<%BANNER%>

Aquatic Vertebrate Usage of Littoral Habitat Prior to Extreme Habitat Modification in Lake Tohopekaliga, Florida

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 E20110115_AAAACT INGEST_TIME 2011-01-15T17:52:30Z PACKAGE UFE0008580_00001
AGREEMENT_INFO ACCOUNT UF PROJECT UFDC
FILES
FILE SIZE 218713 DFID F20110115_AABYDU ORIGIN DEPOSITOR PATH muench_a_Page_023.jpg GLOBAL false PRESERVATION BIT MESSAGE_DIGEST ALGORITHM MD5
d263bc4682a4fa791dd2f445f9228328
SHA-1
65d409254e11631ef72737776e633557dcd9a8e5
1051954 F20110115_AABYEI muench_a_Page_038.jp2
42a96ff079e3646889392a12da5b0a84
c877a8def24e5b75fd0928e12c51b58beffd33bf
25389 F20110115_AABYDV muench_a_Page_103thm.jpg
a84890f05a7d2d378d9c4e27558ff33d
4d75840cda592c8a2ffce747f63100b6d57c7460
25271604 F20110115_AABYEJ muench_a_Page_065.tif
aaca2b3ecbe326544ec7445c26f55fdb
71ee44fd86418736eec6f71aa85055af745b1424
2094 F20110115_AABYEK muench_a_Page_032.txt
b7a045757d4ad9ae3341120f591a7997
7a99d8daf5fa7f3d2f5a6045cfbeaf46a744eec4
8423998 F20110115_AABYDW muench_a_Page_041.tif
4f72378363834b01f0c8d9c6c87ad46e
692eaf5b6039546546c89015bff003a9b009d097
F20110115_AABYEL muench_a_Page_008.tif
0309fec96933687be104fe0a0d5a53e4
1e538c9fb6a4741e320f7889d509cb7404d9932f
21203 F20110115_AABYDX muench_a_Page_001.QC.jpg
d05af78387d9bb319af866657447859f
7cdd8eab5c6b1ee4cf43dcee465fca00ee1cb450
1053954 F20110115_AABYFA muench_a_Page_101.tif
43debbb81bd862e033a0230fb7dbb9a7
fd9e9e3c02a4690c93d065ade22e12c0792604af
24366 F20110115_AABYEM muench_a_Page_102thm.jpg
4e447c205c43f4e34f80be9ce2197d75
18e2ec33def12cd12cc97f024e0c92fa19d61be4
2224 F20110115_AABYDY muench_a_Page_108.txt
ffc1fbaa3bc2f576c7d8714dcebe2192
d7cd3d5a0e0663d418030a7cf17e037df0727006
106706 F20110115_AABYFB muench_a_Page_019.jp2
d01cca0a085002b2ac44e9a4fa20c8d9
8f08c1bd5a3da30337ef8f3d9b211fa3639deb9b
11084 F20110115_AABYEN muench_a_Page_094.pro
1d3692c9138800bcfb75ab7522f1db7e
6bfe9d782ee97525aef29268809c105db40ab201
49205 F20110115_AABYDZ muench_a_Page_102.pro
7fc54e101c41e185270c4c9fd76ea774
80b2387395a44a221266cd85e87d48fd07c18bcf
F20110115_AABYFC muench_a_Page_096.tif
83cd1f4ae2dd48c7a5cf593e108036e0
1bed5764daada9403f61df000ca0c611b5099426
F20110115_AABYEO muench_a_Page_047.tif
0f7389fcf1a0ac84cb854df1f17005f3
90af87f3a01491c6029360df09ef44b6f5e8c1f9
1842 F20110115_AABYFD muench_a_Page_045.txt
406773e33cc057135d3adb4e830e8a38
fadf823fe2b7e2ebbbaee32c88e587a40f3956f5
110033 F20110115_AABYEP muench_a_Page_104.jp2
208c297c7487666193bccfc3f81235ee
319bf660f204f0a04a58d785a654eb90e8157357
F20110115_AABYFE muench_a_Page_080.tif
a4d5ef9e9e12427816f6c8ecf7dc555b
1f6f4edfe7e28a5372be24909646afe0ae1acd96
74696 F20110115_AABYEQ muench_a_Page_025.QC.jpg
ebd319e492c7a7a0b2656f9f96b49af3
f54285680e9e9ec6a6c87397173f1127ec71d746
78674 F20110115_AABYFF muench_a_Page_087.QC.jpg
647a22665d7147fa3ff98fbdebf18bf2
a49c241de4c88fb6ea8ed4a3de89eacfd948dee9
85072 F20110115_AABYER muench_a_Page_114.QC.jpg
ca4eb82dbe3a8ef63f53753635d57194
b14434e27058ebb18e0b3281191feaf0861f6a00
33642 F20110115_AABYFG muench_a_Page_027.jp2
4de6a33c8ecb745a126b80e3b2e363b1
b41eb26d41b03d9723e7ca29ca8e564efc842e20
327808 F20110115_AABYES muench_a_Page_061.jp2
13c09b73c36065af899a115a6a234aa5
4effa2acb1b88875c6381936ce543a4a3149d80f
44230 F20110115_AABYFH muench_a_Page_011.pro
68692dbc26a6f33fe85ddf44ced89eb0
80132058ff58cc3c55421736faaa33c7fa825c1c
F20110115_AABYET muench_a_Page_112.tif
c0a8f041c6e3944a63b4dd6a0203d27f
dee9b6c24e6866f8b7ddaf55d1cb1566325eb141
93156 F20110115_AABYFI muench_a_Page_004.jp2
72fc9fccb41d361494646ba9d464221b
2c202336ca3562e9b7b75ce11cac267efdeb83de
211339 F20110115_AABYEU muench_a_Page_015.jpg
c1b448991ea4efc85c9526c9a6d79046
1fd10c9917ae45485cd6546565dfa803664479c1
81141 F20110115_AABYFJ muench_a_Page_031.QC.jpg
ed1e8c7d1967600d0395fd51dc39b68b
4892d04feb746d04f159e59b2f51e414614ec107
21085 F20110115_AABYEV muench_a_Page_012thm.jpg
8eda7d5474b1099071ea6a5d42f22b32
12602cb78144e77c49be364aede75585cefbdebd
114 F20110115_AABYFK muench_a_Page_002.txt
59b0c1c298dc9690d68cdb80a8822e68
4f089a64988c8bd3a97536b6e43ade44d90a5a73
9559 F20110115_AABYEW muench_a_Page_003.QC.jpg
09ef0982a69529343f271662d2ab0087
992dcad3c9b509a6f6d1ec5c8e107ed5436ed3ff
215645 F20110115_AABYFL muench_a_Page_030.jpg
00d636467cdacda0fc8af23f29125890
49c969d569ea5ac11c40e863cf8a6860fa0d44a1
105841 F20110115_AABYGA muench_a_Page_033.QC.jpg
db77a2d288b3068a369ec980a7d12a08
161d295c74e7907d1d7c35ccfd9afec50e7fb8f4
35401 F20110115_AABYFM muench_a_Page_068thm.jpg
518a33d564a5c084348edb798c18b3e5
27f97016219346d1df7ae94018330ff5f369fea2
295740 F20110115_AABYEX muench_a_Page_007.jpg
e9fc541ad6139b17459e6b15e4a0f5ed
14a66a4a6507cb8e45e71442181e9fda3a7c3b1e
1946 F20110115_AABYGB muench_a_Page_055.txt
1324835fd079ff5814c14426a1702b01
3dfca75fe21dbe04653df47fc65fe29ca4ea57d6
794 F20110115_AABYFN muench_a_Page_067.txt
8e65b802c8e56810803907316412f364
174f569b0c1590ed6760a5724e70b74502f12188
F20110115_AABYEY muench_a_Page_048.tif
2b23bd9bf42ec289a81234c5aa23f658
069210e233b80ddf6673200981080ffb43cf31ca
F20110115_AABYGC muench_a_Page_043.tif
138223227f13a9c9814ebf86601b9bb8
ae88ef3bf3121d81ff52e84317b71ab2f33e61cb
54650 F20110115_AABYFO muench_a_Page_108.pro
