanks to Dr. Edmond J. Dunne for his guidance and mentoring in the field, laborat ory and throughout the de velopment of this thesis, and most of all, his friendship. Dr. Clark and Dr. D unne are outstanding scientists, but even better persons. My committ ee members, Dr. K. Ramesh Reddy and Dr. L. Hartwell Allen, were also very influential in th e development and review of this research. Lastly, I would like to thank my parents, Sarah Anderson and many friends, from near and far, for fostering my goals and providing unconditional support. Many other important people are listed in Appendix D, Table D-1.

PAGE 5

v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES...........................................................................................................viii LIST OF FIGURES...........................................................................................................xi ABSTRACT.....................................................................................................................xi v CHAPTER 1. INTRODUCTION........................................................................................................1 Problem........................................................................................................................ .2 Regional Characteristics........................................................................................3 Everglades......................................................................................................3 Lake Okeechobee’s watershed.......................................................................5 Phosphorus loading to Lake Okeechobee......................................................7 Policy and Planning......................................................................................................9 Lake Okeechobee Legislation.....................................................................................10 Phosphorus Best Management Practices....................................................................11 Hydrologic Restoration of Isolated Wetlands.....................................................12 Phosphorus in wetland soils.........................................................................13 Alternative forage crops...............................................................................14 Thesis Objectives........................................................................................................15 2. PHOSPHORUS ASSIMILATION BY ISOLATED WETLAND VEGETATION..17 Introduction.................................................................................................................17 Factors Influencing Phosphorus Retention..........................................................19 Research Objectives............................................................................................21 Research Questions and Hypotheses...................................................................22 Materials and Methods...............................................................................................23 Study Sites...........................................................................................................23 Sampling..............................................................................................................25 Sample processing...............................................................................................26 Laboratory Analysis............................................................................................27 Data and Statistics Analysis................................................................................27 Results........................................................................................................................ .28 Species Composition along a Hydrologic Gradient............................................28 Ecosystem Phosphorus Storage...........................................................................31 Standing Biomass................................................................................................33 Standing Biomass by Individual Species............................................................36

PAGE 6

vi Phosphorus Storage in Biomass..........................................................................37 Standing Biomass by Individual Species............................................................41 Discussion...................................................................................................................42 Hydrology............................................................................................................42 Ecosystem storage...............................................................................................43 Biomass in Pasture Wetlands..............................................................................44 Disturbance Effects on Biomass..........................................................................45 Phosphorus Concentrations.................................................................................47 Phosphorus Storage.............................................................................................48 Conclusions.................................................................................................................49 3. FACILATATING WETLAND HYDR OLOGIC RESTORATION WHILE MAINTAINING FORAGE PRODUCTI ON: HYDROLOGIC TOLERANCES OF PASPALUM NOTATUM AND HEMARTHRIA ALTISSIMA ..............................51 Introduction.................................................................................................................51 Background..........................................................................................................51 Research Objectives............................................................................................53 Research Questions and Hypotheses...................................................................53 Materials and Methods...............................................................................................54 Experimental Design...........................................................................................54 Treatments...........................................................................................................56 Sampling..............................................................................................................57 Soil...............................................................................................................58 Above ground biomass sampling.................................................................58 Below ground biomass.................................................................................60 Laboratory analysis......................................................................................60 Results........................................................................................................................ .60 Initial characterization.........................................................................................60 Forage Production...............................................................................................61 Bahiagrass forage production.......................................................................63 Limpograss forage production.....................................................................64 Species comparison......................................................................................65 Total Biomass......................................................................................................68 Root to Shoot Ratios............................................................................................69 Phosphorus Assimilation.....................................................................................71 Phosphorus tissue concentrations.................................................................71 Phosphorus storage.......................................................................................73 Phosphorus storage root to shoot ratios........................................................76 Discussion...................................................................................................................77 Forage Production...............................................................................................77 Flood Tolerance...................................................................................................78 Phosphorus Uptake..............................................................................................80 Conclusions.................................................................................................................81 4. SUMMARY AND CONCLUSIONS.........................................................................83

PAGE 7

vii Summary.....................................................................................................................83 Objective I: Biomass Production a nd Phosphorus Storage in Wetlands.............83 Vegetation Stress.................................................................................................84 Objective II: Facilitating Landuse and Wetland Restoration.............................84 Unexpected Results.............................................................................................85 Implications for Restoration.......................................................................................85 Conclusions.................................................................................................................88 Unanswered Questions and Need for Further Research.............................................89 APPENDIX A. SUPPLEMENTAL BACKGROUND INFORMATION...........................................90 B. SUPPLEMENTAL FIELD DATA.............................................................................92 C. SUPPLEMENTAL MESOCOSM DATA................................................................102 D. SUPPLEMENTAL ACKNOWLEDGEMENTS......................................................138 LIST OF REFERENCES.................................................................................................139 BIOGRAPHICAL SKETCH...........................................................................................146

PAGE 8

viii LIST OF TABLES Table page 1-1 Okeechobee watershed land use by percent of total land...........................................6 2-1 Mean and standard deviation of hydroperiods at each site......................................29 2-2 Mean species hydroperiod of both sites...................................................................29 2-3 Root to shoot ratios by zone.....................................................................................36 2-4 Below ground biomass concentrations by zone.......................................................38 2-5 Above ground biomass P concentrations by zone....................................................40 2-6 BGB to AGB P storage ratios..................................................................................41 3-1 Sampling dates and details.......................................................................................58 3-2 Total biomass after 163 and 375 days......................................................................68 3-3 Root to shoot ratios..................................................................................................71 3-4 Total P storage species comparison.........................................................................74 3-5 Root to shoot P storag e ratios with statistics............................................................77 4-1 Estimation of P export concentrations to tributaries from various land-uses..........87 A-1 Summary of Okeechobee Basins BMPs...................................................................90 A-2 Total P loads to Lake Okeechobee 1991-2003.........................................................91 B-1 Phosphorus storage by com ponents, site and zone..................................................92 B-2 Biomass production by com ponents, site and zone..................................................92 B-3 Species hydroperiod.................................................................................................93 B-4 Total biomass production.........................................................................................95 B-5 Below ground biomass production...........................................................................95

PAGE 9

ix B-6 Above ground biomass production..........................................................................95 B-7 Total biomass P storage............................................................................................97 B-8 Below ground biomass P storage.............................................................................97 B-9 Above ground biomass P storage.............................................................................97 B-10 Phosphorus concentratio n in above ground biomass.............................................100 B-11 Phosphorus storage in above ground biomass........................................................101 C-1 Nutrient concentrations on day 0...........................................................................102 C-2 Species comparison of fora ge production per harvest............................................103 C-3 Species comparison of cumulative forage production...........................................103 C-4 Overall below ground bioma ss – all treatments combined....................................103 C-5 Forage production per harvest................................................................................104 C-6 Cumulative forage by treatment and day...............................................................105 C-7 Below ground biomass species comparison...........................................................106 C-8 Residual biomass harvested on days 163 and 375.................................................106 C-9 Below ground biomass pr oduction time comparison.............................................106 C-10 Bahiagrass forage production pe r harvest treatment comparison........................109 C-11 Bahiagrass cumulative forage pr oduction treatment comparison........................110 C-12 Limpograss forage production pe r harvest treatment comparison.......................111 C-13 Cumulative limpograss forage pr oduction treatment comparison.......................112 C-14 Bahiagrass BGB production – treatment comparison............................................113 C-15 Limpograss BGB production – treatment comparison...........................................113 C-16 Forage P concentratio ns – species comparison......................................................114 C-17 Bahiagrass forage P concentrations treatment comparison..................................115 C-18 Limpograss forage P concentrations treatment comparison................................116 C-19 Below ground biomass P concentrations – species comparison............................117

PAGE 10

x C-20 Bahiagrass BGB P concentra tions treatment comparison...................................117 C-21 Limpograss BGB P concentra tions treatment comparison..................................118 C-22 Phosphorus storage in forage species comparison per harvest............................119 C-23 Cumulative P storage in fo rage species comparison............................................120 C-24 Bahiagrass forage P storage per harvest.................................................................121 C-25 Bahiagrass cumulativ e forage P storage.................................................................122 C-26 Limpograss forage P storage per harvest...............................................................123 C-27 Cumulative limpograss forage P storage................................................................124 C-28 Below ground biomass P st orage species comparison...........................................127 C-29 Bahiagrass below groun d biomass P storage.........................................................127 C-30 Limpograss below groun d biomass P storage........................................................127 C-31 Climatic conditions from day 1 to 375...................................................................129 D-1 Thanks....................................................................................................................1 38

PAGE 11

xi LIST OF FIGURES Figure page 1-1 Phosphorus concentrations in Lake Okeechobee.......................................................3 1-2 Historic, current, and future flow pattern of the Everglades......................................4 1-3 Land-use map of four priority basi ns of the Lake Okeechobee watershed................8 1-4 Wetland coverage in the priority basins.....................................................................9 2-1 Mechanisms driving P cycling.................................................................................21 2-2 Map of land use in the 4 priority basins...................................................................24 2-3 Isolated wetlands select ed for long term monitoring...............................................24 2-4 Stratified sampling zones: center, edge and upland.................................................25 2-5 Logistics fit of species..............................................................................................30 2-6 Logistics fit of species by site..................................................................................31 2-7 Phosphorus storage components..............................................................................32 2-8 Comparison of AGB and BGB co mponents at Beaty and Larson...........................32 2-9 Total biomass at Beaty and Larson wetlands...........................................................33 2-10 Below ground biomass at B eaty and Larson wetlandss...........................................34 2-11 Above ground biomass at Beaty and Larson wetlands............................................34 2-12 Biomass partitioning AGB vs. BGB........................................................................35 2-13 Above ground biomass by species for all zones......................................................37 2-14 Total biomass P storage............................................................................................38 2-15 Phosphorus storage in BGB at Beaty and Larson wetlands.....................................39 2-16 Phosphorus storage in AGB at Beaty and Larson wetlands.....................................40

PAGE 12

xii 2-17 Above and below ground biomass P storage...........................................................41 2-18 Phosphorus storage by species.................................................................................42 2-19 Nutrient storage and growth in plants......................................................................47 3-1 Study site at University of Florida, Gainesville, Florida..........................................55 3-2 Mesocosm diagram..................................................................................................56 3-3 Inverse relationship of water depth and redox.........................................................57 3-4 Harvesting procedure...............................................................................................59 3-5 Forage production per harvest for ea ch species all treatments combined..............62 3-6 Cumulative forage production with all treatments combined..................................62 3-7 Bahiagrass treatment comparisons...........................................................................63 3-8 Limpograss treatment comparisons..........................................................................65 3-9 Forage production per harvest by treatment.............................................................66 3-10 Cumulative forage pr oduction by treatment.............................................................67 3-11 Below ground biomass production...........................................................................69 3-12 Above and below ground biomass production after 375 days.................................70 3-13 Mean P concentrations for bahiagrass forage by harvest day and by treatments.....72 3-14 Mean P concentrations for limpograss forage by harvest day and by treatments....73 3-15 Total P storage (AGB + BGB) after 375 days.........................................................74 3-16 Cumulative P harvested in forage............................................................................75 3-17 Relative comparison of root a nd shoot P storage after 375 days.............................77 B-1 Species distribution by hydroperiod.........................................................................94 B-2 Above ground biomass by species and zone............................................................96 B-3 Phosphorus concentrations by species.....................................................................98 B-4 Phosphorus storage by zone.....................................................................................99 C-1 Relative root and shoot biomass after 163 days.....................................................107

PAGE 13

xiii C-2 Total biomass production after 163 days...............................................................107 C-3 Total biomass production after 375 days...............................................................108 C-4 Bahiagrass total biomass and P storage..................................................................125 C-5 Limpograss total biomass and P storage................................................................125 C-6 Bahiagrass BGB and P storage...............................................................................126 C-7 Limpograss BGB and P storage.............................................................................126 C-8 Root to shoot P storage ratios.................................................................................128 C-9 Total P storage (AGB +BGB) at 163 days.............................................................128

PAGE 14

xiv Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science PHOSPHORUS STORAGE DYNAMICS IN WETLAND VEGETATION AND FORAGE GRASS SPECIES: FACI LITATING WETLAND HYDROLOGIC RESTORATION IN THE LAKE OKEECHOBEE WATERSHED By Jeffrey D. Smith August, 2006 Chair: Mark Clark Major Department: Soil and Water Science Nutrient export from agricultural activities in the Lake Okeechobee watershed has contributed to eutrophication of the Lake and regulatory implementation of a phosphorus (P) Total Maximum Daily Load (TMDL) rule Historically, anth ropogenic manipulation of hydrology lowered water tables, creating im proved conditions for upland forage grass production. This action also in creased runoff rates and P load ing to the Lake. Hydrologic restoration of historically is olated wetlands within the wa tershed is a proposed best management practice (BMP) to increase P retention capacities of these wetlands. However, longer hydroperiods could potentially decrease pasture pr oductivity, and as a consequence, adversely affect the economic viab ility of the cattle i ndustry in the region. Previous studies have shown that soils under longer hydroperiods in the Okeechobee watershed have greater P storage potential than surrounding upland soils. This research primarily focuses on the vegetative component of P storage in pasture wetlands. The objectives were to evaluate biomass produc tion and P storage dynamics in vegetation under various hydroperiods and to determine the efficacy of using an alternative forage grass species to maintain pasture productivity after wetland restoration.

PAGE 15

xv Four isolated wetlands in Okeechobee Count y, Florida were sampled in November, 2004; March, 2005; and July, 2005. In this stud y, total P storage in wetlands (with a 50m upland buffer) included soil (10 cm dept h), below ground biomass (BGB), above ground biomass (AGB) and litter components. Soil was the primary P storage component representing greater th an 88% of the total P stored in the wetlands, while BGB, AGB and litter represented 8%, 3%, and 1% respect ively. Total biomass (AGB+BGB) production and P storage in biomass were inversely re lated to hydroperiod in wetlands at the more intensively managed pasture, while P storage in biomass was positively related to hydroperiod in wetlands at the less intensively managed pastur e. Management intensity (i.e., cattle density and pasture maintenance) may be influencing P storage capacities of vegetative, and affecting the relations hip between hydroperiod and P storage. In a separate mesocosm study in Gainesville, Florida, Paspalum notatum Flgg (bahiagrass) and Hemarthria altissima ‘Floralta’ (limpograss), a wet-tolerant forage grass, were evaluated under five different hydrologic treat ments. Water levels were stabilized at 10, 0, -10, and -15 cm relative to the soil surface, wh ile the control only received rain water and was allowed to drai n completely. Limpograss had greater forage (AGB) production and P assimilation than ba hiagrass in all treatments. However, bahiagrass had greater total biomass ( AGB+BGB) production in all but the 10 cm inundated treatment. Bahiagrass to tal P storage was only greater than limpograss in the -10 cm water level. This indicates that lim pograss has a greater hydrol ogic tolera nce than bahiagrass and similar P storage potential. Therefore, to maintain pasture carrying capacity and vegetative P storage during BMP implementation, limpograss may be a more suitable forage in restored pastures wetlands.

PAGE 16

1 CHAPTER 1 INTRODUCTION Nutrient export from agricultural activities in the Lake Okeechobee watershed has contributed to eutrophication of the Lake and regulatory implementation of a phosphorus (P) Total Maximum Daily Load (TMDL) rule Historically, anthropogenic manipulation of hydrology drained wetlands and lowered the water table, creating improved conditions for upland forage grass production. This action increased runoff rates and P loading to the Lake. Four priority basins occupy 12% of th e watershed’s area, but export 35% of the P load entering the Lake (FDEP, 2001). Hydrol ogic restoration of historically isolated wetlands is a proposed best management practice (BMP) to increase P retention capacities of these wetland ecosystems, t hus decreasing P loads entering the Lake. However, longer hydroperiods could potentially decrease pasture pr oductivity, and as a consequence, adversely affect the economic viab ility of the cattle i ndustry in the region. Previous studies have shown that soil s under longer hydroperiods in the Lake Okeechobee Basin have greater P storage potential than surrounding upland soils (McKee, 2005). This research primarily focuses on the vegetative component of P storage in pasture wetlands. The objectives were to eval uate biomass production and P storage dynamics in vegetation under various hyd roperiods and to determine the efficacy of using an alternative forage grass species to maintain pasture pr oductivity after wetland restoration.

PAGE 17

2 Problem In 1998, Lake Okeechobee was listed as a water quality limit ed (WQL) water body (FDEP, 2001). This condition was the result of over 50 years of excessive pollutant loading. Nutrients, dissolved oxygen, unionized ammonia, chlorides, coliforms and iron have threatened numerous societal and envir onmental values of the Lake, contributing to eutrophication, resultant algal blooms, and s ubsequent alterations of flora and fauna species composition (SFWMD et al., 2004). Phosphorus is often considered the limiting nutrient in freshwater aquatic systems (Reddy et al., 1999b). The development of agri culture (predominately animal operations) in the mid 1900’s, along with anthropogenic manipulation of hydrology in the watershed has drastically increased P loading to th e Lake (Havens et al., 2005; Reddy & Debusk, 1987). Between 1968 and 2004, the P concentrati on in the pelagic zone of the Lake (Figure 1-1) increa sed from ~40 g L-1 to over 120 g L-1 (Flaig & Reddy, 1995; Havens et al., 2005). The measured P load entering the Lake in 2004 was 548 metric tons, while the five year rolling average between 20002004 was 528 metric tons P (Havens et al., 2005). In 2001, 14 of the 29 drainage basins with in the Lake’s watershed were exceeding their P loading targets (FDEP, 2001). Severa l BMPs intended to reduce P export from agricultural lands within the watershed have been or are in the process of being implemented (Table A-1, Appendix A). In the 1980’s the implem entation of BMPs significantly reduced P concentra tions in tributaries (Gunsalus et al., 1992). However, P concentrations and loads continue to ex ceed established target loads (Table A-2, Appendix A).

PAGE 18

3 Figure 1-1. Phosphorus concentrations in Lake Okeechobee (Lake Okeechobee Issue Team & South FLorida Ecosystem Restoration Working Group, 1999) Regional Characteristics Everglades Historically, Lake Okeechobee was the cen tral component of the Everglades Ecosystem, interconnecting the Kissimmee Ri ver Basin in Central Florida to the Everglades in South Florida. Central Florida’s Upper Chain of Lakes were the headwaters of the Everglades. Water migr ated south through the Kissimmee River Basin into the northern part of Lake Okeechobee. The Lake was a natural detention basin, releasing water over its southern banks duri ng times of high water. Overflow water migrated south through sawgrass prairies in to a vast ridge and slough system, before eventually seeping into Florida Bay. Currently, as a result of anthropogenic fl ood control and land development, Central and South Florida hydrology is drastically different (Figure 1-2). In 1928, over 2000 people were killed when the storm surge from a major hurricane flooded hundreds of acres surrounding Lake Okeechobee (Kleinbe rg, 2003). That spurred the effort to

PAGE 19

4 construct the Herbert Hoover Dike around the Lake for flood control. This, in turn, reduced the size of the Lake to 650 squa re miles and altered natural hydrologic fluctuations by creating a depende nce on a vast system of water control structures. In the decades that followed, the Kissimmee River was channelized, the Caloosahatchee River was dredged and St. Lucie canal was built to divert water from the Lake out to the Atlantic Ocean and Gulf of Mexico. The hydrology of the Everglades also became systematically controlled with the constructi on of several dikes, levies, pump stations, roads and canals. Figure 1-2. Historic (left), current (middle), and future (right) flow pattern of the Everglades ( Second Louisiana--Florida Ecosys tem Restoration Information Exchange 2001) As a result, the Everglades ecosystem is no longer a continuous River of Grass (Douglas, 1947), but rather, se veral compartmentalized aqua tic systems manipulated by anthropogenic water control structures. Ag ricultural and urban development throughout South Florida drained wetlands thus, increasing anthropogeni c control of hydrology. In the early 1900’s, the sawgrass prairies south of the Lake were drained for agriculture

PAGE 20

5 development, converting the area into what is currently known as the Everglades Agricultural Area (EAA). The paradigm that drove these vast hydr ologic alterations in South Florida was based on the need for flood prevention in popul ated areas and cultiv atable ag ricultural land. Development and wetland drainage has spawned numerous resource dilemmas. Over fertilization in agricultural sectors has been identified as a primary source of excess nutrient loading to Lake Okeechobee and th e Everglades’ Water Conservation Areas (WCA). As a result freshw ater quality and supplies have been compromised. Large quantities of water are diverted from Lake Okeechobee to the coasts to prevent high P loads from entering the Everglades. However, this has degraded estuarine ecosystems on both coasts. In addition drainage has cause d rapid mineralization of organic soils and accreted nutrients, which are ultimately transported to Lake Okeechobee or the Everglades. The Everglades is a low nutrien t ecosystem, with ambient P concentrations of ~10 g L-1. Thus, discharging water with high P concentrations from the Lake into the Everglades will be detrimental to the ecosy stem. This reinforces the importance of reducing P loading to Lake Okeechobee. Lake Okeechobee’s watershed Lake Okeechobee is the second largest fres hwater lake within the contiguous United States with a surface area of 1,890 km2 and contributing watershed of over 6,000 km2. It provides numerous societal and envir onmental values including water supply for agriculture and urban sectors, flood protection and a multi million dollar sport and commercial fishing industry (SFWMD, 2004). Agriculture is the primary land use with in the watershed, occupying over half of the land area, while wetlands and terres trial ecosystems are categorically second

PAGE 21

6 (SFWMD, 1995). Improved pasture, sugar cane, upland forest, rangeland, unimproved pasture and citrus make up the largest percentage of agricult ural land area (Table 1-1). Table 1-1. Okeechobee watershed la nd use by percent of total land. Lake Okeechobee Watershed Landuse Hectares Percentage Improved Pasture 182,590 24.02% Wetlands 128,884 16.96% Water 127,339 16.75% Sugar Cane 91,409 12.03% Upland Forest 69,479 9.14% Rangeland 50,283 6.62% Unimproved Pasture 30,829 4.06% Citrus 22,941 3.02% Urban and built-up 22,325 2.94% Barren 6,284 0.83% Dairies 5,209 0.69% Row Crops 4,926 0.65% Transportation, Communication, and Utilities 4,619 0.61% Field Crops 4,147 0.55% Woodland Pasture 3,935 0.52% Fallow Crop Land 1,756 0.23% Sod Farms 941 0.12% Tree Nursery 907 0.12% Horse Farm 397 0.05% Fruit Orchards 378 0.05% Aquaculture 165 0.02% Other 164 0.02% Ornamentals 74 0.01% Other Grove 29 0.00% Floriculture 8 0.00% Table created from land use data (SFWMD, 1995) Large areas of wetlands in the watershe d have been drained for agricultural development. Riparian and non-riparian (isolated) wetlands are abundant throughout Lake Okeechobee’s watershed, covering 17% (1,290 km2) of the area (Table 1-1). The landscape in the northern part of the watershed was historically characterized by the presence of numerous isolated wetlands. Th ese wetlands are referred to as ‘historically’ isolated because they lacked a surface wate r connection to tributar ies, except overland flow in times of flooding. Extensive drainage efforts established vast networks of ditches

PAGE 22

7 and canals, short circuiting the natural water re tention and nutrient as similative capacity of the landscape. Most isolated wetlands in the northern watershed were drained in mid1900’s to support the rapidly incr easing beef cattle and dairy industry; creating improved pasture conditions for upland forage species (Flaig & Havens, 1995). Draining wetlands was a trend that occurred throughout the countr y, ultimately depleting more than half of the wetlands within the contiguous Un ited States (Mitsch & Gosselink, 2000). Phosphorus loading to Lake Okeechobee Agricultural activities are responsible fo r 98% of all P imported to the watershed, the majority of which is pasture fertilizer a nd dairy feed (Fluck et al., 1992). There is a high correlation between P imports to the watershed and P loading to Lake Okeechobee (Boggess et al., 1995). Non-point so urce runoff from agriculture, particularly, beef cattle and dairy operations, is recognized as a primar y source of P loading to the Lake (Flaig & Havens, 1995) Four priority basins within the Lake ’s watershed, S-65D, S-65E, S-154, and S-191 (Figure 1-3), have been identified as “hot spots” based on land us e intensity and high P discharge. These four basins occupy 12% of the land, and export as much as 35% of the total P load entering the Lake (FDEP, 2001). Within these priority basins, 61% of the land area supports agricultural ac tivities (47% improved pasture, 14% dairy) (Figure 1-3), while 11% of the land is occupied by partia lly drained or otherwise impacted wetlands (Figure 1-4). In fact, 45% of isolated wetlands in the priority basins have been at least partially drained.

PAGE 23

8 Figure 1-3. Land-use map of four priority basins of the Lake Okeechobee watershed (McKee, 2005)

PAGE 24

9 Figure 1-4. Wetland coverage in th e priority basins (McKee, 2005) Policy and Planning While flood control is still a priorit y, the water retention and contaminate assimilative capacity of wetlands are now wide ly recognized. In the face of increasing population and water demand in South Florida, major efforts are underway to carry out some of the largest ecological restoration projects ever undertaken. The Comprehensive Everglades Restoration Plan (CERP) (SFWMD & USACE, 2005), the Lake Okeechobee Protection Plan (LOPP) (SFWMD et al., 2004), and the Kissimmee River Restoration Program (KRR) are umbrella programs designe d to preserve and protect water resources in Central and South Florida.

PAGE 25

10 CERP is an $8 billion restoration project comprised of over 60 major projects with the underlying objective to preserve and proj ect the quality and supply of freshwater resources. Currently, an average of 1.7 billi on gallons of fresh water is diverted from Lake Okeechobee out to sea annually (SFW MD, 2005). CERP will attempt to capture and store most of that water in new reser voirs and Aquifer Storage and Recovery (ASR) wells. Lake Okeechobee Legislation A water body is considered WQL or impair ed when its pollutant load exceeds water quality standards for its designated use. Lake Okeechobee is designated as a Class I or potable water supply (FAC). In complia nce with Section 303(d) of the Clean Water Act (CWA) the establishment of TMDL is required for all impaired water bodies (FDEP, 2001). Since excessive P loading is primarily responsible for eutrophication of Lake Okeechobee (Havens et al., 1995), a TMDL of 140 metric tons yr-1 P was developed to achieve the target concentration of 40 ppb P within the Lake’s pelagic zone by 2050. Phosphorus is currently the only pollutant wi th a required TMDL requirement for Lake Okeechobee. Water quality problems in Lake Okeechobee have been widely recognized since the late 1960’s (Allen et al., 1975; Flaig & Havens, 1995; Fluck et al., 1992; Gunsalus et al., 1992; Gustafson & Wang, 2002). In 1987, the Surface Water Improvement and Management (SWIM) Act (Florida St atutes, Sections 373.451 and 373.4595) was developed to focus on preservati on and restoration of some of Florida’s most significant water bodies. Lake Okeechobee was named in th at act, specifically mandating a 40% reduction of P loads in order to achieve a P c oncentration of 40 ppb in the pelagic zone. It regulated P sources from dairies by im plementing farm buyout programs, BMPs and

PAGE 26

11 structural retrofits to control systems (i.e. lagoon systems). As a result, P loads entering the tributaries between the late 1980’s and 1990’s were reduced (Gunsalus et al., 1992). However, by the mid-1990’s loads still exceeded SWIM targets and the load reduction trend was no longer declining (LOAP, 1999). Th is resulted in legislative action that called for more aggressive action than mandated by SWIM. The Lake Okeechobee Protection Act (LOPA) (Chapter 00-130, La ws of Florida) was passed in 2000. LOPA mandated the implem entation of a restoration and protection program which includes a P TMDL and BMPs to reduce nutrient loading to the Lake. The Lake Okeechobee Protection Program (L OPP) was developed to achieve and maintain compliance with Florida water quality standards. It involves the implementation of a P TMDL along with ot her research and monitoring objectives required by LOPA. (SFWMD, 2004). In addition, the Lake Okeechobee Watershed Project (LOWP) is a component of CERP th at aims to reduce P loading to the Lake, attenuate peak flows and restore riparian and isolated wetland habitat. LOPP and LOWP contain similar P source contro l programs through the implementation of “voluntary” and cost-share BMPs; however LOPP addresses re gional projects not included in CERP. Phosphorus Best Management Practices Best Management Practices are conser vation guidelines developed using Best Available Technology (BAT) to reduce point and non-point source water pollution while maintaining economically viable agricultura l productivity (Bottcher et al., 1995). However, the success of a BMP is only as eff ective as its level of acceptance. In fact, overcoming social and political obstacles ma y be more challenging than the fundamental science supporting the BMP, t hus innovative educational appro aches that facilitate an understanding of potential cost s and benefits associated w ith implementing the BMP is

PAGE 27

12 necessary (Bottcher et al., 1995). The su ccess and cost-effectiveness of BMPs are dependent on regional goals and BATs. Howeve r, in all cases, mutual awareness of potential outcomes and willingness to compromi se by all parties is essential to achieve BMP objectives and maintain the economic viability of the land-use. Many P BMPs targeting dairy and cattle se ctors have been implemented in the Lake Okeechobee drainage basin (Table A1, Appendix A). Required implementation of P BMPs as part of the Rural Clean Wate r Program (RCWP) and the Okeechobee Dairy Rule (FAC, 1996) have effectively reduced P discharge from dairies by 50%, thus improving discharge water quality (Gunsalus et al., 1992). However, the discharge reductions are relative to previous discharges which may have been several times greater than acceptable concentrations and have not been enough to lower the overall P load entering the Lake. Hydrologic Restoration of Isolated Wetlands Hydrologic restoration of hi storically isolated wetlands is a potential BMP that could play a significant role in meet ing the P TMDL of 140 metric tons yr-1. Wetlands are known to assimilate and immobilize nutrien ts and other contamin ates in living and dead (detritus) plant biomass and in soils. Phosphorus storage in wetland soils is dependent on the P concentrations of the ove rlying water column and the sediment pool. Organic matter accumulation and the abundance of iron and aluminum oxides within the soil influences sediment P flux with the ove rlying water column (Reddy et al., 1999b). Anaerobic decomposition is a slow process th at facilitates the accumulation of organic material. The accumulation of organic material immobilizes P in the process, thus acting as a P sink as long as anaerobic conditions persist. Phosphorus assimilation in living biomass is short term process which can re -release labile nutrien ts back into the

PAGE 28

13 environment upon senescence and decomposition. However, P accretion in plant biomass has been shown to account for 12-73% of total P removal from nutrient enriched waters (Reddy & Debusk, 1985). Phosphorus in wetland soils Mckee (2005) conducted a survey of 118 wetlands within the four priority basins to determine P storage in soils of isolated wetlands. Wetlands on dairy, improved and unimproved pasture land-uses were divided in to center, edge, upland and ditch zones and sampled. Physical parameters, such as or ganic matter content and bulk density between wetland center and upland were significantly di fferent when compared between like-land uses. Total P (TP) analysis showed signifi cantly higher concentrations in wetland centers compared to uplands for all three land use t ypes. There were also significantly higher TP concentrations in wetland centers compared with edge or ditch soils for improved and unimproved pasture land use types. Across land-uses, Mckee found signifi cantly greater center and edge TP concentrations in dairy wetla nd soils than in improved a nd unimproved pasture wetland soils. However, no significant differen ce was found between improved and unimproved pasture land-use types. Cent er and edge soils from diffe rent wetland types were also significantly different. Forest ed swamp soils had significantl y higher center TP values than emergent marshes and open water emergent marsh soils, while edge concentrations were significantly greater in scrub-shrub swamps compared to emergent marsh and open water emergent marshes. There were also si gnificantly greater P c oncentrations in edge soils of forested swamps compared to emergent marshes. McKee’s results theoretica lly support a hydrologic restor ation BMP of isolated wetlands as a means to increase P retention in the watershed and decrease P loads to the

PAGE 29

14 Lake. Hydrologic restoration would raise the water table, and increase the zone of inundation; maximizing the potential of the wetlands to accrete P in soils. However, this would also cause a shift in species com position, likely decreasing upland forage grass production, and pasture carrying. Thus, a c onsequence of restoration may adversely affect the economic viability of cattle opera tions as productive pa sture area would likely be reduced. Alternative forage crops BMPs with the potential to negatively impact economic viability should not be considered BMPs, unless altern ative funds are available to subsidize their implementation (Bottcher et al., 1995). However, along with hydrologic restoration, alternative practices could be implemented to minimize forage lo ss or even enhance pasture productivity. Wet cropping systems have been suggested as pot ential means of reduci ng P imports to the watershed and concentrations in dairy wastew ater by utilizing a vegetative species with high P assimilative capacity and sufficient fora ge value (Reddy et al., 2003). A forage species could remove the majority of P in a treatment wetland, while a periphyton cell would act as a polishing mechanism to further reduce P concentrations. Since, P storage in vegetative biomass is short-term, wet cropping systems utilize recycled nutrients to produce a forage, t hus reducing the need for P imports to the watershed. Wet cropping could also be a removal mechanism by harvesting and exporting forage and assimilated P out of the wate rshed to be utilized in other agricultural operations. Removal of sod from pastures is an effective way of export P because it is an economically valuable product that can also re duce the cost of past ure renovations when converting to more productive grasses.

PAGE 30

15 Another option to maintain pasture carry ing capacity after hyd rologic restoration is to utilize alternative fora ge species that have high productivity under wet conditions. Hydrologic restoration will alter the wate r table, potentially, creating unsuitable conditions for the existing dominant forage grass, Paspalum notatum Flgg (bahiagrass). Utilization of the wet tolerant forage grass species, Hemarthria altissima, ‘ Floralta ’ (limpograss) may be a beneficial subs titute for bahiagrass, potentially reducing the pasture area that would otherwise be lo st if no alternative is implemented with hydrologic restoration. This has potentially positive economic implications for implementing a hydrologic restoration BMP. If limpograss ha s comparable or highe r forage production and quality then this BMP may not only be ecologically benefici al, but it may also provide an economic incentive to implement it. Thesis Objectives This research is part of a collaborative effort to evaluate the effectiveness of hydrologic restoration of histor ically isolated wetlands as a BMP to enhance and utilize the P storage potential within the Lake’s watershed and reduce nutrient loading to the Lake. More specifically, one objective of th is research is to evaluate the role of vegetation in wetland P storage. Since the im plementation of wetland BMPs are, in part, dependent on their level of acceptance, addressing landowner concerns for lost pasture productivity is necessary. Another objective of this research investigates the efficacy of using the wet tolerant forage species, Hemarthria altissima ‘Floralta’ (limpograss), in upland areas, adjacent to wetlands, to alleviat e the potential loss of productive pasture due to hydrologic restoration. The th esis objectives are as follows:

PAGE 31

16 I. Assess biomass production and P assi milation by wetland vegetation and forage grasses under various hydroperiods. II. Determine the efficacy of establishing a wet tolerant forage grass in wetland transition zones before hydrologic rest oration to minimize loss of productive pasture Chapter II focuses on standing biomass and P storage of various vegetative components. Chapter III describes a me socosm study that tested the hydrologic tolerances of bahiagrass and limpograss. Chapter IV summarizes results from both studies, discusses implications of wetland rest oration and presents conclusions from this study.

PAGE 32

17 CHAPTER 2 PHOSPHORUS ASSIMILATION BY ISOLATED WETLAND VEGETATION Introduction Nutrient export, primarily phosphorus (P), from non-point source agricultural activities in the Lake Okeechobee watershed has contributed to near hyper-eutrophic conditions in the Lake (Reddy et al., 1999b). As a result, a Total Maximum Daily Load (TMDL) rule for P and associated Best Management Practices (BMP) have been implemented to reduce nutri ent loading to Lake Okeec hobee (Bottcher et al., 1995; FDEP, 2001; Havens et al., 2005; SFWM D,2004). Many voluntary BMPs have effectively lowered P exports from improved pasture and dairies in high P export basins (Gunsalus et al., 1992) and in the Everglades Agricultural Area (EAA) (Flaig & Havens, 1995). However, in-Lake P concentra tions currently average 120 g L-1; three times the TMDL target concentration of 40 g L-1. Water column total nitrogen (TN) to TP ratios in the Lake are 13:1, which favors cyanob acteria dominance (Havens et al., 2005) Between 1994 and 1998, two of the Lake’s northern tributaries, the Lower Kissimmee River (LKR) and Taylor Creek/N ubin Slough (TCNS), supplied 43% of the water, and 56% of the total P load entering the Lake. The ratio of water supplied to P load for these two tributaries is disproportio nate, LKR actually supplies 33% of the water and 32% of the P load, while TCNS supplies 10% water and 24% of the P load (FDEP, 2001). The high P discharge from these tributarie s is primarily the result of four “priority” drainage basins with in their watersheds. These four basins (Figure 2-2) occupy

PAGE 33

18 12% of the Lake’s watershed, and export as much as 35% of the total P load entering the Lake. Within the priority basins 68% of the land area supports agricultural activities (45% improved pasture, 4% da iry), while 15% of the land is contains wetlands (SFWMD, 1995). The historical extent of wetlands is unknown; however, within the priority basins 45% of isolated wetlands have been at le ast partially drained (SFWMD, 2004). Ditches that drain these wetlands act as a conduit for transporting dissolved P directly to the Lake. Cattle ranching and agriculture have been the primary land uses in the watershed since the mid 1800’s. Beef cattle populations rapidly increased in the early to mid 1900’s, spurring the drainage and transfor mation of native range lands into high production improved pastures. From 1940 to 1970 the area of improved pasture increased from 34,000 to 170,000 ha (Flaig & Havens, 1995) and by 1995 it occupied 183,000 ha of the watershed; ~24 % (Table 11). The vast network of drainage canals exacerbated nutrient loss from the landscap e by lowering the water table and hydraulic retention times (HRT), thus d ecreasing P assimilative potential of historically isolated wetlands and perpetuating the need to impor t more nutrients (Flaig & Havens, 1995). Extensive wetland drainage further intensif ied cattle production and increased P imports to the watershed in the forms of cattle feed and fertilizer Studies indicate that there is a strong co rrelation between P imports to watershed and P loading to the Lake (Boggess et al., 1995; Hiscock et al., 2003). Continual net imports of P have created an excess of bioavaila ble P. Soils in the northern watershed are poorly drained and have limited P binding capacity, however, low topographic relief limits runoff and subsequent P exports from uplands (Flaig & Havens, 1995). Boggess et al., (1995) estimated that 90% of P importe d between 1985 and 1989 was retained in the

PAGE 34

19 watershed, while more recent estimates from 1997 to 2001 indicate an 83% retention of imported P (Hiscock et al., 2003). In both stud ies, the majority of imported P was stored in uplands (71% and 74%) however, of the portion that was loaded to wetlands, the percent assimilated decreased over time from 60% to 32%. Hiscock et al., (2003) attributes the reduction in st orage to decreased assimilativ e potential, not decreased wetland area. This suggests that many wetlands may already be saturated with P, and even if imports to the watershed decrease or stop, they may become a source rather than a sink. Therefore, reducing water flow, in addi tion to P imports, may be the most effective way to reduce P loading to the Lake. Factors Influencing Phosphorus Retention Phosphorus is an essential nutrient for primary producers and is limiting in most freshwater ecosystems. However, many agricult ural wetlands are not limited by P, due to its relative abundance and bioge ochemical stability (Mitsch & Gosselink, 2000). In the Okeechobee watershed, Reddy et al., (1995) found that nitrogen (N) and P concentrations in aquatic macrophytes’ tissue are generally hi gh, indicating that ne ither nutrient is limiting plant growth. Other studies have determ ined that wetland plants with N:P ratios below 14 are N limited (Koerselman & Meuleman, 1996). Despite the apparent abundance of both nutrients, the N:P ratios in tributary macrophytes were between 4 and 6 (Reddy et al., 1995), suggesti ng that P is more abundant than N. Although, generally speaking, wetland plants in the Okeechobee wate rshed are not considered to be limited by P or N availability. The physical, chemical and biological mech anisms controlling P assimilation in wetland ecosystems has been well document ed (Braskerud, 2002; Flaig & Reddy, 1995; Gilliam, 1995; Kadlec, R H, 1999; Kadlec & Knight, 1996; Mitsch & Gosselink, 2000;

PAGE 35

20 Reddy et al., 1999a; Reddy et al., 1999b; Ri chardson, 1985; Sharpley, 1995). Unlike the biogeochemical cycles of nitrogen, carbon, su lfur and oxygen, P does not have a naturally occurring gaseous phase. It is accreted in wetlands by immobiliz ation, adsorption and precipitation processes (Figure 2-1). The re lative portion of inorga nic and organic forms depends on soil, vegetation, hydrology and land use characteristics (Reddy et al., 1999a). Adsorption and precipitation are abiotic processe s that occur in the soil and are indirectly controlled by pH. Immobilization is a te mporary biotic process by which dissolved inorganic P is assimilated in vegetative or microbial biomass as organic P. Vegetative biomass has a high rate of turnover; often seve ral times a year in warmer climates. After senescence, a portion of labile P leaches back into the water column as the detrital material breaks down. A small portion of P in recalcitrant detritus is accreted in the soil as organic P. Phosphorus assimilation in vegetation is dependent on species productivity and turnover rates, nutrient availability, landuse intensity, hydrology, and biochemical and physicochemical properties (Reddy et al., 1999 a). Biomass is not be considered a sustainable long-term P removal mechanism in wetlands because it is a short-term storage that releases as much as 80% of assim ilated P back into the water column after senescence (Reddy et al., 1995). However, ac cretion of recalcitrant biomass residuals (detritus) is the only sustainable long -term storage mechanism for P removal by biological means (Kadlec & Knight, 1996; Richardson, 1985).

PAGE 36

21 A. B. Figure 2-1. Mechanisms driving P cycli ng. A) Mechanisms driving forms of P (Sharpley, 1995). B). Phosphor us cycling in wetlands. Research Objectives The use of constructed and restored is olated wetlands in Lake Okeechobee’s watershed has been suggested as a potentia l means of decreasing P loads entering the Lake (Flaig & Reddy, 1995; Havens et al ., 2005; LOPA, 1999; Re ddy et al., 2003; Reddy et al., 1996; SFWMD, 2004). Si nce TMDLs are established based on both concentration

PAGE 37

22 and flow, restoring the natural hydrology to is olated wetlands could reduce storm water runoff while increasing wetland HRT and P accr etion in residual organic material. Previous studies in the four priority basi ns have shown significantly greater soil P concentrations and organic material content in wetland centers than in adjacent uplands (McKee, 2005). These findings suggest that hydrologic restor ation could increase on-site P storage in soils by increasing wetland area. A key component responsible for increased P storage capacity in wetland centers is biomass production. High biomass and anoxic conditions foster residual biomass and P accr etion, stabilize soil porewater, and reduce concentrations in surface water (Reddy et al., 1999b) Biomass production and P assimilation are the primary focus of this research. Research Questions and Hypotheses 1. What role does vegetative bioma ss play in total wetland P storage? H1: Biomass P storage will have a lesse r role when compared to surface soil P storage. 2. Does biomass differ along a hydrologic gradient? H2: Biomass is higher in the center of the wetland 3. Does total P storage in standing bi omass differ along a hydrologic gradient? H3: Total biomass P storage will be higher in wetlands than uplands 4. Where is P partitioned in vegetation? H4: More P will be stored in above ground biomass (AGB) than below ground biomass (BGB) While P export rates from various land-uses have been broadly established (Flaig & Havens, 1995) the compounded influence of hydrology and grazing pressure on biomass production and subsequent organic P storage in wetlands in the Okeechobee basin has not been extensively studied. This chapter fo cuses on P storage in above and below ground

PAGE 38

23 standing biomass and vegetative assemblage s along hydrologic gr adients in pasture wetlands. Data presented in this chapter will be compared to vegetation data after hydrologic restoration to evalua te the effect on total wetl and P storage and vegetation dynamics. Methods focus on above ground biomass (AGB) and below ground biomass (BGB) sample collection, processing and labor atory analysis. In addition, soil and litter samples were collected and pre-sampling photographs of each quadrate were taken. Materials and Methods Study Sites Four wetlands from two different ranches located in the priority basins were selected for long-term monitoring. Selection criteria were based on land use intensity and proximity of two similarly sized, hydrologi cally modified wetlands. The Larson site, located in basin S-154, is more intensely managed then the Be aty site, located in basin S65D (Figure 2-2). Management intensity wa s subjectively determined based on land-use history, pasture maintenance re gime and grazing pressure. The two wetlands at the Larson site, Larson East (LE) (8056’28.08” W, 2720’56.06” N) and Larson West (LW) (80 56’47.49” W, 2720’59.27” N), are roughly 2.5 ha each, while the Beaty wetlands, B eaty North (BN) (8056’54.50” W, 2724’41.41” N) and Beaty South (BS) (8056’43.21” W, 2724’27.53” N), are roughly 1.3 and 1.4 ha respectfully (Figure 2-3). Wetland size was calculated in ArcGIS. The perimeter was delineated on site with GPS tracking by wa lking along vegetation community transitions between upland forage grass ( Paspalum notatum ) and unconsolidated wetland species ( Juncus effusis ).