3b8c86be6981fa8c167702055a5f26d2
6b588b1a8a6bd1bea46e9acc21e9c3aa05533432
113199 F20110115_AABYEZ muench_a_Page_026.jp2
83c24ef6b8121751570b36f3dbd7a5f0
bd68e606691b9a957f219531f25e02b86fb9942c
23958 F20110115_AABYGD muench_a_Page_055thm.jpg
75ddd6969dfe80a36f2422f14c006677
36b8bbaa0fb88bd88d4e134008464a51420d78f5
477182 F20110115_AABYFP muench_a_Page_092.jp2
4d0f3d585b038205c55de042f9e74ebe
65b5776e3d63a4d2643f98de5152a77942ee5432
879 F20110115_AABYGE muench_a_Page_060.txt
f1643ff23388b50d229777b6f8ba8869
163ccde3e2076c40191061b3aeaf92617e9709d6
36715 F20110115_AABYFQ muench_a_Page_063thm.jpg
c215d3b3754834c9f4f728ef27e944f0
176bf970118a19086fbdaa7f8bc3e5d0760c34ef
F20110115_AABYGF muench_a_Page_074.tif
faf58aa5aeaf4ddcecf684c94cac721e
73fdd5208db853c2e922a3b3e7b6e319a31b35c5
45701 F20110115_AABYFR muench_a_Page_073.QC.jpg
02b9441c7c2ab700c5f2bf4536acd2e1
48dd28913f6ca91c004b6273d259abd01b4567f7
25234 F20110115_AABYGG muench_a_Page_056thm.jpg
9e9708ec5015da2d91b1c12035eb76f4
90130a14e482a7b42c8dbf0bf3929d36b1eb25d8
108042 F20110115_AABYFS muench_a_Page_089.jp2
1618bc4e14f4eb14814893636e48f13f
76b4049cbf41bb03811ddd027b1fae05bea0dfc6
51559 F20110115_AABYGH muench_a_Page_023.pro
2038d48c3c1d3e9b8469d2c65e3a4f36
413326e22cccced89d779d730fedf673ee6bd0d4
F20110115_AABYFT muench_a_Page_072.tif
73d4ee04f87b53c60fbb2303fd76ea9e
578e52f0096146f9bf44845bfc7812f0e4fae931
20238 F20110115_AABYGI muench_a_Page_090thm.jpg
f4832d1239eefd6a6e5448d896aa6a7e
f7e7e5643eaab12f5eeb212d0fb7419675948b88
2018 F20110115_AABYFU muench_a_Page_057.txt
5690b755932bb2357fc520dc912778e7
99621f985e7d69cb9b57c5839d0d3ecca4561656
1976 F20110115_AABYGJ muench_a_Page_035.txt
d267c5917b958dce4e72015f957ccb2d
a3abc0359b85ea99024dc33f3296851c61015adc
50333 F20110115_AABYFV muench_a_Page_081thm.jpg
3b52e85957e194f989c049f8db1d8351
476dc56e12803b4dcdb654285c38a4d2439701cb
2955 F20110115_AABYGK muench_a_Page_041.txt
9662fe2b035c04a25db7b29e0f4c5166
53f0876402dbb5d4f2f294c5f81c3b6460f23fe1
113539 F20110115_AABYFW muench_a_Page_040.jp2
912dc1fd2fbcb085ba9d4d3bd5e7a8f2
3f16fb95ed7468f962f2893ce77172834a246a1b
1677 F20110115_AABYGL muench_a_Page_075.txt
64157ed14b7a2fe68d2442a005815f22
2262e8ca74a9567dae91efb3ea776f786e7c019b
52692 F20110115_AABYFX muench_a_Page_031.pro
b9b5adb2e0980c5297ba452fd7dd34b5
98878bb161cd5cd99eb6f1f49584f04439b89567
51463 F20110115_AABYGM muench_a_Page_057.pro
7220cf2820a1e422ae42d799aa9491c7
6cc4b160b11a6e823aaba39cfc2872fac741a612
2017 F20110115_AABYHA muench_a_Page_040.txt
51f2c062aa00e5cf4a32ddac26c69c4a
ba731cd59b79abe137ce551e0727c568be4aa408
105790 F20110115_AABYGN muench_a_Page_053.jp2
d9bfa5985e01c5282007b5c637d18821
12f11cc3d0c7b895a8ef0abbd45af271f3d74506
78469 F20110115_AABYFY muench_a_Page_104.QC.jpg
a1b657b320342b4b5c403425a3014bb5
651592de23e59a7aec1b29761e515d69f9bdb358
79365 F20110115_AABYHB muench_a_Page_084.QC.jpg
a820987eef5fdd92580099c196b0402f
8186e3c23e905df14bbc595d667a8cc72330a327
36135 F20110115_AABYGO muench_a_Page_020.pro
1f7d08e8c8bbb69332c00662c6bce7f5
a85e8d3a7f0ddc46b5f9deb22fa3711d6e7a781f
83062 F20110115_AABYFZ muench_a_Page_106.QC.jpg
bb1bdd3c2d6b9ce98430d6ef620c94ca
94ec68b42ef80acf4727bc321d376f132acd7222
208169 F20110115_AABYHC muench_a_Page_084.jpg
a19dcd81cac3d1919b60caa26677dea3
a2fe6e9b3a7f1cc3784f7af222713ebd5aca69d8
555871 F20110115_AABYGP muench_a_Page_074.jp2
060a231a7633c8abd5653055790224f6
b350f566716281bb749d02105a44815d96406e84
49106 F20110115_AABYHD muench_a_Page_016.pro
6d0a550e85acc7cfd37b25c18159c0a3
fbe11510192a064a790cac0d0a5d037245106bb3
F20110115_AABYGQ muench_a_Page_108.tif
e6b35d0269bc7b3167dc9e4365180e80
1c07954f1b9ae20762cb5218a9e445a54cf9e6db
109471 F20110115_AABYHE muench_a_Page_078.jp2
6774f8dbff3f16ed952dcf96fd20ee41
bf42c4cdf23babc5ed62bfce8065e5171e7c4fd9
1927 F20110115_AABYGR muench_a_Page_053.txt
2fda65f2bc84056ded17bf8d7bf1c7a8
1812add88097b96bf959f42a7bfb2455760c6aac
26044 F20110115_AABYHF muench_a_Page_087thm.jpg
096740936cf0e465b0bd821cdfcfb37a
d131e0651d0bc3d7433274ec7c2cca0d0e967ba9
2000 F20110115_AABYGS muench_a_Page_071.txt
8c814ddca9cf8ec098bc14b78a1d0b5b
619e316af5a3b044dd3955930b5f50e59cbf7930
104297 F20110115_AABYHG muench_a_Page_024.jp2
3228f4b560b46b7855b5af83d07b7b3b
90430f7eace0bde1bb2e047f65f7f5f49c47cb16
2671 F20110115_AABYGT muench_a_Page_109.txt
d856088c4e25b3ada7f811cfbaedc9d1
85bf50d831b6f2690666ffa5883c2ec5ee1f15fc
99768 F20110115_AABYHH muench_a_Page_059.QC.jpg
0b1a6904613565e4a1b6045ef3d0c38f
cbd2fbea7280ea04dbec60af7665a2d7820bbcab
26498 F20110115_AABYGU muench_a_Page_110thm.jpg
33b0fbd79167148b1bda8da2bdbd5e48
a36aab66afac187927298ab73a9ce756afee2c25
48776 F20110115_AABYHI muench_a_Page_053.pro
85b983bb2bf924029b5c82901dd5e0d5
0330aa9830dec828e8de152575ed4a4346ea8d04
10547 F20110115_AABYGV muench_a_Page_061.pro
ee8e1d051c1795decca5bd36411699d5
fff34116b78239b33f3f48dfe2aac1e4729943d6
F20110115_AABYHJ muench_a_Page_076.tif
398dcbdad45e6d569e4d03b54bd7741c
c5163fdab7153f189cd6f5e9018bc2badda3460d
1805 F20110115_AABYGW muench_a_Page_054.txt
087669aba45a9eedd2c13ef33e5cccea
c6937246e75b96ad92347902dc5e140c9205dfca
19143 F20110115_AABYHK muench_a_Page_010thm.jpg
e5a7c45d992601f2af7014f902698734
56bd946dec1db9edeab876419f897190e5aae32d
46958 F20110115_AABYGX muench_a_Page_025.pro
2f2fec9f06ca38c15717eb83b8f97bb4
4aac759d871443937a4aa6a2d837d8fe5c45888a
1665 F20110115_AABYHL muench_a_Page_010.txt
d758bc429d3ebecb02ca7c4453d40db1
1b78be3b0fe5cbcf496a1b3f424155b60094eb61
15153 F20110115_AABYGY muench_a_Page_063.pro
ed4cda9f8cad2955f2aa6a64bd9851fa
53dfdffd28ba04d74e5dabeb4342b73e129c608d
F20110115_AABYIA muench_a_Page_089.tif
d3726f474755650bcb96b537743b39cd
0bd0ce83a916ec89da118a12ea4dc3825b07436f
35907 F20110115_AABYHM muench_a_Page_098.QC.jpg
d206e1d4f3bb930201ac24134973992e
19641eb6ae7f4c0fd7bb1a047e4aa6f8fc92b643
65309 F20110115_AABYIB muench_a_Page_082.pro
1d0eba1e0f4cbc8b9ebadf912b9f6a1b
205692a64f455ea99f95098e8493d55ead7733b4
202111 F20110115_AABYHN muench_a_Page_019.jpg
b8d503fc20ff96a469ccc1d2a603fca4
191a5b6688701f9d6b2a8d3eb54a47bbca1492d1
105531 F20110115_AABYGZ muench_a_Page_013.QC.jpg
45e409ce3744d2d4e47a30d4fceb8047
eee1bd8d502b9bac10b4afc9704abd5d67c6feb2
24186 F20110115_AABYIC muench_a_Page_084thm.jpg
5fc599bf36cc65e9b43179486d9785c3
d149e27ee3911475da6cfca8dba79ba3708996d7
124922 F20110115_AABYHO muench_a_Page_092.jpg
f142d9a8992b727d30e3d8a22677cdd1
174cdaf28266211367651c6a1d97e4195091f674
62557 F20110115_AABYID muench_a_Page_042.QC.jpg
df48138231ea46ab7804d0f4a8be62d1
5914b1a445b701086fe4b8ccca23c3638cd6cbb7
45459 F20110115_AABYHP muench_a_Page_039.pro
765984e3cd9528ab6bd03c713c0e8fd3
6b886ebdffd144c011766512b5bfe565e6277e1e
635 F20110115_AABYIE muench_a_Page_061.txt
92cc02f65efa1511c1cc1596eef33bce
8594d419e21e519afede178cdabbc99f799383a5
25405 F20110115_AABYHQ muench_a_Page_029thm.jpg
d6970846e82f59aa05a575ce794b7c58
7f078d90558ff94348036223d4bf773f875ac167
8142 F20110115_AABYIF muench_a_Page_115thm.jpg
da5193ff0861a24bbfb1dc20ca550f5e
f6cc88d1fc9063a01673b33e19ed7699519e5390
19719 F20110115_AABYHR muench_a_Page_097thm.jpg
1cdf0c667a433c6df8763166116540ea
ae8af04cb8372c84288a9a04c4e247d078af4cdd
49163 F20110115_AABYIG muench_a_Page_084.pro
abe695b34ba163842f912a590e4a4eff
d6d2d030a41c1e8c5c06962ea9f39f5232df9db7
80677 F20110115_AABYHS muench_a_Page_029.QC.jpg
fd2eb8af179096e1c5836c20bf203be8
06b14aebdd198a9acd991fa81cfff83d03570ec4
54326 F20110115_AABYIH muench_a_Page_020.QC.jpg
eb4dd0d9b0594238b98aac3a86d2cf0d
b464a1e809d2a11d3f60611b3e207779588ecf9f
274245 F20110115_AABYHT muench_a_Page_059.jpg
1b3d68b4570fcdde866200e6c347ec2c
aa787d4f50526b6f5488a74a8177ee3ee30f31b5
41789 F20110115_AABYII muench_a_Page_042.pro
297f7d2d2841ea9d5a060b36261e90a3
e3dcedb840c74d5a9a22347d044b62ca31bc31de
39354 F20110115_AABYHU muench_a_Page_058thm.jpg
e9aec7ab27748030f3e6849fe49413ec
eb10dbdeebf0bbc8bf405271f17e3762f74300aa
130657 F20110115_AABYIJ muench_a_Page_114.jp2
501affdd8a4fff59d138967ba1f1a3c7
6e0f3542141bdbfc9f21ce33fc66cb22e5cf122c
F20110115_AABYHV muench_a_Page_071.tif
a8ed4e2e90a1c806d94dd8db37223ac5
d6adfadc3c70bd1b530056b2bd3c0279a5df573f
F20110115_AABYIK muench_a_Page_058.tif
47692aac07f27b2672a98842454cc831
0b0cef8ee9b51da596844b206ee3b6047ea3075e
49101 F20110115_AABYHW muench_a_Page_055.pro
2381ed8ac23aa6e815f2b717b68e4a9f
506a2b324db2fa484c27e0e9286d8533ad637850
100103 F20110115_AABYIL muench_a_Page_011.jp2
5e85099b91e72c5a132a60747d1e0a92
a3ceb7db73841d6a06f6fd6d7205acb23b9045f2
23147 F20110115_AABYHX muench_a_Page_039thm.jpg
4dfee5aaf1338a81da5472edc0ba4580
4ac6e54f7feda0e19ef54630b0b886d268a4e184
168953 F20110115_AABYJA muench_a_Page_107.jpg
dfd5fb62955a21fea625e034da33e987
4a94da9f708cefb0a45bb7bea30d03cd44c4e444
F20110115_AABYIM muench_a_Page_066.tif
90230f788e940e7259f5dbd0bd74c2b8
c022671f01749f7fa183e3f87306a7b7292c4788
51932 F20110115_AABYHY muench_a_Page_079.pro
a20a56f82e91f28c84db569aaeaa3535
9777df3ad2ab5ea23b6e2c53f9307addc9b78b53
118727 F20110115_AABYIN muench_a_Page_116.jpg
be9fb244d8856ef575d78a4ecf1e2ffd
dc0198a7d0c0946e5dd86c733c42bc5ba9bd8e9a
76609 F20110115_AABYHZ muench_a_Page_072.jpg
083e89cbd50f2ec9ed31658a1fc92b14
8945f0dcfed122d020d5f85062469af0e4191e28
1314 F20110115_AABYJB muench_a_Page_093.txt
e51f951383b42f73dab93a2d843f9fff
eba4a9859e3f02d993b11d5fd80ce788dc064554
F20110115_AABYIO muench_a_Page_031.tif
35eea72b67ee253c260ed4d9d999d3e8
1f93e715d1a8423bd2d3a275a9606a9fbe0bc6a1
41667 F20110115_AABYJC muench_a_Page_004.pro
c58002fc9b85472b94e3cdfcac539c12
5af10edee5d83198e003cc53c6a5cd07bf3dbed6
100501 F20110115_AABYIP muench_a_Page_101.jp2
4907657f303449e1a35ba69d9dff23bc
3ba98a3885dc586e42bf92f7efd276397548c275
1051961 F20110115_AABYJD muench_a_Page_079.jp2
e0daec11375618ec3064c199295bf68e
8fee4887f6112326734862cee5d1b791fc84c258
24204 F20110115_AABYIQ muench_a_Page_036thm.jpg
f84a1f65acf6c2cf71e6c69272d63b7e
8cb8d7d0d2fb3a9bdcc526bdc7a05c6b6e475dbd
214932 F20110115_AABYJE muench_a_Page_103.jpg
55fae836a24ab20a309617f097bdfdea
ff732dd0f20534af9e1c139145c483c4ab36271e
F20110115_AABYIR muench_a_Page_077.tif
456b4962b93a6333ddd42634475836b7
c87c0308e4da1542f1fc30d36fc326e15507063a
99088 F20110115_AABYJF muench_a_Page_050.QC.jpg
75706ab3706ecec483e93f24b8665d45
4474fecf8d98a7c944864675528e064c1c9cdf1e
32921 F20110115_AABYIS muench_a_Page_082thm.jpg
001ebba50c763c6cdb49dd768846f637
4d571ed683007609f55a0f3c8033b0785013a5b7
94834 F20110115_AABYJG muench_a_Page_012.jp2