PAGE 39

24 Figure 2-2. Map of land use in the 4 prior ity basins. S-191 is the Taylor Creek/Nubbin Slough (TCNS) basin and S-65D, S-65E and S-154 part of the Lower Kissimmee River (LKR) Basin The Larson si te is located in basin S-154. The Beaty site is located in S-65D. a. b. Figure 2-3. Isolated wetlands selected for long term monitoring. (a) Beaty Ranch wetlands; top left wetland is referred to as Beaty North (BN) and the bottom right wetland is Beaty South (BS). (b) Larson Ranch wetlands; wetland to the left is referred to as Larson East (LE) and the wetland on right is Larson West (LW)

PAGE 40

25 Sampling Data from three sampling events: November 19-21, 2004, March 25-26, 2005 and July 14-16, 2005 were collected in this st udy. Based on results from McKee, 2005 who found significantly greater soil P concentratio ns in wetland center zones than adjacent uplands, a stratified random sampling scheme was used to sample wet marsh-(center), transitional-(edge) and forage-(upl and) zones (Figure 2-4). Re spective zone data from all sampling dates were combined and analyzed to minimize temporal va riability. Five 1 m2 quadrates in each zone were located with GPS using predetermined random coordinates from ArcGIS. Figure 2-4. Stratified sampling z ones: center, edge and upland. Five randomly placed 1 m2 quadrates (not drawn to scale) were sampled in each zone. Beaty North shown as an example.

PAGE 41

26 In the upland quadrates, AGB was cu t as close to the ground as possible (approximately 1 to 3 cm above the ground su rface) using electric gr ass clippers, while hand clippers were used in edge and center zones. All removable AGB from individual quadrates was collected. After AGB was clippe d, three BGB cores were extracted from random locations within the quadrate with a 15 cm diameter aluminum cylinder, to a depth of 20 cm. The majority of the soil was dry-shaken or wet washed in the field using a 1 cm2 mesh sieve (depe nding on the whether water was present). Since it was not possible to collect 100% of AGB in the quadrate by the initial clipping, the remaining residual AGB (the stubb le left over after clipping) was removed from the BGB cores and placed in a separate bag. The amount of residual AGB per coresurface area was extrapolated to estimate the total residual AGB not collected in the field from the 1 m2 quadrate. This number was later added to the live biomass component of AGB. Sample processing All samples were transported from Okeec hobee County to Gainesville, Florida for post-collection processing. Ra ther than homogenizing all biomass within the quadrate, AGB of each species was sorted into living and senesced life stages. Both life stages of individual species were sort ed, weighed and analyzed sepa rately as components of the total biomass in the quadrate. Relative dominance of each species was determined based on the quantity of living and senesced biomass re lative to other species in the quadrate. Above ground biomass was sorted into primary, secondary, tertiary, et c, and residual or unidentifiable species. Due to the larg e quantity of AGB per quadrate, large homogeneous samples, such as upland forage species, were sub-sampled and sorted by life stage. Using the ratio of living to sene sced biomass from sub-samples and the total

PAGE 42

27 biomass of the homogeneous sample, the live and senesced portions were calculated without processing all of the biomass. Below ground biomass was washed and sieved to remove any remaining soil. All sorted AGB and washed BGB samples were dried at 70C for 72 hours, and weighed. Below ground biomass per m2 was calculated by extrapolating the biomass per coresurface area up to 1 m2. All AGB that was sub-sampled was weighed and discarded. All other samples were rough ground, sub-sample d and fine ground to pass through a # 40 sieve. Laboratory Analysis All samples were analyzed for Total Phosphorus (TP), Total Carbon (TC), and Total Nitrogen (TN), although, since P dynamics are the primary focus in these studies, only P data is presented in this chapter. Total P was extracted from 0.2-0.5 g of plant tissue using the ignition method (Andersen, 1976). Data and Statistics Analysis Data were averaged by ranch (site), and th erefore are averages from two wetlands. Statistical comparisons were made between zones at each site. Sites were not statistically compared. JMP Statistical Software was us ed to perform data analyses. For mean comparisons of more than two parameters the Tukey-Kramer HSD (honestly significant difference) test was used (JMP, 1989-2005). Outliers greater th an four standard deviations from the mean were excluded from data analysis. All quadrate values, except outliers, were included in the calculation a nd statistical comparisons of total biomass, total P storage, AGB and BGB. Calculations and comparisons of root-to-shoot ratios only included quadrates that contai ned both BGB and AGB values.

PAGE 43

28 Hydroperiods were determined for each qua drate elevation using stage data from a pressure transducer located in the center of each wetland. To determine the length of time the water level was above the soil surface, a bench mark elevation, which correlated to the transducer level in the well, was esta blished at the ground surface by the well. All quadrate elevations were corrected relati ve to the benchmark elevation and the transducer. The transducer r ecorded stage every half hour. The number of half hours the water level was above the ground was counted and converted to days. A regression of days vs. elevation (stage) was developed for each wetland. Corrected quadrate elevations were entered into the re gression equation and the co rresponding hydroperiod was returned. Zone hydroperiods were an average of all quadrates within each specified zone at each site (i.e., Beaty center hydroperiod wa s the average of all quadrates within the center zones of both wetlands). Results Appendix B contains numerous tables and figures of supplemental data and statistical comparisons. Species Composition along a Hydrologic Gradient Center, edge and upland zones within each wetland were determined visually using vegetative community compositions and aerial im ages. Overall, center, edge and upland zones at the Beaty site had l onger hydroperiods than zones at the Larson site (Table 2-1). The average hydroperiods of center and edge zo nes at the Beaty site were ~122 and ~96 days longer than the same zones at Lars on. In fact, Beaty edge zones had similar hydroperiods as Larson center zones.

PAGE 44

29 Table 2-1. Mean and standard devia tion of hydroperiod s at each site. Hydroperiod Site Zone n (days) *Difference p value Center 38269 48.9 a Edge 38141 67.4 b Beaty Upland 3831.2 64.9 c < 0.01 Center 33147 63.9 a Edge 3444.5 28.4 b Larson Upland 479.11 15.3 c < 0.01 Mean comparisons using Tukey-Kramer HSD test. Table 2-2. Mean species hydroperiod of both sites. Species Hydroperiod by Site Site Species n Indicator Days Range Difference p value Andropogon 4 FAC 77.7 68.2 0-128 BC Baccopa 1 OBL 150 150-150 ABC Eleocharis 1 OBL 312 312-312 ABC Juncus 30 OBL 155 74.3 0-302 B Ludwigia repens 3 OBL 239 67.7 161-283 AB Luziola + P. acuminatum 3 FACW 154 6.35 150-161 ABC Micranthemum 1 OBL 161 161-161 ABC Other 32 178 95.7 0-314 B P. notatum 39 UPL 45.3 70.9 0-304 C Panicum 31 OBL 250 86.5 0-323 A Polygonum 14 OBL 215 88.8 0-314 AB Pontederia 19 OBL 269 38.7 171-315 A Sagittaria 1 OBL 297 297-297 ABC Beaty Utricularia 1 OBL 298 298-298 ABC < 0.01 Alternanthera 10OBL 56.9 38.5 16-121 YZ Eleocharis 2 OBL 43.5 12 35-52 XYZ Juncus 8 OBL 54.4 27.5 0-87 YZ Ludwigia repens 1 OBL 67 67-67 XYZ Luziola + P. acuminatum 25 FACW 127 72.2 16-240 X Other 30 66.1 68.9 0-282 Y P. notatum 42 UPL 11.4 17.8 0-66 Z Panicum 2 OBL 66 72.1 15-117 XYZ Polygonum 16 OBL 93.6 52.6 15-195 XY Larson Pontederia 7 OBL 164 54.2 70-227 X < 0.01 Shaded areas represent species that were pr esent at both ranches. Indicators: (OBL) Obligate Wetland, (FACW) Facultative We tland, (FAC) Facultative, (UPL) Upland Species (Tobe et al., 1998).

PAGE 45

30 Average hydroperiods of species present at both sites were longer at the Beaty wetlands than at Larson (Tab le 2-2). These differences are similar to the zone hydroperiod differences between sites (Table 2-1). For instance, the average hydroperiod of Juncus effusis Pontederia cordata, and Polygonum hydropiperoides were 100, 105 and 121 days longer at Beaty than at Larson. Figure 2-5. Logistics fit of species. Negative log-likelihood or uncer tainty relative to hydroperiod for all wetlands. This figur e illustrates the re lative dominance of species as a percentage of the total species pres ent at a given hydroperiod Community biodiversity is greatest between upland and center zones (R2=0.096) The logistics fit of species (Figures 2-5 and 2-6) account for the likelihood that a species will be present under a given hydroperi od. It quantifies domin ance of individual species relative to other species at the same hydroperiod based on frequency of occurrence. It does not quantify biomass. Figure 2-5 shows the species distribution relative to hydroperiod in all wetlands. The Beaty we tlands, overall, had greater

PAGE 46

31 biodiversity than Larson as measured by the num ber of species present. At both sites, community biodiversity was greatest in the transitional-edge zones between upland and center zones (Figures 2-6 A & B). A. B. Figure 2-6. Logistics fit of species by site (-log-likelihood). These figures illustrate the relative dominance of species as a percenta ge of the total species present at a given hydroperiod. A). Species distribution at the Beaty (R2=0.17) site is dominated by bahiagrass in the upland and Panicum and Pontederia in the center. B). Larson (R2=0.19) species distributi on is also dominated by bahiagrass in the upland, however, a mix of Luziola fluitans and Paspalum acuminatum dominate the center zones. Ecosystem Phosphorus Storage For the purpose of this study, total P storage in wetlands (with a 50 m upland buffer) included soil (10cm depth), AGB, BGB a nd litter components. At both sites, soil was the primary P storage component, repres enting greater than 88% of the total P storage in the wetlands, while BGB, AGB and litter represented 8%, 3%, and 1% respectively (Figure 2-7). Harves ted BGB was significantly greater ( = 0.05) than

PAGE 47

32 standing AGB at the Beaty site (Figure 2-8 A), however, there were no significant P storage differences between BGB and AGB at either site (Figure 2-8 B). 0 2,000 4,000 6,000 8,000 10,000 12,000 14,000 16,000 18,000 20,000 BGBLitterSoilAGB ComponentTP ( mg m -2 ) Beaty Larson Figure 2-7. Phosphorus storage co mponents. Soil with in the top 10 cm stores more than 88% of the total P stored in these four components, while BGB, litter and AGB roughly account for 8%, 1% and 3%, respectively. Table B-1 in Appendix B contains P storage tota ls by site, zone and component. A. B. 0 500 1,000 1,500 2,000 2,500 BGBAGB Com p onentBiomass ( g m -2 ) Beaty Larson 0 500 1,000 1,500 2,000 2,50 0 BGBAGB ComponentTP ( mg m -2 ) Beaty Larson Figure 2-8. Comparison of AG B and BGB components at B eaty and Larson. A). Mean biomass harvested from all zones. B) Mean P harvested from all zones

PAGE 48

33 Standing Biomass Total biomass (AGB+BGB) in upland zones, which mainly consisted of forage grass, were similar at both sites, ~1,900 g m-2. There were no signifi cant differences in total biomass between zones at the Beaty site, howeve r Larson edge zones were significantly greater ( = 0.05) than centers, and upland zones were significantly greater ( = 0.05) than edges (Figure 2-9). The sa me relationship was true for BGB, which accounted for 68-93% of total biomass (Fi gure 2-10). Upland BGB and AGB were similar at both sites; ~1700 and ~240 g m-2 respectively. Mean BGB was larger than AGB in all zones at both s ites (Figure 2-12). Center a nd edge zones at Beaty had significantly greater ( = 0.05) AGB than upland zones, while Larson AGB was greater in upland zones than in edge zones (Figure 2-11). A. B. Figure 2-9. Total biomass at Beaty (A) and Larson (B) wetlands. Different lower case letters indicate significa nt differences. Note the difference in scales.

PAGE 49

34 A. B. Figure 2-10. Below ground biomass at Beaty (A) and Larson (B) wetlands. Different lower case letters indicate significant differences. A. B. Figure 2-11. Above ground biomass at Beaty (A) and Larson (B) wetlands. Different lower case letters indicate significant differences. Note the difference in scales.

PAGE 50

35 A. B. (3,000) (2,500) (2,000) (1,500) (1,000) (500) 0 500 CenterEdgeUpland Beaty Zone and SiteBiomass ( g m -2 ) (3,000) (2,500) (2,000) (1,500) (1,000) (500) 0 500 CenterEdgeUpland Larson Zone and SiteBiomass ( g m -2 ) Figure 2-12. Biomass partitioning AGB vs. BGB. At both sites, Beaty (A) and Larson (B) there was more BGB than AGB in all zones Approximately 20% of the quadrates di d not have measurable biomass for both AGB and BGB components. Therefore, since to tal biomass is the sum of AGB and BGB, one component made up 100% of the total bi omass for ~20% of all quadrates. This occurred when the quantity (mass) of bioma ss within individual quadrates was below the harvestable threshold. In some zones st anding AGB was limited by grazing, while the quantity of harvested BGB was dependent on the types of species present within individual quadrates. It is possible for the total biomass of a quadrate to be composted of 100% AGB and no BGB. For example, sp reading ground cover species such as P. hydropiperoides, Panicum hemitomon, Luziola fluitans and Paspalum acuminatum may have been rooted outside of the quadrate, but AGB from plants may have grown into the quadrate. Since BGB root to shoot rati os (Table 2-3) were only calculated in quadrates that contained both AGB and BGB co mponents, they are slightly different from relative AGB and BGB zonal averages (Figure 2-12). Ratios at both sites were greater than one in all

PAGE 51

36 zones, indicating that BGB was greater than AGB. Ratios at Beaty were lower than Larson and were not significan tly different by zone. Larson edge zones had significantly greater ( = 0.05) ratios than the upl and zones (Table 2-3). Table 2-3. Root to shoot ratios by zone. BGB:AGB Site Zone n Ratio *Difference p value Center 31 5.73 6.21 a Edge 29 8.36 22.2 a Beaty Upland 31 10.4 16.3 a 0.52 Center 25 23.4 40.5 a,b Edge 28 149 340 a Larson Upland 39 17.5 48.5 b 0.02 Mean comparisons using Tukey-Kramer HSD test. Standing Biomass by Individual Species Unidentifiable AGB species were collectivel y labeled as “other”, and represent a combination of multiple species. The “oth er” category often yielded similar biomass values as identifiable species. Overall J. effusis had the most AGB at the Beaty site, while Paspalum notatum (bahiagrass) was greatest at La rson (Figure 2-13). At both sites bahiagrass had the greatest AGB in upland zone s. Biomass in edge and center zones at Beaty was dominated by J. effusis, P. hemitomon, P. hydropiperoides, and P. cordata while Larson edge and cent er zones were dominated by J. effusis, P. hydropiperoides, and P. cordata (Figure B-2, Appendix B)

PAGE 52

37 A. B. 0 50 100 150 200 250 300 350 400 450 500 550 600 650 700Andropogon Baccopa Eleocharis Juncus Ludwigia L uziola + P. acuminatum Micranthemum Other P. notatum Panicum Polygonum Pontedaria Sagittaria Utricularia Biomass ( g m -2 ) 0 50 100 150 200 250 300 350 400 450 500Alternanthera Eleocharis Juncus Ludwigia Luziola + P. acuminatum Other P. notatum Panicum Polygonum Pontedaria Biomass ( g m -2 ) Figure 2-13. Above ground biomass by species fo r all zones (center, edge and upland). A) Beaty wetlands. B) Larson wetlands Phosphorus Storage in Biomass Phosphorus storage in total biomass ( AGB + BGB) was positively related to hydroperiod at Beaty, while Larson wetlands had an inverse relationshi p (Figure 2-14). Phosphorus storage in edge and upland zone s did not differ much between sites, but Beaty center zones stored ~1000 mg m-2 more P in total bioma ss than Larson centers. Center zones stored significantly more ( = 0.05) P than uplands at Beaty, while the opposite trend existed at Larson (Figure 2-14).

PAGE 53

38 A. B. Figure 2-14. Total biomass P storage. A) Beaty wetlands were positively related to hydroperiod, while Larson had an inversely relationship ( = 0.05). Different lower case letters indicate significant differences between zones at the same ranch sites Below ground biomass P concentrations (Tab le 2-4) and storage (Figure 2-15 A) did not differ between zones at the Beaty we tlands. At Larson, P concentrations in upland BGB were significantly lower ( = 0.05) than center and edge concentrations (Table 2-4). However, there was still a gene ral trend of decreasing P storage from center to upland (Figure 2-15 B), where upland a nd edge BGB stored significantly more ( = 0.05) P than center zones. Phosphorus concentrations we re significantly greater in AGB than BGB in all zones at both sites. At Beaty, P concentrations were greater in center zones than in edge and uplands. At Larson all zones were significantly different ( = 0.05); exhibiting a positive relationship with hydroperiod (Table 25). The center and edge zones at Beaty stored more P in AGB than upland zones, wh ile Larson center zones stored more P than

PAGE 54

39 edge zones (Figure 2-16). P hosphorus storage in BGB is larg er than AGB in all zones at Beaty and in edge and upland zones at Lars on. While BGB made up the largest portion of total biomass in all zones, AGB P concentr ations drastically influenced total biomass P storage. This was exhibited in Larson centers where AGB P storage was greater than BGB (Figure 2-17 B) despite the fact that harvested BGB was great er than harvested AGB. Table 2-4. Below ground bioma ss concentrations by zone Below Ground TP Concentration Site Zone n (mg/kg) *Difference p value Center 34 765 210 a Edge 34 719 285 a Beaty Upland 34 674 134 a 0.23 Center 28 781 136 a Edge 29 802 168 a Larson Upland 40 678 145 b 0.002 Mean comparisons using Tukey-Kramer HSD test. A. B. Figure 2-15. Phosphorus storage in BGB at Beaty (A) and Larson (B) wetlands. Different lower case letters indicate significant differences = 0.05.

PAGE 55

40 Table 2-5. Above ground biomass P concentrations by zone. Above Ground TP Concentration Site Zone n (mg/kg) *Difference p value Center 122 1830 976 a Edge 118 1340 702 b Beaty Upland 68 1340 623 b < 0.01 Center 69 3150 1010 a Edge 80 2690 1060 b Larson Upland 89 1650 647 c < 0.01 Mean comparisons using Tukey-Kramer HSD test. A. B. Figure 2-16. Phosphorus storage in AGB at Beaty (A) and Larson (B) wetlands. Different lower case letters indi cate significant differences ( = 0.05). The P storage root-to-shoot ratios were ca lculated the same way as the harvested biomass root-to-shoot ratios; only quadrates that contai ned both BGB and AGB were used in the calculation. All ratios were greater than one a nd did not differ significantly by zone, meaning P storage was greatest in BG B. These ratios contradict mean AGB and BGB values in Larson centers. Overall, ratio data suggests that AGB stores more P than BGB in Larson centers (Table 2-6). The differe nce, once again, is that the ratios (Table 2-6) are an average of indivi dual quadrate ratios within ea ch respective zone, which only

PAGE 56

41 included quadrates that had bot h AGB and BGB, where as P st orage values (Figure 2-17) of each component were averages of all harvested AGB and BGB with each respective zone. A. B. (2,500) (2,000) (1,500) (1,000) (500) 0 500 1,000 1,500 CenterEdgeUpland Beaty Zone and SiteTP ( mg m -2 ) (2,000) (1,500) (1,000) (500) 0 500 1,000 1,500 CenterEdgeUpland Larson Zone and SiteTP ( mg m -2 ) Figure 2-17. Above and below ground biomass P storage. Table 2-6. BGB to AGB P storage ratios BGB:AGB P Storage Site Zone n Ratio *Difference p value Center 31 3.28 3.94 a Edge 29 6.22 14.8 a Beaty Upland 31 6.30 9.86 a 0.45 Center 23 6.50 10.2 a Edge 28 28.4 65.6 a Larson Upland 39 6.76 15.3 a 0.0548 Mean comparisons using Tukey-Kramer HSD test. Standing Biomass by Individual Species Compared to other species, P. hydropiperoides, which was predominately present in center and edge zones, stored the largest amount of P in AGB at both sites (Figures 218). J. effusis, P. hemitomon and “other” species were secondary AGB P storage species in center and edge zones at Beaty (Fi gure B-4, Appendix B). “Other” AGB was a secondary storage in Larson center and edge zones. Bahiagrass stored the most P in

PAGE 57

42 uplands at both sites. At Larson, bahiagrass and J. effusis had similar P storage in upland and center zones. A. B. 0 100 200 300 400 500 600 700 800Andropogon Baccopa Eleocharis Juncus Ludwigia Luziola + P. acuminatum Micranthemum Other P. notatum Panicum Polygonum Pontedaria Sagittaria Utricularia TP ( mg m -2 ) 0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400Alternanthera Eleocharis Juncus Ludwigia Luziola + P. acuminatum Other P. notatum Panicum Polygonum Pontedaria TP ( mg m -2 ) Figure 2-18. Phosphorus storage by species A). Beaty ranch. B). Larson ranch. Discussion Hydrology Hydrology controls many physicochemical mech anisms in wetlands and is the most important determinant of wetland type a nd class (Kadlec & Knight, 1996; Mitsch & Gosselink, 2000). Hydroperiod represents th e number of days a year a wetland is inundated. The hydro-pattern or hydrologic regi me is characterized by five components: 1) duration, 2) frequency, 3) depth, 4) flow, and 5) timing or season of flooding. All of these components influence the establishmen t of vegetation and nutrient availability

PAGE 58

43 along hydrologic gradients. The wetlands in this study were under different hydroperiods and presumably, different hydrol ogic regimes as the effectiv eness of the ditches varied between wetlands and sites. Many external factors infl uence hydrology making it difficult to compare wetlands under different management intensities. For instance, ditches at the Larson site effectively drained more wetland area than di tches at Beaty. As a result, Beaty wetlands were inundated most of the year, while the Larson wetlands were only inundated roughly half of a year. Anaerobic conditions create an environment that is conducive for organic matter accumulation and P immobilization. Thus, Beaty has more potential to accumulate organic matter. Species biodiversity was greatest in tran sitional edge zones, between upland and center zones. Relatively stable environmen ts under short or long hydroperiods favor the establishment of obligate species communities (i.e. bahiagrass in the upland or Panicum and Pontederia in wetland centers). While under moderate hydroperiods in the edge zones (ecotones), fluctuating hydrologic conditions continuously eliminate and regenerate species, decreasing mono-domin ance and increasing facultative species richness. Ecosystem storage Although P storage in soil is dependent on various site physicochemical characteristics, it is often the primary st orage component in wetland and terrestrial ecosystems (Dolan et al., 1981; Richardson, 1985). Results from this study support H1: the role of vegetation in total P storage is significantly less than soil P storage in wetlands. Although vegetation plays a lesser ro le in terms of ac tive P storage, its

PAGE 59

44 importance can not be overstat ed. It indirectly increases the P storage capacity of wetlands as residual biomass decomposes and becomes incorporated into the soil. Biomass in Pasture Wetlands Biomass production studies in pasture we tlands are limited, probably due to the intrinsically high variability of disturbances to these eco systems. Different land use intensity, hydrologic regimes and grazing pressu re create high vari ability, and make it difficult to compare wetlands at different sites. Many studies have identified relationships between envi ronmental gradients (i.e hydrology) and vegetation community establishment under stable conditio ns (Seabloom et al., 2001; Van der Valk, A G, 1981; Van der Valk, A G et al., 1994; Wellings et al., 1988; Whigham et al., 2002). However, species distribution and annual biom ass are generally variable over time and reflect a balance of current environmental c onditions and historical recruitment events (Seabloom et al., 2001; Whigham et al., 2002). Our study minimizes temporal variability over one year, but does not represent long term seasonal variability that may be present in response to climatic variability. Generally, environmental gradients create pa tterns of biomass distribution within wetlands. A three year study of restored ag ricultural wetlands in Maryland found AGB was inversely related to hydroperiod (Whigham et al., 2002). In that study, wetlands were divided into submersed, emergent/seasonal, a nd temporary zones; similar to the center, edge and upland zones in our study, however, it appears that their zones my have been under longer hydroperiods. Biomass was signi ficantly greater in temporary zones [uplands] than in the emerge nt/seasonal zones [edge] and the emergent/seasonal zones were significantly greater than the submersed zo nes [center] in each of the three years. Even if their zones were under longer hydrope riods, our AGB results do not support any

PAGE 60

45 relationship to hydroperiod, indica ting that other factors, such as management intensity, are influencing AGB. Overall, BGB was the largest portion of to tal biomass, regardless of zone. This was not surprising since emergent wetland a nd terrestrial macrophytes generally have greater BGB than AGB due to extensive netw orks of roots and rhizomes (Reddy et al., 1999a). The inverse relationship between BGB and hydroperiod at the Larson site may be the result of more intensive grazing pressu re. There is an observable difference in grazing intensity between the two ranches. While cows do not graze BGB, heavily grazed AGB directly affects BGB production a nd plant survivability. Total biomass at Larson had the same inverse relationship to hydroperiod. Larson wetland zones were more heavily grazed and dominated by low-growing, unidentified annuals, Eleocharis spp. and L. fluitans Intense grazing followed by prolonged inundation discourages recruitment of perennial species. Thus dur ing flooding events, low growing species do not survive inundated conditions. Ultimately, when water levels recede, bare soil is exposed and subject to mineralization and er osion. The combination of intensive grazing and flooding primarily favors annua l species in Larson wetlands. Disturbance Effects on Biomass Harvested biomass results do not support H2: Biomass at Larson and Beaty can not be correlated with hydroperiod. Low AGB and la rge, highly variable ratios in edge zones are likely the result of the compounding eff ects of the hydrologic regime, intense pasture management and higher grazing density. Once ag ain this suggests a significant effect of grazing on biomass and P storage. External disturbances occur at multiple scales and can confound relationships between environmental gradients and wetla nd structure (Magee & Kentula, 2005; Van

PAGE 61

46 der Valk, A G et al., 1994). Direct imp acts of grazing on wetlands often include herbivory of vegetation, nutrient inputs, and so il trampling; all of which directly or indirectly alter species co mposition (Clary, 1995; Steinman et al., 2003). Grazing was not measured in this study; however, it is evident, based on these results and visual observations, that grazing has a dramatic affect on species composition and standing AGB. Bohlen et al. (2004) found differences in plant species assemblages in improved and semi-improved pastures. In their study, the less intensively grazed, semi-improved pastures were dominated by P. hemitomon which has high forage value, while intensively grazed, improved pastures s upported more diverse plant communities including J. effusis In addition, they found that ca ttle exclusions within improved pastures lead to an increase in P. hemitomon coverage. This suggests that preferential grazing of P. hemitomon may actually foster species bi odiversity incl uding unpalatable species such as J. effusis (Bohlen et al., 2004). Biomass results from this study suggest th at there is high vari ability within the same land use classification. Although all we tlands are in improve d pastures, wetland center and edge biomass results are not even similar between ranches Thus, one limitation of this study, was a lack of replic ation among sites. To minimize variability, site data contained mean values of both wetla nds at each ranch. Whigham et al., (2002) also found high variability between sites. This suggests that different management intensities within the same land use can be highly variable between sites and may not represent biomass dynamics within individu al wetlands. Thus, comparisons between sites should take into account manage ment intensity and land use type.

PAGE 62

47 Phosphorus Concentrations Higher AGB P concentrations in center zones at both Larson and Beaty were similar to the trend found by Whigham et al., (2002). In two out of the three years, AGB P concentrations in the Mary land wetlands had a positive re lationship with hydroperiod; opposite of the standing AGB trend. They conclu ded that nutrient cy cling processes are less variable than sp atial and temporal biomass diffe rences (Whigham et al., 2002). Another study found that vegetation in wetla nds receiving treate d sewage effluent showed increased P concentrations in AGB in response to both increased water levels and nutrient additions (Bay ley et al., 1985). Figure 2-19. Nutrient storage and growth in plants. Growth is typically maximized at lower nutrient supplies than the maximu m tissue storage po tential (Reddy & Debusk, 1987) It is hypothesized that incr eased P availability in wetla nds centers may facilitate “luxury uptake” of P by obligate wetland specie s. This occurs when plants take up P beyond their required needs for growth (Figur e 2-19). Biomass production is usually maximized at lower nutrient supplies, while nutrient uptake by plants is maximized at

PAGE 63

48 higher nutrient levels. The difference between the growth and nutrient uptake rates is the P storage potential (Reddy & Debusk, 1987). Phosphorus Storage Tissue P concentrations are also temporal ly and spatially variable and are not a reliable indicator of long-t erm P storage. They can vary with plant age, season and nutrient availability. For instan ce, P concentrations are typi cally higher in younger plants than in mature plants (Reddy & Debusk, 1987). Phosphorus storage potential in plants is a function of both tissue con centrations and the maximum standing crop (Reddy et al., 1995; Reddy & Debusk, 1987). The maximum st anding crop is often the primary determinant of P storage. For example, Sagittaria latifolia had the greatest P concentration of any species; however, because it was not prevalent in the wetlands, the amount of P stored was relatively small. Whigham et al. (2002) found that P storage varied between wetlands, but exhibited simila r patterns of distribu tion as standing AGB. Overall, total P storage at Beaty wa s positively related to hydroperiod, while Larson was inversely related. The influence of biomass as the primary component of total biomass P storage is evident at the La rson site. The total biomass (Figure 2-9 B) and P storage graphs (Figure 2-14 B) exhibi t similar general trends between zones. However, vegetation in Beaty cen ters stored significantly more P than upland zones, even though biomass results (Figure 29 A) did not differ by zone. Therefore, differences in P concentrations along a hydrologi c gradient are also influe ncing total P storage. Reddy and Debusk (1987) found greater than 50% of the nutrients in emergent macrophytes were stored in BGB portions of plan ts. Results from this study suggest that the relative roles of biomass and concentra tion in P storage may vary between AGB and

PAGE 64

49 BGB components. Since BGB makes up the majority of the total biomass in all zones at both sites, it was expected to store the most P. Although it was not statistically significant, AGB in Larson centers stored more P than BGB. Thus, high P concentrations in AGB had a greater influence on P storage th an biomass in the center zones at Larson. Phosphorus storage results do not conclusively support either H3: total biomass P storage would be greater in wetlands than uplands, or H4: more P will be stored in AGB than BGB. Since the Beaty wetlands stor ed more P in center zones, and the Larson wetlands showed the opposite trend, th ere is no conclusive trend and H3 was rejected. Since BGB stored significantly more P than AGB in all zones except Larson centers, H4 was also rejected. Conclusions Based on biomass results that lack speci fic trends, and opposite P storage trends along a hydrologic gradient, it is hypothesi zed that altered hydrology, management intensity and grazing may be influencing environmental gradients in the Okeechobee wetlands. The positive relationship between total biomass P storag e and hydroperiod at the Beaty site may be the combined result of longer hydroperiods and lower management intensity (including grazing pre ssure) relative to Larson. Despite ongoing disturbances (grazing) to these wetlands, P concentration gradients in vegetation, which were positively related to hydroperiod at both si tes, are similar to those found in other studies. However, P storage in vegetation is short-term, highly variable, and represents less th an 10% of total P storage in wetlands. Soil stores the majority of the total P in these wetlan ds; up to 90%. Below ground biomass and P storage is greater than AGB in all zones with the exception of P storage in Larson centers.

PAGE 65

50 Historically high net P imports to the wa tershed have saturated the P assimilative capacity of some wetlands, making them P sources rather than sinks. Hydrologic restoration would increase HRT, anaerobic conditions and organic matter accumulation. Presumably, over a prolonged period of time, if hydrology were restored, P imports were significantly decreased, and grazing pressure was minimized, wetland P assimilative capacity would increase, thus reducing P exports to the Lake. In addition to reducing P loads to the Lake, restoration also stores water in the landscape, which potentially reduces the Lake stage and discharg e of fresh water to the coasts.

PAGE 66

51 CHAPTER 3 FACILATATING WETLAND HYDROL OGIC RESTORATION WHILE MAINTAINING FORAGE PRODUCTION: HYDROLOGIC TOLERANCES OF PASPALUM NOTATUM AND HEMARTHRIA ALTISSIMA Introduction Background Hydrologic restoration of hi storically isolated wetla nds in the Lake Okeechobee watershed is considered a Best Management Practice (BMP) to decrease Phosphorus (P) loading to the Lake. The watershed has low geographic relief and many isolated wetlands have been drained to create im proved conditions for upland forage grass species. Restoration of draine d isolated wetlands involves blocking ditches or installing water control structures to raise the water tabl e back to historical levels, thus retaining water and P within these wetlands. An incr ease in wetland stage could greatly expand wetland footprints and zones of inundation, thus changing hydrope riods and hydrologic regimes of restored wetlands. Long-term flooding with decreased stage fluctuations would likely alter existing vegetative comm unities along hydrologic gradients, decreasing upland forage productivity in areas adjacent to wetlands. Since hydr ologic restoration of isolated wetlands reveres the current mana gement objective, landowner acceptance of this BMP may depend on the introduction of alternative forage grass species that are tolerant of prolonged hydr operiods and less frequent stage fluctuations. The most commonly used forage species in Florida is Paspalum notatum Flgg (‘Pensacola’ bahiagrass). Native to Central and South America, bahiagrass is a deep rooted, warm-season perennial grass that was originally planted for forage and soil

PAGE 67

52 stabilization in the southern United States(V ioli, 2000). Bahiagrass is a resilient, low maintenance species that is tolerant of a wide range of hydrologic and soil conditions; however, it is best adapted to moist, sandy so ils. It forms tough sod mats with a vast network of stolons and roots, often to a de pth of seven feet. (Chambliss & Adjei, 2006; Violi). Ninety percent of its forage pr oduction occurs between April and September (Mislevy, 2002). While bahiagrass does not se em to invade established communities, it does dominate habitats and resists invasion from other species (Violi, 2000). Once established, it is difficult to remove. It ha s been estimated that bahiagrass stolons can store enough nutrients to remain viable for tw o to three years (Chambliss & Adjei, 2006). Hemarthria altissima ‘Floralta’ (limpograss) is a fo rage species that has gained popularity since it was introdu ced (USDA Plant Introduction 364888) in 1984. Native to South Africa, limpograss was originally select ed for its winter hardiness, producing as much as 35% of its total annual production between November and March (Pate, 1998). Limpograss was specifically selected for its persistence under grazing (Quesenberry et al., 1984). It was the fourth limpograss cultivar released in Florida and is currently the only one recommended for pasture establishmen t (Pate, 1998; Sollenberg er et al., 2006). Contrary to bahiagrass, it is best adapted to poorly drained sandy soils and is not recommended for droughty sands (Pate, 1998; Solle nberger et al., 2006). In fact, it grows well in wet areas that are often continuously flooded during the wet season (Pate, 1998). Both bahiagrass and limpograss are exotic species as defined by the Florida Exotic Pest Plant Council. Ba hiagrass is a naturaliz ed exotic that was once listed as a Category I invasive exotic but has since been removed from the list. Limpograss is a listed as a Category II invasive exotic; mean ing that it shows the potential to disrupt

PAGE 68

53 native plant communities but has not yet incr eased in abundance and frequency to be considered a major nuisan ce species. (FLEPPC, 2005). Research Objectives Previous studies have evaluated the fora ge quality of and animal performance on bahiagrass and limpograss (Holderbaum et al., 1991; Holderbaum et al., 1992; Kalmbacher, R. S. et al., 1984; Kalmbacher R. et al., 1998; Long et al., 1986; Newman et al., 2002a; Newman et al., 2002b; Pate, 1998; Quesenberry et al., 1984; Sollenberger et al., 1988; Sollenberger et al., 1989), however for the purpose of this study the primary objectives were to evaluate survivability, productivity and P storage under different hydrologic conditions. Limpograss has been recommended for use in moist sites in Fl orida (Sollenberger et al., 2006), however, its sp ecific hydrologic tolerance has not been evaluated. Bahiagrass has shown short term tolera nce to flooding (David, 1999), however, ultimately over time it gets out competed by wetland species. There are multiple environmental factors that determine ideal ha bitats for species, such as grazing intensity, hydrologic regime, competition, and soil conditions. Hydrology can influence competition and physicochemical soil interactions. It is often considered one of the most influential determinants of establishment and persistence of wetlands plants (Mitsch & Gosselink, 2000). The objective of this research was to evaluate the role of hydrology on bahiagrass and limpograss in non-competitive mesocosm studies. Research Questions and Hypotheses 1. Which species has greater forage production? H1: Limpograss will have greater forage production than bahiagrass

PAGE 69

54 2. What are the hydrologic tolerances of bahiagrass and limpograss? H2: Bahiagrass will have greater total biomass production in drier treatments and limpograss will have greater total biomass production in the wetter treatments. 3. Which species assimilates more P? H3: Limpograss will have higher P storage 4. Where is P partitioned within the plant? H4: Root to shoot P storage ratio s for bahiagrass will be >1, and limpograss will be <1 This chapter compares the effect of five different water leve l treatments on below ground biomass (BGB) and above ground biom ass (AGB) production and P storage of both species in non-competitive mesocosm studies. Materials and Methods Experimental Design This experiment was designed to evaluate the response of two forage grass species to a range of hydrologic conditions typical of the transitional zone between isolated wetland and improved upland pasture. To determine how hydrology affects productivity and nutrient uptake, limpograss and bahiagra ss were evaluated in fifteen non-competitive mesocosms (1.33 m x 0.81 m x 0.76 m polyethylene tubs). Mesocosms were located in Gainesville, Florida (29.6 N 82.3 W). The experiment consisted of five treatmen ts +10, 0, -10, and -15 (water levels in centimeters, relative to the soil surface), and a control (rain water only and well drained), which are discussed in the next section. Th ere were three replicate mesocosms for each treatment. Each mesocosm contained three sub-replicates (pots) of each species. Subreplicate samples were combin ed together into one composite sample of each species.

PAGE 70

55 The sub-replicates were grown in 3 ga llon (25 cm diameter x 20 cm deep) polyethylene pots. Soil was collected from a pasture in Okeechobee, Florida, and homogenized before being dispensed into pots. Propagules of both limpograss and bahiagrass were harvested from pasture plots at the Range Cattle Research Center in Ona, Florida. Soil was washed from the propagules and seven bare-root sp rigs were planted in each pot to establish monocultures of each spec ies. Fifty-two pots of bahiagrass and 61 pots of limpograss were established 90 days pr ior to treatment. During the grown-in phase, both species were watered regularl y and pruned uniformly to stimulate new growth. a. b. Figure 3-1. Study site at Universi ty of Florida, Gainesville, Florida: (a) mesocosms were aligned in two rows and randomly assigne d a treatment. (b) Tubs receiving water were hooked up to the potable water line between the rows. Mesocosms were aligned in two rows and randomly assigned a treatment. Tanks receiving water were hooked up to a potable water supply and extern al overflow stand pipes, were used to maintain water levels fo r each treatment (Figure 3-1). Forty-five of the healthiest (determined visually) pots of each species were selected and three pots of each species were randomly placed into the 15 mesocosms. Each mesocosm contained three pots of limpograss and three pots of bahiagrass. To simulate field conditions, regulate temperature, and prevent oxygen pr oduction by photosynthetic algae in an open

PAGE 71

56 water column, mason sand was used to fill the remaining space between pots (Figure 32). Figure 3-2. Mesocosm diagram. Each meso cosm contained three pots of each species embedded in mason sand. Water level in four of the five treatments was maintained by a drip irrigation system and an external overflow-standpipe. Treatments Four of the five hydrologic treatments were maintained at a constant stage by drip irrigation emitters and external overflow-standpipes, while the fifth treatment, the control, only received rain water and wa s allowed to drain completely. Treatments receiving water included an inundation treat ment (+10 cm), where the water level was maintained 10 cm above the soil surface, and three saturation treatments (0 cm, -10 cm, 15 cm), where the water level was maintain ed 0, 10 and 15 cm, respectively, below the soil surface. Rainfall data is listed in Appendix C Table C-31. These treatments will hereinafter be referred to as +10, 0, -10, -15 and control (C). The study was initiated (day zero) on July 1st, 2004. One week prior to this date, water levels were gradually raised to their treatment levels. The timing coincides with the approximate beginning of the wet season in central Florida. Soil redox was measured in randomly selected pots of each treatment at a depth of 10 cm below the soil surface.