e762c1525a9cf26e30e3bcc3da5d6462
d1f18f2eb6b684d474773072c3555d7dad4c29a6
F20110115_AABYIT muench_a_Page_010.tif
7ffd89ebe80201a52f5ad5b89b79deff
e4b5246f47a9a975765e1ea53fc9e9c88031f589
1051984 F20110115_AABYJH muench_a_Page_008.jp2
e8bc1d93929055bcaf1f0faa84145f5e
04a6b3dd056c23f399bb976e6ff8ec93af6caaad
23553 F20110115_AABYIU muench_a_Page_024thm.jpg
b0d5206aa66fcd91f7938caf5ad7c3b1
81106e6e667a659c477e238ef9cc9a5b07bd7a1d
24731 F20110115_AABYJI muench_a_Page_115.QC.jpg
3320d9e21bc9b9c9f42fbbfb187ef47a
2eb081ba459a629e46dc74e928b30ff8bf4a69c4
168856 F20110115_AABYIV muench_a_Page_075.jpg
7fc5a74b7ca55a6cc7c2b247d40969f4
afdd3b8d6e879f8a97d7f03e3d3f65582b01211d
80700 F20110115_AABYJJ muench_a_Page_078.QC.jpg
e55bb7c8ba82878a7ee28e4887068bd2
05a29034243d256f5ef09b2262e5e788d3737e11
212273 F20110115_AABYIW muench_a_Page_018.jpg
ab57d80d80c2933e8b4de9b4509ce130
f05a5b25f6858be315cdecc2767852cd52c94672
F20110115_AABYJK muench_a_Page_023.tif
0c85d653e1897d7c106fa97ed6494497
0b471094410b3acb40825478f27675c15adeb7f1
50812 F20110115_AABYIX muench_a_Page_086.pro
7339bdaa53f3707489f5adb6a25688d1
6749c03e1455da7b2bee4128b95a4c382d2d319d
F20110115_AABYJL muench_a_Page_059.tif
d299f2b5da17e8d598ab9ac73ba47cb4
bc99a120a2e31f03c0fbf23e128ace9198291714
84885 F20110115_AABYIY muench_a_Page_113.QC.jpg
63ea457acc479a80f075f9fd87f2936d
ba36b8d1319db07cbce812fa824c672d0dd039c5
24077 F20110115_AABYKA muench_a_Page_022thm.jpg
9bf41ebcb2c54ce308f657f6299d3f96
97e3b71f152ac789562e2d4a1855eab45910db2d
209857 F20110115_AABYJM muench_a_Page_017.jpg
9b68616b00cf8730d03c311fb40983a5
e6795616022a460f8346b71fe255d77e203b19a2
39895 F20110115_AABYIZ muench_a_Page_107.pro
13c23a6281d4aaebd86accdc0523abc2
836cdf0f51a89ee3f20e898f537ce9580b0a3bff
2440 F20110115_AABYKB muench_a_Page_111.txt
1176ccfc0104290177873351b5f1fc5f
e6ae3bf4dc9d4766169f5ec7a5eb019ec1f9a933
F20110115_AABYJN muench_a_Page_046.tif
89ebebdb69ccc10afac1090175a417a8
46a6e87de09f3c2a8c910ba2251a062b3d221507
110470 F20110115_AABYKC muench_a_Page_015.jp2
6199cc7b28cdb69f7234cb23daea2ae8
de8b188c637186edcf8aa3eb554a8d90a4e58f54
49148 F20110115_AABYJO muench_a_Page_085.pro
69e2f6855e2385e703be9c9827820c1b
77faf7684261aecf8c9613e8457835d8b3b2ed57
267340 F20110115_AABYKD muench_a_Page_112.jpg
4ec33cc2f5f9d090639c4b2f8e278b39
ca81e4c3384214c4fae968a76de02a88ff85606c
43773 F20110115_AABYJP muench_a_Page_012.pro
f60a5dc587c8b12f120c213a7ce7d409
73a79385d125014fd6aea3bb82b6b7b040717936
90393 F20110115_AABYKE muench_a_Page_109.QC.jpg
ca27b7cab846977754fdfd45e9861aa7
9c810428e6bda3067ae373d422d876cceab8668a
14397 F20110115_AABYJQ muench_a_Page_116thm.jpg
c98e4a7e0664c82bced59251e409918e
7776bd79552bbee4dd7af2d201f50d60c8555954
295491 F20110115_AABYKF muench_a_Page_094.jp2
445ec39992636a6bff164d3a0adcf1f4
6c3e6847711d9b54ddd94d4d974a4b6a1de7f461
F20110115_AABYJR muench_a_Page_012.tif
b834846cde2783f61c561eade1d705f5
7fe51401586240832f2661404b6b0ffe8ab83118
20677 F20110115_AABYKG muench_a_Page_028thm.jpg
6fc63db9e6abaff61e752d5957dffbc5
2471d7c0ca7432932dff769490b6c0f4dbb03b1d
46114 F20110115_AABYJS muench_a_Page_071.QC.jpg
8224b3d16a1b92f16d28dd9548e5a423
562966d69ed273b8550648a9725fe38dd3bf0920
67248 F20110115_AABYKH muench_a_Page_012.QC.jpg
fc6689d84b0de6b5cc232c82d86e6da5
2a91fc02f005673c054c5a96d08796c6c75075bf
34899 F20110115_AABYJT muench_a_Page_062thm.jpg
47522edeac4dacd57f945fd680d577cf
e21d6819da9bfd9da97a03503cea670602af7bff
243338 F20110115_AABYKI muench_a_Page_013.jpg
65a550435ea449c1257cb3549363954b
3ee0e9bf0a4de26ce7bcef2d00bd11f4e084780a
62798 F20110115_AABYJU muench_a_Page_044.pro
3a6cc08c9b348cfc5629ade85778ae21
b9d3e4b7d1345495f037fc9470f9087b3f71aa65
F20110115_AABYKJ muench_a_Page_116.tif
55c8a14804b4e648b6747f44c3dabd12
0a9265aea2a78938b5b0498881a1fc8858de9cf9
56075 F20110115_AABYJV muench_a_Page_047.pro
01f14e96008b60670a777dc3764421d1
1f8bff40a37a6755d496be2a16b0afabd8a92317
132942 F20110115_AABYKK muench_a_Page_112.jp2
7b21a3279abc67c2c5eed3611b89e54b
f8f465189184cb5783a901dbe36b8e01622b1b20
73282 F20110115_AABYJW muench_a_Page_034.QC.jpg
cdb88e5808a6347c5d7928fba9421d55
341431a5201c9ec8369b7a636b6a6bef620b2478
2245 F20110115_AABYKL muench_a_Page_048.txt
88f19f910203288d679f142e3c2c2243
3d64e81e5196d564f82da712db4d9a4a57cd9c04
190154 F20110115_AABYJX muench_a_Page_011.jpg
f8714e62635b728f3de2b63e7459ec7d
ec5b9d3f691fe515c078ecf3820c93fd5d52e16b
79167 F20110115_AABYLA muench_a_Page_026.QC.jpg
8d976992a91bb77dc96c933eb4268446
1e7dd918bb1c811b381bf676555a081f5b5c9cb0
F20110115_AABYKM muench_a_Page_042.tif
c5fed19598a63ceaa537ed37c67e16a2
c26ae6c5fba66fd0ee07043cdd614d5633eefe14
111319 F20110115_AABYJY muench_a_Page_103.jp2
df8fa48249f0ab4b336e22a4d8e0ebca
b0b0f82d551e5f35a6ec5f348dddb9973e593edf
483 F20110115_AABYLB muench_a_Page_001.txt
872b85f0f7363e51c7b2d27c5ab4b90c
1014ba81e6a0729de9586c26c413e265e92ea917
1950 F20110115_AABYKN muench_a_Page_017.txt
3b1a2d3483923fb1b6280768f82daa18
6b7837505eea4ef24e18511757b5462dd8233ebc
89640 F20110115_AABYJZ muench_a_Page_067.jpg
858996f8e80c8345594957bc0c78e818
99eb9ddfb56a57b67f8474c1f5d46def17e39041
F20110115_AABYLC muench_a_Page_110.tif
d3ca29d319424c52d6f35af0ee925638