PAGE 72

57 Redox values were inversely related to water depths suggesting the e ffect of saturation and inundation reduced oxygen availability a nd increased anaerobic conditions in the soils (Figure 3-3). Figure 3-3. Inverse relationship of water de pth and redox. This diagram illustrates the inverse relationship of treatment wate r depth and measured redox potential within the five treatments. Soil redox was measured 10 cm below the soil surface. Sampling Soil, BGB, and two components of AGB were sampled over the course of one year. The components of AGB were forage, c onsisting of all biomass above 15 cm, and residual biomass (RB), the biomass that remained from 0-15 cm after harvest. Soil was sampled at the beginning of the experiment (d ay 1), at the end of the first growing season

PAGE 73

58 (day 163) and at the end of th e experiment (day 375). Forage was harvested periodically to evaluate temporal differences in biomass production P concentration and P assimilation. Forage samples were colle cted on days 27, 55, 83, 163, 305 and 375. At the end of the first growing season (day 163), two of the three sub-replicate pots of each species in each mesocosm were harveste d to determine BGB and RB production and P storage. Forage, RB and BGB were harvested from the remaining pots in each mesocosm on day 375. Sampling dates and biomass compon ents harvested are li sted in Table 3-1. Table 3-1. Sampling dates and details. Date Day Pots per composite sample Component Sampled 7/1/2004 1 3 Soil, Forage (for nutrient baseline) 7/28/2004 27 3 Forage 8/25/2004 5 3 Forage 9/22/2004 83 3 Forage 12/11/2004 163 3 Soil, Forage 12/11/2004 163 2 BGB, RB 5/2/2005 305 1 Forage 7/11/2005 375 1 Soil, Forage, RB, BGB This table summarizes the sampling even ts, corresponding components sampled and number of sub-replicates in composite samp les. All pots in each tank where averaged by species and pot (i.e., for biomass – g pot-1 tank-1 species-1 = average g pot-1 of each pot in each tank of each species). Soil Two soil cores (1.8 cm diameter x 20 cm depth) from pots of the same species within the same mesocosm were combined into one composite sample. Table 3-1 contains the number of pots in each compos ite. Roots and litter were removed from the soil before being dried at 70 C for 72 hours. The soil was than machine (ball) ground, sieved through a #40 mesh sieve and stored at room temperature. Above ground biomass sampling Forage sampling was designed to s imulate flash grazing, by periodically harvesting all biomass over 15 cm. This height was established 5 cm above the highest

PAGE 74

59 water level treatment to enable atmospheric gas exchange with the residual biomass for all treatments. Composite samples of each species from each meso cosm were collected using grass shears, a 15 cm-tall grated st and and a shop vacuum to ensure accurate collection. The grate was set over a pot to es tablish the clipping he ight and the vacuum was used to pull the grass th rough the grate and gather c lipped material within the vacuum (Figure 3-4). The vacuum was emptied after each composite sample per mesocosm. The post-harvest processing pr ocedure involved drying vegetation in a drying room at 70 C for 72 hours. Dry forage was than ground in a Wiley Mill, passed through a #40 mesh sieve and st ored at room temperature. a. b. c. Figure 3-4. Harvesting procedure. (a) Clipping grass with 15 cm stand and vacuum. (b) Close up view of clipping processes. (c) Vacuum was emptied after each composite sample per mesocosm. At the end of the first growing season (day 163) the RB was harvested from two of the three pots within each mesocosm. Th e post-harvest processing procedure was the same for all vegetative components. Resi dual biomass from each mesocosm was added to the respective cumulative forage pr oduction for each species and treatment to determine total AGB production g pot-1 after 163 days. The same procedure was carried out on day 375 to determine total cu mulative AGB production after 375 days.

PAGE 75

60 Below ground biomass Below ground biomass included all roots, rhizomes and stolons below the crown of the AGB shoot. Once both components of AGB were harvested (days 163 and 375) the root ball was removed from the pot and fl ushed with water to remove all soil. The same post-harvest sample processing procedure used with AGB was also used with BGB. Since harvesting BGB was a dest ructive process, sampling wa s only preformed twice. All BGB data is a cumulative total and presented as BGB production or P storage after 163 or 375 days. Laboratory analysis Soil and biomass samples were analyzed for Total Phosphorus Ash (TP) using the Ignition Method (Andersen, 1976), Total Nitr ogen (TN) and Total Carbon (TC) as described in Chapter II methods. Composite biomass and P storage were averaged by treatments and reported on a grams per pot basis. In addition, soil was also analyzed for plant available P using the Mehlich I d ilute concentration strong acid extraction procedure (Kuo, 1996). Results Supplemental tables and figures containing all data and statistical comparisons are reported in Appendix C. Initial characterization Daily environmental conditions including air and soil temperature, rainfall and humidity are listed in Table C-31 of Appendix C. Total P ash (TP), total nitrogen (TN) and total carbon (TC) tissue c oncentrations at day zero ar e listed in Table C-1 in Appendix C. Soil TP, TN and TC concentrations on day 0 averaged 0.003%, 0.092%, and 1.65 % respectively. Forage tissue concentrations ranged from 0.15-0.18%, 1.25-

PAGE 76

61 1.42%, and 42-44 % for TP, TN and TC respectively. The focus of this thesis relates to P storage in vegetation, therefore, only TP data are reported beyond the initial conditions. All production data for forage, RB and BGB are expressed either as production per harvest or cumulative production (sum of net ha rvests over time) in grams of biomass per pot. Total AGB is the sum of cumulativ e forage production and residual biomass normalized on a grams per pot basis. Forage Production Data in this section is presented as over all production by each species regardless of treatment. Overall, limpogra ss had significantly greater ( = 0.05) forage production per harvest than bahiagrass on all sampling days except day 83 (Figure 35). In the first 83 days, both species exhibited a decline in forage production. However, by the end of the first growing season (day 163), limpograss continued to producing forage while bahiagrass was essentially dormant until the beginning of the following growing season. Both species increased forage productivity between early-May and mid-July of the second growing season. On a cumulative ba sis, limpograss had significantly greater forage production on all sampling days (Figur e 3-6). After 375 days, bahiagrass and limpograss had produced 9.52 2.73 and 32.4 14.7 g pot-1 of forage respectively.

PAGE 77

62 Figure 3-5. Forage production pe r harvest for each species with all treatments combined. Day 0 is July 1st 2004. Limpograss had significan tly greater forage production per harvest on all sampling days exp ect day 83 (Table C-2, Appendix C). Figure 3-6. Cumulative forage production with all trea tments combined. Each consecutive harvest was added to previ ous harvests. Limpograss produced more forage than bahiagrass on a 375 da y period. (Table C-3, Appendix C).

PAGE 78

63 Bahiagrass forage production To determine the response of each speci es to various water level treatments comparisons were made between treatments of each species using the Tukey-Kramer HSD test. There were differences in forage production between harvests (Figure 3-7A). Throughout the experiment, the +10 and c ontrol treatments had similar forage production. Between the second and third ha rvest the +10 treatment had significantly greater forage production than the 0, -10, a nd -15 cm treatments. In addition, on day 375, the control had significantly greater forage production than the 0 cm treatment. On a cumulative basis, forage production across a ll five treatments ranged between 7.88 1.49 to 12.1 3.61 g pot-1 after 375 days and did not differ significantly between treatments. All treatments exhibited similar cumulative pr oduction curves over time (Figure 3-7B). A. B. Figure 3-7. Bahiagrass treatment comparisons A) Biomass production per harvest; +10 cm treatments significantly greater than the 0, -10, -15 cm treatments on days 83, while the Control was gr eater than the. B) Cumu lative forage production; no significant differences between treatments. (Tables C-10 & C-11, Appendix C).

PAGE 79

64 Limpograss forage production Limpograss exhibited significant treatment effects after day 27. Initially, on day 55, the control had significantly greater fo rage production than the +10, 0 and -10 cm treatments. However, on day 163 the control was the only treatment with a lower net harvest than its previous harvest on da y 83 (Figure 3-8A, Table C-12, Appendix C). Although the span of time be tween harvests (days 55-83 and 83-163) are different, by day 163, the +10 and -10 treatments roughly doubled the amount of forage produced between days 55 and 83. On day 305, the +10 cm treatment had greater forage production than the 0, -10, and -15 cm, wh ile on day 375 the control had greater production than the 0, -10, and -15 cm treatments. The differences in limpograss production per harvest did not cause significant differences in cumulative forage producti on between treatments until days 305 and 375. On day 305, the production per harvest tr eatment differences were mirrored by cumulative forage production. The +10 tr eatment had greater cumulative production (27.0 2.16 g pot-1) than the 0, -10, and -15 cm treatments after 305 days. On day 375, like the individual harvest differences, the control had greater production (50.2 16.5 g pot-1) than the 0, -10 and -15 cm treatments. While the +10 cm treatment produced significantly more forage (44.5 3.98 g pot-1) than the -10 cm treatment (21.1 3.31 g pot-1) after 375 days, it was not statistically gr eater than the 0, and -15 cm treatments, despite a power value of 0.95.

PAGE 80

65 A. B. Figure 3-8. Limpograss treatment comparisons A) Forage production per harvest. B) Cumulative forage production (T ables C-12 & C-13, Appendix C). Species comparison This section compares biomass production of limpograss and bahiagrass within the same hydrologic treatments. Limpograss had si milar or greater fora ge production than bahiagrass on all harvest days, in all trea tments (Figure 3-9). By the first sampling, limpograss had significantly greater forage pr oduction than bahiagrass in all treatments. Both species exhibited a decline in forage pr oduction per harvest in all treatments after day 27 (Figure 3-9), where only the control and -15 cm limpograss treatments were significantly greater than bahiagrass. However, limpograss rebounded and produced significantly more forage than bahiagrass in all treatments by the final harvest of the growing season (day 163). The same tre nd continued in the second growing season, where limpograss had significantly greater fora ge production per harv est than bahiagrass with the exception of the -15 cm treatment on day305. As a result limpograss had significantly greater cumulative forage produc tion after every harv est day (Figure 3-10)

PAGE 81

66 Figure 3-9. Forage production pe r harvest by treatment. A) +10 cm treatment. B) 0 cm treatment. C) -10 cm treatment. D) -15 cm treatment. E) Control. (Table C-5 Appendix C)

PAGE 82

67 Figure 3-10. Cumulative forage production by treatment. A) +10 cm treatment. B) 0 cm treatment. C) -10 cm treatment. D) -15 cm treatment. E) Control. (Table C-6, Appendix C).

PAGE 83

68 Total Biomass After 375 days, bahiagrass total biomass (AGB + BGB) was significantly greater than limpograss in the 0 and -10 cm treatments (Table 3-2). This was not consistent with total biomass results in the first 163 days, wh ere bahiagrass had signifi cantly greater total biomass production than limpograss in all tr eatments. This is primarily due to significantly greater bahiagrass BGB in all treatments. There were no significant in creases in BGB production in any treatments between days 163 to 375 for either species. However, there were significant BGB decreases in bahiagrass +10 and -15 treatments ( = 0.05), and in limpograss +10 cm ( = 0.08) and control ( = 0.05) (Figure 3-11 and Table C-9, Appendix C). In addition, forage production increased more in limpograss than bahiagrass during the same time period (Appendix C, Table C-5). This offset th e differences in total biomass production between species on day 163 to insignificant levels in the +10, -15 cm and control treatments by day 375. Table 3-2. Total biomass (AGB + BGB) after 163 and 375 days. Total Biomass (g/pot) Days Treatment nBahia Floralta p value +10 385.0 10.8 <57.2 1.47 0.01 0 3109 13.0 <63.0 6.73 0.01 -10 3113 2.78 <72.0 10.7 < 0.01 -15 3115 3.59 <65.2 7.88 < 0.01 163 C 3105 14.3 <77.7 4.30 0.03 +10 376.6 3.00 <84.5 6.51 0.13 0 3112 18.6 <76.5 2.92 0.03 -10 3112 16.0 <70.6 6.62 0.01 -15 385.4 11.6 <76.0 21.1 0.54 375 C 3113 17.7 <99.6 20.8 0.45 It is counter intuitive that cumulative biomass could decrease, but since BGB was only harvested twice, these data only repr esent the net BGB afte r 163 and 375 days, not

PAGE 84

69 the variability within those time periods. Therefore, the quantity of bahiagrass BGB that died was greater than the forage prod uced between 163 and 375 days, resulting in negative net total production. A. B. Figure 3-11. Below ground biomass produc tion. A) Bahiagrass BGB production B) limpograss BGB production (Table C-9, Appendix C). Under constant inundation, both species will survive for at least 375 days. However, cumulative biomass production for ba hiagrass actually decreased between days 163 and 375, while limpograss increased. Both species appear to have been in an acclimation phase between days 163 and 375. The l ack of significant differences in total biomass between species after 375 days in th e +10, -15 and control treatments (Table 32) indicate that those treatments were in fluencing total biomass productivity for both species. Although not statistically significant, bahiagrass still had more total biomass in the -15 cm and control treatments after 375 da ys, while limpograss had more in the +10 treatment. Root to Shoot Ratios In general BGB production had an invers e relationship AG forage production for both species. The average BGB production for all bahiagrass pots, regardless of

PAGE 85

70 treatment, was 85.6 14.0 g pot-1 after 163 days and 79.6 20.8 g pot-1 after 375 days. Limpograss BGB production was 38.0 9.07 g pot-1 after 163 days and 29.5 8.5 g pot-1 after 375 days. Bahiagrass maintained significantly more BGB than limpograss in all treatments after 163 and 375 days. Relative portions of total AGB (residual biomass + forage) and BGB for each treatment and species are gra phed in Figure 3-12. In all treatments, bahiagrass had significantly great er root to shoot ratios th an limpograss after 163 days. After 375 days, all root to s hoot ratios for limpograss were less than one while bahiagrass ratios were greater than one (Table 3-3). Thus, after 375 days limpograss produced more AGB than BGB while bahiagrass produced more BGB than AGB. Figure 3-12. Above and below ground bi omass production after 375 days. Above ground biomass (top) is the sum of cumu lative forage production and residual biomass. Below ground biomass (bottom) is all biomass harvested below the soil surface after 375 days.

PAGE 86

71 Table 3-3. Root to shoot ratios. Root:Shoot Between Species Day Treatment Bahia Floralta p value +10 3.55 0.56 >0.94 0.19 < 0.01 0 5.06 2.27 >1.17 0.22 0.04 -10 4.25 0.85 >1.58 0.40 0.01 -15 5.35 0.80 >1.42 0.34 < 0.01 163 C 4.22 1.19 >1.49 0.15 0.02 +10 2.11 0.21 >0.31 0.07 < 0.01 0 5.48 2.59 >0.81 0.23 0.04 -10 5.76 0.43 >0.90 0.39 < 0.01 -15 4.26 0.76 >0.95 0.19 < 0.01 375 C 3.91 1.21 >0.36 0.15 0.01 Phosphorus Assimilation Phosphorus tissue concentrations Phosphorus concentrations varied by speci es and by treatment. On day 0, the only significant difference in tissue concentrations between species was in the 0 cm treatment where limpograss (1790 331 mg kg-1) had a significantly greater forage P concentration than bahiagrass (1530 58.0 mg kg-1). Both species exhibited a decline in P concentrations by the first harvest (day 27). All limpograss treatments (965 220 to 1260 152 mg kg-1) and the 0, -10 cm and control (1180 60.0 to 1220 37.0 mg/kg) bahiagrass treatments had significantly lowe r forage P concentrations by the first sampling on day 27 (Figures 3-13 and 3-14) The bahiagrass +10 treatment had significantly greater forage P concentrations than limpograss on days 27, 55 and 375. In addition, P concentrations of the bahiagrass fo rage control treatment were greater than limpograss on day 375, although on day 163 the lim pograss forage control treatment was greater than the bahiagrass.

PAGE 87

72 On day 0, there were no significant differen ces in forage P concentration between bahiagrass treatments. However, on da ys 27, 55, 83, and 163, the wettest bahiagrass treatment (+10 cm) had significantly greater P concentrations than all other treatments (Figure 3-13). By the fina l harvest (day 375) the bahiag rass +10 treatment had greater forage P concentrations than the 0 and -10 treatments. Limpograss, on the other hand did not have many differences between treatme nts. The only difference was on day 163 when the +10 treatment was greater than the -15 cm (Figure 3-14). Figure 3-13. Mean P concentrations (mg/kg) for bahiagrass forage by harvest day and by treatments (Table C-17, Appendix C).

PAGE 88

73 Figure 3-14. Mean P concentrations (mg/kg) for limpograss forage by harvest day and by treatments (Table C-18, Appendix C). Below ground biomass concentrations (303-668 mg kg-1) were relatively low compared to forage. On day 163, limpograss had greater BGB P c oncentrations than bahiagrass in the 0 cm and control treatments. However, on day 375, there were no significant differences. Nor were there any significant differences for either species when treatments were compared. Phosphorus storage Phosphorus assimilation (mg pot-1) is a function of concentration and biomass production. After 375 days, th e only significant difference in total P storage (AGB +BGB) between species was in the 0 cm tr eatment, where bahiagrass was greater than limpograss (Figure 3-15). This was not consiste nt with total P storag e at the end of the first growing season (day 163) where bahiag rass was greater than limpograss in all treatments except the control (Table 3-4). This change over time in total P storage can

PAGE 89

74 be attributed to negative net BGB production in bahiagrass and greater forage production in limpograss between days 163 and 375. 0 10 20 30 40 50 60 70 80 90 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC Treatment and SpeciesTP (mg pot -1 ) Figure 3-15. Total P storage (AGB + BGB) after 375 days. Table 3-4. Total P storage ( AGB + BGB) species comparison. Total P Storage (mg/pot) Day Treatment Bahia Floralta p value +10 50.9 4.06 >40.5 1.32 0.01 0 53.7 5.56 >41.1 4.69 0.04 -10 56.9 4.48 >39.6 4.49 0.01 -15 56.0 5.73 >38 5.43 0.02 163 C 54.1 10.9 <57.5 2.38 0.62 +10 47.8 4.84 <50.2 2.67 0.49 0 50.5 13.1 >37.3 2.05 0.16 -10 46.8 5.22 >34.2 5.43 0.04 -15 36.0 5.13 >36.1 5.69 0.99 375 C 62.8 16.4 >55.3 9.09 0.53 Total P storage has the same general tre nd as biomass production. The same is true for BGB production and BGB P storage. In addition, there is a positive relationship between P partitioning and biomass allocation. Bahiagrass had greater P storage in BGB than in forage, while limpograss had the greater P storage in forage. Overall, for each species P storage in BG B was not significantly different between treatments after 375 days. However, there wa s a significantly differe nt treatment effect

PAGE 90

75 on day 163. After 163 days the limpograss cont rol stored significantly more P in BGB than all other limpograss treatments. Limpograss had greater P harvested in forage than bahiagrass in the all but the +10 treatment by day 27. By day 55, P harvested in limpograss forage was greater in all treatments except the -15 cm and control treatments, while by day 83 limpograss was only greater in the control treatment. By days 163 and 305, limpograss forage took up more P than bahiagrass in all but the 15 cm treatment, while on day 375, limpograss had greater P storage in all treatments ex cept the -15 and -10 cm treatments. On a cumulative basis, there was a str ong relationship between forage P storage (Figure 3-16) and cumulative forage producti on (Figure 3-7B, 3-12B). Limpograss had greater forage P storage than bahiagrass in all treatments except the +10 cm treatment on days 27, 55, 83 and 163. However, after day 163, limpograss had greater P storage in all treatments. A. B. Figure 3-16. Cumulative P harvested in forage. A) Bahiagrass P harvested B) Limpograss P harvested (Table C-23, Appendix C) There were no differences in harvested P between bahiagrass treatments on days 27, 55 and 305. However, on days 83 and 163, the +10 cm treatment had significantly

PAGE 91

76 more P harvested than all other treatments. The +10 cm and control treatments had greater P harvested than the 0 and -10 cm tr eatments on day 375. On a cumulative basis, on all days after 55, there was significantly more P harvested in the +10 treatment than in the 0, -10 and -15 cm treatments. Like bahiagrass, there were no differen ces in harvested P between limpograss treatments on day 27. However, by day 55, the driest treatments had assimilated the most P. The control had significantly more P harv ested than all other tr eatments, and the -15 cm treatment had more than the -10 cm treat ment. By day 83, the influence of biomass production on P storage began to emerge as P harvested in the cont rol was greater than the 0, -10 and -15 cm treatments. In the la tter and earlier parts of the growing season (days 163 and 305), there was more P harvested in the +10 cm treatment than all other bahiagrass treatments, while on day 375, ther e was more P harvested in the +10 and control than in the all other treatments. On a cumulative basis, by day 83, the bahiagrass control had assimilated significantly more P than the -10 cm treatment. By day 163, the control stored more than the 0 and -15 cm (in addition to the -10 cm ) treatments. By day 305 the +10 treatment had significantly greater cumulative P storag e than the 0, -10 and 15 cm treatments. By day 375, the +10 cm and control had greater cu mulative storage than the 0, -10, and -15 cm treatments. Phosphorus storage r oot to shoot ratios Root to shoot ratios for P storage had a positive relationship to biomass ratios. Figure 3-17 shows relative porti ons of AGB and BGB P storag e. After 375 days, the +10 treatment was the only bahiagrass treatment with a ratio less than one. Limpograss P storage ratios were all less th an one and positively relate d to biomass ratios. All

PAGE 92

77 bahiagrass treatments had significantly gr eater P storage ratios than limpograss treatments after 375 days (Table 3-5) Figure 3-17. Relative comparison of root and shoot P storage after 375 days. Table 3-5. Root to shoot P st orage ratios with statistics. Root:Shoot P Storage Day Treatment Bahia Floralta p value +10 1.27 0.27 >0.69 0.22 0.04 0 2.45 0.23 >1.01 0.13 < 0.01 -10 2.46 0.67 >1.10 0.28 0.03 -15 2.95 0.40 >0.95 0.05 < 0.01 163 C 1.95 0.10 >1.17 0.05 < 0.01 +10 0.70 0.12 >0.15 0.04 < 0.01 0 2.58 1.35 >0.39 0.09 0.05 -10 2.29 0.14 >0.48 0.17 < 0.01 -15 1.42 0.33 >0.55 0.32 0.03 375 C 1.35 0.34 >0.22 0.08 < 0.01 Discussion Forage Production Bahiagrass and limpograss are physiologically different. Bahiagrass typically has less forage production than limpograss because it allocates ~50% of its energy to root and stolon production (Chambliss & Adjei, 2006). Th is was evident from the bahiagrass root

PAGE 93

78 to shoot ratios, where BGB ranged from 2.11 to 5.76 times higher than AGB (Table 3-3). In addition bahiagrass is a long day plant that is strongly influenced by photoperiod (Marousky & Blondon, 1995). Thus its annual production is lower because it has a shorter growing season. On the other hand, lim pograss tends to allocate more energy to AG forage production, as root to shoot bi omass ratios were less than one in all treatments. Limpograss is known to support relatively high cattle stocking rates and for having superior late fall and early sp ring production compared to bahiagrass (Sollenberger et al., 2006). Results from th is study support that statement. Overall limpograss had greater forage production than ba hiagrass in the latter and earlier parts of the growing season in all treatments. In the first 83 days, both species exhibited a decline in production between harvests, regardless of tr eatment (Figure 3-5). This was likely the result of the combined effects of harvest stress, temperature a nd light effects in the latter part of the peak growing season. Harvest st ress was evident after th e first harvest (day 27) as production was significantly less for bot h species by the second harvest. After 83 days, limpograss rebounded, while bahiagrass production continued to decrease. The bahiagrass decline after day 83 is likely the result of decrea sed photoperiod during shorter days and not harvest stress. Ev en during the peak of the growing season, limpograss had greater forage production than bahiagrass. Forage production results support H1 limpograss has greater cumulative forage production than bahiagrass in all treatments. Flood Tolerance Water levels did not appear to have an e ffect on forage production for either species until after the first harvest. There were no statistical differences between bahiagrass

PAGE 94

79 treatments until the third harvest. This suggests that bahiagrass forage production may not be affected by water levels as deep as 10 cm above the soil surface for up to 55 days. The wettest bahiagrass treatment actually produced more forage than the othe r treatments toward the end of the in fi rst growing season. The same was true for the limpograss +10 treatment. In addition the limpograss +10 had the greatest forage production in the earlier part of the following growing season. The wettest treatment seems to start forage production earlier and extend it later in the growing season fo r both species, but more for limpograss. The reasoning for this may be related to a temperature buffering effect caused by standing water. Thus, flooding ma y create an artificial environment that decreases diurnal temperatur e fluctuations, thus prolongi ng the growing season. After 375 days, forage production per harvest was sim ilar in the wettest and driest treatments for both species, while the intermediate tr eatments generally produced less forage. Bahiagrass is resilient to many environm ental conditions. However, under longer hydroperiods bahiagrass is not as resilient and may be out competed by facultative or obligate wetland species. Efforts to re store native wetland species in bahiagrass dominated pastures have been challenging. While mechanically removing the sod and applying herbicide has been the most effectiv e way to remove bahiagrass (Violi, 2000), prolonged flooding will also eliminate bahiagrass and enable wetland species to establish (David, 1999). David (1999) examined the di stribution and density of bahiagrass and other wetlands species in hydrologically re stored wetlands and found that bahiagrass persisted for 2 years after inundation, before dying off by the fourth year. In addition, wetland species such as Panicum hemitomon and Pontederia cordata increased in frequency of occurrence under longer hydroperi ods. Clearly both species will survive

PAGE 95

80 375 days in 10 cm of water in a non-compe titive environment. However, even in a competitive environment it may take up to three years for different vegetation to establish between normal and high water after a change in hydrologic regime (Van der Valk, A G et al., 1994). Between days 83 and 163, both species a ppeared to acclimate to treatment conditions. Initially, the cont rols of both species had th e greatest forage production however by day 163 there was no difference between the wettest and the driest treatments. This suggests that in the short-term, hydrology alone in a non-competitive environment may not cause a sh ift in species. Over the du ration of this investigation, bahiagrass did not have significantly greater total bioma ss production in drier treatments, nor did limpograss have signi ficantly greater total biomass in the wetter treatments. Based on treatment effect comparisons, H2 can not be completely accepted. Phosphorus Uptake After 375 days, total P storage was sim ilar for both species in all treatments expect the -10 cm where bahiagrass stored more than limpograss. Therefore, H3 is rejected in favor of the null hypothesis th at limpograss does not store more P than bahiagrass. Despite, similar assimilative capaci ties, P stored in bahiagrass is relatively more stable than P stored in limpograss because the majority of P stored in bahiagrass is in BGB. Above ground forage is typically mo re labile and is subject to grazing. As discussed in Chapter II, vegeta tion is a short-term P storage mechanism. In addition, if the vegetation is continually grazed, nutrien ts in digested plant tissue are more bioavailable than senesced vegetation in wetla nds. Therefore, P can be mobilized from the soil into the water column by way of grazing.

PAGE 96

81 Phosphorus partitioning in vegetation had a si milar trend as biomass allocation. All bahiagrass treatments except the +10 stored mo re P in BGB than in AGB. The higher P concentrations in the bahiagrass +10 treatm ent caused higher P storage in AGB than in BGB. This was consistent with all limpogr ass treatments more P was assimilated in AGB than BGB. Therefore, H4 is only partially accepted. All bahiagrass ratios except the +10 cm treatment were greater than one and all limpograss P ratios were less than one. Inundation actually increased P storage in both species by increasing tissue P concentrations in the +10 treatment. The re latively high P tissue concentrations in the +10 treatment are consistent with results found in wetland center zones as described in Chapter II results and Whigham (2002). Reddy and Debusk (1985) observed lower tissu e concentrations in summer months and suggested that higher pr oductivity in the summe r likely diluted concentrations. In addition they suggested that slow growth and luxury uptake likely caused increased concentrations in the winter (Reddy & De busk, 1985). The same line of reasoning may also explain the elevated P concentrations in the +10 treatments. Biomass production decreased in the bahiagrass +10 treatment, how ever P concentrations were significantly greater than the other treatments. Wher e nutrients are readily available, tissue concentrations may have a direct relationshi p to biomass productivity. Future research may look into the possibility of specific plan t species’ rates of nutrient uptake over time vs. resultant tissue concentrations. Conclusions Limpograss appears to thrive in the wettest and driest mesocosm treatments. In fact the limpograss +10 and control treat ments had the greatest production of all

PAGE 97

82 treatments. Although bahiagrass survived under inundated conditions for 375 days, more than likely it would not survive competiti on from other plant species, trampling, grazing and water stress in situ Both species had similar total P storage in all treatments except the -10 cm treatment. The majority of P stored in bahiagrass is in BGB, while most P assimilated in limpograss is stored in forage. Thus util izing limpograss for P removal from wetlands may be best optimized by harvesting and e xporting hay and assimilated P away from the wetlands. Overall, after 375 days limpograss had great er forage production than bahiagrass in all treatments, a greater hydrologic tolerance and similar P storage potential. Therefore, in order to maintain pasture carrying cap acity and vegetative P storage during BMP implementation, limpograss may be a more suitabl e forage in restored pastures wetlands even under higher water levels and extended hydroperiods.

PAGE 98

83 CHAPTER 4 SUMMARY AND CONCLUSIONS Summary The overall goal of this research was to evaluate the biomass production and P storage potential of vegetation in historically isolated past ure wetlands and determine the efficacy of using a wet tolerant forage speci es to minimize the loss of improved pasture area as a result of hydrologic restoration. Objective I: Biomass Production a nd Phosphorus Storage in Wetlands I. Assess biomass production and P assi milation by wetland vegetation and forage grasses under various hydroperiods. Results from Chapter II and McKee (2005) indicate that wetland soils in the Okeechobee basin store more P per unit area than surrounding upland soils and vegetative components. The direct role of vege tation in active total P storage is relatively small, short-term, and highly variable compared to the physical storage capacity of soil. However, the presence of vegetation is an important component of ecosystem P storage because it increases the total P rete ntion capacity of wetland soils. Phosphorus storage in vegetation alo ng hydrologic gradie nts was variable depending on the type of species present, land-use intensity, grazing pressure and hydrology. There were, however, similar trends in AGB P storage. In general, wetland zones at both sites stored more P in AGB than in upland zones. In addition, total P storage (AGB + BGB) had a positive rela tionship to hydroperiod at Beaty while the

PAGE 99

84 opposite trend existed at Larson. This wa s likely related to higher AGB tissue P concentrations in wetland centers than uplands and differences in land-use intensity. Vegetation Stress Hydrology is often the primary determinant of vegetation composition within wetlands. However, in pasture wetlands, the stress of grazing likely influences vegetation community establishment and persistence. Although grazing was not measured in this experiment, vegetation pa tterns, biodiversity of species, and large hydroperiod ranges for the same species at diffe rent sites suggests that hydrology is not solely responsible for species distribution w ithin pasture wetlands. In addition, grazing may have a significant effect on wetland P storage capacity. Objective II: Facilitating La nd-use and Wetland Restoration II. Determine the efficacy of establishing a wet tolerant forage grass in wetland transition zones before hydrologic rest oration to minimize loss of productive pasture Overall limpograss had greater cumulative fo rage production than bahiagrass in all hydrologic treatments. This was primarily due to its ability to produc e forage earlier and later in the growing season. More importa ntly, limpograss production was similar in the wettest and driest treatments, producing significantly more fo rage than the intermediate water level treatments. The wette st and driest bahiagrass trea tments also had the greatest production relative to th e other treatments. Both species will survive for 375 days under non-competitive, inundated soil conditions as long as the biomass is not comp letely submerged. However, it is unlikely that bahiagrass would be competitive under wet conditions with its low productivity.

PAGE 100

85 Unexpected Results One unexpected result that was consistent between both species in the mesocosm experiment was that the greatest biomass pr oduction and subsequent P storage occurred in the wettest and driest treatments. It was hypothesized that biomass would have a negative parabolic shape when treatments we re aligned on the X-axis in order from wettest to driest. Essentially, it was thought that both species would have similar curves, expect limpograss’s curve would be shifte d more toward the wet treatments and bahiagrass’s more toward the dry treatments. However, as described in Chapter III, the results were opposite. Each species had a positive parabolic shape. Another unexpected result found in both th e field and mesocosm studies was the increased tissue P concentrations under flooded conditions. Above ground biomass tissue P concentrations in wetland center zones a nd in the +10 inundated mesocosm treatment were greater than uplands and drier mesoco sm treatments. Higher concentrations of plants in wet conditions we re likely caused by slow grow th and luxury uptake. Implications for Restoration Bottcher et al., (1995) defines BMPs as on-farm activities to reduce nutrient exports to water bodies and tributaries to environmenta lly acceptable levels, while simultaneously maintaining an economically viable farming operation. In addition, BMPs that adversely affect the economic vi ability of a farming operation should be subsidized to maintain profita bility (Bottcher et al., 1995). Many BMPs are considered voluntary; howev er in a watershed where a TMDL has been mandated, water quality compliance is requi red. The term “voluntary” refers to the choice of options ranchers ha ve to comply with TMDL goals. In the Okeechobee basin, ranchers have the option to monitor the nutrien t discharge from their property to ensure

PAGE 101

86 that it is meeting water quality standard s or implement recommended BMPs for the designated land-use, which presumes that discharge water quality standards are being met. As with any industry, it is advantageous fo r ranchers to strive for economies of scale in order to maximize profitability (i.e the marginal cost per cow decreases while total production increases). However, econom ies of scale in agriculture tend to be limited compared to other industries because of finite production space and time. A hydrologic restoration BMP could potentially affect the economic viability of a cattle operation by increasing marginal costs of pr oduction. Hypothetical ly, if a rancher was stocking a 10 ha bahiagrass pastur e at a density of two cows ha-1 prior to restoration, and after restoration the usable pa sture area decreased to 9 ha, the cost per cow increases as two fewer cows support the ranch’s overhead fixed costs. One study evaluating cattle productivity in pastures with high, medium and low stocking rates, representative of relative stocking rates in the Okeechobee watershed, found that a decrease in stocking rate had a one-to-one relationship with ranch revenues (Arthington et al., 2003 ). Another component of that study evaluated water quality impacts at the same stocking rates. The researchers found that cattle density did not significantly influence concentra tions or loads of nutrients in runoff waters (Bohlen et al., 2004). Despite, the fact that wetl ands in pastures with the hi ghest cattle traffic had the highest P concentration, they concluded that cattle impacts may be more related to the activity of the cattle wi thin wetlands rather than the past ure stocking density. In addition, they suggest BMPs that reduce stocking rates or remove cattle from pastures do not appear to be an effective way to reduce nut rient runoff. Further, BMPs that focus on

PAGE 102

87 preventing net P imports to the watershed or decreasing runoff to tributaries have the greatest potential to reduce nutrient load s to the Lake (Bohlen et al., 2004). Bohlen et al. (2004) also f ound that improved pastures e xported five to seven times more P than semi-improved pastures. They attribute the difference to a historical accumulation of P in improved pastures from fertilizer applications prior to 1987, while the semi-improved pastures had never been fe rtilized. Bottcher et al. (1995) modeled runoff concentrations using water quality da ta from various tributaries of known landuses. They also found differences in runo ff concentrations between improved and semiimproved pastures (Table 4-1). However, they attribute the difference to animal densities (Bottcher et al., 1995). Table 4-1. Estimation of P export concentratio ns to tributaries from various land-uses. Table from (Bottcher et al., 1995) It is evident that hydrolo gic restoration would reduce P exports in runoff waters, potentially meeting water quality standards. However, in bahiagrass pastures, carrying capacity would likely decrease as a result of hydrologic rest oration. Planting limpograss around pasture wetlands before restoration ma y minimize the loss of usable pasture and help facilitate BMP acceptance. Since limpogr ass has greater production than bahiagrass across a range of hydrologic conditions from + 10 cm flooded, to well-drained conditions, it has the potential to mainta in usable pasture space afte r hydrologic restoration.

PAGE 103

88 Conclusions It is difficult to assess the physical P storage potential of vegetation in pasture wetlands because of temporal variability, hydrolo gic stress, land-use variability and continual grazing. It is evident from this research that cattle grazing activities are influencing biomass production and P storage along hydrologic gradients. Total P storage in vegetation at the two ranches exhibited opposite trends along hydr ologic gradients. However, elevated tissue P concentrations under flooded conditions were unexpected results, consistent in both field and mesoco sm studies. Results from this research suggest that the capacity of ve getation to physically store P is highly variable in improved pasture wetlands. However, the presence of vegetation remains an important component of the total assimilative potential of wetland ecosystems. Limpograss had greater production than ba hiagrass under all hydrologic treatments in non-competitive mesocosm studies. In fact limpograss thrived in th e wettest and driest treatments. Its growth characteristics are similar to the native Panicum hemitomon Although bahiagrass survived for 375 days in 10 cm flooded conditions, it is unlikely that it would be able to survive a nd compete with other species in situ Forage quality was not evaluated in this study, however, literat ure indicates that animal performance is similar between both species (Sollenberger et al., 1988). However, proper management of limpograss pastures can carry more animals and produce greater gain ha-1 than bahiagrass pastures (Sollenberger et al., 1989). While the majority of nutrients within th e watershed are stored in uplands (Reddy et al., 1996), wetlands component s (soil, litter, AGB, BGB) store more P per unit area. Thus, hydrologic restoration would retain wa ter, increasing the z one of inundation and subsequent P assimilative capacity.

PAGE 104

89 The mechanism responsible for transporting P to the lake is the vast infrastructure of ditches and canals that short circuit the landscapes natural ability to assimilate nutrients and contaminates. In addition, net P imports to the watershed are highly correlated with P loading to the Lake (Hiscock et al., 2003). Even t hough the Institute of Food and Agriculture Sciences (IFAS) at the University of Florid a adopted a zero P fertilizer recommendation on bahiagrass pastures a nd a maximum of 40 kg P ha-1 for other forage grasses (Kidder et al., 2000), P imports to the wa tershed continue to exceed P exports. Thus, reducing P imports to the watershed and decreasing the quantity of runoff water to the Lake is likely the most effective short-term solution to meet TMDL goals. Unanswered Questions and Need for Further Research The higher P concentrations in wetla nd centers and in the +10 cm flooded mesocosm treatment are hypothesized to be the result of luxury uptake, induced by flooded conditions. Water level fluctuations and inundated soil conditions alter biotic and abiotic interactions, change redox potential and create concentra tion gradients. This affects bioavailability of nutrients and the ener gy required to mine for nutrients. Nutrient availability under reduced conditions likely plays an important role in plant tissue concentrations. Few studies have examined the biotic mechanisms of luxury uptake of nutrients by plants.

PAGE 105

90 APPENDIX A SUPPLEMENTAL BACKGROUND INFORMATION Table A-1. Summary of Okeechobee Basins BMPs Source: (Bottcher et al., 1995)

PAGE 106

91 Table A-2. Total P loads (M tons) to Lake Okeechobee 1991-2003 Source: (SFWMD, 2004)

PAGE 107

92 APPENDIX B SUPPLEMENTAL FIELD DATA Table B-1. Phosphorus storage by components, site and zone. Site Zone n BGB TP (mg/m2) Litter TP (mg/m2) Soil TP (mg/m2) AGB TP (mg/m2) Center 34 1210 869 248 185 13000 4580 632 450 Edge 34 907 576 192 150 13800 7190 476 369 Beaty Upland 34 1060 391 161 121 10100 2270 258 140 Center 30 400 402 320 305 18200 5180 509 669 Edge 29 890 449 238 238 11400 4730 250 309 Larson Upland 43 1150 589 275 177 13300 5270 344 212 Table B-2. Biomass production by components, site and zone. Site Zone n BGB (g/m2) Litter (g/m2) AGB (g/m2) Center 34 1570 1040 283 211 433 261 Edge 34 1390 912 219 167 443 284 Beaty Upland 34 1620 617 161 115 251 119 Center 30 508 523 304 271 178 233 Edge 29 1170 679 113 118 91.5 114 Larson Upland 43 1740 866 205 118 230 151

PAGE 108

93 Table B-3. Species hydroperiod. Species Hydroperiod Species n Indicator Days Range Difference p value Alternanthera 10OBL 56.9 38.5 16-121 BC Andropogon 4 FAC 77.7 68.2 0-128 ABC Baccopa 1 OBL 150 150-150 ABC Eleocharis 3 OBL 133 155 35-312 ABC Juncus 38 OBL 134 78.8 0-302 B Ludwigia 4 OBL 196 102 67-283 AB Luziola + P. acuminatum 28 FACW 130 68.6 16-240 B Micranthemum 1 OBL 161 161-161 ABC Other 62 124 100 0-314 B P. notatum 81 UPL 27.5 53 0-304 C Panicum 33 OBL 238 96 0-323 A Polygonum 30 OBL 148 92.8 0-314 B Pontederia 26 OBL 240 63.6 70-315 A Sagittaria 1 OBL 297 297-297 ABC Utricularia 1 OBL 298 298-298 ABC < 0.01 Indicates species present at both sites

PAGE 109

94 A B. Figure B-1. Species distribu tion by hydroperiod. Shaded area s represent stratified zones correlated to hydroperiod on the x-axis Bars represent the range of hydroperiods under which the corresponding species were observed. A). Species present at the Beaty site. B). Species present at the Larson site.

PAGE 110

95 Table B-4. Total biomass production. Total Biomass Site Zone n g/m2 *Difference p value Center 31 1950 1060 a Edge 29 1840 1160 a Beaty Upland 31 1940 598 a 0.89 Center 25 745 557 c Edge 28 1270 681 b Larson Upland 39 1870 805 a < 0.01 Mean comparisons using Tukey-Kramer HSD test. Table B-5. Below gr ound biomass production. Below Ground Biomass Site Zone n (g/m2) *Difference p value Center 331570 1040 a Edge 331390 912 a Beaty Upland 341620 617 a 0.55 Center 29508 523 c Edge 291170 679 b Larson Upland 411740 866 a < 0.01 Mean comparisons using Tukey-Kramer HSD test. Table B-6. Above gr ound biomass production. Above Ground Biomass Site Zone n (g/m2) *Difference p value Center 34433 261 a Edge 36443 284 a Beaty Upland 35251 119 b < 0.01 Center 28178 233 a,b Edge 3391.5 114 b Larson Upland 43230 151 a 0.003 Mean comparisons using Tukey-Kramer HSD test.

PAGE 111

96 A. B. Figure B-2. Above ground biomass by species and zone. A). Beaty AGB in upland, edge and center zones. B). Larson AGB in upland, edge and center zones.

PAGE 112

97 Table B-7. Total biomass P storage. Total Biomass P Storage Site Zone n (mg/m2) *Difference p value Center 31 1810 969 a † Edge 29 1440 749 a,b † Beaty Upland 31 1370 399 b † 0.05 Center 23 901 750 b Edge 28 1130 581 a,b Larson Upland 39 1400 554 a 0.01 Mean comparisons using Tukey-Kramer HSD test † Indicates = 0.10. All other comparisons = 0.05. Table B-8. Below ground biomass P storage. Below Ground P Storage Site Zone n (mg/m2) *Difference p value Center 33 1210 869 a Edge 33 907 576 a Beaty Upland 34 1060 391 a 0.16 Center 28 400 402 b Edge 29 890 449 a Larson Upland 40 1150 589 a < 0.01 Mean comparisons using Tukey-Kramer HSD test. Table B-9. Above ground biomass P storage. Above Ground Biomass P Storage Site Zone n (mg/m2) *Difference p value Center 34 632 450 a Edge 36 476 369 a Beaty Upland 35 258 140 b < 0.01 Center 27 509 669 a Edge 33 250 309 b Larson Upland 43 344 212 a,b 0.05 Mean comparisons using Tukey-Kramer HSD test.

PAGE 113

98 A. B. Figure B-3. Phosphorus con centrations by species.

PAGE 114

99 A. B. Figure B-4. Phosphorus storage by zone. A) Beaty B) Larson

PAGE 115

100 Table B-10. Phosphorus concentr ation in above ground biomass AGB Species TP Concentration by Site Site Species n Indicator (mg/kg) Difference p value Andropogon 10 FAC 854 549 CD Baccopa 2 OBL 3340 ABCD Eleocharis 3 OBL 1360 BCD Juncus 89 OBL 1020 477 D Ludwigia 5 OBL 1880 305 BCD Luziola + P. acuminatum 6 FACW 2260 341 BCD Micranthemum 2 OBL 3300 ABCD Other 92 1480 721 C P. notatum 119 UPL 1370 560 CD Panicum 89 OBL 1550 763 C Polygonum 39 OBL 2190 678 B Pontederia 54 OBL 2140 1090 B Sagittaria 2 OBL 5880 A Beaty Utricularia 2 OBL 4240 AB < 0.01 Alternanthera 22 OBL 4360 844 W Eleocharis 3 OBL 3040 WXYZ Juncus 24 OBL 1470 514 Z Ludwigia 2 OBL 1790 XYZ Luziola + P. acuminatum 53 FACW 3360 731 X Other 84 2510 784 Y P. notatum 123 UPL 1640 633 Z Panicum 4 OBL 3180 689 WXYZ Polygonum 43 OBL 3130 1040 X Larson Pontederia 19 OBL 2990 1100 XY < 0.01

PAGE 116

101 Table B-11. Phosphorus storag e in above ground biomass AGB Species P storage by Site Site Species n Indicator (mg/m2) Difference p value Andropogon 4 FAC 19.4 4.26 A Baccopa 1 OBL 22.5 A Eleocharis 1 OBL 39.2 A Juncus 30 OBL 317 389 A Ludwigia 3 OBL 223 176 A Luziola + P. acuminatum 3 FACW 125 185 A Micranthemum 1 OBL 13.3 A Other 32 256 278 A P. notatum 39 UPL 261 145 A Panicum 31 OBL 324 331 A Polygonum 14 OBL 429 388 A Pontederia 19 OBL 182 219 A Sagittaria 1 OBL 37.2 A Beaty Utricularia 1 OBL 54.0 A 0.31 Alternanthera 10OBL 12.7 10.1 Y Eleocharis 2 OBL 149 XY Juncus 8 OBL 158 210 Y Ludwigia 1 OBL 103 XY Luziola + P. acuminatum 25 FACW 72.8 70.1 Y Other 30 261 278 Y P. notatum 42 UPL 341 208 XY Panicum 2 OBL 34.8 XY Polygonum 16 OBL 611 767 X Larson Pontederia 7 OBL 170 179 XY < 0.01

PAGE 117

102 APPENDIX C SUPPLEMENTAL MESOCOSM DATA Table C-1. Nutrient co ncentrations on day 0. Day 0 Concentrations Type Species Treatment nTP % TN % TC % Sand 80.004 0.003 . +10 30.003 0.001 0.087 0.004 1.52 0.106 0 30.003 0.001 0.092 0.005 1.69 0.064 -10 30.003 0.001 0.098 0.004 1.76 0.105 -15 30.003 0.001 0.088 0.004 1.56 0.088 Bahia C 30.004 0.001 0.089 0.005 1.60 0.110 +10 30.003 0.001 0.090 0.006 1.57 0.173 0 30.004 0.001 0.093 0.006 1.69 0.116 -10 30.004 0.000 0.094 0.004 1.66 0.077 -15 30.003 0.001 0.094 0.005 1.71 0.101 Soil Floralta C 30.004 0.001 0.093 0.003 1.71 0.078 +10 30.151 0.008 1.32 0.110 42.6 1.200 0 30.153 0.006 1.32 0.074 42.6 0.470 -10 30.149 0.004 1.26 0.044 42.9 0.880 -15 30.150 0.007 1.28 0.066 43.1 0.347 Bahia C 30.145 0.010 1.34 0.048 44.0 1.340 +10 30.166 0.015 1.35 0.079 42.7 0.174 0 30.179 0.033 1.33 0.188 41.9 0.755 -10 30.181 0.021 1.42 0.145 42.3 0.470 -15 30.180 0.001 1.40 0.096 42.3 0.389 Forage Floralta C 30.171 0.014 1.34 0.153 40.8 1.680

PAGE 118

103 Table C-2. Species comparison of forage production per harvest Forage Production Per Harvest (g/pot) Days n Bahia Floralta p value 27 15 4.02 0.79 <7.52 0.99 < 0.01 55 15 1.40 0.42 <1.97 0.66 0.01 83 15 0.92 0.44 <1.13 0.46 0.22 163 15 0.32 0.15 <1.55 0.54 < 0.01 305 15 0.56 0.39 <6.96 4.63 < 0.01 375 15 2.30 1.38 <13.3 10.1 < 0.01 Mean of species forage production regardless of treatment. Table C-3. Species comparison of cumulative forage production. Cumulative Forage Production (g/pot) Days n Bahia Floralta p value 27 15 4.03 0.79 <7.52 0.99 < 0.01 55 15 5.42 0.97 <9.49 0.98 < 0.01 83 15 6.34 1.21 <10.6 1.28 < 0.01 163 15 6.66 1.31 <12.2 1.39 < 0.01 305 15 7.22 1.46 <19.1 5.49 < 0.01 375 15 9.52 2.73 <32.3 14.7 < 0.01 Mean of species forage production regardless of treatment Table C-4. Overall below ground bi omass – all treatments combined Below Ground Biomass (g/pot) Days n Bahia Floralta p value 163 15 85.6 14.0 >38.0 9.07 < 0.01 375 15 79.6 20.8 >29.5 8.52 < 0.01

PAGE 119

104 Table C-5. Forage production per harvest. Forage Production Per Harvest (g/pot) Days Treatment nBahia Floralta p value +10 33.93 0.90 <8.12 1.39 0.01 0 34.06 0.45 <7.56 0.9 < 0.01 -10 33.87 0.27 <8.03 0.14 < 0.01 -15 33.59 0.80 <6.86 0.42 < 0.01 27 C 34.67 1.30 <7.02 1.41 0.10 +10 31.84 0.70 >1.73 0.34 0.81 0 31.21 0.42 <1.54 0.44 0.40 -10 31.28 0.29 <1.35 0.05 0.71 -15 31.24 0.09 <2.34 0.47 0.02 55 C 31.43 0.23 <2.92 0.11 < 0.01 +10 31.61 0.30 >1.15 0.34 0.15 0 30.76 0.25 <1.04 0.37 0.35 -10 30.68 0.35 <0.81 0.27 0.63 -15 30.58 0.22 <0.81 0.33 0.37 83 C 30.96 0.04 <1.82 0.13 < 0.01 +10 30.51 0.20 <2.36 0.39 < 0.01 0 30.27 0.04 <1.24 0.29 < 0.01 -10 30.20 0.11 <1.47 0.14 < 0.01 -15 30.32 0.07 <1.27 0.65 0.07 163 C 30.30 0.13 <1.42 0.31 < 0.01 +10 30.39 0.21 <13.7 2.67 < 0.01 0 30.43 0.40 <4.02 1.32 0.01 -10 30.28 0.06 <3.95 1.76 0.02 -15 30.77 0.30 <4.59 3.55 0.14 305 C 30.93 0.57 <8.56 4.38 0.04 +10 33.44 0.55 <17.5 1.84 < 0.01 0 31.21 0.25 <7.58 2.27 0.01 -10 31.58 0.82 <5.44 2.06 0.04 -15 31.44 0.29 <7.34 3.56 0.05 375 C 33.81 1.76 <28.4 11.3 0.02

PAGE 120

105 Table C-6. Cumulative fora ge by treatment and day. Cumulative Forage Production (g/pot) Days Treatment nBahia Floralta p value +10 33.93 0.90 <8.12 1.39 0.01 0 34.06 0.45 <7.56 0.90 < 0.01 -10 33.87 0.27 <8.03 0.14 < 0.01 -15 33.59 0.8 <6.86 0.42 < 0.01 27 C 34.67 1.30 <7.02 1.41 0.10 +10 35.77 1.40 <9.84 1.72 0.03 0 35.26 0.50 <9.09 0.46 < 0.01 -10 35.15 0.56 <9.38 0.11 < 0.01 -15 34.83 0.83 <9.19 0.84 < 0.01 55 C 36.10 1.34 <9.94 1.38 0.03 +10 37.38 1.70 <11.0 2.06 0.08 0 36.03 0.73 <10.1 0.39 < 0.01 -10 35.84 0.67 <10.2 0.38 < 0.01 -15 35.4 0.63 <10.0 1.1 < 0.01 83 C 37.06 1.32 <11.8 1.51 0.02 +10 37.89 1.88 <13.4 2.05 0.03 0 36.30 0.77 <11.4 0.66 < 0.01 -10 36.03 0.74 <11.7 0.51 < 0.01 -15 35.72 0.69 <11.3 0.84 < 0.01 163 C 37.37 1.30 <13.2 1.33 0.01 +10 38.28 1.85 <27.0 2.16 < 0.01 0 36.73 0.86 <15.4 1.96 < 0.01 -10 36.30 0.77 <15.6 1.28 < 0.01 -15 36.49 0.98 <15.9 3.81 0.01 305 C 38.30 1.85 <21.7 5.21 0.01 +10 311.7 2.36 <44.5 3.98 < 0.01 0 37.94 1.06 <23.0 4.24 < 0.01 -10 37.88 1.49 <21.1 3.31 < 0.01 -15 37.94 1.25 <23.2 7.33 0.02 375 C 312.1 3.61 <50.2 16.5 0.02

PAGE 121

106 Table C-7. Below ground bi omass species comparison. Below Ground Biomass (g/pot) Days Treatment nBahia Floralta p value +10 366.0 7.37 >27.5 3.30 < 0.01 0 389.9 16.0 >34.0 6.33 < 0.01 -10 391.1 5.40 >44.1 10.6 < 0.01 -15 396.9 4.33 >38.1 7.99 < 0.01 163 C 384.2 13.3 >46.3 0.60 0.01 +10 351.9 0.98 >19.8 4.52 < 0.01 0 393.7 21.1 >33.9 6.63 0.01 -10 395.2 13.6 >32.0 3.63 < 0.01 -15 369.0 11.2 >36.6 9.76 0.02 375 C 388.3 12.8 >24.9 6.91 < 0.01 Table C-8. Residual biomass harvested on days 163 and 375. Residual Biomass (g/pot) Days Treatment nBahia Floralta p value +10 311.1 2.66 <16.3 0.31 0.03 0 312.9 4.14 <17.7 2.79 0.17 -10 315.9 4.21 <16.3 1.75 0.89 -15 312.6 3.01 <15.9 2.8 0.24 163 C 313.4 4.09 <18.2 2.71 0.17 +10 313.0 0.82 <20.2 4.17 0.04 0 310.6 4.91 <19.6 2.53 0.05 -10 38.71 1.19 <17.5 6.98 0.10 -15 38.41 1.06 <16.2 5.13 0.06 375 C 312.3 5.31 <24.5 5.92 0.06 Table C-9. Below ground biomass production time comparison. Below Ground Biomass Production (g/pot) Species Treatment Day 163 Day 375 p value +10 66.0 7.37 >51.9 0.98 0.03 0 89.9 16.0 <93.7 21.1 0.81 -10 91.1 5.40 <95.2 13.6 0.65 -15 96.9 4.33 >69.0 11.2 0.02 Bahia C 84.2 13.3 <88.3 12.8 0.72 +10 27.5 3.30 >19.8 4.52 0.08 0 34.0 6.33 >33.9 6.63 0.99 -10 44.1 10.6 >32.0 3.63 0.13 -15 38.1 7.99 >36.6 9.76 0.86 Floralta C 46.3 0.60 >24.9 6.91 0.01

PAGE 122

107 BGB:AGB (163 Days) (110) (95) (80) (65) (50) (35) (20) (5) 10 25 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC S p ecies and TreatmentBiomass (g pot -1 ) Figure C-1. Relative root and shoot biomass after 163 days. Total Biomass (163 Days) 0 20 40 60 80 100 120 140 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC Treatment and SpeciesBiomass ( g pot -1 ) Figure C-2. Total biomass production after 163 days.

PAGE 123

108 Total Biomass (375 Days) 0 20 40 60 80 100 120 140 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC Treatment and SpeciesBiomass ( g pot -1 ) Figure C-3. Total biomass production after 375 days.

PAGE 124

109 Table C-10. Bahiagrass forage production per harvest treatment comparison using Tukey-Kramer HSD Test Bahia Forage Production Per Ha rvest (Tukey-Kramer HSD Test) Days Treatment nBahia Difference p value +10 33.93 0.90 A 0 34.06 0.45 A -10 33.87 0.27 A -15 33.59 0.80 A 27 C 34.67 1.30 A 0.61 +10 31.84 0.70 A 0 31.21 0.42 A -10 31.28 0.29 A -15 31.24 0.09 A 55 C 31.43 0.23 A 0.34 +10 31.61 0.30 A 0 30.76 0.25 B -10 30.68 0.35 B -15 30.58 0.22 B 83 C 30.96 0.04 AB < 0.01 +10 30.51 0.20 A 0 30.27 0.04 A -10 30.20 0.11 A -15 30.32 0.07 A 163 C 30.30 0.13 A 0.10 +10 30.39 0.21 A 0 30.43 0.40 A -10 30.28 0.06 A -15 30.77 0.30 A 305 C 30.93 0.57 A 0.19 +10 33.44 0.55 AB 0 31.21 0.25 A -10 31.58 0.82 AB -15 31.44 0.29 AB 375 C 33.81 1.76 B 0.01 In this table Bahia was compared to itself not to the other species.

PAGE 125

110 Table C-11. Bahiagrass cumulative forage production treatment comparison using Tukey-Kramer HSD Test Cumulative Bahia Forage Production (Tukey-Kramer HSD Test) Days Treatment nBahia Difference p value +10 33.93 0.90 A 0 34.06 0.45 A -10 33.87 0.27 A -15 33.59 0.8 A 27 C 34.67 1.30 A 0.60 +10 35.77 1.40 A 0 35.26 0.50 A -10 35.15 0.56 A -15 34.83 0.83 A 55 C 36.10 1.34 A 0.57 +10 37.38 1.70 A 0 36.03 0.73 A -10 35.84 0.67 A -15 35.4 0.63 A 83 C 37.06 1.32 A 0.21 +10 37.89 1.88 A 0 36.30 0.77 A -10 36.03 0.74 A -15 35.72 0.69 A 163 C 37.37 1.30 A 0.19 +10 38.28 1.85 A 0 36.73 0.86 A -10 36.30 0.77 A -15 36.49 0.98 A 305 C 38.30 1.85 A 0.25 +10 311.7 2.36 A 0 37.94 1.06 A -10 37.88 1.49 A -15 37.94 1.25 A 375 C 312.1 3.61 A 0.07 In this table Bahia was compared to itself not to the other species.

PAGE 126

111 Table C-12. Limpograss forage production per harvest treatment comparison using Tukey-Kramer HSD Test Floralta Forage Production Per Harvest (Tukey-Kramer HSD Test) Days Treatment nFloralta Difference p value +10 38.12 1.39 A 0 37.56 0.9 A -10 38.03 0.14 A -15 36.86 0.42 A 27 C 37.02 1.41 A 0.46 +10 31.73 0.34 AC 0 31.54 0.44 AC -10 31.35 0.05 A -15 32.34 0.47 BC 55 C 32.92 0.11 B < 0.01 +10 31.15 0.34 AB 0 31.04 0.37 AB -10 30.81 0.27 B -15 30.81 0.33 B 83 C 31.82 0.13 A 0.01 +10 32.36 0.39 B 0 31.24 0.29 A -10 31.47 0.14 AB -15 31.27 0.65 A 163 C 31.42 0.31 AB 0.03 +10 313.7 2.67 A 0 34.02 1.32 B -10 33.95 1.76 B -15 34.59 3.55 B 305 C 38.56 4.38 AB 0.01 +10 317.5 1.84 AB 0 37.58 2.27 B -10 35.44 2.06 B -15 37.34 3.56 B 375 C 328.4 11.3 A < 0.01 In this table Floralta was compared to itself not to the other species.

PAGE 127

112 Table C-13. Cumulative limpograss forage production treatment comparison using Tukey-Kramer HSD Test Cumulative Floralta Forage Production (Tukey-Kramer HSD Test) Days Treatment nFloralta Difference p value +10 38.12 1.39 A 0 37.56 0.90 A -10 38.03 0.14 A -15 36.86 0.42 A 27 C 37.02 1.41 A 0.46 +10 39.84 1.72 A 0 39.09 0.46 A -10 39.38 0.11 A -15 39.19 0.84 A 55 C 39.94 1.38 A 0.82 +10 311.0 2.06 A 0 310.1 0.39 A -10 310.2 0.38 A -15 310.0 1.1 A 83 C 311.8 1.51 A 0.43 +10 313.4 2.05 A 0 311.4 0.66 A -10 311.7 0.51 A -15 311.3 0.84 A 163 C 313.2 1.33 A 0.16 +10 327.0 2.16 A 0 315.4 1.96 B -10 315.6 1.28 B -15 315.9 3.81 B 305 C 321.7 5.21 AB < 0.01 +10 344.5 3.98 BC 0 323.0 4.24 AC -10 321.1 3.31 A -15 323.2 7.33 AC 375 C 350.2 16.5 B < 0.01 In this table Floralta was compared to itself not to the other species.

PAGE 128

113 Table C-14. Bahiagrass BGB production – treatment comparison using Tukey-Kramer HSD Bahia B.G. Production (T ukey-Kramer HSD Test) Days Treatment nBahia Difference p value +10 366.0 7.37 A 0 389.9 16.0 AB -10 391.1 5.40 AB -15 396.9 4.33 B 163 C 384.2 13.3 AB 0.04 +10 351.9 0.98 A 0 393.7 21.1 B -10 395.2 13.6 B -15 369.0 11.2 AB 375 C 388.3 12.8 B 0.01 Table C-15. Limpograss BGB production – treatment comp arison using Tukey-Kramer HSD Floralta B.G. Production (Tukey-Kramer HSD Test) Days Treatment nFloralta Difference p value +10 327.5 3.30 A 0 334.0 6.33 AB -10 344.1 10.6 AB -15 338.1 7.99 AB 163 C 346.3 0.60 B 0.04 +10 319.8 4.52 A 0 333.9 6.63 A -10 332.0 3.63 A -15 336.6 9.76 A 375 C 324.9 6.91 A 0.06

PAGE 129

114 Table C-16. Forage P concentr ations – species comparison. Forage Phosphorus Concentrations (mg/kg) Day Treatment Bahia Floralta p value +10 1,510 83.0 <1,660 143 0.68 0 1,530 58.0 <1,790 331 0.04 -10 1,490 36.0 <1,810 210 0.13 -15 1,500 73.0 <1,800 13.0 0.75 0 C 1,454 98.0 <1,780 135 0.43 +10 1,600 24.0 >965 220 0.01 0 1,220 62.0 >1,010 146 0.08 -10 1,150 11.0 >967 212 0.21 -15 1,180 60.0 >1,080 72 0.12 27 C 1,210 55.0 <1,260 152 0.65 +10 1,520 117 >1,110 103 0.01 0 1,180 57.0 <1,360 110 0.06 -10 1,230 37.0 <1,280 95 0.48 -15 1,200 15.0 <1,230 88 0.56 55 C 1,280 92.0 <1,320 108 0.62 +10 1,670 34.0 >1,550 204 0.35 0 1,160 80.0 <1,360 149 0.13 -10 1,190 34.0 <1,420 189 0.10 -15 1,190 28.0 <1,440 54.0 < 0.01 83 C 1,190 66.0 <1,490 204 0.07 +10 1,400 195 <1,410 64.0 0.98 0 1,030 129 <1,030 248 0.98 -10 877 28.0 <1,010 150 0.22 -15 942 100 >789 154 0.22 163 C 817 74.0 <1,070 112 0.03 +10 1,150 105 >712 41.0 < 0.01 0 478 80.0 <708 155 0.09 -10 718 44.0 >683 75.0 0.51 -15 976 295 >580 41.0 0.08 375 C 1,130 121 >587 73.0 < 0.01

PAGE 130

115 Table C-17. Bahiagrass forage P concentra tions treatment comparison using TukeyKramer HSD. Bahia Forage Phosphorus Conc. (mg/kg) Tukey-Kramer Day Treatment Bahia Difference p value +10 1,510 83.0 A 0 1,530 58.0 A -10 1,490 36.0 A -15 1,500 73.0 A 0 C 1,454 98.0 A 0.73 +10 1,600 24.0 A 0 1,220 62.0 B -10 1,150 11.0 B -15 1,180 60.0 B 27 C 1,210 55.0 B < 0.01 +10 1,5120 117 A 0 1,180 57.0 B -10 1,230 37.0 B -15 1,200 15.0 B 55 C 1,280 92.0 B < 0.01 +10 1,670 34.0 A 0 1,160 80.0 B -10 1,190 34.0 B -15 1,190 28.0 B 83 C 1,190 66.0 B < 0.01 +10 1,400 195 A 0 1,030 129 B -10 877 28.0 B -15 942 100 B 163 C 817 74.0 B < 0.01 +10 1,150 105 A 0 478 80.0 B -10 718 44.0 BC -15 976 295 AC 375 C 1,130 121 AC < 0.01 In this table Bahia was compared to itself not to the other species.

PAGE 131

116 Table C-18. Limpograss forage P concentra tions treatment comparison using TukeyKramer HSD. Floralta Forage Phosphorus Conc. (mg/kg) Tukey-Kramer Day Treatment Floralta Difference p value +10 1,660 143 A 0 1,790 331 A -10 1,810 210 A -15 1,800 13.0 A 0 C 1,780 135 A 0.84 +10 965 220 A 0 1,010 146 A -10 967 212 A -15 1,080 72 A 27 C 1,260 152 A 0.26 +10 1,110 103 A 0 1,360 110 A -10 1,280 95 A -15 1,230 88 A 55 C 1,320 108 A 0.10 +10 1,550 204 A 0 1,360 149 A -10 1,420 189 A -15 1,440 54.0 A 83 C 1,490 204 A 0.71 +10 1,410 64.0 A 0 1,030 248 AB -10 1,010 150 AB -15 789 154 B 163 C 1,070 112 AB 0.01 +10 712 41.0 A 0 708 155 A -10 683 75.0 A -15 580 41.0 A 375 C 587 73.0 A 0.23 In this table Floralta was compared to itself not to the other species

PAGE 132

117 Table C-19. Below ground biomass P c oncentrations – species comparison. BGB Phosphorus Concentrations (mg/kg) Day Treatment Bahia Floralta p value +10 430 45.0 <592 115 0.08 0 432 77.0 <609 35.0 0.03 -10 441 56.0 <472 23.0 0.42 -15 433 68.0 <505 157 0.53 163 C 430 98.0 <668 11.0 0.01 +10 377 42.0 >320 48.0 0.23 0 375 60.0 >304 16.0 0.12 -10 343 26.0 >333 15.0 0.70 -15 303 17.0 <345 149 0.62 375 C 395 41.0 >383 29.0 0.67 Table C-20. Bahiagrass BGB P concentrati ons treatment comparison using TukeyKramer HSD. Bahia BGB Phosphorus Conc. (mg/kg) Tukey-Kramer Day Treatment Bahia Difference p value +10 430 45.0 A 0 432 77.0 A -10 441 56.0 A -15 433 68.0 A 163 C 430 98.0 A 0.99 +10 377 42.0 A 0 375 60.0 A -10 343 26.0 A -15 303 17.0 A 375 C 395 41.0 A 0.12

PAGE 133

118 Table C-21. Limpograss BGB P concentratio ns treatment comparison using TukeyKramer HSD. Floralta BGB Phosphorus Conc. (mg/kg) Tukey-Kramer Day Treatment Floralta Difference p value +10 592 115 A 0 609 35.0 A -10 472 23.0 A -15 505 157 A 163 C 668 11.0 A 0.12 +10 320 48.0 A 0 304 16.0 A -10 333 15.0 A -15 345 149 A 375 C 383 29.0 A 0.72

PAGE 134

119 Table C-22. Phosphorus storage in fora ge species comparison per harvest. Forage P Assimilation per Harvest (mg/pot) Day Treatment nBahia Floralta p value +10 36.28 1.50 <7.72 1.71 0.34 0 34.95 0.34 <7.60 0.95 0.01 -10 34.44 0.28 <7.77 1.70 0.03 -15 34.21 0.74 <7.39 0.84 0.01 27 C 35.62 1.28 <8.72 0.95 0.03 +10 32.81 1.16 >1.93 0.48 0.29 0 31.41 0.44 <2.06 0.45 0.15 -10 31.58 0.34 <1.72 0.15 0.53 -15 31.49 0.12 <2.86 0.35 < 0.01 55 C 31.84 0.39 <3.85 0.21 < 0.01 +10 32.69 0.55 >1.75 0.47 0.09 0 30.88 0.25 <1.37 0.38 0.14 -10 30.81 0.43 <1.15 0.38 0.36 -15 30.68 0.25 <1.16 0.47 0.20 83 C 31.15 0.07 <2.70 0.28 < 0.01 +10 30.70 0.25 <3.30 0.42 < 0.01 0 30.29 0.08 <1.26 0.36 0.01 -10 30.17 0.09 <1.47 0.10 < 0.01 -15 30.30 0.09 <0.98 0.43 0.57 163 C 30.24 0.08 <1.51 0.35 < 0.01 +10 30.44 0.21 <9.79 2.24 < 0.01 0 30.20 0.18 <2.72 0.25 < 0.01 -10 30.20 0.04 <2.71 1.22 0.02 -15 30.80 0.53 <2.69 2.14 0.21 305 C 31.03 0.63 <4.82 2.13 0.04 +10 33.98 1.03 <12.4 1.43 < 0.01 0 30.59 0.22 <5.15 0.48 < 0.01 -10 31.13 0.55 <3.77 1.66 0.06 -15 31.47 0.76 <4.24 2.11 0.10 375 C 34.24 1.90 <16.1 5.03 0.02

PAGE 135

120 Table C-23. Cumulative P storage in forage species comparison Cumulative Forage Tissue Phosphorus Asssimilation (mg/pot) Day Treatment nBahia Floralta p value +10 36.28 1.50 <7.72 1.71 0.34 0 34.95 0.34 <7.60 0.95 0.01 -10 34.44 0.28 <7.77 1.70 0.03 -15 34.21 0.74 <7.39 0.84 0.01 27 C 35.62 1.28 <8.72 0.95 0.03 +10 39.09 2.41 <9.65 2.12 0.78 0 36.36 0.32 <9.66 0.94 0.00 -10 36.02 0.62 <9.49 1.84 0.04 -15 35.69 0.78 <10.3 1.02 < 0.01 55 C 37.47 1.45 <12.6 0.76 0.01 +10 311.8 2.96 >11.4 2.52 0.87 0 37.24 0.57 <11.0 1.31 0.01 -10 36.83 0.76 <10.6 2.07 0.04 -15 36.38 0.57 <11.4 1.27 < 0.01 83 C 38.61 1.50 <15.3 0.64 < 0.01 +10 312.5 3.15 <14.7 2.15 0.37 0 37.52 0.64 <12.3 1.04 0.00 -10 37.00 0.81 <12.1 2.00 0.01 -15 36.68 0.65 <12.4 0.84 < 0.01 163 C 38.85 1.49 <16.8 0.63 < 0.01 +10 312.9 3.16 <24.5 3.53 0.01 0 37.72 0.63 <15.0 1.14 < 0.01 -10 37.20 0.82 <14.8 2.73 0.01 -15 37.48 1.17 <15.1 1.67 0.00 305 C 39.89 2.12 <21.6 2.66 0.00 +10 316.9 4.11 <36.9 4.94 0.01 0 38.31 0.84 <20.2 1.27 < 0.01 -10 38.32 1.25 <18.6 4.34 0.02 -15 38.95 1.93 <19.3 3.78 0.01 375 C 314.1 4.01 <37.7 7.64 0.01

PAGE 136

121 Table C-24. Bahiagrass fora ge P storage per harvest Forage P Assimilation per Harvest Tukey-Kramer (mg/pot) Day Treatment nBahia Difference p value +10 36.28 1.50 A 0 34.95 0.34 A -10 34.44 0.28 A -15 34.21 0.74 A 27 C 35.62 1.28 A 0.12 +10 32.81 1.16 A 0 31.41 0.44 A -10 31.58 0.34 A -15 31.49 0.12 A 55 C 31.84 0.39 A 0.09 +10 32.69 0.55 A 0 30.88 0.25 B -10 30.81 0.43 B -15 30.68 0.25 B 83 C 31.15 0.07 B < 0.01 +10 30.70 0.25 A 0 30.29 0.08 B -10 30.17 0.09 B -15 30.30 0.09 B 163 C 30.24 0.08 B 0.01 +10 30.44 0.21 A 0 30.20 0.18 A -10 30.20 0.04 A -15 30.80 0.53 A 305 C 31.03 0.63 A 0.09 +10 33.98 1.03 A 0 30.59 0.22 B -10 31.13 0.55 B -15 31.47 0.76 AB 375 C 34.24 1.90 A < 0.01 In this table Bahia was compared to itself not to the other species.

PAGE 137

122 Table C-25. Bahiagrass cumulative forage P storage. Cumulative Forage P Assimilation Tukey-Kramer (mg/pot) Day Treatment nBahia Difference p value +10 36.28 1.50 A 0 34.95 0.34 A -10 34.44 0.28 A -15 34.21 0.74 A 27 C 35.62 1.28 A 0.12 +10 39.09 2.41 A 0 36.36 0.32 A -10 36.02 0.62 A -15 35.69 0.78 A 55 C 37.47 1.45 A 0.06 +10 311.8 2.96 A 0 37.24 0.57 B -10 36.83 0.76 B -15 36.38 0.57 B 83 C 38.61 1.50 AB 0.01 +10 312.5 3.15 A 0 37.52 0.64 B -10 37.00 0.81 B -15 36.68 0.65 B 163 C 38.85 1.49 AB 0.01 +10 312.9 3.16 A 0 37.72 0.63 B -10 37.20 0.82 B -15 37.48 1.17 B 305 C 39.89 2.12 AB 0.02 +10 316.9 4.11 A 0 38.31 0.84 B -10 38.32 1.25 B -15 38.95 1.93 B 375 C 314.1 4.01 AB 0.01 In this table Bahia was compared to itself not to the other species.

PAGE 138

123 Table C-26. Limpograss forage P storage per harvest. Forage P Assimilation per Harvest Tukey-Kramer (mg/pot) Day Treatment nFloralta Difference p value +10 37.72 1.71 A 0 37.60 0.95 A -10 37.77 1.70 A -15 37.39 0.84 A 27 C 38.72 0.95 A 0.76 +10 31.93 0.48 AC 0 32.06 0.45 AC -10 31.72 0.15 A -15 32.86 0.35 C 55 C 33.85 0.21 B < 0.01 +10 31.75 0.47 AB 0 31.37 0.38 B -10 31.15 0.38 B -15 31.16 0.47 B 83 C 32.70 0.28 A < 0.01 +10 33.30 0.42 A 0 31.26 0.36 B -10 31.47 0.10 B -15 30.98 0.43 B 163 C 31.51 0.35 B < 0.01 +10 39.79 2.24 A 0 32.72 0.25 B -10 32.71 1.22 B -15 32.69 2.14 B 305 C 34.82 2.13 B < 0.01 +10 312.4 1.43 A 0 35.15 0.48 B -10 33.77 1.66 B -15 34.24 2.11 B 375 C 316.1 5.03 A < 0.01 In this table Floralta was compared to itself not to the other species.

PAGE 139

124 Table C-27. Cumulative limpograss forage P storage Cumulative Forage P Assimilation Tukey-Kramer (mg/pot) Day Treatment nFloralta Difference p value +10 37.72 1.71 A 0 37.60 0.95 A -10 37.77 1.70 A -15 37.39 0.84 A 27 C 38.72 0.95 A 0.76 +10 39.65 2.12 A 0 39.66 0.94 A -10 39.49 1.84 A -15 310.3 1.02 A 55 C 312.6 0.76 A 0.12 +10 311.4 2.52 AB 0 311.0 1.31 AB -10 310.6 2.07 B -15 311.4 1.27 AB 83 C 315.3 0.64 A 0.04 +10 314.7 2.15 AB 0 312.3 1.04 B -10 312.1 2.00 B -15 312.4 0.84 B 163 C 316.8 0.63 A < 0.01 +10 324.5 3.53 A 0 315.0 1.14 BC -10 314.8 2.73 B -15 315.1 1.67 BC 305 C 321.6 2.66 AC < 0.01 +10 336.9 4.94 A 0 320.2 1.27 B -10 318.6 4.34 B -15 319.3 3.78 B 375 C 337.7 7.64 A < 0.01 In this table Floralta was compared to itself not to the other species.

PAGE 140

125 A. B. Figure C-4. Bahiagrass total biomass and P storage. A) biomass B) P storage. A. B. Figure C-5. Limpograss total biomass and P storage. A) biomass B) P storage.

PAGE 141

126 A. B. Figure C-6. Bahiagrass BG B and P storage. A) biomass, B) P storage A. B. Figure C-7. Limpograss BGB and P stor age. A) biomass, B) P storage

PAGE 142

127 Table C-28. Below ground biomass P storage species comparison BGB Tissue Phosphorus Asssimilation (mg/pot) Day Treatment Bahia Floralta p value +10 28.2 1.57 >16.3 3.80 0.01 0 38.1 4.74 >20.7 3.60 0.01 -10 40.0 4.37 >20.7 4.12 0.01 -15 41.8 5.47 >18.5 2.89 < 0.01 163 C 35.8 7.86 >30.9 0.73 0.35 +10 19.5 1.88 >6.30 1.43 < 0.01 0 35.9 14.2 >10.4 2.34 0.04 -10 32.5 2.98 >10.6 0.78 < 0.01 -15 21.1 4.70 >12.4 5.30 0.10 375 C 35.0 6.72 >9.49 2.28 < 0.01 Table C-29. Bahiagrass belo w ground biomass P storage BGB Tissue Phosphorus Asssimilation (mg/pot) Day Treatment Bahia Difference p value +10 28.2 1.57 A 0 38.1 4.74 A -10 40.0 4.37 A -15 41.8 5.47 A 163 C 35.8 7.86 A 0.07 +10 19.5 1.88 A 0 35.9 14.2 A -10 32.5 2.98 A -15 21.1 4.70 A 375 C 35.0 6.72 A 0.06 Table C-30. Limpograss belo w ground biomass P storage BGB Tissue Phosphorus Asssimilation (mg/pot) Day Treatment Floralta Difference p value +10 16.3 3.80 A 0 20.7 3.60 A -10 20.7 4.12 A -15 18.5 2.89 A 163 C 30.9 0.73 B < 0.01 +10 6.30 1.43 A 0 10.4 2.34 A -10 10.6 0.78 A -15 12.4 5.30 A 375 C 9.49 2.28 A 0.20

PAGE 143

128 TP Storage (163 Days) (50) (30) (10) 10 30 50 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC Treatment and SpeciesTP ( mg pot -1 ) Figure C-8. Root to shoot P storage ratios TP Storage (163 Days) 0 10 20 30 40 50 60 70 BahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloraltaBahiaFloralta +10+1000-10-10-15-15CC Treatment and SpeciesTP ( mg pot -1 ) Figure C-9. Total P storag e (AGB +BGB) at 163 days.

PAGE 144

129 Table C-31. Climatic conditions from day 1 to 375. Climatic Conditions (FAWN) Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 7/1/2004 1 24.8 19.2 33.9 28. 3 27.3 30.3 1.38 84% 7/2/2004 2 26.4 19.6 34.0 28. 6 26.0 31.5 0.00 77% 7/3/2004 3 25.0 20.2 32.1 28. 7 27.1 30.3 0.11 85% 7/4/2004 4 25.2 19.9 33.2 28. 4 26.3 30.9 0.00 83% 7/5/2004 5 27.3 20.6 34.0 29. 2 26.6 31.8 0.00 77% 7/6/2004 6 28.1 21.4 35.2 30. 0 27.7 32.4 0.00 74% 7/7/2004 7 26.1 20.1 34.0 29. 8 27.9 31.6 1.51 80% 7/8/2004 8 23.9 19.7 32.1 28. 0 26.5 30.2 0.15 88% 7/9/2004 9 26.4 18.2 34.2 28. 5 25.7 31.5 0.00 77% 7/10/2004 10 27.5 20.1 35.3 29.7 27.2 32.4 0.00 74% 7/11/2004 11 25.3 20.7 33.8 29.1 27.3 31.7 1.81 83% 7/12/2004 12 24.8 20.3 34.1 28.2 26.1 30.8 0.01 87% 7/13/2004 13 25.3 20.7 33.0 28.4 26.6 30.8 0.00 85% 7/14/2004 14 27.5 21.4 32.9 29.2 26.7 31.9 0.00 76% 7/15/2004 15 25.9 21.5 31.8 28.9 27.9 29.9 0.28 84% 7/16/2004 16 25.3 21.4 33.2 28.6 26.7 31.4 0.35 82% 7/17/2004 17 24.7 21.1 31.3 28.2 26.6 30.1 0.33 87% 7/18/2004 18 23.9 22.5 26.4 27.1 26.6 27.8 0.74 92% 7/19/2004 19 25.3 22.3 31.8 27.7 26.0 30.1 0.56 87% 7/20/2004 20 25.9 22.0 33.1 28.3 26.6 30.6 0.57 86% 7/21/2004 21 26.7 21.0 33.4 28.8 26.7 31.0 0.00 76% 7/22/2004 22 26.4 18.5 34.0 28.7 26.4 30.9 0.00 74% 7/23/2004 23 27.0 20.6 34.2 29.0 27.1 30.7 0.00 77% 7/24/2004 24 26.4 22.2 35.1 29.0 27.6 30.9 0.02 82% 7/25/2004 25 26.9 21.1 33.5 28.8 27.1 30.7 0.00 76% 7/26/2004 26 27.1 19.8 34.7 29.2 27.2 31.3 0.49 74% 7/27/2004 27 26.1 21.3 34.9 29.2 27.5 31.8 0.94 83% 7/28/2004 28 25.6 21.0 32.7 28.8 27.2 30.8 0.67 87% 7/29/2004 29 26.9 21.8 33.5 28.6 27.3 30.0 0.00 79% 7/30/2004 30 27.0 21.5 33.1 28.7 27.3 30.4 0.00 76% 7/31/2004 31 25.4 20.0 31.8 28.2 26.9 29.3 0.00 85% 8/1/2004 32 26.1 21.7 33.6 28. 3 27.1 29.6 0.04 85% 8/2/2004 33 26.8 22.7 32.8 28. 7 27.7 30.6 0.84 85% 8/3/2004 34 28.0 23.3 34.1 29. 2 27.6 31.4 0.00 78% 8/4/2004 35 27.3 23.3 33.8 29. 5 27.9 31.3 0.00 81% 8/5/2004 36 28.1 22.9 34.0 29. 6 27.9 31.5 0.00 77% 8/6/2004 37 26.8 22.7 33.7 29. 6 28.4 31.1 0.39 83% 8/7/2004 38 23.8 20.4 26.9 27. 9 27.3 29.0 0.00 75% 8/8/2004 39 25.0 21.8 29.3 27. 4 26.5 28.3 0.00 79% 8/9/2004 40 25.7 22.3 30.7 27. 9 26.8 29.1 0.00 80% 8/10/2004 41 24.5 20.1 29.8 27.6 26.6 28.9 0.92 86% 8/11/2004 42 25.2 20.2 31.6 27.6 26.2 29.1 0.01 85% 8/12/2004 43 25.0 22.6 28.2 27.4 26.9 27.8 0.49 88%

PAGE 145

130 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 8/13/2004 44 23.1 21.2 24.7 26.4 25.9 27.2 1.48 94% 8/14/2004 45 23.2 20.2 26.6 25.8 25.2 26.5 0.03 92% 8/15/2004 46 24.8 21.3 29.9 26.4 25.6 27.5 0.00 86% 8/16/2004 47 26.3 20.0 33.4 27.2 25.6 28.8 0.00 80% 8/17/2004 48 26.8 21.7 34.4 28.1 26.9 29.7 0.44 82% 8/18/2004 49 26.0 20.0 33.5 28.0 26.6 29.4 0.31 83% 8/19/2004 50 27.3 21.2 34.5 28.4 26.9 30.0 0.00 78% 8/20/2004 51 26.9 22.1 34.9 28.7 27.5 30.3 0.00 79% 8/21/2004 52 24.2 21.2 31.7 27.7 27.2 28.5 0.51 89% 8/22/2004 53 24.8 21.1 32.8 27.4 26.4 28.2 0.13 89% 8/23/2004 54 25.1 21.3 31.8 27.6 26.5 29.0 0.13 87% 8/24/2004 55 24.9 20.2 32.6 27.5 26.4 28.9 0.00 86% 8/25/2004 56 26.2 22.5 32.4 28.0 26.9 29.3 0.00 82% 8/26/2004 57 26.1 20.9 33.4 28.1 26.9 29.3 0.13 83% 8/27/2004 58 26.5 21.9 32.8 28.2 27.0 29.5 0.00 81% 8/28/2004 59 26.3 20.4 33.3 28.1 26.8 29.2 0.00 79% 8/29/2004 60 26.8 21.4 33.2 28.3 27.0 29.5 0.00 79% 8/30/2004 61 26.6 21.9 33.2 28.3 27.1 29.8 0.00 78% 8/31/2004 62 24.8 21.1 31.5 27.9 27.1 28.7 0.00 88% 9/1/2004 63 26.2 20.3 35.1 28. 2 26.7 30.1 0.02 79% 9/2/2004 64 24.6 20.9 33.0 27. 9 27.0 29.4 0.85 87% 9/3/2004 65 25.9 22.3 30.5 27. 6 26.6 28.8 0.19 84% 9/4/2004 66 26.6 23.4 31.6 27. 7 26.8 28.7 0.02 82% 9/5/2004 67 24.8 22.2 26.7 26. 6 25.5 27.6 2.28 86% 9/6/2004 68 24.0 23.0 25.1 25. 2 24.9 25.5 2.20 90% 9/7/2004 69 25.5 23.3 29.8 25. 5 24.8 26.7 0.94 87% 9/8/2004 70 27.2 24.5 32.0 26. 8 25.7 28.2 0.00 82% 9/9/2004 71 26.4 23.3 33.8 27. 7 26.5 29.3 0.71 85% 9/10/2004 72 24.6 22.3 29.4 27.3 26.8 27.9 0.18 91% 9/11/2004 73 25.6 23.1 30.4 27.3 26.6 28.1 0.00 86% 9/12/2004 74 24.2 22.4 29.4 27.0 26.4 27.4 0.27 90% 9/13/2004 75 24.2 21.6 30.1 26.7 26.1 27.3 0.23 90% 9/14/2004 76 23.4 23.0 25.6 26.3 26.1 26.6 0.00 93% 9/20/2004 77 21.7 20.8 23.2 25.0 24.6 25.3 0.00 79% 9/21/2004 78 21.7 19.3 26.7 24.8 24.1 25.6 0.21 87% 9/22/2004 79 23.9 19.6 30.7 25.3 24.3 26.6 0.00 78% 9/23/2004 80 23.8 17.3 31.2 25.6 24.6 26.8 0.00 79% 9/24/2004 81 22.3 15.5 29.1 25.1 23.9 26.2 0.00 80% 9/25/2004 82 23.9 22.0 28.1 25.3 24.7 26.0 0.06 84% 9/26/2004 83 24.1 22.2 26.2 24.7 24.4 25.1 4.63 89% 9/27/2004 84 23.1 22.7 23.4 24.2 24.0 24.3 0.06 89% 9/28/2004 85 28.3 25.5 29.8 27.3 27.1 27.3 0.00 62% 9/29/2004 86 26.1 20.2 31.0 26.8 25.1 28.0 0.00 75% 9/30/2004 87 23.9 18.4 31.1 26.3 24.8 27.7 0.00 82% 10/1/2004 88 24.3 19.1 32.6 26.3 25.0 27.9 0.00 85% 10/2/2004 89 25.2 19.9 31.8 26.4 25.2 27.6 0.00 84%