c2c47824113f3b03bd0e287cd23d6c7f97845c5b
F20110115_AABYKO muench_a_Page_011.tif
cc01cdd6ac9290e2aa191f28d2f04016
ecafe70d2f36354395724b5304ea34d0b225e26a
69461 F20110115_AABYLD muench_a_Page_004.QC.jpg
7afa32a039b82ab00b77509f271f638b
e835126914756904de0fcc76d5d9088cbebcaf83
1051939 F20110115_AABYKP muench_a_Page_013.jp2
8a833437250cce882efd783099916713
85f3a78d79f9099fbc4adace26366f32fd3a80fa
114931 F20110115_AABYLE muench_a_Page_088.jp2
907a50bc3dc1d5475bf57129e002af24
1c468056be28b44454006644b39fe624a1a312c3
25332 F20110115_AABYKQ muench_a_Page_017thm.jpg
e90e765cd7c54c3a3480540f075bf6af
c5cd25e913af98cabd8e36eff71ecef348044e6d
81322 F20110115_AABYLF muench_a_Page_088.QC.jpg
ed19acac65d88f9582b8c28aa30883c2
b6e7a9bc3f31658193f0c06e1310b91fe2d5746b
25641 F20110115_AABYKR muench_a_Page_113thm.jpg
7b4e44d25f55637b9ad494e262205d77
4a542c37a886975569af736943069db6f1d93e86
40363 F20110115_AABYLG muench_a_Page_090.pro
a34cf6674bd215b1ec5db74606b3a6ba
e83e34268ec07d57f86d6a92328c6d670b7fc0b8
47364 F20110115_AABYKS muench_a_Page_009.pro
e131f03d994bc5dfb1cb863e977b9eda
96b4fb998541bbabc7494382c27152511ffa335c
2577 F20110115_AABYLH muench_a_Page_046.txt
82f81d8e7e1a09497f8b8b33f41b65c3
f70c5d66d2457edc73ddc715f52ce4fc8de2f26d
3050 F20110115_AABYKT muench_a_Page_044.txt
654da89055d5494c1e21c2200908b512
62dfb5e3d6dd875d909e91a9756da7ef23cf4828
8721 F20110115_AABYLI muench_a_Page_001.pro
1089afaa6d92e88bc6bf76a5bd471df7
274578581485b04b31bf29a31bcee61689a9aa94
111291 F20110115_AABYKU muench_a_Page_008.QC.jpg
12e30e6a763ccf1bd8bcf52e0cb6d84f
9e15a4126a5b198da6cb6df552fbd79c1e2e4b44
44147 F20110115_AABYLJ muench_a_Page_005thm.jpg
77c5098ba1fd046db3782c495541e598
d022962888fefa8ae288404970f35ead5e399fdb
105347 F20110115_AABYKV muench_a_Page_081.QC.jpg
2fe00c5716498acac0ab4048fa33fefd
cf80002764fde30b181612aa5d63a367eff95d9b
1897 F20110115_AABYLK muench_a_Page_052.txt
933eb9131d782d4ddf4bae8ddba6b712
52b284e1a3773662bdf183ba3c5653310a2a102f
107210 F20110115_AABYKW muench_a_Page_038.QC.jpg
ab4a4ab3ec109a99c2f1a8467476d61a
9532091443a008c7739e91d17d6288918c714bb0
54569 F20110115_AABYLL muench_a_Page_049.pro
736bf04fd9436c48a706b8f4c35153e9
a3c09c3431ca5ce80e9fc0b19cf17cf52c6d5bae
170050 F20110115_AABYKX muench_a_Page_090.jpg
40f073b6a546feef6188075bf6af37f8
747f9a65400cb957f6e9ab1274529c14d2eb8b96
106883 F20110115_AABYLM muench_a_Page_102.jp2
a344edc26d15554222a8ecdc21afcdec
511badaeb749f33e3211839ab87d9ba130e3f6a1
14278 F20110115_AABYKY muench_a_Page_071.pro
efa15488e0f51e95759b1a281de5aa54
bfe0ecaeba7beab58a539fc8558230f6a4654526
317778 F20110115_AABYMA muench_a_Page_067.jp2
2311997a8e94f36f20b26691bfa27ea2
3a2ee0f8489dda874c873914413b19081b70b076
49924 F20110115_AABYLN muench_a_Page_048.pro
96abdcebe7fbe40a63317b97c5729cd4
6aae56695285c023d27808cfc99c06d67b419264
135422 F20110115_AABXPK muench_a_Page_109.jp2
245631fae7e6edbc92b076ad300a09e9
05bb937ff28808f4ba72c881c6ca902b23782401
111082 F20110115_AABYKZ muench_a_Page_030.jp2
6a48d8d243dc3f871cdd51687e9bcb1c
8c335c667fe0a48d8a716c26451e38a29219d655
345504 F20110115_AABYMB muench_a_Page_069.jp2
ec6e06ebde7d22de559ef64f4ae12b69
66933578dd06998f6c4d23356eb4229c348de66a
F20110115_AABYLO muench_a_Page_060.tif
e58d6864a827dbb42544a3c84fea3cc6
7d2ca0fc039ed9156a33e210c8d573753a5dd53a
F20110115_AABXPL muench_a_Page_017.tif
7a13d18bdbd587bb49bc3da0fa86a784
51762322fe675cf8f5e767329f210a3d0437857c
59863 F20110115_AABYMC muench_a_Page_111.pro
b1300b70b2e523c80731dbe06dcdfdb8
acb7ae884e3039182ffb0dd8c8fa786c26ccb8d3
24383 F20110115_AABXQA muench_a_Page_089thm.jpg
50fedea8dafdaa9461d2e4911170ffd3
d9151d9b54b79b44f32ee343e8118cd9657c59b7
109309 F20110115_AABYLP muench_a_Page_079.QC.jpg
ce1003e4d374aad64ef350222f5a361d
01d7f32db856ec6b9f0283ee8595f706732d0b03
F20110115_AABXPM muench_a_Page_001.tif
c2e67fd0dcd0ef9cdb535a4c7596b77c
3708c5873164af2c2204accc1a1846be2958d650
1759 F20110115_AABYMD muench_a_Page_011.txt
bb8c0e42637803a98fd799c945f03242
d096feb931f5760050434a12241d7d1b58cc75fb
54308 F20110115_AABXQB muench_a_Page_063.QC.jpg
6ec88ec5afb607cf4bd82ae9d787461a
876963880057cb53ac2b2cf4afcbc847a60334ea
102113 F20110115_AABYLQ muench_a_Page_066.jpg
b41be24a33f1e9538e137cf9c5a30e66
84e0d369a8edde946c82bc440c1fdbd0bc2892c3
191502 F20110115_AABXPN muench_a_Page_100.jpg
062cceb94cc5a140952c9f73c84e5da4
70a96c162661b08639ece857631743692ca3f2a1
50508 F20110115_AABYME muench_a_Page_038thm.jpg
eae620ad7bdb29f5ea2b1e505b466b8f
fe8f6e14c519e8033d0c618d1ecf7be62274396f
F20110115_AABXQC muench_a_Page_097.tif
7abd765b69bced9eb657e26511fb37ff
a0d8475f5647fa7b1c2f2cf6887a359c7c8bdde0
F20110115_AABYLR muench_a_Page_111.tif
a333613363b6ab06a0135b08806bf78b
bbdffe1221739368a342e857540e3c3d7b0bb8f3
F20110115_AABXPO muench_a_Page_049.tif
a00fb6f1d23aea91d9c7a5eb258c6d04
f12269d86ad8ba65685a88224020507a22ee2214
184727 F20110115_AABYMF muench_a_Page_012.jpg
21814c050f4372f174f5aaeaa045bd02
44df24222028f0d445783df9eb794c8e571c3f24
24555 F20110115_AABXQD muench_a_Page_016thm.jpg
d381703c7c3a2b069d386cafe0f62ce9
89dfa4b4b6e58473b2106535a5b5dd076703fafc
244022 F20110115_AABYLS muench_a_Page_033.jpg
8ae8e6d1eadb3fd44794b1f2da22f5aa
2be8eaf4612884f803351ada2d52bec377ee2bee
50360 F20110115_AABXPP muench_a_Page_015.pro