PAGE 146

131 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 10/3/2004 90 25.2 20.7 32.0 26.9 25.8 28.0 0.00 82% 10/4/2004 91 24.3 18.9 30.8 26.3 25.0 27.5 0.00 82% 10/5/2004 92 23.3 18.6 30.1 26.2 25.3 27.3 0.00 80% 10/6/2004 93 22.9 18.5 29.5 25.3 24.6 26.1 0.00 78% 10/7/2004 94 21.5 17.4 27.9 24.6 23.9 25.5 0.04 76% 10/8/2004 95 21.6 17.1 28.1 24.2 23.1 25.2 0.00 80% 10/9/2004 96 23.1 18.1 28.8 24.5 23.6 25.5 0.00 80% 10/10/2004 97 23.5 19.4 28.9 24.8 24.0 25.7 0.00 80% 10/11/2004 98 21.0 17.8 22.8 24.4 23.7 24.9 0.26 92% 10/12/2004 99 22.1 17.1 28.7 24.1 23.0 25.4 0.00 86% 10/13/2004 100 21.7 15.2 28.4 24.5 23.5 26.0 0.00 80% 10/14/2004 101 19.8 11.8 27.0 23.6 22.0 25.1 0.07 75% 10/15/2004 102 16.5 8.0 20.1 23.3 21.9 24.4 0.61 79% 10/16/2004 103 16.8 7.2 26.4 21.7 19.9 23.6 0.00 75% 10/17/2004 104 20.3 12.9 29.3 22.4 20.8 24.3 0.00 79% 10/18/2004 105 20.4 13.6 27.7 22.7 21.5 23.9 0.00 82% 10/19/2004 106 23.4 16.7 30.8 23.4 22.0 25.0 0.00 83% 10/20/2004 107 24.0 19.9 29.8 24.7 23.7 26.0 0.18 86% 10/21/2004 108 21.1 17.0 26.6 24.0 23.5 24.9 0.00 91% 10/22/2004 109 19.9 15.7 26.2 23.3 22.4 24.4 0.00 83% 10/23/2004 110 18.5 12.4 26.3 22.6 21.6 23.7 0.00 78% 10/24/2004 111 18.8 10.9 27.7 22.3 20.7 24.0 0.00 82% 10/25/2004 112 20.4 15.4 28.2 23.1 21.8 24.6 0.00 82% 10/26/2004 113 19.0 13.7 26.9 22.6 21.5 23.8 0.00 82% 10/27/2004 114 18.8 12.6 27.2 22.2 20.9 23.6 0.00 81% 10/28/2004 115 20.2 13.8 27.9 22.4 21.1 23.8 0.00 86% 10/29/2004 116 23.0 19.0 29.7 23.6 22.4 25.3 0.08 85% 10/30/2004 117 22.6 18.0 30.2 24.1 23.1 25.6 0.00 87% 10/31/2004 118 22.0 16.6 29.6 23.8 22.6 25.1 0.00 86% 11/1/2004 119 22.6 16.0 29.3 23.6 22.4 24.8 0.00 82% 11/2/2004 120 23.1 18.5 30.0 24.0 22.9 25.3 0.00 81% 11/3/2004 121 22.2 16.4 30.4 23.7 22.5 25.0 0.00 81% 11/4/2004 122 22.1 15.3 29.2 23.5 22.1 24.7 0.15 84% 11/5/2004 123 15.3 9.3 21.6 22.4 21.0 23.7 0.00 76% 11/6/2004 124 13.6 7.0 23.6 20.4 19.0 21.9 0.00 78% 11/7/2004 125 13.3 5.2 24.1 19.6 18.1 21.2 0.00 73% 11/8/2004 126 15.6 6.9 26.6 19.4 17.7 21.2 0.00 76% 11/9/2004 127 16.0 11.1 22.1 19.5 18.8 20.0 0.00 74% 11/10/2004 128 17.2 11.4 26.0 19.4 18.2 20.7 0.00 76% 11/11/2004 129 17.7 10.7 25.6 19.7 18.5 20.9 0.00 82% 11/12/2004 130 19.4 14.9 26.5 20.5 19.6 21.5 0.04 85% 11/13/2004 131 19.4 15.4 24.6 20.8 20.0 21.9 0.00 87% 11/14/2004 132 15.1 13.8 16.4 19.8 19.1 20.7 0.00 86% 11/15/2004 133 15.2 9.9 21.8 19.0 18.0 20.0 0.00 79% 11/16/2004 134 13.5 6.5 23.7 18.7 17.4 20.1 0.00 77% 11/17/2004 135 12.5 4.9 22.9 18.1 16.8 19.4 0.00 74%

PAGE 147

132 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 11/18/2004 136 13.7 4.6 24.6 17.8 16.3 19.2 0.00 79% 11/19/2004 137 17.2 9.3 26.5 18.7 17.3 20.3 0.00 79% 11/20/2004 138 19.8 16.1 23.8 19.9 19.2 20.8 0.02 84% 11/21/2004 139 20.9 14.1 28.1 20.7 19.5 22.1 0.00 79% 11/22/2004 140 21.7 15.5 27.0 20.9 19.2 21.8 0.00 78% 11/23/2004 141 19.1 14.1 26.7 20.7 19.6 22.0 0.00 83% 11/24/2004 142 20.8 14.8 25.9 20.6 19.8 21.5 1.75 84% 11/25/2004 143 16.3 6.6 21.8 20.6 19.1 21.7 0.10 73% 11/26/2004 144 9.0 1.7 16.0 17.5 16.5 19.0 0.00 77% 11/27/2004 145 14.5 7.9 22.5 17.4 16.3 18.6 0.57 87% 11/28/2004 146 13.4 6.0 16.6 18.1 17.2 19.0 0.00 91% 11/29/2004 147 13.5 4.0 23.7 17.0 15.4 18.4 0.00 84% 11/30/2004 148 17.6 13.5 26.0 18.4 17.4 19.8 0.00 86% 12/1/2004 149 15.9 7.6 20.7 18.7 18.0 19.6 0.00 84% 12/2/2004 150 11.5 3.7 21.6 17.2 15.7 18.6 0.00 78% 12/3/2004 151 10.1 4.3 17.6 16.5 15.5 17.5 0.00 73% 12/4/2004 152 9.7 3.5 14.6 16.0 15.3 16.7 0.02 86% 12/5/2004 153 10.7 1.7 20.0 15.1 13.7 16.4 0.00 83% 12/6/2004 154 17.1 9.8 26.5 16.9 15.4 18.9 0.00 85% 12/7/2004 155 20.6 15.8 26.5 18.3 17.2 19.8 0.00 85% 12/8/2004 156 20.9 15.3 27.8 19.6 18.3 21.3 0.00 86% 12/9/2004 157 21.5 15.0 27.5 19.8 18.7 21.1 0.00 84% 12/10/2004 158 20.8 15.1 24.4 20.4 19.3 21.1 0.74 76% 12/11/2004 159 10.4 4.3 15.6 17.8 16.2 19.2 0.00 67% 12/12/2004 160 8.1 2.1 16.0 15.5 14.3 16.8 0.00 73% 12/13/2004 161 13.9 4.4 22.2 15.4 13.9 17.0 0.00 82% 12/14/2004 162 6.5 1.1 11.5 14.8 13.4 16.1 0.00 50% 12/15/2004 163 2.1 -3.1 10.2 12.4 11.4 13.4 0.00 61% 12/16/2004 164 7.8 -0.1 14.6 12.2 11.0 13.7 0.00 72% 12/17/2004 165 10.1 3.3 17.4 13.3 12.1 14.6 0.00 84% 12/18/2004 166 10.6 4.1 19.3 14.1 12.9 15.6 0.00 81% 12/19/2004 167 8.6 0.9 17.5 13.3 11.9 14.6 0.00 66% 12/20/2004 168 2.8 -2.4 9.6 12.1 11.1 13.1 0.00 50% 12/21/2004 169 7.0 -3.2 19.3 11.7 10.1 13.6 0.00 77% 12/22/2004 170 14.9 4.3 23.9 13.4 11.9 15.3 0.00 74% 12/23/2004 171 18.7 11.0 23.9 16.0 14.9 17.4 0.20 82% 12/24/2004 172 6.3 4.6 10.7 14.6 13.2 16.5 0.13 87% 12/25/2004 173 5.0 3.9 5.7 12.2 11.4 13.2 0.50 95% 12/26/2004 174 3.5 -1.6 7.9 11.0 10.2 11.6 0.23 89% 12/27/2004 175 5.1 -2.7 15.3 10.3 8.8 11.8 0.00 73% 12/28/2004 176 9.3 1.3 18.8 11.2 9.6 12.9 0.00 81% 12/29/2004 177 11.8 2.8 21.9 12.7 11.1 14.4 0.00 81% 12/30/2004 178 12.5 4.6 22.0 13.4 11.9 14.8 0.00 77% 12/31/2004 179 12.6 6.2 21.7 13.7 12.5 14.9 0.00 87% 1/1/2005 180 15.6 8.0 23.3 14. 5 13.1 16.1 0.00 82% 1/2/2005 181 16.7 10.2 24.8 15. 9 14.9 17.1 0.00 81%

PAGE 148

133 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 1/3/2005 182 15.2 9.0 25.0 15. 7 14.3 17.3 0.00 84% 1/4/2005 183 15.4 8.6 25.0 15. 9 14.6 17.4 0.00 83% 1/5/2005 184 16.3 9.8 25.6 16. 3 14.8 18.0 0.00 82% 1/6/2005 185 18.4 10.9 26.2 17. 1 15.9 18.7 0.00 78% 1/7/2005 186 17.9 13.2 24.3 17. 6 16.5 19.0 0.00 89% 1/8/2005 187 18.9 13.8 27.0 18. 1 16.9 19.5 0.01 84% 1/9/2005 188 18.7 12.6 26.7 18. 3 17.0 20.0 0.00 86% 1/10/2005 189 17.2 11.4 24.0 18.4 17.8 19.4 0.00 91% 1/11/2005 190 16.1 10.8 23.4 17.8 16.7 19.1 0.00 89% 1/12/2005 191 18.7 13.0 25.6 18.2 17.2 19.5 0.00 80% 1/13/2005 192 21.0 12.5 27.1 18.3 17.3 19.4 0.00 81% 1/14/2005 193 15.3 10.6 22.6 18.4 17.0 19.2 1.29 87% 1/15/2005 194 10.6 8.2 12.3 15.9 14.8 17.0 0.00 80% 1/16/2005 195 8.9 2.8 15.3 14.5 13.7 15.6 0.00 68% 1/17/2005 196 3.9 -2.0 10.6 12.8 11.8 13.8 0.00 63% 1/18/2005 197 3.3 -3.0 11.8 11.4 10.3 12.6 0.00 65% 1/19/2005 198 6.1 -2.2 15.1 11.6 9.9 13.5 0.00 63% 1/20/2005 199 10.9 6.3 17.8 12.9 11.6 14.8 0.00 70% 1/21/2005 200 12.6 3.3 22.9 13.2 11.4 15.2 0.00 81% 1/22/2005 201 16.7 14.4 21.2 15.0 14.2 16.0 0.21 90% 1/23/2005 202 5.5 -2.4 18.5 13.9 11.7 15.5 0.01 48% 1/24/2005 203 2.2 -6.4 12.8 11.0 9.4 12.5 0.00 50% 1/25/2005 204 8.3 -1.9 18.6 11.4 9.5 13.4 0.00 55% 1/26/2005 205 15.4 8.9 22.7 13.3 11.8 15.1 0.00 81% 1/27/2005 206 16.1 11.0 23.6 15.2 14.1 16.9 0.00 79% 1/28/2005 207 9.8 8.6 11.4 14.0 13.3 15.2 0.00 82% 1/29/2005 208 12.5 9.1 17.0 13.7 13.0 14.6 0.35 89% 1/30/2005 209 12.8 4.2 18.7 15.0 14.2 16.7 0.08 87% 1/31/2005 210 5.5 1.1 11.7 12.8 12.1 14.3 0.00 90% 2/1/2005 211 9.2 1.9 16.7 12. 4 11.2 13.9 0.00 82% 2/2/2005 212 10.1 8.4 14.8 12. 9 12.4 13.4 0.26 87% 2/3/2005 213 12.7 7.2 20.5 13. 9 12.8 15.4 0.08 92% 2/4/2005 214 8.1 1.2 14.8 13. 5 12.5 14.5 0.00 76% 2/5/2005 215 7.7 -1.2 17.9 12.5 10.7 14.3 0.00 72% 2/6/2005 216 12.1 3.2 21.1 13. 1 11.3 14.9 0.00 74% 2/7/2005 217 15.0 8.7 23.4 14. 5 13.4 16.1 0.00 73% 2/8/2005 218 14.3 5.3 25.2 15. 0 13.1 17.2 0.00 71% 2/9/2005 219 16.2 6.9 25.0 15. 8 13.9 17.8 0.00 75% 2/10/2005 220 15.0 6.0 19.0 16.6 14.9 17.9 0.21 60% 2/11/2005 221 6.8 -0.7 15.3 13.8 12.2 15.4 0.00 45% 2/12/2005 222 8.5 -2.0 20.1 13.2 11.1 15.5 0.00 62%

PAGE 149

134 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 2/13/2005 223 12.1 3.3 23.1 14.0 12.1 16.2 0.00 65% 2/14/2005 224 15.1 5.4 22.6 14.5 13.1 15.9 0.02 77% 2/15/2005 225 17.3 11.5 24.4 16.2 14.9 18.0 0.00 86% 2/16/2005 226 17.7 9.7 25.8 16.8 15.0 18.7 0.00 80% 2/17/2005 227 16.3 8.3 22.4 17.2 16.4 18.3 0.00 75% 2/18/2005 228 9.2 0.6 17.8 15.5 14.0 17.1 0.00 43% 2/19/2005 229 9.7 -1.7 21.6 14.5 12.6 16.5 0.00 61% 2/20/2005 230 13.5 4.0 24.1 15.4 13.5 17.4 0.00 70% 2/21/2005 231 17.8 10.9 23.9 16.1 14.9 17.3 0.00 77% 2/22/2005 232 20.9 18.1 26.6 18.1 16.8 20.0 0.00 82% 2/23/2005 233 21.0 18.0 26.9 19.3 18.1 20.9 0.06 78% 2/24/2005 234 19.5 16.3 23.6 19.2 18.5 19.9 0.02 83% 2/25/2005 235 13.3 10.0 16.2 17.4 16.1 18.7 0.27 92% 2/26/2005 236 12.6 9.3 17.6 16.0 15.2 16.9 0.00 84% 2/27/2005 237 15.7 13.0 21.1 16.2 15.8 16.8 1.18 93% 2/28/2005 238 15.4 10.4 21.9 17.0 15.7 18.6 0.00 76% 3/1/2005 239 11.8 4.8 18.8 15. 8 14.2 17.4 0.00 58% 3/2/2005 240 7.3 -2.1 15.6 14.8 12.8 16.8 0.00 44% 3/3/2005 241 6.1 1.2 9.0 13. 5 13.0 14.5 0.17 86% 3/4/2005 242 10.0 2.0 19.2 14. 2 12.2 16.8 0.00 71% 3/5/2005 243 11.8 1.8 22.0 14. 8 12.5 17.1 0.00 68% 3/6/2005 244 13.3 2.5 22.4 15. 5 13.3 17.9 0.00 54% 3/7/2005 245 15.8 4.4 25.1 16. 5 14.3 18.7 0.00 63% 3/8/2005 246 15.8 6.4 19.8 17. 4 16.2 18.9 0.79 59% 3/9/2005 247 7.8 3.1 9.8 14. 4 13.6 16.1 0.00 60% 3/10/2005 248 10.1 0.8 18.2 14.4 12.3 17.0 0.00 65% 3/11/2005 249 13.0 3.1 20.9 15.2 12.9 17.4 0.00 70% 3/12/2005 250 14.0 2.0 21.8 15.7 13.6 17.8 0.00 54% 3/13/2005 251 18.7 11.7 25.9 17.0 14.9 19.4 0.00 72% 3/14/2005 252 21.1 18.6 25.5 18.3 17.6 19.6 0.00 82% 3/15/2005 253 15.5 13.1 18.7 17.8 17.0 18.7 0.15 89% 3/16/2005 254 17.9 13.5 23.8 17.6 16.7 18.8 0.66 93% 3/17/2005 255 13.6 8.1 17.2 17.2 16.1 17.8 0.09 94% 3/18/2005 256 7.9 6.1 9.9 15.3 14.8 16.1 0.00 83% 3/19/2005 257 9.5 0.1 19.8 15.3 12.8 18.0 0.00 75% 3/20/2005 258 14.9 3.6 24.7 16.4 13.8 19.1 0.00 66% 3/21/2005 259 18.3 13.7 24.1 17.6 16.5 18.9 0.17 68% 3/22/2005 260 21.4 16.9 28.9 18.8 17.6 20.4 0.16 84% 3/23/2005 261 20.6 14.7 25.3 19.3 18.5 20.4 0.13 71% 3/24/2005 262 19.6 9.6 28.6 19.2 16.6 22.0 0.00 68% 3/25/2005 263 19.8 17.3 21.3 19.5 19.2 20.1 1.87 88% 3/26/2005 264 19.2 17.9 20.3 19.3 19.1 19.5 1.12 94%

PAGE 150

135 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 3/27/2005 265 22.7 17.6 28.9 19.9 18.5 21.4 0.00 82% 3/28/2005 266 18.3 14.2 23.2 20.5 19.3 21.7 0.08 67% 3/29/2005 267 18.4 11.7 27.0 19.8 17.5 22.4 0.00 60% 3/30/2005 268 17.8 7.8 27.9 20.2 17.6 22.8 0.00 64% 3/31/2005 269 20.2 12.5 28.6 21.0 19.0 22.9 0.17 77% 4/1/2005 270 21.4 15.3 29.3 21. 4 19.7 23.0 1.35 79% 4/2/2005 271 16.8 8.0 21.2 21. 2 20.0 23.2 0.27 61% 4/3/2005 272 13.5 3.9 22.3 19. 4 17.1 21.8 0.00 53% 4/4/2005 273 16.2 5.6 27.3 19. 5 17.0 21.9 0.00 62% 4/5/2005 274 18.7 7.3 27.9 19. 9 17.8 21.8 0.00 60% 4/6/2005 275 18.7 12.3 27.7 19. 9 18.7 22.4 0.00 76% 4/7/2005 276 19.4 16.9 23.3 19. 9 19.4 20.4 1.36 86% 4/8/2005 277 18.2 13.1 22.5 20. 0 18.5 21.5 0.00 80% 4/9/2005 278 18.8 11.8 26.6 20. 7 18.6 22.9 0.00 75% 4/10/2005 279 17.0 7.5 24.9 20.5 19.1 22.0 0.00 69% 4/11/2005 280 18.1 7.1 27.5 20.2 17.9 22.5 0.00 70% 4/12/2005 281 20.2 13.5 26.0 21.0 19.6 22.4 0.19 75% 4/13/2005 282 19.6 12.4 24.3 22.1 20.4 24.2 0.00 68% 4/14/2005 283 14.5 7.1 20.5 20.8 19.4 22.2 0.00 78% 4/15/2005 284 13.2 5.3 22.0 19.6 17.8 21.7 0.00 63% 4/16/2005 285 5.7 5.7 5.7 18.9 18.9 18.9 0.00 93% 4/17/2005 286 20.1 9.9 25.7 20.5 17.6 21.6 0.00 45% 4/18/2005 287 17.5 6.2 26.7 20.1 17.7 22.4 0.00 63% 4/19/2005 288 18.2 9.3 26.8 21.0 18.9 23.3 0.00 68% 4/20/2005 289 19.2 9.5 28.3 21.5 19.4 23.6 0.00 65% 4/21/2005 290 19.5 10.2 28.1 21.8 19.8 23.7 0.00 70% 4/23/2005 291 19.1 12.5 25.4 21.6 20.7 23.0 0.05 71% 4/24/2005 292 12.5 5.7 19.1 20.6 18.6 22.7 0.00 53% 4/25/2005 293 13.7 2.6 23.2 20.0 17.8 22.4 0.00 61% 4/26/2005 294 18.8 12.9 22.7 20.3 19.6 20.9 0.59 83% 4/27/2005 295 19.9 12.9 27.0 21.1 19.1 24.0 0.00 64% 4/28/2005 296 17.7 8.0 26.5 21.2 18.7 23.7 0.00 58% 4/29/2005 297 19.2 8.1 28.6 21.7 19.2 24.3 0.00 71% 4/30/2005 298 21.3 14.7 28.8 22.2 20.8 23.6 0.29 77% 5/1/2005 299 18.4 14.1 21.0 21. 4 20.8 22.0 0.03 88% 5/2/2005 300 19.5 12.7 27.4 22. 0 19.8 24.9 0.00 74% 5/3/2005 301 19.2 9.4 27.9 22. 4 19.9 25.0 0.00 64% 5/4/2005 302 17.3 14.0 18.8 21. 2 20.3 22.9 0.88 92% 5/5/2005 303 18.1 15.5 20.0 20. 3 20.0 20.7 1.17 93% 5/6/2005 304 16.6 11.7 22.4 20. 2 19.4 21.5 0.00 81% 5/7/2005 305 18.1 9.5 26.9 21. 1 18.4 24.0 0.00 72% 5/8/2005 306 19.5 10.9 27.7 22. 1 19.6 24.5 0.00 71% 5/9/2005 307 20.1 12.1 28.6 22. 8 20.3 25.7 0.00 71% 5/10/2005 308 21.4 12.3 29.6 23.4 20.9 25.8 0.00 68% 5/11/2005 309 21.0 16.1 28.8 23.7 22.2 25.2 0.01 81% 5/12/2005 310 22.2 13.5 30.4 23.8 21.5 26.1 0.00 70% 5/13/2005 311 21.6 13.1 29.1 24.1 22.3 25.8 0.00 71%

PAGE 151

136 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 5/14/2005 312 21.4 12.7 29.7 24.1 22.1 26.1 0.00 70% 5/15/2005 313 21.9 13.4 30.3 24.4 22.5 26.1 0.00 73% 5/16/2005 314 22.6 14.7 31.5 24.8 22.8 26.8 0.00 70% 5/17/2005 315 22.0 14.9 30.2 24.8 23.3 26.1 0.00 78% 5/18/2005 316 22.3 15.3 30.3 24.6 23.1 26.1 0.00 77% 5/19/2005 317 23.3 16.3 30.0 24.8 23.5 26.2 0.00 72% 5/20/2005 318 23.4 15.2 30.6 25.1 23.4 26.7 0.00 74% 5/21/2005 319 21.8 17.6 28.5 24.8 23.9 25.7 0.89 85% 5/22/2005 320 21.2 14.5 29.3 24.4 22.2 26.8 0.00 76% 5/23/2005 321 23.5 15.4 30.6 25.3 22.8 28.0 0.00 73% 5/24/2005 322 26.5 21.9 32.1 26.3 24.6 28.3 0.00 75% 5/25/2005 323 21.9 14.3 28.1 26.1 24.5 27.5 0.00 65% 5/26/2005 324 20.6 11.5 28.8 25.4 23.6 27.3 0.00 64% 5/27/2005 325 22.4 12.8 30.5 25.2 23.7 26.6 0.00 70% 5/28/2005 326 23.5 15.7 31.1 25.7 24.1 27.4 0.00 70% 5/29/2005 327 23.9 16.0 32.1 25.8 24.2 27.3 0.00 73% 5/30/2005 328 25.0 18.9 31.7 26.0 24.8 27.4 0.00 76% 5/31/2005 329 23.5 20.6 28.6 25.8 24.4 26.9 1.49 86% 6/1/2005 330 24.3 21.9 28.6 25. 6 24.7 26.7 0.01 81% 6/2/2005 331 24.4 21.4 29.2 25. 8 24.7 27.1 0.11 83% 6/3/2005 332 24.1 21.1 28.5 25. 7 24.7 26.7 1.59 84% 6/4/2005 333 24.2 21.5 28.2 25. 4 24.5 26.5 0.00 81% 6/5/2005 334 25.5 18.6 32.7 26. 3 24.3 28.5 0.00 76% 6/6/2005 335 26.0 19.9 32.8 27. 2 25.3 29.2 0.01 77% 6/7/2005 336 25.6 19.4 33.7 27. 6 25.6 29.8 0.00 79% 6/8/2005 337 25.3 20.5 33.8 28. 0 26.4 30.1 0.00 82% 6/9/2005 338 25.7 20.2 32.4 27. 8 26.2 29.6 0.02 79% 6/10/2005 339 25.1 23.8 28.0 26.8 26.2 27.6 0.08 84% 6/11/2005 340 25.7 23.9 29.3 26.3 25.6 27.3 0.69 85% 6/12/2005 341 25.6 23.6 30.3 26.8 25.9 27.8 0.02 88% 6/13/2005 342 25.2 22.0 33.0 27.1 25.8 28.7 0.18 87% 6/14/2005 343 25.9 21.4 33.5 27.5 25.9 29.7 0.81 84% 6/15/2005 344 27.6 24.2 33.5 28.3 26.8 30.2 0.00 81% 6/16/2005 345 26.5 22.0 33.9 28.7 27.1 31.0 0.10 83% 6/17/2005 346 27.2 22.1 32.8 28.8 26.9 31.0 0.00 71% 6/18/2005 347 24.1 20.5 30.8 27.9 27.1 29.4 0.43 83% 6/19/2005 348 25.3 18.9 31.0 27.7 25.8 29.8 0.00 77% 6/20/2005 349 24.1 18.3 30.7 27.7 26.0 29.6 0.00 74% 6/21/2005 350 24.1 18.1 31.3 27.6 25.8 29.6 0.00 78% 6/22/2005 351 25.0 19.1 32.6 28.0 26.3 30.0 0.00 77% 6/23/2005 352 25.1 17.8 32.8 28.0 26.2 30.0 0.00 71% 6/24/2005 353 23.8 17.7 30.2 27.4 26.3 28.6 0.06 77% 6/25/2005 354 24.4 21.6 28.7 27.4 26.3 28.6 0.00 84% 6/26/2005 355 26.4 22.7 33.4 28.1 26.7 30.0 0.00 80% 6/27/2005 356 25.5 22.0 32.9 27.9 27.1 29.4 2.00 85% 6/28/2005 357 25.0 22.5 30.5 27.2 26.5 28.1 0.13 88%

PAGE 152

137 Table C-31. Continued Air Temperature (C) Soil Temperature (C) Rain Humidity Date Day Avg. Min. Max. Avg. Min. Max. Inches Percent 6/29/2005 358 25.2 22.7 30.5 27.2 26.4 28.1 1.36 89% 6/30/2005 359 24.8 23.4 28.7 27.1 26.7 27.6 0.19 90% 7/1/2005 360 26.5 23.3 31.7 27. 7 26.4 29.3 0.13 83% 7/2/2005 361 26.2 23.1 31.8 28. 1 27.2 29.0 0.25 85% 7/3/2005 362 25.7 22.7 31.0 27. 9 27.0 28.6 1.21 87% 7/4/2005 363 27.6 22.5 33.1 28. 5 26.9 30.3 0.01 80% 7/5/2005 364 28.0 21.2 34.7 29. 2 27.5 31.0 0.00 74% 7/6/2005 365 27.7 22.0 34.0 29. 5 27.9 31.0 0.00 75% 7/7/2005 366 27.9 22.5 34.0 29. 5 28.0 31.2 0.00 75% 7/8/2005 367 27.6 22.0 35.2 29. 5 28.0 31.3 0.00 78% 7/9/2005 368 26.9 22.7 31.7 29. 0 28.5 29.7 0.57 82% 7/10/2005 369 25.1 22.5 28.5 27.2 26.8 28.4 1.58 86% 7/11/2005 370 26.0 23.5 32.1 27.6 26.8 28.7 1.10 86% 7/12/2005 371 26.5 21.5 33.4 28.1 26.6 29.7 0.00 81% 7/13/2005 372 26.6 21.9 34.8 28.7 27.3 30.5 0.02 81% 7/14/2005 373 26.6 21.7 33.4 28.8 27.4 30.1 0.00 81% 7/15/2005 374 25.9 22.5 32.7 28.8 27.8 30.0 0.00 82% 7/16/2005 375 27.9 21.6 34.8 28.8 27.3 30.4 0.00 73% 7/17/2005 376 27.4 21.7 34.7 29.0 27.6 30.5 0.05 79% 7/18/2005 377 27.4 21.7 34.2 29.2 27.8 30.9 0.00 76% 7/19/2005 378 27.4 22.1 34.0 29. 3 28.0 30.7 0.00 78%

PAGE 153

138 APPENDIX D SUPPLEMENTAL ACKNOWLEDGEMENTS Table D-1. Thanks. Many thanks to Those who have helped me along the way Brian Abrams Cecilia Kennedy Dr. Hartwell Allen Sylvia Lang-Josan Marty Anderson John Leader Sarah Anderson Chris Lewis Natlie Balcer Hui X Lu Charles Bohall Kathleen Mckee Dr. Patrick Bohlen Dr. Paul Mislevy Jeremy Bright Jeremy Paris Dr. Mark Brown Ryan Penton Charles Campbell Dan Perkins Yubao Cao Kevin Ratkus Cory Catts Dr. Ramesh Reddy Dr. Mark Clark Caroline Reis-Hamilton Dr. Matt Cohen Casey Schmidt Dr. Bree Darby Dr. Thomas Sinclair Dr. Edmond Dunne Gunner Smith Dr. Mitch Flinchum Erik Spalvins Adrienne Frisbee Ms Yu Wang Kevin Grace Ondine Wells Xiao Wei Gu Bill White Dr. Willie Harris Dr. Ann Wilkie Eric Jorczak Gavin Wilson Dr. Rob Kalmbacher This been a great experience both academica lly and personally. There have been so many people, from East Lansing to Gainesville that have directly or indirectly helped guide me along on this journey. I am grateful to have had the opportunity to cross so many paths along the way, and hope to someday cross them again. Peace.

PAGE 154

139 LIST OF REFERENCES Allen, L. H. Jr, E. H. Stewart, W. G. Jr. Knisel, & R. A. Slack. (1975). Seasonal Variation in Runoff and Water Quality from th e Taylor Creek Watershed, Okeechobee County, Florida. Paper presented at the Soil and Crop Science Society of Florida. Andersen, J. M. (1976). An Ignition Method for Determination of Total Phosphorus in Lake Sediments. Water Research, 10 329-331. Arthington, John, Patrick Bohl en, & Fritz Roka. (2003). Effect of Stocking Rate on Measures of Cow-Calf Productivity and Nutrient Loads in Surface Water Runoff (Document An141) Retrieved 5/16/2006, from http://edis.ifas.ufl.edu/AN141 Bayley, Suzanne E, Jr John Zoltek, Albert J Hermann, Thomas J Dolan, & Louis Tortora. (1985). Experimental Manipul ation of Nutrients and Water in a Freshwater Marsh: Effects on Biomass, Decomp osition, and Nutrient Accumulation. Limnology and Oceanography, 30 (3), 500-512. Boggess, C F, E G Flaig, & R C Fluck. ( 1995). Phosphorus Budget-Basin Relationship for Lake Okeechobee Tributary Basins. Ecological Engineering, 5 143-162. Bohlen, Patrick, Ken Campbell, John Cap ece, John Earman, J. Jeffrey Mullahey, Don Graetz, et al. (2004). Agro-Ecosystem Indi cators of Sustainability as Affected by Cattle Stocking Density in Ranch Management System. MacArthur Agro-ecology Research Center 2002-2003 Biennial Re port for the John D. and Catherine T. MacArthur Foundation Bottcher, A B, Terry K Tremwel, & Kenne th L Campbell. (1995). Best Management Practices for Water Quality Improvement in the Lake Okeechobee Watershed. Ecological Engineering, 5 341-356. Braskerud, B C. (2002). Factors Affecting P hosphorus Retention in Small Constructed Wetlands Treating Agricultural Non-Point Source Pollution. Ecological Engineering, 19 41-61. Chambliss, C. G., & M. B. Adjei. (2006). Bahiagrass (SS-AGR-36) Retrieved May, 2006, from http://edis.ifas.ufl.edu/AA184 Clary, Warren P. (1995). Vegetation and Soil Responses to Grazing Simulation on Riparian Meadows. Journal of Range Management, 48 18-25.

PAGE 155

140 David, Peter G. (1999). Response of Exotics to Restored Hydroperiod at Dupuis Reserve, Florida. Restoration Ecology, 7 (4), 407-410. Dolan, Thomas J, Suzanne E Bayley, Jr John Zoltek, & Albert J Hermann. (1981). Phosphorus Dynamics of a Florida Fr eshwater Marsh Receiving Treatment Wastewater. Journal of Applied Ecology, 18 205-219. Douglas, Marjory Stoneman. (1947). The Everglades; River of Grass Sarasota: Pineapple Press, Inc. FAC, Florida Administrative Code. (1996). Feedlot and Dair y Wastewater Treatment and Management Requirements. Chapter 62-670. FAC, Florida Administrative Code. (2000). Classification of Surface Waters, Usage, Reclassification, Classifi ed Waters. Chapter 62-302.400. FDEP, Florida Department of Environmenta l Protection. (2001). Total Maximum Daily Load for Total Phosphorus Lake Okeechobee, Florida. Flaig, Eric G, & K R Reddy. (1995). Fate of Phosphorus in the Lake Okeechobee Watershed, Florida, USA: Ov erview and Recommendations. Ecological Engineering, 5 127-142. Flaig, Eric G., & Karl Have ns. (1995). Historical Tre nds in the Lake Okeechobee Ecosystem. Arch. Hydrobiol./Suppl., 107 (1), 1-24. FLEPPC, Florida Exotic Pe st Plant Council. (2005). List of Florida's Invasive Species Retrieved March, 2006, from http://www.fleppc.org/list/05List.htm Fluck, R. C., C. Fonyo, & E. Flaig. (1992). Land-Use-Based Phosphorus Balances for Lake Okeechobee, Florida, Drainage Basins. American Society of Agricultural Engineers, 8 (6), 813-820. Gilliam, J W. (1995). Phosphor us Control Strategies. Ecological Engineering, 5 405414. Gunsalus, Boyd, Eric G Flaig, & Gary Ritter. (1992). Effectiveness of Agricultural Best Management Practices Implemented in the Taylor Creek/Nubbin Slough Watershed and the Lower Kissimmee River Basin. Paper presented at the National RCWP Symposium, SFWMD, West Palm Beach, FL. Gustafson, Shelley, & Deane Wang. (2002). Eff ects of Agriculture Runoff on Vegetation Composition of a Priority Conservation Wetland, Vermont, USA. Journal of Environmental Quality, 31 350-357.

PAGE 156

141 Havens, Karl E, Mark Brady, Erin Colbor n, Steffany Gornak, Susan Gray, R Thomas Janes, et al. (2005). Lake Okeechobee Prot ection Program State of the Lake and Watershed. In 2005 South Florida Environmental Report Havens, Karl E, Jr. Victor J. Bierman, Eric G. Flaig, Charles Hanlon, R. Thomas James, Bradley L. Jones, et al. (1995). Hist orical Trends in the Lake Okeechobee Ecosystem Vi. Synthesis. Arch. Hydrobiol./Suppl., 107 101-111. Hiscock, Jeffrey G, C Scott Thourot, & J oyce Zhang. (2003). Phosphorus Budget Land Use Relationships for the Northern Lake Okeechobee Watershed, Florida. Ecological Engineering, 21 63-74. Holderbaum, J F, L E Sollenberger, J E Moore, W E Kunkle, D B Bates, & A C Hammond. (1991). Protein Supplementati on of Steers Grazing Limpograss Pasture. Journal of Production Agriculture, 4 (3), 437-441. Holderbaum, J F, L E Sollenberger, K H Qu esenberry, J E Moore, & C S Jones, Jr. (1992). Canopy Structure and Nutritive Value of Limpograss Pastures During Mid-Summer to Early Autumn. Agronomy Journal, 84 11-16. Kadlec, R H. (1999). The Limits of Phosphorus Removal in Wetlands. Wetlands Ecology and Management, 7 165-175. Kadlec, Robert H., & Robert J. Knig ht. (1996). Chapter 14: Phosphorus. In Treatment Wetlands Boca Raton: Lewis Publishers. Kalmbacher, R S, K R Long, M K Johns on, & F G Martin. (1984). Botanical Composition of Diets of Cattle Grazing South Florida Rangeland. Journal of Range Management, 37 (4), 334-340. Kalmbacher, Rob, Jeff Mullahey, & Kevin H ill. (1998). Limpograss and Hymenachne Grown on Flatwoods Range Pond Margins. Journal of Range Management, 51 (282-287). Kidder, G., C. G. Chambliss, & R. Mylavarapu. (2000). UF/IFAS Standardized Fertilization Recommendations for Ag ronomic Crops (Fact Sheet Sl-129) Retrieved March, 2006, from http://edis.ifas.ufl.edu Kleinberg, Eliot. (2003). Black Cloud; the Great Florida Hurricane of 1928 New York: Carroll & Graf Publishers. Koerselman, Willem, & Arthur F. M. Me uleman. (1996). The Ve getation N:P Ratio: A New Tool to Detect the Natu re of Nutrient Limitation. Journal of Applied Ecology, 33 1441-1450.

PAGE 157

142 Kuo, S. (1996). Phosphorus Availability Indices. In J. M. Bigham (Ed.), Methods of Soil Analysis. Part 3. Chemical Methods (pp. 893). Madison, Wisconsin: Soil Science Society of America, Inc. Lake Okeechobee Issue Team, & South Flor ida Ecosystem Restoration Working Group. (1999). Lake Okeechobee Action Plan. In Un ited States Environmental Protection & South Florida Water Management Distri ct (Ed.). West Palm Beach, Florida. Leopold, Aldo. (1966). A Sand County Almanac. New York: Ballantine. Long, K R, R S Kalmbacher, & F G Martin. (198 6). Diet Quality of Steers Grazing Three Range Sites in South Florida. Journal of Range Management, 39 (5), 389-392. Magee, Teresa K, & Mary E. Kentula. (2005). Response of Wetland Plant Species to Hydrologic Conditions. Wetlands Ecology and Management, 13 163-181. Marousky, F. J., & F. Blondon. (1995). Red and Far-Red Light Influence Carbon Partitioning, Growth and Flowering of Bahiagrass ( Paspalum Notatum ). Journal of Agricultural Science, 125 355-359. McKee, Kathleen. (2005). Predicting Soil Phosphorus Storag e in Historically Isolated Wetlands within the Lake Okeechobee Priority Basins. Unpublished Master's Thesis, University of Florida, Gainesville, FL. Mislevy, P. (2002). Forage Alternatives for Florida Cattle. Paper presented at the Annual Beef Cattle Short Course Proceedings, Gainesville, FL. Mitsch, William J., & James G. Gosselink. (2000). Wetlands (Third ed.). New York: John Wiley & Sons, Inc. Newman, Y C, L E Sollenberger, W E Kunkle, & D B Bates. (2002a). Crude Protein Fractionation and Degradation Para meters of Limpograss Herbage. Agronomy Journal, 94 1381-1386. Newman, Y C, L E Sollenberger, W E K unkle, & C G Chambliss. (2002b). Canopy Height and Nitrogen Supplementation Eff ects on Performance of Heifers Grazing Limpograss. Agronomy Journal, 94 1375-1380. Pate, Findlay. (1998). 'Floralta' Hemarthria, a Popular Grass for South Florida Retrieved September, 2004, from http://rcrec-ona.ifas.ufl.edu/PRFAR6.html Quesenberry, K H, W R Ocumpaugh, O C Ru elke, L S Dunavin, & P Mislevy. (1984). 'Floralta': A Limpgrass Selected for Yi eld and Persistence in Pastures (Circular S-312) Retrieved September, 2004, from http://rcrec-ona.ifas.ufl.edu/cirs312.html

PAGE 158

143 Reddy, K R, Mark Clark, J Jawitz, T DeBu sk, M Annable, W Wise, et al. (2003). Phosphorus Retention and Storage by Isol ated and Constructed Wetlands in the Okeechobee Drainage Basin. In Soil a nd Water Science Department-UF-IFAS, Environmental Engineering Sciences De partment-UF & DB Environmental Labs Inc (Eds.). Reddy, K R, O A Diaz, L J Scinto, & M Ag ami. (1995). Phosphorus Dynamics in Selected Wetlands and Streams of the Lake Okeechobee Basin. Ecological Engineering, 5 183-207. Reddy, K R, E G Flaig, & D A Graetz. (1996). Phosphorus Storage Capacity of Uplands, Wetlands and Streams of the Lake Okeechobee Watershed, Florida. Agriculture, EcoSystems and Environment, 59 203-216. Reddy, K R, R H Kadlec, E G Flaig, & P M Gale. (1999a). Phosphorus Retention in Streams and Wetlands: A Review. In Critical Reviews in Environmental Science and Technology (Vol. 29, pp. 83-146). Reddy, K R, E Lowe, & T Fontaine. (1999b). Phosphorus in Florida's Ecosystems: Analysis of Current Issues. In K. R. Reddy, G. A. O'Connor & C. L. Schelske (Eds.), Phosphorus Biogeochemis try in Subtropical Ecos ystems: Florida as a Case Example (pp. 111-141): CRC/Lewis Publ. Reddy, K. R., & W. F. Debusk. (1985). Nutrient Removal Potential of Selected Aquatic Macrophytes. Journal of Environmental Qualty, 14 (4), 459-463. Reddy, K. R., & W. F. Debusk. (1987). Nutrie nt Storage Capabilities of Aquatic and Wetland Plants. In K. R. Reddy & W. H. Smith (Eds.), Aquatic Plants for Water Treatment and Resource Recovery (pp. 337-357): Magnolia Publishing. Richardson, Curtis J. (1985). Mechanisms C ontrolling Phosphorus Retention Capacity in Freshwater Wetlands. Science, 228 (4706), 1424-1427. SAS Institute, Inc. (1989-2005). Jmp Statistical Software Cary, NC. Seabloom, Eric W., Kirk A. Moloney, & Arnol d G. Van der Valk. (2001). Constraints on the Establishment of Plants Along a Flucuating Water-Depth Gradient. Journal of Ecology, 82 (8), 2216-2232. Second Louisiana--Florida Ecosystem Restoration Information Exchange (2001). Retrieved 5/4/2006, from http://www.lacoast.gov/education/ever glades/New_Orleans_Exchange_Mar_2001 _files/frame.htm

PAGE 159

144 SFWMD, South Florida Water Ma nagement District. (1995). Land Use of Lake Okeechobee Retrieved March, 2006, from http://www.sfwmd.gov/org/wrp/wrp_ok ee/2_wrp_okee_info/maps/landuse.html SFWMD, South Florida Water Ma nagement District. (2006). Lake Okeechobee: Watershed Overview Retrieved March, 2006, from http://www.sfwmd.gov/org/wrp/wrp _okee/2_wrp_okee_h20shed/2_wrp_okee_h2 0shed.html SFWMD, South Florida Wate r Management District, Florida Department of Environmental Protection, & Florida Depa rtment of Agriculture and Consumer Services. (2004). Lake Okeechobee Prot ection Plan. South Florida Water Management District, West Palm Beach, Florida. SFWMD, South Florida Water Ma nagement District, & U.S. Army Corps of Engineers. (2005). Comprehensive Everglades Rest oration Plan. Master Implementation Sequencing Plan. Sharpley, Andrew N. (1995). Soil Phosphorus Dynamics: Agronomic and Environmental Impacts. Ecological Engineering, 5 261-279. Sollenberger, L E, C G Chambliss, W E Kunkle, W F Brown, & K H Quesenberry. (2006). Floralta Limpograss (H emarthria Altissima) Retrieved 5/12/2006, from http://edis.ifas.ufl.edu/BODY_AA218 Sollenberger, L E, W R Ocumpaugh, V P B Eu clides, J E Moore, K H Quesenberry, & C S Jones, Jr. (1988). Animal Performance on Continuously Stocked 'Pensacola' Bahiagrass and 'Floralta' Limpograss Pastures. Journal of Production Agriculture, 1 (3), 216-220. Sollenberger, L E, G A Rusland, C S Jones, Jr, K A Albrecht, & K L Gieger. (1989). Animal and Forage Responses on Rotationally Grazed Floralta' Limpgrass and Pensacola' Bahiagrass Pastures. Agronomy Journal, 81 760-764. Steinman, Alan D, Julie Conklin, Patrick J Bohlen, & Donald G. Uzarski. (2003). Influence of Cattle Grazing and Pasture Land Use on Macroinvertebrate Communities in Freshwater Wetlands. Wetlands, 23 (4), 877-889. Tobe, John D., Kathy Craddock Burks, Richar d W. Cantrell, Mark A. Garland, Maynard E. Sweeley, David W. Hall, et al. (1998). Florida Wetland Plants; an Identification Manual Tallahassee, FL: Florida Department of Environmental Protection. Van der Valk, A G. (1981). Succession in Wetlands: A Gleasonian Approach. Journal of Ecology, 62 (3), 688-696.

PAGE 160

145 Van der Valk, A G, L Squires, & C H We lling. (1994). Assessing the Impacts of an Increase in Water Level on Wetland Vegetation. Ecological Applications, 4 (3), 525-534. Violi, Helen. (2000). Element Stewardship Abstract for Paspalum Notatum Flgg Arlington, Virginia: The Nature Conservancy. Wellings, Charles H, Roger l Pederson, & Ar nold G Van der Valk. (1988). Recruitment from Seed Bank and the Development of Zonation of Em ergent Vegetation During a Drawdown in a Prairie Wetland. Journal of Ecology, 76 483-496. Whigham, Dennis, Mary Pittek, Kirsten H Ho fmockel, Thomas Jordan, & Antoninette L Pepin. (2002). Biomass and Nutrient D ynamics in Restored Wetlands on the Outer Coastal Plain of Maryland, USA. Wetlands, 22 (3), 562-574.

PAGE 161

146 BIOGRAPHICAL SKETCH Jeffrey D. Smith grew up on a small cash-crop farm in Byron, Michigan, a small town near Flint, Michigan. He earned a B achelor of Science in horticulture/landscape design with an agribusiness specialization from Michigan State University in 2002. Before graduating, Jeff started and operated a successful landscape design/build firm in Lansing, Michigan, for about 3 years, before working with the Mich igan Department of Agriculture’s Groundwater Monitoring Program---a desired transition into an environmental monitoring related field. In 2003, he moved to Florida to pursue a master’s degree in wetland ecology-a long time goal, which produced this thesis. He does not know what the future has in store pr ofessionally or academ ically, but feels the opportunities are endless.


Permanent Link: http://ufdc.ufl.edu/UFE0015200/00001

Material Information

Title: Phosphorous Storage Dynamics in Wetland Vegetation and Forage Grass Species: Facilitating Wetland Hydrologic Restoration in the Lake Okeechobee Watershed
Physical Description: Mixed Material
Copyright Date: 2008

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0015200:00001

Permanent Link: http://ufdc.ufl.edu/UFE0015200/00001

Material Information

Title: Phosphorous Storage Dynamics in Wetland Vegetation and Forage Grass Species: Facilitating Wetland Hydrologic Restoration in the Lake Okeechobee Watershed
Physical Description: Mixed Material
Copyright Date: 2008

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0015200:00001


This item has the following downloads:


Full Text












PHOSPHORUS STORAGE DYNAMICS IN WETLAND VEGETATION AND
FORAGE GRASS SPECIES: FACILITATING WETLAND HYDROLOGIC
RESTORATION IN THE LAKE OKEECHOBEE WATERSHED













By

JEFFREY D. SMITH


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2006

































Copyright 2006

by

Jeffrey David Smith



























This thesis is dedicated to all who strive to educate themselves, and those who support
education.





"The question is, does the educated citizen know he is only a cog in an
ecological mechanism? That if he will work with that mechanism his mental
wealth and his material wealth can expand indefinitely? But that if he refuses
to work with it, it will ultimately grind him to dust? If education does not
teach us these things, then what is education for?"(Leopold, 1966).















ACKNOWLEDGMENTS

I would like to thank Dr. Mark W. Clark for his contagious enthusiasm for science,

and life in general, and for his guidance and support throughout the course of this

research. I would also like to give special thanks to Dr. Edmond J. Dunne for his

guidance and mentoring in the field, laboratory and throughout the development of this

thesis, and most of all, his friendship. Dr. Clark and Dr. Dunne are outstanding scientists,

but even better persons. My committee members, Dr. K. Ramesh Reddy and Dr. L.

Hartwell Allen, were also very influential in the development and review of this research.

Lastly, I would like to thank my parents, Sarah Anderson and many friends, from near

and far, for fostering my goals and providing unconditional support. Many other

important people are listed in Appendix D, Table D-1.











TABLE OF CONTENTS




A C K N O W L E D G M E N T S ................................................................................................. iv

LIST OF TA BLE S .................. .................................. .... .. ........ ........ ....... viii

LIST OF FIGURES ......... ......................... ...... ........ ............ xi

ABSTRACT .............. .......................................... xiv

CHAPTER

1. IN TR OD U CTION ............................................... .. ......................... ..

P rob lem ............................................................ ................ .. 2
R egion al C h aracteristics ............................................................. .....................3
E v erglades .................................................... ................ .. 3
Lake Okeechobee's watershed ............. ................................ ...............5
Phosphorus loading to Lake Okeechobee ............................................... 7
Policy and Planning .................. .............................. .. ...... ................ .9
Lake Okeechobee Legislation........................................................ 10
Phosphorus Best Management Practices ........................................... .................11
Hydrologic Restoration of Isolated W wetlands ................................................12
Phosphorus in w etland soils ..................................................................... 13
A alternative forage crops .................................................... .. .... ......... 14
Thesis Objectives .............. ..... .... .... ...... ...... ............................. 15

2. PHOSPHORUS ASSIMILATION BY ISOLATED WETLAND VEGETATION ..17

In tro du ctio n ....... ......... ......................................................... 17
Factors Influencing Phosphorus Retention........................................................ 19
Research Objectives .............................. ....... ... ............ .. .........21
Research Questions and Hypotheses.............. ............................................... 22
M materials and M methods ....................................................................... ..................23
Study Sites ..........................................23.............................
Sam pling ........... ............................. ................................... 25
Sam ple processing .................... .... .... .... .. .... ........ ........ .............. 26
L laboratory A analysis ............................................ .. ........ .... ...........27
D ata and Statistics A analysis ........................................ .......................... 27
R e su lts .................. ....... .... ................... ................... ...................... 2 8
Species Composition along a Hydrologic Gradient ........................................28
Ecosystem Phosphorus Storage .................................................. ............... 31
Standing Biomass ............................ ............. .......... .................. 33
Standing Biom ass by Individual Species .................................... ............... 36


v









Phosphorus Storage in Biom ass ........................................ ....... ............... 37
Standing Biom ass by Individual Species ................................. ................ 41
D isc u ssio n ............................................................................................................. 4 2
H y d ro lo g y ...................................................................................................4 2
E cosystem storage ....................... ...... .......... ............... .... ..... .. 43
Biom ass in Pasture W wetlands ........................................ ......................... 44
D isturbance Effects on B iom ass................................... .................................... 45
Phosphorus Concentrations ........................................ ........................... 47
P hosphoru s Storage .............................. ........................ .. ........ .... ............4 8
C o n c lu sio n s........................................................................................................... 4 9

3. FACILATATING WETLAND HYDROLOGIC RESTORATION WHILE
MAINTAINING FORAGE PRODUCTION: HYDROLOGIC TOLERANCES
OF PASPALUM NOTATUM AND HEMARTHRIA AL TISSIMA ............................ 51

Introduction ............... ......... .......................51
Background ............ ......... .. ......... ..........51
Research Objectives ................................ ...... .............. .. .........53
Research Questions and Hypotheses............ ........................... ...............53
M materials and M methods ....................................................................... ..................54
E x p erim ental D esign ........................................ ............................................54
T re a tm e n ts ..................................................................................................... 5 6
Sampling .......... .. .. ......... ............... 57
S o il ......................... ..................5 8
A bove ground biom ass sam pling ........................................ ............... 58
Below ground biom ass ....... .. ...................................................... ....... 60
L laboratory analysis .............................................. ............... 60
Results ............... ..... ............ ............. ............... 60
Initial characterization ...................... ......... ...............60
F o rag e P ro du ctio n ......................................................................................... 6 1
B ahiagrass forage production............................................. 63
Limpograss forage production ......................... ........ ............... 64
Species comparison .............. ..... .......................65
Total Biomass ................. .. ...... ..................68
Root to Shoot Ratios ..... .................... ....... ........69
Phosphorus Assimilation ................................. .......................... ...... 71
Phosphorus tissue concentrations ............... ........... ........... .................71
Phosphorus storage.........................................73
Phosphorus storage root to shoot ratios.................................. ............. 76
D isc u ssio n ..............................................................................................7 7
F o rag e P ro du ctio n ......................................................................................... 7 7
Flood Tolerance ............... ......... .......... ........78
P h o sp h oru s U ptak e ........................................................................................ 80
C conclusions ................................................ 81

4. SUMMARY AND CONCLUSIONS ........................ ............... 83









Su m m ary ................... ... ............. ...................................... ................. 83
Objective I: Biomass Production and Phosphorus Storage in Wetlands............83
V vegetation Stress .................................................................... .. 84
Objective II: Facilitating Land-use and Wetland Restoration.............................84
U expected R results ......................... .. ...................... ...... ........... 85
Im plications for R restoration ............................................................ .....................85
C on clu sion s ............... .. .... .. ..... ... ....... ........... .. ............................... 8 8
Unanswered Questions and Need for Further Research...........................................89

APPENDIX

A. SUPPLEMENTAL BACKGROUND INFORMATION ..........................................90

B. SUPPLEMENTAL FIELD DATA.................................... ..................... 92

C. SUPPLEMENTAL MESOCOSM DATA............... ............................ 102

D. SUPPLEMENTAL ACKNOWLEDGEMENTS................ ...... ............... 138

L IST O F R E F E R E N C E S .................................................... ................... .................... 139

BIOGRAPHICAL SKETCH ................................................ ......... ............... 146
















LIST OF TABLES

Table page

1-1 Okeechobee watershed land use by percent of total land ................ ..................6

2-1 Mean and standard deviation of hydroperiods at each site. ................................29

2-2 Mean species hydroperiod of both sites. ...................................... ............... 29

2-3 R oot to shoot ratios by zone .................................. ............... ............... 36

2-4 Below ground biomass concentrations by zone ................................................. 38

2-5 Above ground biomass P concentrations by zone............................................40

2-6 BGB to A GB P storage ratios ............................................................................ 41

3-1 Sam pling dates and details. ...... ........................... ....................................... 58

3-2 Total biomass after 163 and 375 days. ............................... ............................... 68

3-3 R oot to shoot ratios. ......................................... ... .... ........ ..... .... 71

3-4 Total P storage species comparison. ............................................. ............... 74

3-5 Root to shoot P storage ratios with statistics................................. .................77

4-1 Estimation of P export concentrations to tributaries from various land-uses. .........87

A- Summary of Okeechobee Basins BMPs........................................ ............... 90

A-2 Total P loads to Lake Okeechobee 1991-2003.................................... ................ 91

B-l Phosphorus storage by components, site and zone. ............................................92

B-2 Biomass production by components, site and zone......................... ............... 92

B -3 Species hydroperiod. ...................... .. ...... ................ ... ...... .. ...............93

B -4 T otal biom ass production. ........................................... ........................................95

B-5 Below ground biomass production........ .................................................95









B-6 Above ground biomass production. .............................................. ............... 95

B -7 T otal biom ass P storage......................................... .............................................97

B-8 Below ground biomass P storage. ........................................ ........................ 97

B-9 Above ground biomass P storage. ........................................ ......................... 97

B-10 Phosphorus concentration in above ground biomass ..........................................100

B-11 Phosphorus storage in above ground biomass....................................................101

C-1 Nutrient concentrations on day 0. ........................................ ....... ............... 102

C-2 Species comparison of forage production per harvest ........................... ........103

C-3 Species comparison of cumulative forage production. ........................................ 103

C-4 Overall below ground biomass all treatments combined ..................................103

C-5 Forage production per harvest .............. ................... ......... .. ............... 104

C-6 Cumulative forage by treatment and day. ................................... ............... 105

C-7 Below ground biomass species comparison.......... ............ ............. 106

C-8 Residual biomass harvested on days 163 and 375. ..............................................106

C-9 Below ground biomass production time comparison...................... ..............106

C-10 Bahiagrass forage production per harvest treatment comparison........................109

C-11 Bahiagrass cumulative forage production treatment comparison ........................110

C-12 Limpograss forage production per harvest treatment comparison.....................111

C-13 Cumulative limpograss forage production treatment comparison....................112

C-14 Bahiagrass BGB production treatment comparison.......................... .........113

C-15 Limpograss BGB production treatment comparison...................... ................113

C-16 Forage P concentrations species comparison. ............ .... ................ ..............114

C-17 Bahiagrass forage P concentrations treatment comparison ..............................115

C-18 Limpograss forage P concentrations treatment comparison ............................116

C-19 Below ground biomass P concentrations species comparison. ...........................117









C-20 Bahiagrass BGB P concentrations treatment comparison ....................... 117

C-21 Limpograss BGB P concentrations treatment comparison.............................118

C-22 Phosphorus storage in forage species comparison per harvest. ...........................119

C-23 Cumulative P storage in forage species comparison.................. ............ 120

C-24 Bahiagrass forage P storage per harvest........................................... ...........121

C-25 Bahiagrass cumulative forage P storage................................................. ........... 122

C-26 Limpograss forage P storage per harvest. ................................... ............... 123

C-27 Cumulative limpograss forage P storage..................................................... 124

C-28 Below ground biomass P storage species comparison ................... .............127

C-29 Bahiagrass below ground biomass P storage ............................... ............... .127

C-30 Limpograss below ground biomass P storage .............. ................................. 127

C-31 Climatic conditions from day 1 to 375........ ................. .....................129

D -1 T hanks. ............................................................................. 138
















LIST OF FIGURES


Figure p

1-1 Phosphorus concentrations in Lake Okeechobee............. ........... ...............3

1-2 Historic, current, and future flow pattern of the Everglades..................................4

1-3 Land-use map of four priority basins of the Lake Okeechobee watershed ................8

1-4 Wetland coverage in the priority basins....................... ...................9

2-1 M echanism s driving P cycling. ........................................ .......................... 21

2-2 M ap of land use in the 4 priority basins.................................................. ........24

2-3 Isolated wetlands selected for long term monitoring ............................................24

2-4 Stratified sampling zones: center, edge and upland. ..........................................25

2 -5 L og istics fit of sp ecies ................................................................... ..................... 3 0

2-6 Logistics fit of species by site. ............................................................................ 31

2-7 Phosphorus storage components. ........................................ ......................... 32

2-8 Comparison of AGB and BGB components at Beaty and Larson. ..........................32

2-9 Total biomass at Beaty and Larson wetlands .......................................................33

2-10 Below ground biomass at Beaty and Larson wetlandss.................... ...............34

2-11 Above ground biomass at Beaty and Larson wetlands ........................................34

2-12 Biom ass partitioning AGB vs. BGB ............................................. ............... 35

2-13 Above ground biomass by species for all zones ............................................... 37

2-14 Total biom ass P storage......................................... .................. ............... 38

2-15 Phosphorus storage in BGB at Beaty and Larson wetlands.............. ................ 39

2-16 Phosphorus storage in AGB at Beaty and Larson wetlands................................40









2-17 Above and below ground biomass P storage. .................................. .................41

2-18 Phosphorus storage by species. ........................................ .......................... 42

2-19 Nutrient storage and growth in plants ........................................... ............... 47

3-1 Study site at University of Florida, Gainesville, Florida ......................................55

3-2 M esocosm diagram .......................................... ............... .... ....... 56

3-3 Inverse relationship of water depth and redox ...............................................57

3-4 H harvesting procedure ....................................................................... ..................59

3-5 Forage production per harvest for each species all treatments combined..............62

3-6 Cumulative forage production with all treatments combined ................................62

3-7 B ahiagrass treatm ent com prisons ...........................................................................63

3-8 Limpograss treatment comparisons........................................ 65

3-9 Forage production per harvest by treatment ........... .............. ........................66

3-10 Cumulative forage production by treatment....................... .................. 67

3-11 Below ground biom ass production...................................... ......................... 69

3-12 Above and below ground biomass production after 375 days ..............................70

3-13 Mean P concentrations for bahiagrass forage by harvest day and by treatments.....72

3-14 Mean P concentrations for limpograss forage by harvest day and by treatments ....73

3-15 Total P storage (AGB + BGB) after 375 days. .............................. ................74

3-16 Cum ulative P harvested in forage ........................................ ........................ 75

3-17 Relative comparison of root and shoot P storage after 375 days. ............................77

B-1 Species distribution by hydroperiod........................................ 94

B-2 Above ground biomass by species and zone. .................... .................96

B-3 Phosphorus concentrations by species. ........................................ ............... 98

B-4 Phosphorus storage by zone .............................................................................. 99

C-1 Relative root and shoot biomass after 163 days ..................................................107









C-2 Total biomass production after 163 days. ................................... ............... 107

C-3 Total biomass production after 375 days. ................................... ............... 108

C-4 Bahiagrass total biomass and P storage......... ........... ........... ............. 125

C-5 Limpograss total biom ass and P storage ..................................... .................125

C-6 Bahiagrass BGB and P storage........................................ ........................... 126

C-7 Limpograss BGB and P storage ........................................................ ............... 126

C-8 R oot to shoot P storage ratios.......................................... ........................... 128

C-9 Total P storage (AGB +BGB) at 163 days ......................................................128










Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

PHOSPHORUS STORAGE DYNAMICS IN WETLAND VEGETATION AND
FORAGE GRASS SPECIES: FACILITATING WETLAND HYDROLOGIC
RESTORATION IN THE LAKE OKEECHOBEE WATERSHED

By

Jeffrey D. Smith

August, 2006

Chair: Mark Clark
Major Department: Soil and Water Science

Nutrient export from agricultural activities in the Lake Okeechobee watershed has

contributed to eutrophication of the Lake and regulatory implementation of a phosphorus

(P) Total Maximum Daily Load (TMDL) rule. Historically, anthropogenic manipulation

of hydrology lowered water tables, creating improved conditions for upland forage grass

production. This action also increased runoff rates and P loading to the Lake. Hydrologic

restoration of historically isolated wetlands within the watershed is a proposed best

management practice (BMP) to increase P retention capacities of these wetlands.

However, longer hydroperiods could potentially decrease pasture productivity, and as a

consequence, adversely affect the economic viability of the cattle industry in the region.

Previous studies have shown that soils under longer hydroperiods in the Okeechobee

watershed have greater P storage potential than surrounding upland soils. This research

primarily focuses on the vegetative component of P storage in pasture wetlands. The

objectives were to evaluate biomass production and P storage dynamics in vegetation

under various hydroperiods and to determine the efficacy of using an alternative forage

grass species to maintain pasture productivity after wetland restoration.








Four isolated wetlands in Okeechobee County, Florida were sampled in November,

2004; March, 2005; and July, 2005. In this study, total P storage in wetlands (with a 50-

m upland buffer) included soil (10 cm depth), below ground biomass (BGB), above

ground biomass (AGB) and litter components. Soil was the primary P storage component

representing greater than 88% of the total P stored in the wetlands, while BGB, AGB and

litter represented 8%, 3%, and 1% respectively. Total biomass (AGB+BGB) production

and P storage in biomass were inversely related to hydroperiod in wetlands at the more

intensively managed pasture, while P storage in biomass was positively related to

hydroperiod in wetlands at the less intensively managed pasture. Management intensity

(i.e., cattle density and pasture maintenance) may be influencing P storage capacities of

vegetative, and affecting the relationship between hydroperiod and P storage.

In a separate mesocosm study in Gainesville, Florida, Paspalum notatum Fhi,,','

(bahiagrass) and Hemarthria altissima 'Floralta' (limpograss), a wet-tolerant forage

grass, were evaluated under five different hydrologic treatments. Water levels were

stabilized at 10, 0, -10, and -15 cm relative to the soil surface, while the control only

received rain water and was allowed to drain completely. Limpograss had greater forage

(AGB) production and P assimilation than bahiagrass in all treatments. However,

bahiagrass had greater total biomass (AGB+BGB) production in all but the 10 cm

inundated treatment. Bahiagrass total P storage was only greater than limpograss in the

-10 cm water level. This indicates that limpograss has a greater hydrologic tolerance than

bahiagrass and similar P storage potential. Therefore, to maintain pasture carrying

capacity and vegetative P storage during BMP implementation, limpograss may be a

more suitable forage in restored pastures wetlands.














CHAPTER 1
INTRODUCTION

Nutrient export from agricultural activities in the Lake Okeechobee watershed has

contributed to eutrophication of the Lake and regulatory implementation of a phosphorus

(P) Total Maximum Daily Load (TMDL) rule. Historically, anthropogenic manipulation

of hydrology drained wetlands and lowered the water table, creating improved conditions

for upland forage grass production. This action increased runoff rates and P loading to the

Lake. Four priority basins occupy 12% of the watershed's area, but export 35% of the P

load entering the Lake (FDEP, 2001). Hydrologic restoration of historically isolated

wetlands is a proposed best management practice (BMP) to increase P retention

capacities of these wetland ecosystems, thus decreasing P loads entering the Lake.

However, longer hydroperiods could potentially decrease pasture productivity, and as a

consequence, adversely affect the economic viability of the cattle industry in the region.

Previous studies have shown that soils under longer hydroperiods in the Lake

Okeechobee Basin have greater P storage potential than surrounding upland soils

(McKee, 2005). This research primarily focuses on the vegetative component of P

storage in pasture wetlands. The objectives were to evaluate biomass production and P

storage dynamics in vegetation under various hydroperiods and to determine the efficacy

of using an alternative forage grass species to maintain pasture productivity after wetland

restoration.









Problem

In 1998, Lake Okeechobee was listed as a water quality limited (WQL) water body

(FDEP, 2001). This condition was the result of over 50 years of excessive pollutant

loading. Nutrients, dissolved oxygen, unionized ammonia, chlorides, coliforms and iron

have threatened numerous societal and environmental values of the Lake, contributing to

eutrophication, resultant algal blooms, and subsequent alterations of flora and fauna

species composition (SFWMD et al., 2004).

Phosphorus is often considered the limiting nutrient in freshwater aquatic systems

(Reddy et al., 1999b). The development of agriculture (predominately animal operations)

in the mid 1900's, along with anthropogenic manipulation of hydrology in the watershed

has drastically increased P loading to the Lake (Havens et al., 2005; Reddy & Debusk,

1987). Between 1968 and 2004, the P concentration in the pelagic zone of the Lake

(Figure 1-1) increased from -40 [tg L-1 to over 120 [tg L-1 (Flaig & Reddy, 1995; Havens

et al., 2005). The measured P load entering the Lake in 2004 was 548 metric tons, while

the five year rolling average between 2000-2004 was 528 metric tons P (Havens et al.,

2005).

In 2001, 14 of the 29 drainage basins within the Lake's watershed were exceeding

their P loading targets (FDEP, 2001). Several BMPs intended to reduce P export from

agricultural lands within the watershed have been or are in the process of being

implemented (Table A-i, Appendix A). In the 1980's the implementation of BMPs

significantly reduced P concentrations in tributaries (Gunsalus et al., 1992). However, P

concentrations and loads continue to exceed established target loads (Table A-2,

Appendix A).











Lakewater Total Phosphorus
140
120
a. e*
a 100 -
c *- *
o 80 e
60 -
i 40 ..... .......................................
a,
0
S20 Restoration Goal = 40 ppb


68 70 72 74 76 78 80 82 84 86 88 90 92 94 96 98
Year


Figure 1-1. Phosphorus concentrations in Lake Okeechobee (Lake Okeechobee Issue
Team & South FLorida Ecosystem Restoration Working Group, 1999)

Regional Characteristics

Everglades

Historically, Lake Okeechobee was the central component of the Everglades

Ecosystem, interconnecting the Kissimmee River Basin in Central Florida to the

Everglades in South Florida. Central Florida's Upper Chain of Lakes were the

headwaters of the Everglades. Water migrated south through the Kissimmee River Basin

into the northern part of Lake Okeechobee. The Lake was a natural detention basin,

releasing water over its southern banks during times of high water. Overflow water

migrated south through sawgrass prairies into a vast ridge and slough system, before

eventually seeping into Florida Bay.

Currently, as a result of anthropogenic flood control and land development, Central

and South Florida hydrology is drastically different (Figure 1-2). In 1928, over 2000

people were killed when the storm surge from a major hurricane flooded hundreds of

acres surrounding Lake Okeechobee (Kleinberg, 2003). That spurred the effort to









construct the Herbert Hoover Dike around the Lake for flood control. This, in turn,

reduced the size of the Lake to 650 square miles and altered natural hydrologic

fluctuations by creating a dependence on a vast system of water control structures. In the

decades that followed, the Kissimmee River was channelized, the Caloosahatchee River

was dredged and St. Lucie canal was built to divert water from the Lake out to the

Atlantic Ocean and Gulf of Mexico. The hydrology of the Everglades also became

systematically controlled with the construction of several dikes, levies, pump stations,

roads and canals.

















Figure 1-2. Historic (left), current (middle), and future (right) flow pattern of the
Everglades (Second Louisiana--Florida Ecosystem Restoration Information
Exchange, 2001)

As a result, the Everglades ecosystem is no longer a continuous River of Grass

(Douglas, 1947), but rather, several compartmentalized aquatic systems manipulated by

anthropogenic water control structures. Agricultural and urban development throughout

South Florida drained wetlands, thus, increasing anthropogenic control of hydrology. In

the early 1900's, the sawgrass prairies south of the Lake were drained for agriculture









development, converting the area into what is currently known as the Everglades

Agricultural Area (EAA).

The paradigm that drove these vast hydrologic alterations in South Florida was

based on the need for flood prevention in populated areas and cultivatable agricultural

land. Development and wetland drainage has spawned numerous resource dilemmas.

Over fertilization in agricultural sectors has been identified as a primary source of excess

nutrient loading to Lake Okeechobee and the Everglades' Water Conservation Areas

(WCA). As a result freshwater quality and supplies have been compromised. Large

quantities of water are diverted from Lake Okeechobee to the coasts to prevent high P

loads from entering the Everglades. However, this has degraded estuarine ecosystems on

both coasts. In addition drainage has caused rapid mineralization of organic soils and

accreted nutrients, which are ultimately transported to Lake Okeechobee or the

Everglades. The Everglades is a low nutrient ecosystem, with ambient P concentrations

of -10 tg L-1. Thus, discharging water with high P concentrations from the Lake into the

Everglades will be detrimental to the ecosystem. This reinforces the importance of

reducing P loading to Lake Okeechobee.

Lake Okeechobee's watershed

Lake Okeechobee is the second largest freshwater lake within the contiguous

United States with a surface area of 1,890 km2 and contributing watershed of over 6,000

km2. It provides numerous societal and environmental values including water supply for

agriculture and urban sectors, flood protection and a multi million dollar sport and

commercial fishing industry (SFWMD, 2004).

Agriculture is the primary land use within the watershed, occupying over half of

the land area, while wetlands and terrestrial ecosystems are categorically second










(SFWMD, 1995). Improved pasture, sugarcane, upland forest, rangeland, unimproved

pasture and citrus make up the largest percentage of agricultural land area (Table 1-1).

Table 1-1. Okeechobee watershed land use by percent of total land.
Lake Okeechobee Watershed
Landuse Hectares Percentage
Improved Pasture 182,590 24.02%
Wetlands 128,884 16.96%
Water 127,339 16.75%
Sugar Cane 91,409 12.03%
Upland Forest 69,479 9.14%
Rangeland 50,283 6.62%
Unimproved Pasture 30,829 4.06%
Citrus 22,941 3.02%
Urban and built-up 22,325 2.94%
Barren 6,284 0.83%
Dairies 5,209 0.69%
Row Crops 4,926 0.65%
Transportation, Communication, and Utilities 4,619 0.61%
Field Crops 4,147 0.55%
Woodland Pasture 3,935 0.52%
Fallow Crop Land 1,756 0.23%
Sod Farms 941 0.12%
Tree Nursery 907 0.12%
Horse Farm 397 0.05%
Fruit Orchards 378 0.05%
Aquaculture 165 0.02%
Other 164 0.02%
Ornamentals 74 0.01%
Other Grove 29 0.00%
Floriculture 8 0.00%
Table created from land use data (SFWMD, 1995)

Large areas of wetlands in the watershed have been drained for agricultural

development. Riparian and non-riparian (isolated) wetlands are abundant throughout

Lake Okeechobee's watershed, covering 17% (1,290 km2) of the area (Table 1-1). The

landscape in the northern part of the watershed was historically characterized by the

presence of numerous isolated wetlands. These wetlands are referred to as 'historically'

isolated because they lacked a surface water connection to tributaries, except overland

flow in times of flooding. Extensive drainage efforts established vast networks of ditches









and canals, short circuiting the natural water retention and nutrient assimilative capacity

of the landscape. Most isolated wetlands in the northern watershed were drained in mid-

1900's to support the rapidly increasing beef cattle and dairy industry; creating improved

pasture conditions for upland forage species (Flaig & Havens, 1995). Draining wetlands

was a trend that occurred throughout the country, ultimately depleting more than half of

the wetlands within the contiguous United States (Mitsch & Gosselink, 2000).

Phosphorus loading to Lake Okeechobee

Agricultural activities are responsible for 98% of all P imported to the watershed,

the majority of which is pasture fertilizer and dairy feed (Fluck et al., 1992). There is a

high correlation between P imports to the watershed and P loading to Lake Okeechobee

(Boggess et al., 1995). Non-point source runoff from agriculture, particularly, beef cattle

and dairy operations, is recognized as a primary source of P loading to the Lake (Flaig &

Havens, 1995)

Four priority basins within the Lake's watershed, S-65D, S-65E, S-154, and S-191

(Figure 1-3), have been identified as "hot spots" based on land use intensity and high P

discharge. These four basins occupy 12% of the land, and export as much as 35% of the

total P load entering the Lake (FDEP, 2001). Within these priority basins, 61% of the

land area supports agricultural activities (47% improved pasture, 14% dairy) (Figure 1-3),

while 11% of the land is occupied by partially drained or otherwise impacted wetlands

(Figure 1-4). In fact, 45% of isolated wetlands in the priority basins have been at least

partially drained.






























dairies
I improved pastures
unirrproved pastures/ rangelands
0 5 10 20lometers
MENOMOEEEEEEEEEilomneters


Figure 1-3. Land-use map of four priority basins of the Lake Okeechobee watershed
(McKee, 2005)


















5 10 20 4. -
--











0 5 10 2lometers -


Lake Okeechobee ,,

Figure 1-4. Wetland coverage in the priority basins (McKee, 2005)

Policy and Planning

While flood control is still a priority, the water retention and contaminate

assimilative capacity of wetlands are now widely recognized. In the face of increasing

population and water demand in South Florida, major efforts are underway to carry out

some of the largest ecological restoration projects ever undertaken. The Comprehensive

Everglades Restoration Plan (CERP) (SFWMD & USACE, 2005), the Lake Okeechobee

Protection Plan (LOPP) (SFWMD et al., 2004), and the Kissimmee River Restoration

Program (KRR) are umbrella programs designed to preserve and protect water resources

in Central and South Florida.
in Central and South Florida.









CERP is an $8 billion restoration project, comprised of over 60 major projects with

the underlying objective to preserve and project the quality and supply of freshwater

resources. Currently, an average of 1.7 billion gallons of fresh water is diverted from

Lake Okeechobee out to sea annually (SFWMD, 2005). CERP will attempt to capture

and store most of that water in new reservoirs and Aquifer Storage and Recovery (ASR)

wells.

Lake Okeechobee Legislation

A water body is considered WQL or impaired when its pollutant load exceeds

water quality standards for its designated use. Lake Okeechobee is designated as a Class I

or potable water supply (FAC). In compliance with Section 303(d) of the Clean Water

Act (CWA) the establishment of TMDL is required for all impaired water bodies (FDEP,

2001). Since excessive P loading is primarily responsible for eutrophication of Lake

Okeechobee (Havens et al., 1995), a TMDL of 140 metric tons yr-1 P was developed to

achieve the target concentration of 40 ppb P within the Lake's pelagic zone by 2050.

Phosphorus is currently the only pollutant with a required TMDL requirement for Lake

Okeechobee.

Water quality problems in Lake Okeechobee have been widely recognized since the

late 1960's (Allen et al., 1975; Flaig & Havens, 1995; Fluck et al., 1992; Gunsalus et al.,

1992; Gustafson & Wang, 2002). In 1987, the Surface Water Improvement and

Management (SWIM) Act (Florida Statutes, Sections 373.451 and 373.4595) was

developed to focus on preservation and restoration of some of Florida's most significant

water bodies. Lake Okeechobee was named in that act, specifically mandating a 40%

reduction of P loads in order to achieve a P concentration of 40 ppb in the pelagic zone.

It regulated P sources from dairies by implementing farm buyout programs, BMPs and









structural retrofits to control systems (i.e. lagoon systems). As a result, P loads entering

the tributaries between the late 1980's and 1990's were reduced (Gunsalus et al., 1992).

However, by the mid-1990's loads still exceeded SWIM targets and the load reduction

trend was no longer declining (LOAP, 1999). This resulted in legislative action that

called for more aggressive action than mandated by SWIM.

The Lake Okeechobee Protection Act (LOPA) (Chapter 00-130, Laws of Florida)

was passed in 2000. LOPA mandated the implementation of a restoration and protection

program which includes a P TMDL and BMPs to reduce nutrient loading to the Lake.

The Lake Okeechobee Protection Program (LOPP) was developed to achieve and

maintain compliance with Florida water quality standards. It involves the

implementation of a P TMDL along with other research and monitoring objectives

required by LOPA. (SFWMD, 2004). In addition, the Lake Okeechobee Watershed

Project (LOWP) is a component of CERP that aims to reduce P loading to the Lake,

attenuate peak flows and restore riparian and isolated wetland habitat. LOPP and LOWP

contain similar P source control programs through the implementation of "voluntary" and

cost-share BMPs; however LOPP addresses regional projects not included in CERP.

Phosphorus Best Management Practices

Best Management Practices are conservation guidelines developed using Best

Available Technology (BAT) to reduce point and non-point source water pollution while

maintaining economically viable agricultural productivity (Bottcher et al., 1995).

However, the success of a BMP is only as effective as its level of acceptance. In fact,

overcoming social and political obstacles may be more challenging than the fundamental

science supporting the BMP, thus innovative educational approaches that facilitate an

understanding of potential costs and benefits associated with implementing the BMP is









necessary (Bottcher et al., 1995). The success and cost-effectiveness of BMPs are

dependent on regional goals and BATs. However, in all cases, mutual awareness of

potential outcomes and willingness to compromise by all parties is essential to achieve

BMP objectives and maintain the economic viability of the land-use.

Many P BMPs targeting dairy and cattle sectors have been implemented in the

Lake Okeechobee drainage basin (Table A-i, Appendix A). Required implementation of

P BMPs as part of the Rural Clean Water Program (RCWP) and the Okeechobee Dairy

Rule (FAC, 1996) have effectively reduced P discharge from dairies by 50%, thus

improving discharge water quality (Gunsalus et al., 1992). However, the discharge

reductions are relative to previous discharges, which may have been several times greater

than acceptable concentrations, and have not been enough to lower the overall P load

entering the Lake.

Hydrologic Restoration of Isolated Wetlands

Hydrologic restoration of historically isolated wetlands is a potential BMP that

could play a significant role in meeting the P TMDL of 140 metric tons yr Wetlands

are known to assimilate and immobilize nutrients and other contaminates in living and

dead (detritus) plant biomass and in soils. Phosphorus storage in wetland soils is

dependent on the P concentrations of the overlying water column and the sediment pool.

Organic matter accumulation and the abundance of iron and aluminum oxides within the

soil influences sediment P flux with the overlying water column (Reddy et al., 1999b).

Anaerobic decomposition is a slow process that facilitates the accumulation of organic

material. The accumulation of organic material immobilizes P in the process, thus acting

as a P sink as long as anaerobic conditions persist. Phosphorus assimilation in living

biomass is short term process which can re-release labile nutrients back into the









environment upon senescence and decomposition. However, P accretion in plant

biomass has been shown to account for 12-73% of total P removal from nutrient enriched

waters (Reddy & Debusk, 1985).

Phosphorus in wetland soils

Mckee (2005) conducted a survey of 118 wetlands within the four priority basins to

determine P storage in soils of isolated wetlands. Wetlands on dairy, improved and

unimproved pasture land-uses were divided into center, edge, upland and ditch zones and

sampled. Physical parameters, such as organic matter content and bulk density between

wetland center and upland were significantly different when compared between like-land

uses. Total P (TP) analysis showed significantly higher concentrations in wetland centers

compared to uplands for all three land use types. There were also significantly higher TP

concentrations in wetland centers compared with edge or ditch soils for improved and

unimproved pasture land use types.

Across land-uses, Mckee found significantly greater center and edge TP

concentrations in dairy wetland soils than in improved and unimproved pasture wetland

soils. However, no significant difference was found between improved and unimproved

pasture land-use types. Center and edge soils from different wetland types were also

significantly different. Forested swamp soils had significantly higher center TP values

than emergent marshes and open water emergent marsh soils, while edge concentrations

were significantly greater in scrub-shrub swamps compared to emergent marsh and open

water emergent marshes. There were also significantly greater P concentrations in edge

soils of forested swamps compared to emergent marshes.

McKee's results theoretically support a hydrologic restoration BMP of isolated

wetlands as a means to increase P retention in the watershed and decrease P loads to the









Lake. Hydrologic restoration would raise the water table, and increase the zone of

inundation; maximizing the potential of the wetlands to accrete P in soils. However, this

would also cause a shift in species composition, likely decreasing upland forage grass

production, and pasture carrying. Thus, a consequence of restoration may adversely

affect the economic viability of cattle operations as productive pasture area would likely

be reduced.

Alternative forage crops

BMPs with the potential to negatively impact economic viability should not be

considered BMPs, unless alternative funds are available to subsidize their implementation

(Bottcher et al., 1995). However, along with hydrologic restoration, alternative practices

could be implemented to minimize forage loss or even enhance pasture productivity. Wet

cropping systems have been suggested as potential means of reducing P imports to the

watershed and concentrations in dairy wastewater by utilizing a vegetative species with

high P assimilative capacity and sufficient forage value (Reddy et al., 2003). A forage

species could remove the majority of P in a treatment wetland, while a periphyton cell

would act as a polishing mechanism to further reduce P concentrations.

Since, P storage in vegetative biomass is short-term, wet cropping systems utilize

recycled nutrients to produce a forage, thus reducing the need for P imports to the

watershed. Wet cropping could also be a removal mechanism by harvesting and

exporting forage and assimilated P out of the watershed to be utilized in other agricultural

operations. Removal of sod from pastures is an effective way of export P because it is an

economically valuable product that can also reduce the cost of pasture renovations when

converting to more productive grasses.









Another option to maintain pasture carrying capacity after hydrologic restoration

is to utilize alternative forage species that have high productivity under wet conditions.

Hydrologic restoration will alter the water table, potentially, creating unsuitable

conditions for the existing dominant forage grass, Paspalum notatum F1i-,-,

(bahiagrass). Utilization of the wet tolerant forage grass species, Hemarthria altissima,

Tloralta', (limpograss) may be a beneficial substitute for bahiagrass, potentially reducing

the pasture area that would otherwise be lost if no alternative is implemented with

hydrologic restoration.

This has potentially positive economic implications for implementing a

hydrologic restoration BMP. If limpograss has comparable or higher forage production

and quality then this BMP may not only be ecologically beneficial, but it may also

provide an economic incentive to implement it.

Thesis Objectives

This research is part of a collaborative effort to evaluate the effectiveness of

hydrologic restoration of historically isolated wetlands as a BMP to enhance and utilize

the P storage potential within the Lake's watershed and reduce nutrient loading to the

Lake. More specifically, one objective of this research is to evaluate the role of

vegetation in wetland P storage. Since the implementation of wetland BMPs are, in part,

dependent on their level of acceptance, addressing landowner concerns for lost pasture

productivity is necessary. Another objective of this research investigates the efficacy of

using the wet tolerant forage species, Hemarthria altissima 'Floralta' (limpograss), in

upland areas, adjacent to wetlands, to alleviate the potential loss of productive pasture

due to hydrologic restoration. The thesis objectives are as follows:









I. Assess biomass production and P assimilation by wetland vegetation and
forage grasses under various hydroperiods.

II. Determine the efficacy of establishing a wet tolerant forage grass in wetland
transition zones before hydrologic restoration to minimize loss of productive
pasture

Chapter II focuses on standing biomass and P storage of various vegetative

components. Chapter III describes a mesocosm study that tested the hydrologic

tolerances of bahiagrass and limpograss. Chapter IV summarizes results from both

studies, discusses implications of wetland restoration and presents conclusions from this

study.














CHAPTER 2
PHOSPHORUS ASSIMILATION BY ISOLATED WETLAND VEGETATION

Introduction

Nutrient export, primarily phosphorus (P), from non-point source agricultural

activities in the Lake Okeechobee watershed has contributed to near hyper-eutrophic

conditions in the Lake (Reddy et al., 1999b). As a result, a Total Maximum Daily Load

(TMDL) rule for P and associated Best Management Practices (BMP) have been

implemented to reduce nutrient loading to Lake Okeechobee (Bottcher et al., 1995;

FDEP, 2001; Havens et al., 2005; SFWMD,2004). Many voluntary BMPs have

effectively lowered P exports from improved pasture and dairies in high P export basins

(Gunsalus et al., 1992) and in the Everglades Agricultural Area (EAA) (Flaig & Havens,

1995). However, in-Lake P concentrations currently average 120 [tg L-1; three times the

TMDL target concentration of 40 [ag L-1. Water column total nitrogen (TN) to TP ratios

in the Lake are 13:1, which favors cyanobacteria dominance (Havens et al., 2005)

Between 1994 and 1998, two of the Lake's northern tributaries, the Lower

Kissimmee River (LKR) and Taylor Creek/Nubin Slough (TCNS), supplied 43% of the

water, and 56% of the total P load entering the Lake. The ratio of water supplied to P

load for these two tributaries is disproportionate, LKR actually supplies 33% of the water

and 32% of the P load, while TCNS supplies 10% water and 24% of the P load (FDEP,

2001).