a0b4639c2719d6f34e99146c3674e1c5
16c0b9d93765a5c8714a11007717aba25d3b0f5b
26309 F20110115_AABYMG muench_a_Page_088thm.jpg
c03725b91b3588a83922bb4983e74d78
f1cc8d0b2570be428469d8cd797541281983f9b2
210482 F20110115_AABYLT muench_a_Page_055.jpg
2da24dce09430a4be71c0a24ad40cd80
389b1930b51e0835aec395c5f08cad528e23b120
109251 F20110115_AABXPQ muench_a_Page_087.jp2
784e258489821037b2a12ba2ead8d494
43dab62d3390742cf344c177bfc24607aeb154fa
F20110115_AABYMH muench_a_Page_014.tif
31a0cc524151c04122d83d0a9ac95a14
303df424e5140dd5febc814e1e3c48c7dbc4a68d
F20110115_AABXQE muench_a_Page_078.tif
af84f8fa7367f949c25bbee5aa1799ea
ed864ce7056db1edef9766bcc64ad9023f49496d
1960 F20110115_AABYLU muench_a_Page_019.txt
dfab39ac07d95ba6713a5e196a5a135d
6112a4cc6541f497f07a281d024d4f13cd7c0c10
51738 F20110115_AABXPR muench_a_Page_051.pro
2cf6bacce2b02260ec38a70d3436a5b1
fe10a6ed29250cbffc78a26e0803b58ec5586ef3
199879 F20110115_AABYMI muench_a_Page_034.jpg
f53e6f6c947bb8ff42af03dd2af0fe8a
b63d7d27501075c93c7a7f3551441f7d0acb7971
47403 F20110115_AABXQF muench_a_Page_024.pro
257acf4fccad9e903097f801fca5ee4c
132d3bfedbed17cbe40bc8dc7435d0ee18280822
102970 F20110115_AABYLV muench_a_Page_044.QC.jpg
07f37804519b92ae5350ffd17b454575
5ed706094fc9e2be38341e9ae9d49dde5069eeb1
7995 F20110115_AABXPS muench_a_Page_072.pro
1218f5f38a29ca8fe4a4fefa1bb9f610
5da4657317991223e5a5ee8325449c976dbf9e7a
13616 F20110115_AABYMJ muench_a_Page_066.pro
ebd842e71d33335f9b205cf51513e982
8eae4ad5aad9baa02fb3065e5f19f46a42c52721
82735 F20110115_AABXQG muench_a_Page_056.QC.jpg
3ab2b5b2cf831e67eeb0ac2478389688
532fdf0d3aa2bd779b2ca36e60c43674a1e5890f
78124 F20110115_AABYLW muench_a_Page_018.QC.jpg
dab4a15a24fa9a5b7b09da86887263ed
23ccdaca587e0100446a905249e59a50b2f1a7e3
F20110115_AABXPT muench_a_Page_098.tif
93cd593438d3441812856cc9c875fae2
51d5d633fd86bd15a24705a5f1eb42968b0b81ea
81404 F20110115_AABYMK muench_a_Page_077.QC.jpg
80daf137cf332d6a68e89b3b3c360aad
af35575d0c36143a6027bcf3e7f374b6f3d03a56
45534 F20110115_AABXQH muench_a_Page_100.pro
b6490ddc735ae6a3a6a37b8eb149548b
2c51fd0b0764bbbc81cefcbd6d0e45beca956cdc
212553 F20110115_AABYLX muench_a_Page_080.jpg
dc859fc60e745b5539ea65dfcf446175
6d407eb00c54a81a845699358466c70e91dfe9c6
2495 F20110115_AABXPU muench_a_Page_114.txt
e5ccf1e0bfb43fb179d157cace833f3c
06123236d83759ecae643b04c139cd454318ac02
429353 F20110115_AABYML muench_a_Page_063.jp2
23bac4e640341b2c099eb480a9188f8a
22051852ba84367d1d753c785f881ed4b34ab69c
72403 F20110115_AABXQI muench_a_Page_052.QC.jpg
c09836b5b8c33f20f15660dda2d83a6a
afd6c61f7b9b6a2e63d89b9ef4178e27964c5b4b
65353 F20110115_AABYLY muench_a_Page_027.jpg
4ea700497b3faffa869260e6353b9d04
b97bf528c980f7716814aebaea2cb89574492c94
215530 F20110115_AABXPV muench_a_Page_056.jpg
4e644a13cfec4313528a54c2c575b839
85c510f22b71d0e12b0c60d951f28d75536ffc57
268740 F20110115_AABYNA muench_a_Page_049.jpg
b31a619ee3a3f5f6e515356d7cd07338
9f730c44cfd6fefbc37933fdf78d22cf5b2bb3fc
218903 F20110115_AABYMM muench_a_Page_083.jpg
270ec9cc7ecc5f49e58a25c171803c38
7cd9e74af7d008f866bc4489513ca50c3b4b3ec0
2093 F20110115_AABXQJ muench_a_Page_056.txt
3679fa1054ec7fa52dbf7a3ae6114b85
54638966c06a47c3f5aca5c0b4174eca7a1ce936
113549 F20110115_AABYLZ muench_a_Page_023.jp2
bff7090f98cb63989a91d81212a9e9dc
6647dc95781858de7e27247692cddec6eaaebacc
612 F20110115_AABXPW muench_a_Page_072.txt
88aa1ec49904e6a2274f5d200ae2f750
e942fbd02eb888851200b16683ec68e62337fdb6
201034 F20110115_AABYNB muench_a_Page_052.jpg
bf4444199ca23f21e24ded7080ea5c69
da81eb1194855de2c33b3880fc374bb7ca178a62
F20110115_AABYMN muench_a_Page_016.tif
0c2e583929aeecfdd969af176d859187
6ff23605cf68375ca7904fd81acfd54aba20ac37
266999 F20110115_AABXQK muench_a_Page_044.jpg
7ca8de8d5f094615d5e6d51f3f8395b1
73f050541572927003766f5bda2ba66b4bb2df32
78928 F20110115_AABXPX muench_a_Page_020.jp2
fb39778e110fe63014c0689c13e4d5a8
06d8e79612f1939d905a793b892f609161fe4c33
142436 F20110115_AABYNC muench_a_Page_060.jpg
056be8bd924a1f9c2deb54d994a5b09e
e5b5b3cab0cd8e8105d2bb4d5cc6be919e2a578f
109650 F20110115_AABXRA muench_a_Page_035.jp2
59878b3091315f0bcaa7ff509fe2c6b3
2a9cdf48d637d76dbc9535364ddc1f2d84cd4620
174541 F20110115_AABYMO UFE0008580_00001.xml FULL
31266464cc6d2fa02248a755bbc07fd7
38cd7f0e66e0f09ab440f33a888f6f2c3db59ee7
19185 F20110115_AABXQL muench_a_Page_094thm.jpg
cfaef2dadee772bd5bf65cb40b5b5a0b
0732dff70a0d23a5a9236ed4c8f2902cacf2c69f
1051975 F20110115_AABXPY muench_a_Page_046.jp2
3e1660319ef57b2d24139f293f28d382
fbf6865d14ddcfe2b54e899eb20fdd1d4d93e17b
99260 F20110115_AABYND muench_a_Page_062.jpg
6ab37bb7710a80afe7ff454cdb0c8a0f
2b34d641a57142ae77895d4e86de8fdbe17f74cd
F20110115_AABXRB muench_a_Page_082.tif
43e7dbd2bfb5b28df9428d9e7b18e80e
d38d6fe76322ada0b6500b36499fd2e5e0cc8547
454560 F20110115_AABXQM muench_a_Page_070.jp2
ec5313883919040a292dabda3e884221
b1dfa63d541800cd0802c57bad8a8b9e29d9e454
108478 F20110115_AABXPZ muench_a_Page_018.jp2
b9bc5c04170b878397e69103636d5714
3ad38391aa6a27f22d5deec0d3ecabe19fa300bd
105644 F20110115_AABYNE muench_a_Page_063.jpg
0464562cf915cec2ce6c2c278f4a6f24
b946f0b61fdded21233fc4f458acffc85eafb665
87806 F20110115_AABXRC muench_a_Page_107.jp2
dccc372b7294b4ed31f61720d7f75c1b
7196c150c771af7599b6bd8bc15c6067c9462445
50830 F20110115_AABXQN muench_a_Page_006thm.jpg
cbbc8ee55343e3b46f58334fc42b6bc7
22abf859cdf1b299393d7d28b79a5a9939502fb9
100112 F20110115_AABYNF muench_a_Page_065.jpg
600cbdab413f436ec3bd894b7c855d9f
dd292cb639d42bf925188680bf24d8ef21f3ca5c
1900 F20110115_AABXRD muench_a_Page_024.txt
1b195b5fa161c635600be756f1b9dbeb
d798bc538e8db589b09d1dab7318e1c037c623d3
58927 F20110115_AABYMR muench_a_Page_001.jpg
888aa508e4c6f88aea1a1b21f0fb64e0
8e3c8d871014a5a2bb126cbbef6f6de1a1c46d68
48483 F20110115_AABXQO muench_a_Page_050thm.jpg
b3220dd4dc02b9376c8831d4f5a4b0c2
b871fc5ea9be0b6c048a498c0d14a22f58472215
99048 F20110115_AABYNG muench_a_Page_068.jpg
a5bdd80282560467d04eb602ef7967c4
cc685b8b33ebd76fbb111aff06f70cd231090c86
F20110115_AABXRE muench_a_Page_018.tif
a6e8b06b4e810ab081aab59ed587277c
3b870c9544d4fff4070e296d6074c2f78dfcab3f
183274 F20110115_AABYMS muench_a_Page_004.jpg
45e2f85b3cfca6752f19b3122d4a7a1c
174be3c5466b465bdd495731fbd108944f8d408d
F20110115_AABXQP muench_a_Page_026.tif
ad73967e3f7fc5e83f0287d93e67a8bb
dbaaa3721c547f970240819095b0917ae55b9173
109425 F20110115_AABYNH muench_a_Page_070.jpg
f09e1ad3b0355abe753c7d3adfe3705a
1a296d3fc265d4cf989a5ad42f401db6738d876d
211985 F20110115_AABYMT muench_a_Page_022.jpg
b5709c2e41d1f5ac0d7901ef0b2bb0a4
3615b112bb6e5340ae4648adc452d0454f609ef3
51323 F20110115_AABXQQ muench_a_Page_064.QC.jpg
f76efef3a830dee40cf4a28296d19a9a
16bb37c3b3e991ba75390315a220b205818388f8
121893 F20110115_AABYNI muench_a_Page_074.jpg
9d8b4bb9af15da1a658f5088bf54fecb
066457ca85f92fcc8540e2ee07c5498514c1edbf
82050 F20110115_AABXRF muench_a_Page_094.jpg
6dd9f244de98560cf75b3292fbc221f9
35575f905ea2477e1cc33c1374faf37ac00dd97b
214659 F20110115_AABYMU muench_a_Page_029.jpg
f816466bcb24b084b1c6d023cd51df66
23242b8a35fb2add756365a2f0142c21091c9332
F20110115_AABXQR muench_a_Page_113.tif
1cb36648e43241fcf73acdacc0fd1cfb
a810a43e9ecc41367ed0ce5313172e557ab367ce
218281 F20110115_AABYNJ muench_a_Page_077.jpg
d32dcbe7a83867edbb072aeeba2482ff
59c55a73232d06a6bffc602bcd5e2d8ff1ad9930
24158 F20110115_AABXRG muench_a_Page_043thm.jpg
bf77c27c252514e81ae93f18dfc890a7
281e8f4ce6fa7b443ef8bc45509196f3a6e12e02
216973 F20110115_AABYMV muench_a_Page_031.jpg
6f8de794dec60b1c1dc0674b82f9be7b
9e846e24d3ebcea825e6b1696b2a3b9b0ab0af45
79907 F20110115_AABXQS muench_a_Page_030.QC.jpg
8514cbb940265d2f53ec2e9f8af6650d
a5fff7dc2050cb2bfb9e9834b8d119d850ee3b4d
210625 F20110115_AABYNK muench_a_Page_078.jpg
fe2b8454e837f56e9973d1b7686def25
068a3760f297d7529ecabf7525cfbde3d622418f
49639 F20110115_AABXRH muench_a_Page_051thm.jpg
ddaa24dc3a81eeef4e9bf3484dd0c499
8a74adf874bef84f3f8839e99216ba09ed42eae0
217653 F20110115_AABYMW muench_a_Page_032.jpg
f68167b4102ae79a650bbe11c8eef3ff
615cb6233a43a27d516691890c3f03d296d09541
69524 F20110115_AABXQT muench_a_Page_054.QC.jpg
5b049b20b80d6e5fefa045eb88a1be8a
f57e0d0f25ee8dcdb1f4d2df284885dc63bb63d7
244078 F20110115_AABYNL muench_a_Page_079.jpg
2868de3e557cd7af75472a934624a37d
0294ad9e06058864a9e9bfab85b47acae8bd48bb
665 F20110115_AABXRI muench_a_Page_098.txt
2238bf185672079fa15d27c1535864bd
1a02cdba34582f1281dc429ab0182eb6427692a5
209868 F20110115_AABYMX muench_a_Page_035.jpg
2060f82749b1fe8beb1d7b2c45a880a6
797c5a1970596c6c92f40e4dfea7274d38e72ea7
1854 F20110115_AABXQU muench_a_Page_042.txt
6c46d8e0eed0fd57860d4d82d3011252
cb709c19d92fb451cb7943c4687131ef373e041d
111415 F20110115_AABYOA muench_a_Page_029.jp2
bce48053317078c1eca21704deafd410
6ed4375f3e211eb9c08e5b57d5dae830d8f8fcb9
218176 F20110115_AABYNM muench_a_Page_088.jpg
e94904d17a5ef6a7e08e06871925e49b
9e8b0a1447de830c02e93f8ca5e976e8b4756101
22846 F20110115_AABXRJ muench_a_Page_025thm.jpg
2ed07faea37f41fd2edd21008c3ac576
9a3b6c264e3abc18e493acfd9e6bd88772daac82
216378 F20110115_AABYMY muench_a_Page_040.jpg
e6ce42848601df87d08db0c7ba589a57
27e6ac9abd6fd65d87000784fecd0524df9d6b27
F20110115_AABXQV muench_a_Page_104.tif
103431e9db1a56a30304d0ca668cdea7
86e61d8e38ae719f6dfa82671fa7a5b47dad1200
209901 F20110115_AABYNN muench_a_Page_089.jpg
2ac94f178e119ecbd2049bc82d8158b8
1c1b589459f3cd63f540bc486bc7036fe1bacc1d
2587 F20110115_AABXRK muench_a_Page_047.txt
6f8e923dfd72eb0467fad7c312b23ab2
177b03ee44d4ff3a9757243bffcf7284170ac4d9
200164 F20110115_AABYMZ muench_a_Page_045.jpg
d4eb7373f430f2dd1ad8a73d4bcfb0cb
f462640102df2ecd6ee5ea960c0d0fae598bfb2c
F20110115_AABXQW muench_a_Page_009.tif
685121042b8851f949f54344e966246e
cdc4bfc2024b48c5f578539964cb41ab9b3319fd
100507 F20110115_AABYOB muench_a_Page_039.jp2
9234f717629ae96fef7fea68012563c5
458c75abe1636164d46d48eca8a827fbffb284a0
17249 F20110115_AABXSA muench_a_Page_020thm.jpg
82cf16b8cdcca80d18a66cf2a914817b
e44104dc47f4da01fc5ae0aa1f279d51dcdc8dad
86600 F20110115_AABYNO muench_a_Page_097.jpg
6bb2595fbd9119d2cd274a7c8cd74e14
ce8c35c112707024df1617e0dd1c1b363cee299b
219884 F20110115_AABXRL muench_a_Page_106.jpg
8639d891d78f259f7872f5cba0abf6f1
5c432d6369cbcc1295d872e6f2f5bb0e2a852ba5
2015 F20110115_AABXQX muench_a_Page_022.txt
86bda055b29db6b98f8578bb8ea67e05
173a6c2c6700758c9b29cb9cc693bc58e2e1c025
1051958 F20110115_AABYOC muench_a_Page_041.jp2
6e208294858545da1db6fb768c438415
50b2516ad5ab8905a4ebbc3712c2021e56c64f42
49182 F20110115_AABXSB muench_a_Page_105.pro