The high P discharge from these tributaries is primarily the result of four

"priority" drainage basins within their watersheds. These four basins (Figure 2-2) occupy









12% of the Lake's watershed, and export as much as 35% of the total P load entering the

Lake. Within the priority basins 68% of the land area supports agricultural activities

(45% improved pasture, 4% dairy), while 15% of the land is contains wetlands (SFWMD,

1995). The historical extent of wetlands is unknown; however, within the priority basins

45% of isolated wetlands have been at least partially drained (SFWMD, 2004). Ditches

that drain these wetlands act as a conduit for transporting dissolved P directly to the Lake.

Cattle ranching and agriculture have been the primary land uses in the watershed

since the mid 1800's. Beef cattle populations rapidly increased in the early to mid

1900's, spurring the drainage and transformation of native range lands into high

production improved pastures. From 1940 to 1970 the area of improved pasture

increased from 34,000 to 170,000 ha (Flaig & Havens, 1995) and by 1995 it occupied

183,000 ha of the watershed; -24 % (Table 1-1). The vast network of drainage canals

exacerbated nutrient loss from the landscape by lowering the water table and hydraulic

retention times (HRT), thus decreasing P assimilative potential of historically isolated

wetlands and perpetuating the need to import more nutrients (Flaig & Havens, 1995).

Extensive wetland drainage further intensified cattle production and increased P imports

to the watershed in the forms of cattle feed and fertilizer

Studies indicate that there is a strong correlation between P imports to watershed

and P loading to the Lake (Boggess et al., 1995; Hiscock et al., 2003). Continual net

imports of P have created an excess of bioavailable P. Soils in the northern watershed are

poorly drained and have limited P binding capacity, however, low topographic relief

limits runoff and subsequent P exports from uplands (Flaig & Havens, 1995). Boggess et

al., (1995) estimated that 90% of P imported between 1985 and 1989 was retained in the









watershed, while more recent estimates from 1997 to 2001 indicate an 83% retention of

imported P (Hiscock et al., 2003). In both studies, the majority of imported P was stored

in uplands (71% and 74%) however, of the portion that was loaded to wetlands, the

percent assimilated decreased over time from 60% to 32%. Hiscock et al., (2003)

attributes the reduction in storage to decreased assimilative potential, not decreased

wetland area. This suggests that many wetlands may already be saturated with P, and

even if imports to the watershed decrease or stop, they may become a source rather than a

sink. Therefore, reducing water flow, in addition to P imports, may be the most effective

way to reduce P loading to the Lake.

Factors Influencing Phosphorus Retention

Phosphorus is an essential nutrient for primary producers and is limiting in most

freshwater ecosystems. However, many agricultural wetlands are not limited by P, due to

its relative abundance and biogeochemical stability (Mitsch & Gosselink, 2000). In the

Okeechobee watershed, Reddy et al., (1995) found that nitrogen (N) and P concentrations

in aquatic macrophytes' tissue are generally high, indicating that neither nutrient is

limiting plant growth. Other studies have determined that wetland plants with N:P ratios

below 14 are N limited (Koerselman & Meuleman, 1996). Despite the apparent

abundance of both nutrients, the N:P ratios in tributary macrophytes were between 4 and

6 (Reddy et al., 1995), suggesting that P is more abundant than N. Although, generally

speaking, wetland plants in the Okeechobee watershed are not considered to be limited by

P or N availability.

The physical, chemical and biological mechanisms controlling P assimilation in

wetland ecosystems has been well documented (Braskerud, 2002; Flaig & Reddy, 1995;

Gilliam, 1995; Kadlec, RH, 1999; Kadlec & Knight, 1996; Mitsch & Gosselink, 2000;









Reddy et al., 1999a; Reddy et al., 1999b; Richardson, 1985; Sharpley, 1995). Unlike the

biogeochemical cycles of nitrogen, carbon, sulfur and oxygen, P does not have a naturally

occurring gaseous phase. It is accreted in wetlands by immobilization, adsorption and

precipitation processes (Figure 2-1). The relative portion of inorganic and organic forms

depends on soil, vegetation, hydrology and land use characteristics (Reddy et al., 1999a).

Adsorption and precipitation are abiotic processes that occur in the soil and are indirectly

controlled by pH. Immobilization is a temporary biotic process by which dissolved

inorganic P is assimilated in vegetative or microbial biomass as organic P. Vegetative

biomass has a high rate of turnover; often several times a year in warmer climates. After

senescence, a portion of labile P leaches back into the water column as the detrital

material breaks down. A small portion of P in recalcitrant detritus is accreted in the soil

as organic P.

Phosphorus assimilation in vegetation is dependent on species productivity and

turnover rates, nutrient availability, land-use intensity, hydrology, and biochemical and

physicochemical properties (Reddy et al., 1999a). Biomass is not be considered a

sustainable long-term P removal mechanism in wetlands because it is a short-term storage

that releases as much as 80% of assimilated P back into the water column after

senescence (Reddy et al., 1995). However, accretion of recalcitrant biomass residuals

(detritus) is the only sustainable long-term storage mechanism for P removal by

biological means (Kadlec & Knight, 1996; Richardson, 1985).












SLOW
INORGANIC



I PUARYV P I


A.
RAPID CYCUNG ORGANIC & INORGANIC

ANIMAL


SLOW
ORGANIC


Figure 2-1. Mechanisms driving P cycling. A) Mechanisms driving forms of P
(Sharpley, 1995). B). Phosphorus cycling in wetlands.

Research Objectives

The use of constructed and restored isolated wetlands in Lake Okeechobee's

watershed has been suggested as a potential means of decreasing P loads entering the

Lake (Flaig & Reddy, 1995; Havens et al., 2005; LOPA, 1999; Reddy et al., 2003; Reddy

et al., 1996; SFWMD, 2004). Since TMDLs are established based on both concentration









and flow, restoring the natural hydrology to isolated wetlands could reduce storm water

runoff while increasing wetland HRT and P accretion in residual organic material.

Previous studies in the four priority basins have shown significantly greater soil P

concentrations and organic material content in wetland centers than in adjacent uplands

(McKee, 2005). These findings suggest that hydrologic restoration could increase on-site

P storage in soils by increasing wetland area. A key component responsible for increased

P storage capacity in wetland centers is biomass production. High biomass and anoxic

conditions foster residual biomass and P accretion, stabilize soil porewater, and reduce

concentrations in surface water (Reddy et al., 1999b). Biomass production and P

assimilation are the primary focus of this research.

Research Questions and Hypotheses

1. What role does vegetative biomass play in total wetland P storage?

Hi: Biomass P storage will have a lesser role when compared to surface
soil P storage.

2. Does biomass differ along a hydrologic gradient?

H2: Biomass is higher in the center of the wetland

3. Does total P storage in standing biomass differ along a hydrologic gradient?

H3: Total biomass P storage will be higher in wetlands than uplands

4. Where is P partitioned in vegetation?

H4: More P will be stored in above ground biomass (AGB) than below
ground biomass (BGB)

While P export rates from various land-uses have been broadly established (Flaig &

Havens, 1995) the compounded influence of hydrology and grazing pressure on biomass

production and subsequent organic P storage in wetlands in the Okeechobee basin has not

been extensively studied. This chapter focuses on P storage in above and below ground









standing biomass and vegetative assemblages along hydrologic gradients in pasture

wetlands. Data presented in this chapter will be compared to vegetation data after

hydrologic restoration to evaluate the effect on total wetland P storage and vegetation

dynamics. Methods focus on above ground biomass (AGB) and below ground biomass

(BGB) sample collection, processing and laboratory analysis. In addition, soil and litter

samples were collected and pre-sampling photographs of each quadrate were taken.

Materials and Methods

Study Sites

Four wetlands from two different ranches located in the priority basins were

selected for long-term monitoring. Selection criteria were based on land use intensity and

proximity of two similarly sized, hydrologically modified wetlands. The Larson site,

located in basin S-154, is more intensely managed then the Beaty site, located in basin S-

65D (Figure 2-2). Management intensity was subjectively determined based on land-use

history, pasture maintenance regime and grazing pressure.

The two wetlands at the Larson site, Larson East (LE) (8056'28.08" W,

27o20'56.06" N) and Larson West (LW) (80056'47.49" W, 27o20'59.27" N), are roughly

2.5 ha each, while the Beaty wetlands, Beaty North (BN) (8056'54.50" W, 2724'41.41"

N) and Beaty South (BS) (8056'43.21" W, 2724'27.53" N), are roughly 1.3 and 1.4 ha

respectfully (Figure 2-3). Wetland size was calculated in ArcGIS. The perimeter was

delineated on site with GPS tracking by walking along vegetation community transitions

between upland forage grass (Paspalum notatum) and unconsolidated wetland species

(Juncus effusis).


































Figure 2-2. Map of land use in the 4 priority basins. S-191 is the Taylor Creek/Nubbin
Slough (TCNS) basin and S-65D, S-65E and S-154 part of the Lower
Kissimmee River (LKR) Basin The Larson site is located in basin S-154. The
Beaty site is located in S-65D.












a. b.

Figure 2-3. Isolated wetlands selected for long term monitoring. (a) Beaty Ranch
wetlands; top left wetland is referred to as Beaty North (BN) and the bottom
right wetland is Beaty South (BS). (b) Larson Ranch wetlands; wetland to the
left is referred to as Larson East (LE) and the wetland on right is Larson West
(LW)










Sampling

Data from three sampling events: November 19-21, 2004, March 25-26, 2005 and

July 14-16, 2005 were collected in this study. Based on results from McKee, 2005 who

found significantly greater soil P concentrations in wetland center zones than adjacent

uplands, a stratified random sampling scheme was used to sample wet marsh-(center),

transitional-(edge) and forage-(upland) zones (Figure 2-4). Respective zone data from all

sampling dates were combined and analyzed to minimize temporal variability. Five 1 m2

quadrates in each zone were located with GPS using predetermined random coordinates

from ArcGIS.


.... .. .. .. .. .. .. .. .. .. ..







.... .... ... .... .... ...

..pln


Figure 2-4. Stratified sampling zones: center, edge and upland. Five randomly placed
1 m2 quadrates (not drawn to scale) were sampled in each zone. Beaty North
shown as an example.









In the upland quadrates, AGB was cut as close to the ground as possible

(approximately 1 to 3 cm above the ground surface) using electric grass clippers, while

hand clippers were used in edge and center zones. All removable AGB from individual

quadrates was collected. After AGB was clipped, three BGB cores were extracted from

random locations within the quadrate with a 15 cm diameter aluminum cylinder, to a

depth of 20 cm. The majority of the soil was dry-shaken or wet washed in the field using

a 1 cm2 mesh sieve (depending on the whether water was present).

Since it was not possible to collect 100% of AGB in the quadrate by the initial

clipping, the remaining residual AGB (the stubble left over after clipping) was removed

from the BGB cores and placed in a separate bag. The amount of residual AGB per core-

surface area was extrapolated to estimate the total residual AGB not collected in the field

from the 1 m2 quadrate. This number was later added to the live biomass component of

AGB.

Sample processing

All samples were transported from Okeechobee County to Gainesville, Florida for

post-collection processing. Rather than homogenizing all biomass within the quadrate,

AGB of each species was sorted into living and senesced life stages. Both life stages of

individual species were sorted, weighed and analyzed separately as components of the

total biomass in the quadrate. Relative dominance of each species was determined based

on the quantity of living and senesced biomass relative to other species in the quadrate.

Above ground biomass was sorted into primary, secondary, tertiary, etc, and residual or

unidentifiable species. Due to the large quantity of AGB per quadrate, large

homogeneous samples, such as upland forage species, were sub-sampled and sorted by

life stage. Using the ratio of living to senesced biomass from sub-samples and the total









biomass of the homogeneous sample, the live and senesced portions were calculated

without processing all of the biomass.

Below ground biomass was washed and sieved to remove any remaining soil. All

sorted AGB and washed BGB samples were dried at 700C for 72 hours, and weighed.

Below ground biomass per m2 was calculated by extrapolating the biomass per core-

surface area up to 1 m2. All AGB that was sub-sampled was weighed and discarded. All

other samples were rough ground, sub-sampled and fine ground to pass through a # 40

sieve.

Laboratory Analysis

All samples were analyzed for Total Phosphorus (TP), Total Carbon (TC), and

Total Nitrogen (TN), although, since P dynamics are the primary focus in these studies,

only P data is presented in this chapter. Total P was extracted from 0.2-0.5 g of plant

tissue using the ignition method (Andersen, 1976).

Data and Statistics Analysis

Data were averaged by ranch (site), and therefore are averages from two wetlands.

Statistical comparisons were made between zones at each site. Sites were not statistically

compared. JMP Statistical Software was used to perform data analyses. For mean

comparisons of more than two parameters the Tukey-Kramer HSD (honestly significant

difference) test was used (JMP, 1989-2005). Outliers greater than four standard

deviations from the mean were excluded from data analysis. All quadrate values, except

outliers, were included in the calculation and statistical comparisons of total biomass,

total P storage, AGB and BGB. Calculations and comparisons of root-to-shoot ratios

only included quadrates that contained both BGB and AGB values.









Hydroperiods were determined for each quadrate elevation using stage data from a

pressure transducer located in the center of each wetland. To determine the length of

time the water level was above the soil surface, a bench mark elevation, which correlated

to the transducer level in the well, was established at the ground surface by the well. All

quadrate elevations were corrected relative to the benchmark elevation and the

transducer. The transducer recorded stage every half hour. The number of half hours the

water level was above the ground was counted and converted to days. A regression of

days vs. elevation (stage) was developed for each wetland. Corrected quadrate elevations

were entered into the regression equation and the corresponding hydroperiod was

returned. Zone hydroperiods were an average of all quadrates within each specified zone

at each site (i.e., Beaty center hydroperiod was the average of all quadrates within the

center zones of both wetlands).

Results

SAppendix B contains numerous tables and figures of supplemental data and
statistical comparisons.

Species Composition along a Hydrologic Gradient

Center, edge and upland zones within each wetland were determined visually using

vegetative community compositions and aerial images. Overall, center, edge and upland

zones at the Beaty site had longer hydroperiods than zones at the Larson site (Table 2-1).

The average hydroperiods of center and edge zones at the Beaty site were -122 and -96

days longer than the same zones at Larson. In fact, Beaty edge zones had similar

hydroperiods as Larson center zones.










Table 2-1. Mean and standard deviation of hydroperiods at each site.
Hydroperiod
Site Zone n (days) *Difference p value
Center 38 269 48.9 a
Beaty Edge 38 141 67.4 b < 0.01
Upland 38 31.2 64.9 c

Center 33 147 63.9 a
Larson Edge 34 44.5 28.4 b < 0.01
Upland 47 9.11 15.3 c
Mean comparisons using Tukey-Kramer HSD test.


Table 2-2. Mean species hydroperiod of both sites.
Species Hydroperiod by Site
Site Species n Indicator Days Range Difference p value
Andropogon 4 FAC 77.7 68.2 0-128 BC
Baccopa 1 OBL 150 150-150 ABC
Eleocharis 1 OBL 312 312-312 ABC
Juncus 30 OBL 155 74.3 0-302 B
Ludwigia repens 3 OBL 239 67.7 161-283 AB
Luziola + P. acuminatum 3 FACW 154 6.35 150-161 ABC
Micranthemum 1 OBL 161 161-161 ABC
Beaty Other 32 178 95.7 0-314 B < 0.01
P. notatum 39 UPL 45.3 70.9 0-304 C
Panicum 31 OBL 250 86.5 0-323 A
Polygonum 14 OBL 215 88.8 0-314 AB
Pontederia 19 OBL 269 38.7 171-315 A
Sagittaria 1 OBL 297 297-297 ABC
Utricularia 1 OBL 298 298-298 ABC

Alternanthera 10 OBL 56.9 38.5 16-121 YZ
Eleocharis 2 OBL 43.5 12 35-52 XYZ
Juncus 8 OBL 54.4 27.5 0-87 YZ
Ludwigia repens 1 OBL 67 67-67 XYZ
Luziola + P. acuminatum 25 FACW 127 72.2 16-240 X
Larson Other 30 66.1 68.9 0-282 Y <0.01

P. notatum 42 UPL 11.4 17.8 0-66 Z
Panicum 2 OBL 66 72.1 15-117 XYZ
Polygonum 16 OBL 93.6 52.6 15-195 XY
Pontederia 7 OBL 164 54.2 70-227 X
Shaded areas represent species that were present at both ranches. Indicators: (OBL)
Obligate Wetland, (FACW) Facultative Wetland, (FAC) Facultative, (UPL) Upland
Species (Tobe et al., 1998).










Average hydroperiods of species present at both sites were longer at the Beaty

wetlands than at Larson (Table 2-2). These differences are similar to the zone

hydroperiod differences between sites (Table 2-1). For instance, the average hydroperiod

of Juncus effusis, Pontederia cordata, and Polygonum hydropiperoides were 100, 105

and 121 days longer at Beaty than at Larson.


1.00X species
0.90- : > z Alternanthera
0 0 A ^X. Andropogon

.0.70 + XX Eleocharis
I 'o muncus
o 0.60- o ] + + Ludwigia
10.50- oo o ++ 1 Luzioa + P.acumintum
0O.50 o N- + + Micr+nthemm+


l .2- ,PanicMn
0.20- Polygonum
S0.0 x Pontederia
So 7* Sagittaria

0.00- Ultriculari
0 40 80 120 160 200 240 280 320
Hydroperlod (days)

Figure 2-5. Logistics fit of species. Negative log-likelihood or uncertainty relative to
hydroperiod for all wetlands. This figure illustrates the relative dominance of
species as a percentage of the total species present at a given hydroperiod
Community biodiversity is greatest between upland and center zones
(R2=0.096)

The logistics fit of species (Figures 2-5 and 2-6) account for the likelihood that a

species will be present under a given hydroperiod. It quantifies dominance of individual

species relative to other species at the same hydroperiod based on frequency of

occurrence. It does not quantify biomass. Figure 2-5 shows the species distribution

relative to hydroperiod in all wetlands. The Beaty wetlands, overall, had greater










biodiversity than Larson as measured by the number of species present. At both sites,

community biodiversity was greatest in the transitional-edge zones between upland and

center zones (Figures 2-6 A & B).


80 120 160 200
Hydroperiod (days)


1.00
0.90
.0.80
0.70
0.60
10.50
5 0.40
*|0.30
(n0.20
0.10
0.00


, z z z Z 2
0 40 80 120 160 200 240 280
Hydroperiod (days)


Species
z Alternanthera 0 Other
Andropogon a P. notatum
Baccopa t Panicum
Eleocharis Polygonum
o Juncus x Pontederia
Ludwigia Sagittaria
a Luziola + P. acuminatun Utricularia
Figure 2-6. Logistics fit of species by site (-log-likelihood). These figures illustrate the
relative dominance of species as a percentage of the total species present at a
given hydroperiod. A). Species distribution at the Beaty (R2=0.17) site is
dominated by bahiagrass in the upland and Panicum and Pontederia in the
center. B). Larson (R2=0.19) species distribution is also dominated by
bahiagrass in the upland, however, a mix of Luziolafluitans and Paspalum
acuminatum dominate the center zones.

Ecosystem Phosphorus Storage

For the purpose of this study, total P storage in wetlands (with a 50 m upland

buffer) included soil (10cm depth), AGB, BGB and litter components. At both sites, soil

was the primary P storage component, representing greater than 88% of the total P

storage in the wetlands, while BGB, AGB and litter represented 8%, 3%, and 1%

respectively (Figure 2-7). Harvested BGB was significantly greater (a = 0.05) than











standing AGB at the Beaty site (Figure 2-8 A), however, there were no significant P

storage differences between BGB and AGB at either site (Figure 2-8 B).


20,000

18,000

16,000

14,000

12,000

10,000

8,000

6,000

4,000

2,000


BGB Litter Soil AGB
Component


Figure 2-7. Phosphorus storage components. Soil within the top 10 cm stores more than
88% of the total P stored in these four components, while BGB, litter and
AGB roughly account for 8%, 1% and 3%, respectively. Table B-l in
Appendix B contains P storage totals by site, zone and component.

A. B.


2,500


2,000

'E
a 1,500


1,000


500


0


BGB AGB


2,500


2,000


1,500
E

1,000
IL


500o


0


T Beaty U Larson


T T


BGB


AGB


Component Component
Figure 2-8. Comparison of AGB and BGB components at Beaty and Larson. A). Mean
biomass harvested from all zones. B) Mean P harvested from all zones.













Standing Biomass

Total biomass (AGB+BGB) in upland zones, which mainly consisted of forage

grass, were similar at both sites, -1,900 g m-2. There were no significant differences in

total biomass between zones at the Beaty site, however Larson edge zones were

significantly greater (a = 0.05) than centers, and upland zones were significantly greater

(a = 0.05) than edges (Figure 2-9). The same relationship was true for BGB, which

accounted for 68-93% of total biomass (Figure 2-10). Upland BGB and AGB were

-2
similar at both sites; -1700 and -240 g m-2 respectively. Mean BGB was larger than

AGB in all zones at both sites (Figure 2-12). Center and edge zones at Beaty had

significantly greater (a = 0.05) AGB than upland zones, while Larson AGB was greater

in upland zones than in edge zones (Figure 2-11).

A. B.
3,000
a
3,000 a


9a 2,000

500 1,500
E 2,000 E 2 b


o 1,000 1,000
1 ,ooo 1 ,oo
500 500


Center Edge Upland enter Edge Upland
Beaty Larson
Site and Zone Site and Zone
Figure 2-9. Total biomass at Beaty (A) and Larson (B) wetlands. Different lower case
letters indicate significant differences. Note the difference in scales.















T i


Edge Uiand


2,500

2,000
E
1 1,500

S1,000

* 500


Edge -ald


Bey|| Larson
Zone and Sit Zone and Site
Figure 2-10. Below ground biomass at Beaty (A) and Larson (B) wetlands. Different
lower case letters indicate significant differences.


400


"0

I 2W





0


.- I Laman
Zoneand Sie Zneand Site
Figure 2-11. Above ground biomass at Beaty (A) and Larson (B) wetlands. Different
lower case letters indicate significant differences. Note the difference in
scales.


z600

zcxx


S1,000

S500-


0 Ie- I
Center


aoo


600

E
0 400




I a

0


-

-


Center










A. B.


500
0
;' (500)
S(1,000)
S(1,500)
S(2,000)
(2,500)
300 0fl


500
0

(500)
S(1,000)
S(1,500)
S(2,000)
(2,500)
(3.000)


.V lj ytJ i
Center Edge Upland Center Edge Upland

Beaty Larson
Zone and Site Zone and Site

Figure 2-12. Biomass partitioning AGB vs. BGB. At both sites, Beaty (A) and Larson
(B) there was more BGB than AGB in all zones

Approximately 20% of the quadrates did not have measurable biomass for both

AGB and BGB components. Therefore, since total biomass is the sum of AGB and BGB,

one component made up 100% of the total biomass for -20% of all quadrates. This

occurred when the quantity (mass) of biomass within individual quadrates was below the

harvestable threshold. In some zones standing AGB was limited by grazing, while the

quantity of harvested BGB was dependent on the types of species present within

individual quadrates. It is possible for the total biomass of a quadrate to be composted of

100% AGB and no BGB. For example, spreading ground cover species such as P.

hydropiperoides, Panicum hemitomon, Luziola fluitans and Paspalum acuminatum may

have been rooted outside of the quadrate, but AGB from plants may have grown into the

quadrate.

Since BGB root to shoot ratios (Table 2-3) were only calculated in quadrates that

contained both AGB and BGB components, they are slightly different from relative AGB

and BGB zonal averages (Figure 2-12). Ratios at both sites were greater than one in all


T
T






T-









zones, indicating that BGB was greater than AGB. Ratios at Beaty were lower than

Larson and were not significantly different by zone. Larson edge zones had significantly

greater (a = 0.05) ratios than the upland zones (Table 2-3).

Table 2-3. Root to shoot ratios by zone.
BGB:AGB
Site Zone n Ratio *Difference p value
Center 31 5.73 6.21 a
Beaty Edge 29 8.36 22.2 a 0.52
Upland 31 10.4 16.3 a

Center 25 23.4 40.5 a,b
Larson Edge 28 149 340 a 0.02
Upland 39 17.5 48.5 b

Mean comparisons using Tukey-Kramer HSD test.


Standing Biomass by Individual Species

Unidentifiable AGB species were collectively labeled as "other", and represent a

combination of multiple species. The "other" category often yielded similar biomass

values as identifiable species. Overall J. effusis had the most AGB at the Beaty site,

while Paspalum notatum (bahiagrass) was greatest at Larson (Figure 2-13). At both sites

bahiagrass had the greatest AGB in upland zones. Biomass in edge and center zones at

Beaty was dominated by J. effusis, P. hemitomon, P. hydropiperoides, and P. cordata,

while Larson edge and center zones were dominated by J. effusis, P. hydropiperoides,

and P. cordata (Figure B-2, Appendix B)





























IT



& a-u' u E E E E u
00m .c W E Z


n 2
Wc O E


500

450

400

350

300

250

200-

150-



50




0 -0
w n


Figure 2-13. Above ground biomass by species for all zones (center, edge and upland).
A) Beaty wetlands. B) Larson wetlands

Phosphorus Storage in Biomass

Phosphorus storage in total biomass (AGB + BGB) was positively related to

hydroperiod at Beaty, while Larson wetlands had an inverse relationship (Figure 2-14).

Phosphorus storage in edge and upland zones did not differ much between sites, but

-2
Beaty center zones stored -1000 mg m-2 more P in total biomass than Larson centers.

Center zones stored significantly more (a= 0.05) P than uplands at Beaty, while the

opposite trend existed at Larson (Figure 2-14).












A. B.
3,000
a 2,000 a
2,500 a

2,000 b 1,500

E 1,500 E
0) 1,000
E E
1,000 -
500 -
500


Center Edge Upland Center Edge Upland
Beaty Larson
Site andZone Site and Zone

Figure 2-14. Total biomass P storage. A). Beaty wetlands were positively related to
hydroperiod, while Larson had an inversely relationship (a= 0.05). Different
lower case letters indicate significant differences between zones at the same
ranch sites

Below ground biomass P concentrations (Table 2-4) and storage (Figure 2-15 A)

did not differ between zones at the Beaty wetlands. At Larson, P concentrations in

upland BGB were significantly lower (a= 0.05) than center and edge concentrations

(Table 2-4). However, there was still a general trend of decreasing P storage from center

to upland (Figure 2-15 B), where upland and edge BGB stored significantly more (a=

0.05) P than center zones.

Phosphorus concentrations were significantly greater in AGB than BGB in all

zones at both sites. At Beaty, P concentrations were greater in center zones than in edge

and uplands. At Larson all zones were significantly different (a = 0.05); exhibiting a

positive relationship with hydroperiod (Table 2-5). The center and edge zones at Beaty

stored more P in AGB than upland zones, while Larson center zones stored more P than










edge zones (Figure 2-16). Phosphorus storage in BGB is larger than AGB in all zones at

Beaty and in edge and upland zones at Larson. While BGB made up the largest portion

of total biomass in all zones, AGB P concentrations drastically influenced total biomass P

storage. This was exhibited in Larson centers where AGB P storage was greater than

BGB (Figure 2-17 B) despite the fact that harvested BGB was greater than harvested

AGB.

Table 2-4. Below ground biomass concentrations by zone
Below Ground TP Concentration
Site Zone n (mg/kg) *Difference p value
Center 34 765 210 a
Beaty Edge 34 719 285 a 0.23
Upland 34 674 134 a

Center 28 781 136 a
Larson Edge 29 802 168 a 0.002
Upland 40 678 145 b
Mean comparisons using Tukey-Kramer HSD test.


A. B.
a
2,000 2,000
a
1,600 1a a 1,600

1,200 IE 1,200
E E



400400

0 0
Center Edge Upland Center Edge Upland
Beaty Larson
Zone and Site Zone and Site
Figure 2-15. Phosphorus storage in BGB at Beaty (A) and Larson (B) wetlands.
Different lower case letters indicate significant differences a= 0.05.










Table 2-5. Above ground biomass P concentrations by zone.
Above Ground TP Concentration
Site Zone n (mg/kg) *Difference p value
Center 122 1830 976 a
Beaty Edge 118 1340 702 b < 0.01
Upland 68 1340 623 b

Center 69 3150 1010 a
Larson Edge 80 2690 1060 b < 0.01
Upland 89 1650 647 c

Mean comparisons using Tukey-Kramer HSD test.

A. B.
a 1,2o a
1,000
1,000
800










Beaty Larson
Soo

E E 0oo



200 200


Center Edge U0and center Edge and
Beaty Larson
Zone and Site Zne and Site

Figure 2-16. Phosphorus storage in AGB at Beaty (A) and Larson (B) wetlands.
Different lower case letters indicate significant differences (a = 0.05).

The P storage root-to-shoot ratios were calculated the same way as the harvested

biomass root-to-shoot ratios; only quadrates that contained both BGB and AGB were

used in the calculation. All ratios were greater than one and did not differ significantly

by zone, meaning P storage was greatest in BGB. These ratios contradict mean AGB and

BGB values in Larson centers. Overall, ratio data suggests that AGB stores more P than

BGB in Larson centers (Table 2-6). The difference, once again, is that the ratios (Table

2-6) are an average of individual quadrate ratios within each respective zone, which only










included quadrates that had both AGB and BGB, where as P storage values (Figure 2-17)

of each component were averages of all harvested AGB and BGB with each respective

zone.

A. B.
1,500 1,500
1,000 1,000
5001,00
500
0
(05
E (500) E


(1,500) (1 1,000)
(2,000) (1,500)
(2,500) (2,000)
Center Edge Upland Center Edge Upland

Beaty Iarson
Zone and Site Zone and Site
Figure 2-17. Above and below ground biomass P storage.

Table 2-6. BGB to AGB P storage ratios
BGB:AGB P Storage
Site Zone n Ratio *Difference p value
Center 31 3.28 3.94 a
Beaty Edge 29 6.22 14.8 a 0.45
Upland 31 6.30 9.86 a

Center 23 6.50 10.2 a
Larson Edge 28 28.4 65.6 a 0.0548
Upland 39 6.76 15.3 a
Mean comparisons using Tukey-Kramer HSD test.

Standing Biomass by Individual Species

Compared to other species, P. hydropiperoides, which was predominately present

in center and edge zones, stored the largest amount of P in AGB at both sites (Figures 2-

18). J. effusis, P. hemitomon and "other" species were secondary AGB P storage species

in center and edge zones at Beaty (Figure B-4, Appendix B). "Other" AGB was a

secondary storage in Larson center and edge zones. Bahiagrass stored the most P in












uplands at both sites. At Larson, bahiagrass and J. effusis had similar P storage in upland


and center zones.


1400

1300

1200

1100

1000

900

800
E 700
E 600
500

400

300

200

100
0-5


800


700


600


500

E 400
E

300


0O


100


0


~m""EE`E E m
0) 0 CU
0 0 0 o u E

0 W J ~
0
LU 0- 0- 0-+
< CU Io~


L EE



Lu oi
oi
W ii:+


Figure 2-18. Phosphorus storage by species. A). Beaty ranch. B). Larson ranch.


Discussion


Hydrology


Hydrology controls many physicochemical mechanisms in wetlands and is the most


important determinant of wetland type and class (Kadlec & Knight, 1996; Mitsch &


Gosselink, 2000). Hydroperiod represents the number of days a year a wetland is


inundated. The hydro-pattern or hydrologic regime is characterized by five components:


1) duration, 2) frequency, 3) depth, 4) flow, and 5) timing or season of flooding. All of


these components influence the establishment of vegetation and nutrient availability









along hydrologic gradients. The wetlands in this study were under different hydroperiods

and presumably, different hydrologic regimes as the effectiveness of the ditches varied

between wetlands and sites.

Many external factors influence hydrology making it difficult to compare wetlands

under different management intensities. For instance, ditches at the Larson site

effectively drained more wetland area than ditches at Beaty. As a result, Beaty wetlands

were inundated most of the year, while the Larson wetlands were only inundated roughly

half of a year. Anaerobic conditions create an environment that is conducive for organic

matter accumulation and P immobilization. Thus, Beaty has more potential to

accumulate organic matter.

Species biodiversity was greatest in transitional edge zones, between upland and

center zones. Relatively stable environments under short or long hydroperiods favor the

establishment of obligate species communities (i.e. bahiagrass in the upland or Panicum

and Pontederia in wetland centers). While under moderate hydroperiods in the edge

zones ecotoness), fluctuating hydrologic conditions continuously eliminate and

regenerate species, decreasing mono-dominance and increasing facultative species

richness.

Ecosystem storage

Although P storage in soil is dependent on various site physicochemical

characteristics, it is often the primary storage component in wetland and terrestrial

ecosystems (Dolan et al., 1981; Richardson, 1985). Results from this study support Hi:

the role of vegetation in total P storage is significantly less than soil P storage in

wetlands. Although vegetation plays a lesser role in terms of active P storage, its









importance can not be overstated. It indirectly increases the P storage capacity of

wetlands as residual biomass decomposes and becomes incorporated into the soil.

Biomass in Pasture Wetlands

Biomass production studies in pasture wetlands are limited, probably due to the

intrinsically high variability of disturbances to these ecosystems. Different land use

intensity, hydrologic regimes and grazing pressure create high variability, and make it

difficult to compare wetlands at different sites. Many studies have identified

relationships between environmental gradients (i.e. hydrology) and vegetation

community establishment under stable conditions (Seabloom et al., 2001; Van der Valk,

A G, 1981; Van der Valk, A G et al., 1994; Wellings et al., 1988; Whigham et al., 2002).

However, species distribution and annual biomass are generally variable over time and

reflect a balance of current environmental conditions and historical recruitment events

(Seabloom et al., 2001; Whigham et al., 2002). Our study minimizes temporal variability

over one year, but does not represent long term seasonal variability that may be present in

response to climatic variability.

Generally, environmental gradients create patterns of biomass distribution within

wetlands. A three year study of restored agricultural wetlands in Maryland found AGB

was inversely related to hydroperiod (Whigham et al., 2002). In that study, wetlands were

divided into submersed, emergent/seasonal, and temporary zones; similar to the center,

edge and upland zones in our study, however, it appears that their zones my have been

under longer hydroperiods. Biomass was significantly greater in temporary zones

[uplands] than in the emergent/seasonal zones [edge] and the emergent/seasonal zones

were significantly greater than the submersed zones [center] in each of the three years.

Even if their zones were under longer hydroperiods, our AGB results do not support any









relationship to hydroperiod, indicating that other factors, such as management intensity,

are influencing AGB.

Overall, BGB was the largest portion of total biomass, regardless of zone. This

was not surprising since emergent wetland and terrestrial macrophytes generally have

greater BGB than AGB due to extensive networks of roots and rhizomes (Reddy et al.,

1999a). The inverse relationship between BGB and hydroperiod at the Larson site may

be the result of more intensive grazing pressure. There is an observable difference in

grazing intensity between the two ranches. While cows do not graze BGB, heavily

grazed AGB directly affects BGB production and plant survivability. Total biomass at

Larson had the same inverse relationship to hydroperiod. Larson wetland zones were

more heavily grazed and dominated by low-growing, unidentified annuals, Eleocharis

spp., and L. fluitans. Intense grazing followed by prolonged inundation discourages

recruitment of perennial species. Thus during flooding events, low growing species do

not survive inundated conditions. Ultimately, when water levels recede, bare soil is

exposed and subject to mineralization and erosion. The combination of intensive grazing

and flooding primarily favors annual species in Larson wetlands.

Disturbance Effects on Biomass

Harvested biomass results do not support H2: Biomass at Larson and Beaty can not

be correlated with hydroperiod. Low AGB and large, highly variable ratios in edge zones

are likely the result of the compounding effects of the hydrologic regime, intense pasture

management and higher grazing density. Once again this suggests a significant effect of

grazing on biomass and P storage.

External disturbances occur at multiple scales and can confound relationships

between environmental gradients and wetland structure (Magee & Kentula, 2005; Van









der Valk, A G et al., 1994). Direct impacts of grazing on wetlands often include

herbivory of vegetation, nutrient inputs, and soil trampling; all of which directly or

indirectly alter species composition (Clary, 1995; Steinman et al., 2003). Grazing was

not measured in this study; however, it is evident, based on these results and visual

observations, that grazing has a dramatic affect on species composition and standing

AGB. Bohlen et al. (2004) found differences in plant species assemblages in improved

and semi-improved pastures. In their study, the less intensively grazed, semi-improved

pastures were dominated by P. hemitomon, which has high forage value, while

intensively grazed, improved pastures supported more diverse plant communities

including J. effusis. In addition, they found that cattle exclusions within improved

pastures lead to an increase in P. hemitomon coverage. This suggests that preferential

grazing ofP. hemitomon may actually foster species biodiversity including unpalatable

species such as J. effusis (Bohlen et al., 2004).

Biomass results from this study suggest that there is high variability within the

same land use classification. Although all wetlands are in improved pastures, wetland

center and edge biomass results are not even similar between ranches Thus, one

limitation of this study, was a lack of replication among sites. To minimize variability,

site data contained mean values of both wetlands at each ranch. Whigham et al., (2002)

also found high variability between sites. This suggests that different management

intensities within the same land use can be highly variable between sites and may not

represent biomass dynamics within individual wetlands. Thus, comparisons between

sites should take into account management intensity and land use type.






47


Phosphorus Concentrations

Higher AGB P concentrations in center zones at both Larson and Beaty were

similar to the trend found by Whigham et al., (2002). In two out of the three years, AGB

P concentrations in the Maryland wetlands had a positive relationship with hydroperiod;

opposite of the standing AGB trend. They concluded that nutrient cycling processes are

less variable than spatial and temporal biomass differences (Whigham et al., 2002).

Another study found that vegetation in wetlands receiving treated sewage effluent

showed increased P concentrations in AGB in response to both increased water levels and

nutrient additions (Bayley et al., 1985).


z
O
Vmax

I-L


I / Luxury consumption LU
/ I
/>/ -
I- I I
Z
UJI
I A Nutrient Nutrient Nutrient 2
$/4/ I
z deficiencyy sufficiency toxicity

NUTRIENT SUPPLY -


Figure 2-19. Nutrient storage and growth in plants. Growth is typically maximized at
lower nutrient supplies than the maximum tissue storage potential (Reddy &
Debusk, 1987)

It is hypothesized that increased P availability in wetlands centers may facilitate

"luxury uptake" of P by obligate wetland species. This occurs when plants take up P

beyond their required needs for growth (Figure 2-19). Biomass production is usually

maximized at lower nutrient supplies, while nutrient uptake by plants is maximized at









higher nutrient levels. The difference between the growth and nutrient uptake rates is the

P storage potential (Reddy & Debusk, 1987).



Phosphorus Storage

Tissue P concentrations are also temporally and spatially variable and are not a

reliable indicator of long-term P storage. They can vary with plant age, season and

nutrient availability. For instance, P concentrations are typically higher in younger plants

than in mature plants (Reddy & Debusk, 1987). Phosphorus storage potential in plants is

a function of both tissue concentrations and the maximum standing crop (Reddy et al.,

1995; Reddy & Debusk, 1987). The maximum standing crop is often the primary

determinant of P storage. For example, Sagittaria latifolia had the greatest P

concentration of any species; however, because it was not prevalent in the wetlands, the

amount of P stored was relatively small. Whigham et al. (2002) found that P storage

varied between wetlands, but exhibited similar patterns of distribution as standing AGB.

Overall, total P storage at Beaty was positively related to hydroperiod, while

Larson was inversely related. The influence ofbiomass as the primary component of

total biomass P storage is evident at the Larson site. The total biomass (Figure 2-9 B)

and P storage graphs (Figure 2-14 B) exhibit similar general trends between zones.

However, vegetation in Beaty centers stored significantly more P than upland zones, even

though biomass results (Figure 2-9 A) did not differ by zone. Therefore, differences in P

concentrations along a hydrologic gradient are also influencing total P storage.

Reddy and Debusk (1987) found greater than 50% of the nutrients in emergent

macrophytes were stored in BGB portions of plants. Results from this study suggest that

the relative roles of biomass and concentration in P storage may vary between AGB and









BGB components. Since BGB makes up the majority of the total biomass in all zones at

both sites, it was expected to store the most P. Although it was not statistically

significant, AGB in Larson centers stored more P than BGB. Thus, high P concentrations

in AGB had a greater influence on P storage than biomass in the center zones at Larson.

Phosphorus storage results do not conclusively support either H3: total biomass P

storage would be greater in wetlands than uplands, or H4: more P will be stored in AGB

than BGB. Since the Beaty wetlands stored more P in center zones, and the Larson

wetlands showed the opposite trend, there is no conclusive trend and H3 was rejected.

Since BGB stored significantly more P than AGB in all zones except Larson centers, H4

was also rejected.

Conclusions

Based on biomass results that lack specific trends, and opposite P storage trends

along a hydrologic gradient, it is hypothesized that altered hydrology, management

intensity and grazing may be influencing environmental gradients in the Okeechobee

wetlands. The positive relationship between total biomass P storage and hydroperiod at

the Beaty site may be the combined result of longer hydroperiods and lower management

intensity (including grazing pressure) relative to Larson.

Despite ongoing disturbances (grazing) to these wetlands, P concentration gradients

in vegetation, which were positively related to hydroperiod at both sites, are similar to

those found in other studies. However, P storage in vegetation is short-term, highly

variable, and represents less than 10% of total P storage in wetlands. Soil stores the

majority of the total P in these wetlands; up to 90%. Below ground biomass and P

storage is greater than AGB in all zones with the exception of P storage in Larson

centers.









Historically high net P imports to the watershed have saturated the P assimilative

capacity of some wetlands, making them P sources rather than sinks. Hydrologic

restoration would increase HRT, anaerobic conditions and organic matter accumulation.

Presumably, over a prolonged period of time, if hydrology were restored, P imports were

significantly decreased, and grazing pressure was minimized, wetland P assimilative

capacity would increase, thus reducing P exports to the Lake. In addition to reducing P

loads to the Lake, restoration also stores water in the landscape, which potentially

reduces the Lake stage and discharge of fresh water to the coasts.














CHAPTER 3
FACILATATING WETLAND HYDROLOGIC RESTORATION WHILE
MAINTAINING FORAGE PRODUCTION: HYDROLOGIC TOLERANCES OF
PASPAL UM NOTA TUM AND HEMARTHRIA AL TISSIMA

Introduction

Background

Hydrologic restoration of historically isolated wetlands in the Lake Okeechobee

watershed is considered a Best Management Practice (BMP) to decrease Phosphorus (P)

loading to the Lake. The watershed has low geographic relief and many isolated

wetlands have been drained to create improved conditions for upland forage grass

species. Restoration of drained isolated wetlands involves blocking ditches or installing

water control structures to raise the water table back to historical levels, thus retaining

water and P within these wetlands. An increase in wetland stage could greatly expand

wetland footprints and zones of inundation, thus changing hydroperiods and hydrologic

regimes of restored wetlands. Long-term flooding with decreased stage fluctuations

would likely alter existing vegetative communities along hydrologic gradients, decreasing

upland forage productivity in areas adjacent to wetlands. Since hydrologic restoration of

isolated wetlands reveres the current management objective, landowner acceptance of

this BMP may depend on the introduction of alternative forage grass species that are

tolerant of prolonged hydroperiods and less frequent stage fluctuations.

The most commonly used forage species in Florida is Paspalum notatum Fl,,i,

('Pensacola' bahiagrass). Native to Central and South America, bahiagrass is a deep

rooted, warm-season perennial grass that was originally planted for forage and soil









stabilization in the southern United States(Violi, 2000). Bahiagrass is a resilient, low

maintenance species that is tolerant of a wide range of hydrologic and soil conditions;

however, it is best adapted to moist, sandy soils. It forms tough sod mats with a vast

network of stolons and roots, often to a depth of seven feet. (Chambliss & Adjei, 2006;

Violi). Ninety percent of its forage production occurs between April and September

(Mislevy, 2002). While bahiagrass does not seem to invade established communities, it

does dominate habitats and resists invasion from other species (Violi, 2000). Once

established, it is difficult to remove. It has been estimated that bahiagrass stolons can

store enough nutrients to remain viable for two to three years (Chambliss & Adjei, 2006).

Hemarthria altissima 'Floralta' (limpograss) is a forage species that has gained

popularity since it was introduced (USDA Plant Introduction 364888) in 1984. Native to

South Africa, limpograss was originally selected for its winter hardiness, producing as

much as 35% of its total annual production between November and March (Pate, 1998).

Limpograss was specifically selected for its persistence under grazing (Quesenberry et

al., 1984). It was the fourth limpograss cultivar released in Florida and is currently the

only one recommended for pasture establishment (Pate, 1998; Sollenberger et al., 2006).

Contrary to bahiagrass, it is best adapted to poorly drained sandy soils and is not

recommended for drought sands (Pate, 1998; Sollenberger et al., 2006). In fact, it grows

well in wet areas that are often continuously flooded during the wet season (Pate, 1998).

Both bahiagrass and limpograss are exotic species as defined by the Florida

Exotic Pest Plant Council. Bahiagrass is a naturalized exotic that was once listed as a

Category I invasive exotic but has since been removed from the list. Limpograss is a

listed as a Category II invasive exotic; meaning that it shows the potential to disrupt









native plant communities but has not yet increased in abundance and frequency to be

considered a major nuisance species. (FLEPPC, 2005).

Research Objectives

Previous studies have evaluated the forage quality of and animal performance on

bahiagrass and limpograss (Holderbaum et al., 1991; Holderbaum et al., 1992;

Kalmbacher, R. S. et al., 1984; Kalmbacher, R. et al., 1998; Long et al., 1986; Newman

et al., 2002a; Newman et al., 2002b; Pate, 1998; Quesenberry et al., 1984; Sollenberger et

al., 1988; Sollenberger et al., 1989), however for the purpose of this study the primary

objectives were to evaluate survivability, productivity and P storage under different

hydrologic conditions.

Limpograss has been recommended for use in moist sites in Florida (Sollenberger

et al., 2006), however, its specific hydrologic tolerance has not been evaluated.

Bahiagrass has shown short term tolerance to flooding (David, 1999), however,

ultimately over time it gets out competed by wetland species. There are multiple

environmental factors that determine ideal habitats for species, such as grazing intensity,

hydrologic regime, competition, and soil conditions. Hydrology can influence

competition and physicochemical soil interactions. It is often considered one of the most

influential determinants of establishment and persistence of wetlands plants (Mitsch &

Gosselink, 2000). The objective of this research was to evaluate the role of hydrology on

bahiagrass and limpograss in non-competitive mesocosm studies.

Research Questions and Hypotheses


1. Which species has greater forage production?

Hi: Limpograss will have greater forage production than bahiagrass









2. What are the hydrologic tolerances of bahiagrass and limpograss?

H2: Bahiagrass will have greater total biomass production in drier
treatments and limpograss will have greater total biomass production
in the wetter treatments.

3. Which species assimilates more P?

H3: Limpograss will have higher P storage

4. Where is P partitioned within the plant?

H4: Root to shoot P storage ratios for bahiagrass will be >1, and
limpograss will be <1

This chapter compares the effect of five different water level treatments on below

ground biomass (BGB) and above ground biomass (AGB) production and P storage of

both species in non-competitive mesocosm studies.

Materials and Methods

Experimental Design

This experiment was designed to evaluate the response of two forage grass species

to a range of hydrologic conditions typical of the transitional zone between isolated

wetland and improved upland pasture. To determine how hydrology affects productivity

and nutrient uptake, limpograss and bahiagrass were evaluated in fifteen non-competitive

mesocosms (1.33 m x 0.81 m x 0.76 m polyethylene tubs). Mesocosms were located in

Gainesville, Florida (29.60 N 82.30 W).

The experiment consisted of five treatments +10, 0, -10, and -15 (water levels in

centimeters, relative to the soil surface), and a control (rain water only and well drained),

which are discussed in the next section. There were three replicate mesocosms for each

treatment. Each mesocosm contained three sub-replicates (pots) of each species. Sub-

replicate samples were combined together into one composite sample of each species.









The sub-replicates were grown in 3 gallon (25 cm diameter x 20 cm deep) poly-

ethylene pots. Soil was collected from a pasture in Okeechobee, Florida, and

homogenized before being dispensed into pots. Propagules of both limpograss and

bahiagrass were harvested from pasture plots at the Range Cattle Research Center in Ona,

Florida. Soil was washed from the propagules and seven bare-root sprigs were planted in

each pot to establish monocultures of each species. Fifty-two pots of bahiagrass and 61

pots of limpograss were established 90 days prior to treatment. During the grown-in

phase, both species were watered regularly and pruned uniformly to stimulate new

growth.

0+10 C +10 C+10 -10-15






SC 0 -10-15 -10 0 -15
a. b.

Figure 3-1. Study site at University of Florida, Gainesville, Florida: (a) mesocosms were
aligned in two rows and randomly assigned a treatment. (b) Tubs receiving
water were hooked up to the potable water line between the rows.

Mesocosms were aligned in two rows and randomly assigned a treatment. Tanks

receiving water were hooked up to a potable water supply and external overflow stand

pipes, were used to maintain water levels for each treatment (Figure 3-1). Forty-five of

the healthiest (determined visually) pots of each species were selected and three pots of

each species were randomly placed into the 15 mesocosms. Each mesocosm contained

three pots of limpograss and three pots of bahiagrass. To simulate field conditions,

regulate temperature, and prevent oxygen production by photosynthetic algae in an open









water column, mason sand was used to fill the remaining space between pots (Figure 3-

2).
















Figure 3-2. Mesocosm diagram. Each mesocosm contained three pots of each species
embedded in mason sand. Water level in four of the five treatments was
maintained by a drip irrigation system and an external overflow-standpipe.

Treatments

Four of the five hydrologic treatments were maintained at a constant stage by drip

irrigation emitters and external overflow-standpipes, while the fifth treatment, the

control, only received rain water and was allowed to drain completely. Treatments

receiving water included an inundation treatment (+10 cm), where the water level was

maintained 10 cm above the soil surface, and three saturation treatments (0 cm, -10 cm, -

15 cm), where the water level was maintained 0, 10 and 15 cm, respectively, below the

soil surface. Rainfall data is listed in Appendix C Table C-31. These treatments will

hereinafter be referred to as +10, 0, -10, -15 and control (C).

The study was initiated (day zero) on July 1st, 2004. One week prior to this date,

water levels were gradually raised to their treatment levels. The timing coincides with

the approximate beginning of the wet season in central Florida. Soil redox was measured

in randomly selected pots of each treatment at a depth of 10 cm below the soil surface.









Redox values were inversely related to water depths suggesting the effect of saturation

and inundation reduced oxygen availability and increased anaerobic conditions in the

soils (Figure 3-3).



Water Depth




+10-


-10-
-15-

Mesocosm Redox
300
250
200




50 -
0-
10 0 -10 -15 C
Treatment


Figure 3-3. Inverse relationship of water depth and redox. This diagram illustrates the
inverse relationship of treatment water depth and measured redox potential
within the five treatments. Soil redox was measured 10 cm below the soil
surface.

Sampling

Soil, BGB, and two components of AGB were sampled over the course of one year.

The components of AGB were forage, consisting of all biomass above 15 cm, and

residual biomass (RB), the biomass that remained from 0-15 cm after harvest. Soil was

sampled at the beginning of the experiment (day 1), at the end of the first growing season










(day 163) and at the end of the experiment (day 375). Forage was harvested periodically

to evaluate temporal differences in biomass production P concentration and P

assimilation. Forage samples were collected on days 27, 55, 83, 163, 305 and 375. At the

end of the first growing season (day 163), two of the three sub-replicate pots of each

species in each mesocosm were harvested to determine BGB and RB production and P

storage. Forage, RB and BGB were harvested from the remaining pots in each mesocosm

on day 375. Sampling dates and biomass components harvested are listed in Table 3-1.

Table 3-1. Sampling dates and details.
Pots per
Date Day composite Component Sampled
sample
7/1/2004 1 3 Soil, Forage (for nutrient baseline)
7/28/2004 27 3 Forage
8/25/2004 5 3 Forage
9/22/2004 83 3 Forage
12/11/2004 163 3 Soil, Forage
12/11/2004 163 2 BGB, RB
5/2/2005 305 1 Forage
7/11/2005 375 1 Soil, Forage, RB, BGB
This table summarizes the sampling events, corresponding components sampled and
number of sub-replicates in composite samples. All pots in each tank where averaged by
species and pot (i.e., for biomass g pot- tank-1 species-1 = average g pot-' of each pot in
each tank of each species).

Soil

Two soil cores (1.8 cm diameter x 20 cm depth) from pots of the same species

within the same mesocosm were combined into one composite sample. Table 3-1

contains the number of pots in each composite. Roots and litter were removed from the

soil before being dried at 70 OC for 72 hours. The soil was than machine (ball) ground,

sieved through a #40 mesh sieve and stored at room temperature.

Above ground biomass sampling

Forage sampling was designed to simulate flash grazing, by periodically

harvesting all biomass over 15 cm. This height was established 5 cm above the highest









water level treatment to enable atmospheric gas exchange with the residual biomass for

all treatments. Composite samples of each species from each mesocosm were collected

using grass shears, a 15 cm-tall grated stand and a shop vacuum to ensure accurate

collection. The grate was set over a pot to establish the clipping height and the vacuum

was used to pull the grass through the grate and gather clipped material within the

vacuum (Figure 3-4). The vacuum was emptied after each composite sample per

mesocosm. The post-harvest processing procedure involved drying vegetation in a

drying room at 70 OC for 72 hours. Dry forage was than ground in a Wiley Mill, passed

through a #40 mesh sieve and stored at room temperature.










a. b. c

Figure 3-4. Harvesting procedure. (a) Clipping grass with 15 cm stand and vacuum. (b)
Close up view of clipping processes. (c) Vacuum was emptied after each
composite sample per mesocosm.


At the end of the first growing season (day 163) the RB was harvested from two

of the three pots within each mesocosm. The post-harvest processing procedure was the

same for all vegetative components. Residual biomass from each mesocosm was added

to the respective cumulative forage production for each species and treatment to

determine total AGB production g pot- after 163 days. The same procedure was carried

out on day 375 to determine total cumulative AGB production after 375 days.









Below ground biomass

Below ground biomass included all roots, rhizomes and stolons below the crown

of the AGB shoot. Once both components of AGB were harvested (days 163 and 375)

the root ball was removed from the pot and flushed with water to remove all soil. The

same post-harvest sample processing procedure used with AGB was also used with BGB.

Since harvesting BGB was a destructive process, sampling was only preformed twice. All

BGB data is a cumulative total and presented as BGB production or P storage after 163 or

375 days.

Laboratory analysis

Soil and biomass samples were analyzed for Total Phosphorus Ash (TP) using the

Ignition Method (Andersen, 1976), Total Nitrogen (TN) and Total Carbon (TC) as

described in Chapter II methods. Composite biomass and P storage were averaged by

treatments and reported on a grams per pot basis. In addition, soil was also analyzed for

plant available P using the Mehlich I dilute concentration strong acid extraction

procedure (Kuo, 1996).

Results

SSupplemental tables and figures containing all data and statistical comparisons are
reported in Appendix C.

Initial characterization

Daily environmental conditions including air and soil temperature, rainfall and

humidity are listed in Table C-31 of Appendix C. Total P ash (TP), total nitrogen (TN)

and total carbon (TC) tissue concentrations at day zero are listed in Table C-l in

Appendix C. Soil TP, TN and TC concentrations on day 0 averaged 0.003%, 0.092%,

and 1.65 % respectively. Forage tissue concentrations ranged from 0.15-0.18%, 1.25-









1.42%, and 42-44 % for TP, TN and TC respectively. The focus of this thesis relates to P

storage in vegetation, therefore, only TP data are reported beyond the initial conditions.

All production data for forage, RB and BGB are expressed either as production per

harvest or cumulative production (sum of net harvests over time) in grams of biomass per

pot. Total AGB is the sum of cumulative forage production and residual biomass

normalized on a grams per pot basis.

Forage Production

Data in this section is presented as overall production by each species regardless of

treatment. Overall, limpograss had significantly greater (a = 0.05) forage production per

harvest than bahiagrass on all sampling days except day 83 (Figure 3-5). In the first 83

days, both species exhibited a decline in forage production. However, by the end of the

first growing season (day 163), limpograss continued to producing forage while

bahiagrass was essentially dormant until the beginning of the following growing season.

Both species increased forage productivity between early-May and mid-July of the

second growing season. On a cumulative basis, limpograss had significantly greater

forage production on all sampling days (Figure 3-6). After 375 days, bahiagrass and

limpograss had produced 9.52 2.73 and 32.4 14.7 g pot- of forage respectively.





























I 5 5I I5 10I
0 25 50 75 100


Species
* Bahia
* Floralta


125 150 175 200 225 250 275 300 325 350 375 400
Days


Figure 3-5. Forage production per harvest for each species with all treatments combined.
Day 0 is July 1st 2004. Limpograss had significantly greater forage production
per harvest on all sampling days expect day 83 (Table C-2, Appendix C).



60- Species
SBahia

50. Floralta


40-

C.
C30-


20






0 25 50 75 100 125 150 175 200 225 250 275 300 325 350 375 400
Days


Figure 3-6. Cumulative forage production with all treatments combined. Each
consecutive harvest was added to previous harvests. Limpograss produced more
forage than bahiagrass on a 375 day period. (Table C-3, Appendix C).


35-

30-

25-
~2-
Z 20-

15-

10-

5-

0-


I
r






I







63


Bahiagrass forage production

To determine the response of each species to various water level treatments

comparisons were made between treatments of each species using the Tukey-Kramer

HSD test. There were differences in forage production between harvests (Figure 3-7A).

Throughout the experiment, the +10 and control treatments had similar forage

production. Between the second and third harvest the +10 treatment had significantly

greater forage production than the 0, -10, and -15 cm treatments. In addition, on day 375,

the control had significantly greater forage production than the 0 cm treatment. On a

cumulative basis, forage production across all five treatments ranged between 7.88 + 1.49

to 12.1 + 3.61 g pot- after 375 days and did not differ significantly between treatments.

All treatments exhibited similar cumulative production curves over time (Figure 3-7B).

A. B.
71- 1
6- 14-
5- 12-



E E

4-

0 50 100 150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
Elapsed time Elapsed time

A + 10 cm Ocm -10cm + -15cm Control


Figure 3-7. Bahiagrass treatment comparisons. A) Biomass production per harvest; +10
cm treatments significantly greater than the 0, -10, -15 cm treatments on days
83, while the Control was greater than the. B) Cumulative forage production;
no significant differences between treatments. (Tables C-10 & C-l1,
Appendix C).









Limpograss forage production

Limpograss exhibited significant treatment effects after day 27. Initially, on day

55, the control had significantly greater forage production than the +10, 0 and -10 cm

treatments. However, on day 163 the control was the only treatment with a lower net

harvest than its previous harvest on day 83 (Figure 3-8A, Table C-12, Appendix C).

Although the span of time between harvests (days 55-83 and 83-163) are different, by

day 163, the +10 and -10 treatments roughly doubled the amount of forage produced

between days 55 and 83. On day 305, the +10 cm treatment had greater forage

production than the 0, -10, and -15 cm, while on day 375 the control had greater

production than the 0, -10, and -15 cm treatments.

The differences in limpograss production per harvest did not cause significant

differences in cumulative forage production between treatments until days 305 and 375.

On day 305, the production per harvest treatment differences were mirrored by

cumulative forage production. The +10 treatment had greater cumulative production

(27.0 + 2.16 g pot-) than the 0, -10, and -15 cm treatments after 305 days. On day 375,

like the individual harvest differences, the control had greater production (50.2 + 16.5 g

pot-) than the 0, -10 and -15 cm treatments. While the +10 cm treatment produced

significantly more forage (44.5 + 3.98 g pot-) than the -10 cm treatment (21.1 + 3.31 g

pot-) after 375 days, it was not statistically greater than the 0, and -15 cm treatments,

despite a power value of 0.95.










A. B.
40
35- 60-
30- 50-
1'25-
-E 40
20-
151 30-
m 10 + m 20

0* _ __ 10-
I Ii ll I'I I l l
0 50 100 150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
Elapsed time Elapsed time

A + 10 cm 0cm O -10cm + -15cm Control


Figure 3-8. Limpograss treatment comparisons. A) Forage production per harvest. B)
Cumulative forage production (Tables C-12 & C-13, Appendix C).

Species comparison

This section compares biomass production of limpograss and bahiagrass within the

same hydrologic treatments. Limpograss had similar or greater forage production than

bahiagrass on all harvest days, in all treatments (Figure 3-9). By the first sampling,

limpograss had significantly greater forage production than bahiagrass in all treatments.

Both species exhibited a decline in forage production per harvest in all treatments after

day 27 (Figure 3-9), where only the control and -15 cm limpograss treatments were

significantly greater than bahiagrass. However, limpograss rebounded and produced

significantly more forage than bahiagrass in all treatments by the final harvest of the

growing season (day 163). The same trend continued in the second growing season,

where limpograss had significantly greater forage production per harvest than bahiagrass

with the exception of the -15 cm treatment on day305. As a result limpograss had

significantly greater cumulative forage production after every harvest day (Figure 3-10)

























150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
Elapsed time Elapsed time


0 50 100 150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
Elapsed time Elapsed time

40E.
35- Floralta
30- g Bahia
525-
0 -2 0
E1
o /
10-
5-
0
0 50 100 150 200 250 300 350 400
Elapsed time

Figure 3-9. Forage production per harvest by treatment. A) +10 cm treatment. B) 0 cm
treatment. C) -10 cm treatment. D) -15 cm treatment. E) Control. (Table C-5
Appendix C)
















a.25-
0)

c 20-


15



E
5-
10

5-


U I I I I I I I 1
0 50 100 150 200 250 300 350 400
Elapsed time


40
*

-
E 30-
0
m
a)
>20-
Co
E
E 10-
0-


0 50 100 150 200 250 300 350 400
Elapsed time


O
-20-


I 15-


.10-
E
E
S5-


0 50 100 150 200 250
Elapsed time


35-

630-
a

225-
o
2 20-
0
a15-


2
5-


300 350 400
300 350 400


-1-














50 100 150 200 250 300 350 400
Elapsed time


'60-
-
0)
;50-

E 40-

. 30-
C6
S20-
E
3110-


Floralta

I Bahia












50 100 150 200 250 300 350 400
Elapsed time


Figure 3-10. Cumulative forage production by treatment. A) +10 cm treatment. B) 0 cm
treatment. C) -10 cm treatment. D) -15 cm treatment. E) Control. (Table C-6,
Appendix C).


o a
0 0


0-cc


-if I


,,









Total Biomass

After 375 days, bahiagrass total biomass (AGB + BGB) was significantly greater

than limpograss in the 0 and -10 cm treatments (Table 3-2). This was not consistent with

total biomass results in the first 163 days, where bahiagrass had significantly greater total

biomass production than limpograss in all treatments. This is primarily due to

significantly greater bahiagrass BGB in all treatments.

There were no significant increases in BGB production in any treatments between

days 163 to 375 for either species. However, there were significant BGB decreases in

bahiagrass +10 and -15 treatments (a = 0.05), and in limpograss +10 cm (a = 0.08) and

control (a = 0.05) (Figure 3-11 and Table C-9, Appendix C). In addition, forage

production increased more in limpograss than bahiagrass during the same time period

(Appendix C, Table C-5). This offset the differences in total biomass production

between species on day 163 to insignificant levels in the +10, -15 cm and control

treatments by day 375.

Table 3-2. Total biomass (AGB + BGB) after 163 and 375 days.
Total Biomass (g/pot)
Days Treatment n Bahia Floralta p value
+10 3 85.0 10.8 < 57.2 1.47 0.01
0 3 109 13.0 < 63.0 6.73 0.01
163 -10 3 113 2.78 < 72.0 10.7 < 0.01
-15 3 115 3.59 < 65.2 7.88 < 0.01
C 3 105 14.3 < 77.7 4.30 0.03

+10 3 76.6 3.00 < 84.5 6.51 0.13
0 3 112 18.6 < 76.5 2.92 0.03
375 -10 3 112 16.0 < 70.6 6.62 0.01
-15 3 85.4 11.6 < 76.0 21.1 0.54
C 3 113 17.7 < 99.6 20.8 0.45

It is counter intuitive that cumulative biomass could decrease, but since BGB was

only harvested twice, these data only represent the net BGB after 163 and 375 days, not










the variability within those time periods. Therefore, the quantity of bahiagrass BGB that

died was greater than the forage produced between 163 and 375 days, resulting in

negative net total production.

A. B.
120 55
110- o 50-
100- 45


5- + 25-

60- + 20-
50I 15-I I
150 200 250 300 350 400 150 200 250 300 350 400
Elapsed time Elapsed time

A +10 cm 0cm -10cm + -15cm Control

Figure 3-11. Below ground biomass production. A) Bahiagrass BGB production B)
limpograss BGB production (Table C-9, Appendix C).

Under constant inundation, both species will survive for at least 375 days.

However, cumulative biomass production for bahiagrass actually decreased between days

163 and 375, while limpograss increased. Both species appear to have been in an

acclimation phase between days 163 and 375. The lack of significant differences in total

biomass between species after 375 days in the +10, -15 and control treatments (Table 3-

2) indicate that those treatments were influencing total biomass productivity for both

species. Although not statistically significant, bahiagrass still had more total biomass in

the -15 cm and control treatments after 375 days, while limpograss had more in the +10

treatment.

Root to Shoot Ratios

In general BGB production had an inverse relationship AG forage production for

both species. The average BGB production for all bahiagrass pots, regardless of










treatment, was 85.6 14.0 g pot- after 163 days and 79.6 20.8 g pot- after 375 days.

Limpograss BGB production was 38.0 + 9.07 g pot- after 163 days and 29.5 8.5 g pot-'

after 375 days.

Bahiagrass maintained significantly more BGB than limpograss in all treatments

after 163 and 375 days. Relative portions of total AGB (residual biomass + forage) and

BGB for each treatment and species are graphed in Figure 3-12. In all treatments,

bahiagrass had significantly greater root to shoot ratios than limpograss after 163 days.

After 375 days, all root to shoot ratios for limpograss were less than one while bahiagrass

ratios were greater than one (Table 3-3). Thus, after 375 days limpograss produced more

AGB than BGB while bahiagrass produced more BGB than AGB.


85 -

60 -

35 -


(15)
(40)

(65)
(90)

(115)
Bahia Floalta Bahia Floralta Baha Floralta Baha Floralta Bahia Floralta

+10 +10 0 0 -10 -10 -15 -15 C C

Treatment and Species


Figure 3-12. Above and below ground biomass production after 375 days. Above
ground biomass (top) is the sum of cumulative forage production and residual
biomass. Below ground biomass (bottom) is all biomass harvested below the
soil surface after 375 days.












Table 3-3. Root to shoot ratios.

Day Treatment
+10
0
163 -10
-15
C

+10
0
375 -10
-15
C


Root:Shoot Between Species
Bahia Floralta
3.55 0.56 > 0.94 0.19
5.06 2.27 > 1.17 0.22
4.25 0.85 > 1.58 0.40
5.35 0.80 > 1.42 0.34
4.22 1.19 > 1.49 0.15

2.11 0.21 > 0.31 0.07
5.48 2.59 > 0.81 0.23
5.76 0.43 > 0.90 0.39
4.26 0.76 > 0.95 0.19
3.91 1.21 > 0.36 0.15


Phosphorus Assimilation

Phosphorus tissue concentrations

Phosphorus concentrations varied by species and by treatment. On day 0, the only

significant difference in tissue concentrations between species was in the 0 cm treatment

where limpograss (1790 + 331 mg kg-1) had a significantly greater forage P concentration

than bahiagrass (1530 58.0 mg kg-1). Both species exhibited a decline in P

concentrations by the first harvest (day 27). All limpograss treatments (965 220 to

1260 152 mg kg-1) and the 0, -10 cm and control (1180 60.0 to 1220 37.0 mg/kg)

bahiagrass treatments had significantly lower forage P concentrations by the first

sampling on day 27 (Figures 3-13 and 3-14). The bahiagrass +10 treatment had

significantly greater forage P concentrations than limpograss on days 27, 55 and 375. In

addition, P concentrations of the bahiagrass forage control treatment were greater than

limpograss on day 375, although on day 163 the limpograss forage control treatment was

greater than the bahiagrass.


p value
< 0.01
0.04
0.01
< 0.01
0.02

< 0.01
0.04
< 0.01
< 0.01
0.01










On day 0, there were no significant differences in forage P concentration between

bahiagrass treatments. However, on days 27, 55, 83, and 163, the wettest bahiagrass

treatment (+10 cm) had significantly greater P concentrations than all other treatments

(Figure 3-13). By the final harvest (day 375) the bahiagrass +10 treatment had greater

forage P concentrations than the 0 and -10 treatments. Limpograss, on the other hand did

not have many differences between treatments. The only difference was on day 163

when the +10 treatment was greater than the -15 cm (Figure 3-14).


A + 10 cm

0 cm
4 -10 cm

+ 15 cm
* Control

0 50 100 150 200
Elapsed time


250 300 350


Figure 3-13. Mean P concentrations (mg/kg) for bahiagrass forage by harvest day and by
treatments (Table C-17, Appendix C).


1800-


1600-


1400-


,1200-
-
0)
E1000-
I-

800-


600-


400-










2200-
A + 10 cm
2000 -
0 cm
1800- 0- 10 cm

1600 + -15cm

1400 -,a Control
S1400-o
E
1200 -
I-







0 50 100 150 200 250 300 350 400
Elapsed time

Figure 3-14. Mean P concentrations (mg/kg) for limpograss forage by harvest day and by
treatments (Table C-18, Appendix C).

Below ground biomass cneniocentrations (303-668 mg kg) were relatively low

compared to forage. On day 163, limpograss had greater BGB P concentrations than

bahiagrass in the 0 cm and control treatments. However, on day 375, there were no

significant differences. Nor were there any significant differences for either species when

treatments were compared.

Phosphorus storage

Phosphorus assimilation (mg pot-) is a function of concentration and biomass

production. After 375 days, the only significant difference in total P storage (AGB

+BGB) between species was in the 0 cm treatment, where bahiagrass was greater than

limpograss (Figure 3-15). This was not consistent with total P storage at the end of the

first growing season (day 163) where bahiagrass was greater than limpograss in all

treatments except the control (Table 3-4). This change over time in total P storage can










be attributed to negative net BGB production in bahiagrass and greater forage production

in limpograss between days 163 and 375.

90
80
70
60
p 50
S40
30
S20
10
0-
Bahia Foralta Baia Floralta Bahia Foralta Bahia Floralta Bahia Flralta

+10 +10 0 0 -10 -10 -15 -15 C C
Treatmnt and Species

Figure 3-15. Total P storage (AGB + BGB) after 375 days.

Table 3-4. Total P storage (AGB + BGB) species comparison.
Total P Storage (mg/pot)
Day Treatment Bahia Floralta p value
+10 50.9 4.06 > 40.5 1.32 0.01
0 53.7 5.56 > 41.1 4.69 0.04
163 -10 56.9 4.48 > 39.6 4.49 0.01
-15 56.0 5.73 > 38 5.43 0.02
C 54.1 10.9 < 57.5 2.38 0.62

+10 47.8 4.84 < 50.2 2.67 0.49
0 50.5 13.1 > 37.3 2.05 0.16
375 -10 46.8 5.22 > 34.2 5.43 0.04
-15 36.0 5.13 > 36.1 5.69 0.99
C 62.8 16.4 > 55.3 9.09 0.53



Total P storage has the same general trend as biomass production. The same is true

for BGB production and BGB P storage. In addition, there is a positive relationship

between P partitioning and biomass allocation. Bahiagrass had greater P storage in BGB

than in forage, while limpograss had the greater P storage in forage.

Overall, for each species P storage in BGB was not significantly different between

treatments after 375 days. However, there was a significantly different treatment effect










on day 163. After 163 days the limpograss control stored significantly more P in BGB

than all other limpograss treatments.

Limpograss had greater P harvested in forage than bahiagrass in the all but the +10

treatment by day 27. By day 55, P harvested in limpograss forage was greater in all

treatments except the -15 cm and control treatments, while by day 83 limpograss was

only greater in the control treatment. By days 163 and 305, limpograss forage took up

more P than bahiagrass in all but the -15 cm treatment, while on day 375, limpograss had

greater P storage in all treatments except the -15 and -10 cm treatments.

On a cumulative basis, there was a strong relationship between forage P storage

(Figure 3-16) and cumulative forage production (Figure 3-7B, 3-12B). Limpograss had

greater forage P storage than bahiagrass in all treatments except the +10 cm treatment on

days 27, 55, 83 and 163. However, after day 163, limpograss had greater P storage in all

treatments.

A. B.
45
20- 40-
S 235-
015-o 30
E 25-
10o-I 2-



0 50 100 150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
Elapsed time Elapsed time

A+ 10 cm Ocm -10cm + -15cm Control


Figure 3-16. Cumulative P harvested in forage. A) Bahiagrass P harvested B)
Limpograss P harvested (Table C-23, Appendix C)

There were no differences in harvested P between bahiagrass treatments on days

27, 55 and 305. However, on days 83 and 163, the +10 cm treatment had significantly









more P harvested than all other treatments. The +10 cm and control treatments had

greater P harvested than the 0 and -10 cm treatments on day 375. On a cumulative basis,

on all days after 55, there was significantly more P harvested in the +10 treatment than in

the 0, -10 and -15 cm treatments.

Like bahiagrass, there were no differences in harvested P between limpograss

treatments on day 27. However, by day 55, the driest treatments had assimilated the most

P. The control had significantly more P harvested than all other treatments, and the -15

cm treatment had more than the -10 cm treatment. By day 83, the influence ofbiomass

production on P storage began to emerge as P harvested in the control was greater than

the 0, -10 and -15 cm treatments. In the latter and earlier parts of the growing season

(days 163 and 305), there was more P harvested in the +10 cm treatment than all other

bahiagrass treatments, while on day 375, there was more P harvested in the +10 and

control than in the all other treatments.

On a cumulative basis, by day 83, the bahiagrass control had assimilated

significantly more P than the -10 cm treatment. By day 163, the control stored more than

the 0 and -15 cm (in addition to the -10 cm) treatments. By day 305 the +10 treatment

had significantly greater cumulative P storage than the 0, -10 and 15 cm treatments. By

day 375, the +10 cm and control had greater cumulative storage than the 0, -10, and -15

cm treatments.

Phosphorus storage root to shoot ratios

Root to shoot ratios for P storage had a positive relationship to biomass ratios.

Figure 3-17 shows relative portions of AGB and BGB P storage. After 375 days, the +10

treatment was the only bahiagrass treatment with a ratio less than one. Limpograss P

storage ratios were all less than one and positively related to biomass ratios. All










bahiagrass treatments had significantly greater P storage ratios than limpograss

treatments after 375 days (Table 3-5)


50

30 -

,. 10

b (10)

S(30)

(50) -
Bahia Floralta Bahia Fralta Bahia Florala Bahia Floralta Bahia Florata

+10 +10 0 0 -10 -10 -15 -15 C C

Treatment and Species
Figure 3-17. Relative comparison of root and shoot P storage after 375 days.

Table 3-5. Root to shoot P storage ratios with statistics.
Root:Shoot P Storage
Day Treatment Bahia Floralta p value
+10 1.27 0.27 > 0.69 0.22 0.04
0 2.45 0.23 > 1.01 0.13 < 0.01
163 -10 2.46 0.67 > 1.10 0.28 0.03
-15 2.95 0.40 > 0.95 0.05 < 0.01
C 1.95 0.10 > 1.17 0.05 < 0.01

+10 0.70 0.12 > 0.15 0.04 < 0.01
0 2.58 1.35 > 0.39 0.09 0.05
375 -10 2.29 0.14 > 0.48 0.17 < 0.01
-15 1.42 0.33 > 0.55 0.32 0.03
C 1.35 0.34 > 0.22 0.08 < 0.01


Discussion

Forage Production

Bahiagrass and limpograss are physiologically different. Bahiagrass typically has

less forage production than limpograss because it allocates -50% of its energy to root and

stolon production (Chambliss & Adjei, 2006). This was evident from the bahiagrass root









to shoot ratios, where BGB ranged from 2.11 to 5.76 times higher than AGB (Table 3-3).

In addition bahiagrass is a long day plant that is strongly influenced by photoperiod

(Marousky & Blondon, 1995). Thus its annual production is lower because it has a

shorter growing season. On the other hand, limpograss tends to allocate more energy to

AG forage production, as root to shoot biomass ratios were less than one in all

treatments.

Limpograss is known to support relatively high cattle stocking rates and for

having superior late fall and early spring production compared to bahiagrass

(Sollenberger et al., 2006). Results from this study support that statement. Overall

limpograss had greater forage production than bahiagrass in the latter and earlier parts of

the growing season in all treatments. In the first 83 days, both species exhibited a decline

in production between harvests, regardless of treatment (Figure 3-5). This was likely the

result of the combined effects of harvest stress, temperature and light effects in the latter

part of the peak growing season. Harvest stress was evident after the first harvest (day

27) as production was significantly less for both species by the second harvest. After 83

days, limpograss rebounded, while bahiagrass production continued to decrease. The

bahiagrass decline after day 83 is likely the result of decreased photoperiod during

shorter days and not harvest stress. Even during the peak of the growing season,

limpograss had greater forage production than bahiagrass. Forage production results

support H1 limpograss has greater cumulative forage production than bahiagrass in all

treatments.

Flood Tolerance

Water levels did not appear to have an effect on forage production for either species

until after the first harvest. There were no statistical differences between bahiagrass









treatments until the third harvest. This suggests that bahiagrass forage production may

not be affected by water levels as deep as 10 cm above the soil surface for up to 55 days.

The wettest bahiagrass treatment actually produced more forage than the other treatments

toward the end of the in first growing season. The same was true for the limpograss +10

treatment. In addition the limpograss +10 had the greatest forage production in the

earlier part of the following growing season. The wettest treatment seems to start forage

production earlier and extend it later in the growing season for both species, but more for

limpograss. The reasoning for this may be related to a temperature buffering effect

caused by standing water. Thus, flooding may create an artificial environment that

decreases diurnal temperature fluctuations, thus prolonging the growing season. After

375 days, forage production per harvest was similar in the wettest and driest treatments

for both species, while the intermediate treatments generally produced less forage.

Bahiagrass is resilient to many environmental conditions. However, under longer

hydroperiods bahiagrass is not as resilient and may be out competed by facultative or

obligate wetland species. Efforts to restore native wetland species in bahiagrass

dominated pastures have been challenging. While mechanically removing the sod and

applying herbicide has been the most effective way to remove bahiagrass (Violi, 2000),

prolonged flooding will also eliminate bahiagrass, and enable wetland species to establish

(David, 1999). David (1999) examined the distribution and density of bahiagrass and

other wetlands species in hydrologically restored wetlands and found that bahiagrass

persisted for 2 years after inundation, before dying off by the fourth year. In addition,

wetland species such as Panicum hemitomon and Pontederia cordata increased in

frequency of occurrence under longer hydroperiods. Clearly both species will survive









375 days in 10 cm of water in a non-competitive environment. However, even in a

competitive environment it may take up to three years for different vegetation to establish

between normal and high water after a change in hydrologic regime (Van der Valk, A G

et al., 1994).

Between days 83 and 163, both species appeared to acclimate to treatment

conditions. Initially, the controls of both species had the greatest forage production

however by day 163 there was no difference between the wettest and the driest

treatments. This suggests that in the short-term, hydrology alone in a non-competitive

environment may not cause a shift in species. Over the duration of this investigation,

bahiagrass did not have significantly greater total biomass production in drier treatments,

nor did limpograss have significantly greater total biomass in the wetter treatments.

Based on treatment effect comparisons, H2 can not be completely accepted.

Phosphorus Uptake

After 375 days, total P storage was similar for both species in all treatments

expect the -10 cm where bahiagrass stored more than limpograss. Therefore, H3 is

rejected in favor of the null hypothesis that limpograss does not store more P than

bahiagrass. Despite, similar assimilative capacities, P stored in bahiagrass is relatively

more stable than P stored in limpograss because the majority of P stored in bahiagrass is

in BGB. Above ground forage is typically more labile and is subject to grazing. As

discussed in Chapter II, vegetation is a short-term P storage mechanism. In addition, if

the vegetation is continually grazed, nutrients in digested plant tissue are more

bioavailable than senesced vegetation in wetlands. Therefore, P can be mobilized from

the soil into the water column by way of grazing.









Phosphorus partitioning in vegetation had a similar trend as biomass allocation. All

bahiagrass treatments except the +10 stored more P in BGB than in AGB. The higher P

concentrations in the bahiagrass +10 treatment caused higher P storage in AGB than in

BGB. This was consistent with all limpograss treatments more P was assimilated in

AGB than BGB. Therefore, H4 is only partially accepted. All bahiagrass ratios except

the +10 cm treatment were greater than one and all limpograss P ratios were less than

one.

Inundation actually increased P storage in both species by increasing tissue P

concentrations in the +10 treatment. The relatively high P tissue concentrations in the

+10 treatment are consistent with results found in wetland center zones as described in

Chapter II results and Whigham (2002).

Reddy and Debusk (1985) observed lower tissue concentrations in summer months

and suggested that higher productivity in the summer likely diluted concentrations. In

addition they suggested that slow growth and luxury uptake likely caused increased

concentrations in the winter (Reddy & Debusk, 1985). The same line of reasoning may

also explain the elevated P concentrations in the +10 treatments. Biomass production

decreased in the bahiagrass +10 treatment, however P concentrations were significantly

greater than the other treatments. Where nutrients are readily available, tissue

concentrations may have a direct relationship to biomass productivity. Future research

may look into the possibility of specific plant species' rates of nutrient uptake over time

vs. resultant tissue concentrations.

Conclusions

Limpograss appears to thrive in the wettest and driest mesocosm treatments. In

fact the limpograss +10 and control treatments had the greatest production of all









treatments. Although bahiagrass survived under inundated conditions for 375 days, more

than likely it would not survive competition from other plant species, trampling, grazing

and water stress in situ.

Both species had similar total P storage in all treatments except the -10 cm

treatment. The majority of P stored in bahiagrass is in BGB, while most P assimilated in

limpograss is stored in forage. Thus utilizing limpograss for P removal from wetlands

may be best optimized by harvesting and exporting hay and assimilated P away from the

wetlands.

Overall, after 375 days limpograss had greater forage production than bahiagrass in

all treatments, a greater hydrologic tolerance and similar P storage potential. Therefore,

in order to maintain pasture carrying capacity and vegetative P storage during BMP

implementation, limpograss may be a more suitable forage in restored pastures wetlands

even under higher water levels and extended hydroperiods.














CHAPTER 4
SUMMARY AND CONCLUSIONS

Summary

The overall goal of this research was to evaluate the biomass production and P

storage potential of vegetation in historically isolated pasture wetlands and determine the

efficacy of using a wet tolerant forage species to minimize the loss of improved pasture

area as a result of hydrologic restoration.

Objective I: Biomass Production and Phosphorus Storage in Wetlands

I. Assess biomass production and P assimilation by wetland vegetation and
forage grasses under various hydroperiods.

Results from Chapter II and McKee (2005) indicate that wetland soils in the

Okeechobee basin store more P per unit area than surrounding upland soils and

vegetative components. The direct role of vegetation in active total P storage is relatively

small, short-term, and highly variable compared to the physical storage capacity of soil.

However, the presence of vegetation is an important component of ecosystem P storage

because it increases the total P retention capacity of wetland soils.

Phosphorus storage in vegetation along hydrologic gradients was variable

depending on the type of species present, land-use intensity, grazing pressure and

hydrology. There were, however, similar trends in AGB P storage. In general, wetland

zones at both sites stored more P in AGB than in upland zones. In addition, total P

storage (AGB + BGB) had a positive relationship to hydroperiod at Beaty while the









opposite trend existed at Larson. This was likely related to higher AGB tissue P

concentrations in wetland centers than uplands and differences in land-use intensity.

Vegetation Stress

Hydrology is often the primary determinant of vegetation composition within

wetlands. However, in pasture wetlands, the stress of grazing likely influences

vegetation community establishment and persistence. Although grazing was not

measured in this experiment, vegetation patterns, biodiversity of species, and large

hydroperiod ranges for the same species at different sites suggests that hydrology is not

solely responsible for species distribution within pasture wetlands. In addition, grazing

may have a significant effect on wetland P storage capacity.

Objective II: Facilitating Land-use and Wetland Restoration

II. Determine the efficacy of establishing a wet tolerant forage grass in wetland
transition zones before hydrologic restoration to minimize loss of productive
pasture

Overall limpograss had greater cumulative forage production than bahiagrass in all

hydrologic treatments. This was primarily due to its ability to produce forage earlier and

later in the growing season. More importantly, limpograss production was similar in the

wettest and driest treatments, producing significantly more forage than the intermediate

water level treatments. The wettest and driest bahiagrass treatments also had the greatest

production relative to the other treatments.

Both species will survive for 375 days under non-competitive, inundated soil

conditions as long as the biomass is not completely submerged. However, it is unlikely

that bahiagrass would be competitive under wet conditions with its low productivity.









Unexpected Results

One unexpected result that was consistent between both species in the mesocosm

experiment was that the greatest biomass production and subsequent P storage occurred

in the wettest and driest treatments. It was hypothesized that biomass would have a

negative parabolic shape when treatments were aligned on the X-axis in order from

wettest to driest. Essentially, it was thought that both species would have similar curves,

expect limpograss's curve would be shifted more toward the wet treatments and

bahiagrass's more toward the dry treatments. However, as described in Chapter III, the

results were opposite. Each species had a positive parabolic shape.

Another unexpected result found in both the field and mesocosm studies was the

increased tissue P concentrations under flooded conditions. Above ground biomass tissue

P concentrations in wetland center zones and in the +10 inundated mesocosm treatment

were greater than uplands and drier mesocosm treatments. Higher concentrations of

plants in wet conditions were likely caused by slow growth and luxury uptake.

Implications for Restoration

Bottcher et al., (1995) defines BMPs as on-farm activities to reduce nutrient

exports to water bodies and tributaries to environmentally acceptable levels, while

simultaneously maintaining an economically viable farming operation. In addition,

BMPs that adversely affect the economic viability of a farming operation should be

subsidized to maintain profitability (Bottcher et al., 1995).

Many BMPs are considered voluntary; however in a watershed where a TMDL has

been mandated, water quality compliance is required. The term "voluntary" refers to the

choice of options ranchers have to comply with TMDL goals. In the Okeechobee basin,

ranchers have the option to monitor the nutrient discharge from their property to ensure