f7339d725b3c617eda0dab46628ab98e
88e6b035e731bcf013e20ff69dc5cf22cd0f777c
189980 F20110115_AABYNP muench_a_Page_101.jpg
3a2e5353295ab3aa12bfd212c04397bd
b945ced0c550b5fb2e5e018bffbf291014c4a5c7
253068 F20110115_AABXRM muench_a_Page_091.jpg
723e330e6e90097d0f7a9deea6ebc500
43c5e3a855c1b1ec8c5b5f0e5434ed9d4e2b09ef
16892 F20110115_AABXQY muench_a_Page_072thm.jpg
51cc12dc1dcacddc7b006674a832962f
86c76b14066e22a258b9c3f7d2d9d3e19f79260c
985889 F20110115_AABYOD muench_a_Page_045.jp2
90bad7823822d9d287fd6489e29b9da7
f868437db8c4c4e6cd855bc70f7d81e3b5b03b3d
52973 F20110115_AABXSC muench_a_Page_065.QC.jpg
d276413bdfe638b861b94471e13e4430
9b92d4842069d89041a262114578dddc58ea3171
204778 F20110115_AABYNQ muench_a_Page_102.jpg
e897115b93953a236b6b929915ebbf7e
f9bc61ce9c19e1983ece217cb76cacfda299afa8
F20110115_AABXRN muench_a_Page_045.tif
6ccff6d0a300c216464d09961736a22a
5d4d119126cb6be17bbbf347ea44442bf5557aa9
203719 F20110115_AABXQZ muench_a_Page_105.jpg
c1a25c2854ed4321181bba5b83e89ce1
98ba8ec16750e26848e007fcca7a8a6715b0c560
1051953 F20110115_AABYOE muench_a_Page_048.jp2
3e8efb3127801aff384ea4efb4e1f3af
17f6ec2f8f807f123ad5d8cc229b25567e450da8
291150 F20110115_AABXSD muench_a_Page_047.jpg
c67734f230b3f55be74a5fb71bf82747
1cf1ce31deb4e40e6be91dfd644486c899f35fee
263325 F20110115_AABYNR muench_a_Page_110.jpg
9b92be86c37fc0e5253db2d7b5db7f9a
48d902112c9dc830b763df42be247439979aeb5c
109669 F20110115_AABXRO muench_a_Page_017.jp2
cdf32333faa44f23ce125ea9e421b39c
f9e6535dde6c74f597eca25cfaab6563dd14328e
103331 F20110115_AABYOF muench_a_Page_052.jp2
a1b8bf108ea5c0d3b6b3d01bbb932811
48a8418c937f6e99e33770f34b759c1a70e0fe3c
22251 F20110115_AABXSE muench_a_Page_108thm.jpg
57074c2872ff6bd128583570fe570d02
44341a9f73617eb198028f318aa657c4376632ea
76117 F20110115_AABYNS muench_a_Page_115.jpg
50f91753ed956a54e6e465ca219474a0
924e0937e528911f49b5df380915e406a28f1769
39828 F20110115_AABXRP muench_a_Page_028.pro
a21c454044897753f7be368a1a8b9873
63c2c196dd34d79216f75304e53e8bc2cdd515b1
99578 F20110115_AABYOG muench_a_Page_054.jp2
864f0c527031672c4c23d698faf890a0
3b223400d1958f0703825b621f62a0b019c92724
F20110115_AABXSF muench_a_Page_028.tif
879665b3d1cd8426350623163034d836
0c254dd9771a7f05f8ea1285234e6ffde45de63f
5770 F20110115_AABYNT muench_a_Page_002.jp2
4ecae9cf420959e06c7a826b4ac2920d
fe17a955380c6ce073b9422da8fb4ed94e596b53
4009 F20110115_AABXRQ muench_a_Page_006.txt
21b1ba8e4e45ae92eb16bedba73374a8
46489cfa5aadd1885b1add77f74582a8ac9d6135
107665 F20110115_AABYOH muench_a_Page_055.jp2
667ec04799f9eed6e8716b3f6105a652
e0efc9f46ab39df1de245f438b1faaade4420c83
1051962 F20110115_AABYNU muench_a_Page_006.jp2
772e49eac5741e099b1a4450a80b371a
7679d57a03b44dbbcacf62c3f76303dcdc300b58
127996 F20110115_AABXRR muench_a_Page_113.jp2
37d4d6205da288bce4a3579007291ae0
15afd8d833c06d24d0be19fde3ccf46be0dd9787
1051866 F20110115_AABYOI muench_a_Page_058.jp2
6e0507f269da4e78c30e5e61546b2b06
fb97bb3f5f0d5d891aefc6feee34b0a69d2164cd
11791 F20110115_AABXSG muench_a_Page_068.pro
6cdba2591d73081fd70881b625aa2c53
a1af498781f04b8f40e2483ea3542d831c58c6ca
F20110115_AABYNV muench_a_Page_009.jp2
8400d58977d1fa0499e983ee8293f9e1
5a56daaedb3db5fa929aa8edfe22e9117a0e24c3
303998 F20110115_AABXRS muench_a_Page_098.jp2
f45c0c96f71ba50fcb6b9b3173b57396
e0cec14425354078ccb27c1d9eb6728826bd479e
391195 F20110115_AABYOJ muench_a_Page_064.jp2
8f822d0b6b754a539c6a482bfec09aee
16e0f827f5def258c97f6f5e3106e144f32f57aa
1870 F20110115_AABXSH muench_a_Page_009.txt
26d037dc3dde79ccad91e2fd741eabbc
48d3de432830b76ca925ff92c0c0e8c35b952b64
109854 F20110115_AABYNW muench_a_Page_016.jp2
1b75bd515f95761777ba9b1822cc3f34
ec66f161b4d7bca26273a64b0aab60ab3679c687
1688 F20110115_AABXRT muench_a_Page_004.txt
cc9843f296c9e7f210a802dd7ba9de84
5e9b0aa073352d87b437ff1e4269e849cf4ecef3
415743 F20110115_AABYOK muench_a_Page_065.jp2
91fa0402729cc6772c93a953b3b67160
78a8fd1a3c0f96524c5538f4bba60e699c04ab24
52003 F20110115_AABXSI muench_a_Page_047thm.jpg
f5112ed7d49657bbd042ffc4261c2fb6
441259856eb70344a00634159d7ee318c801cc58
90318 F20110115_AABYNX muench_a_Page_021.jp2
a47f90648daebd11b5e85292d8ae9b38
7c0aef53ccde8b2928651fee016014a19aa7983c
F20110115_AABXRU muench_a_Page_056.tif
9cd1b4c0ab22c6e107a780cc4ab92f00
fc506c65bc840a62eb4bacfba1350dae3255cb95
F20110115_AABYPA muench_a_Page_002.tif
ba4ae1ad4722d39737088b05c6ddba3d
a22086e214bdf74de7eec6a1e2218166bd5f7fa7
417639 F20110115_AABYOL muench_a_Page_066.jp2
5130afaaeae20dc1c4c8a1d7a006c126
f993b4b4e764a38442e6cd1c97ffec1e241ffce6
F20110115_AABXSJ muench_a_Page_035.tif
9ef01ff3c7dc0bfba4dc395b2276d4d4
7726b20582af0ce809885fdb74197bb2e593d7af
110699 F20110115_AABYNY muench_a_Page_022.jp2
bb3d26e11aaf0ae16fb6c3f05ae2b875
015649de21a321f0af795560bee5ff33d9b1614d
776016 F20110115_AABXRV muench_a_Page_042.jp2
1f263270ed49b0fa76a71dc01185e465
1697a528a8837940f1761c2dab5500c7f85801a0
F20110115_AABYPB muench_a_Page_005.tif
74afc9d5c224049fd1c293a5e74bc3e1
de72e52bdc2e24bdc0a2b9d30bd71d82e8a87af4
394927 F20110115_AABYOM muench_a_Page_068.jp2
96d4bd03f97931c40a44864fe34d324f
030f08727b1a8444dc6bdb83180db97746f06d30
77936 F20110115_AABXSK muench_a_Page_105.QC.jpg
3715ee33210aa11c1d7d7a2d297ce18d
cf23e4e86502b23997d552e10b54449e2976792f
88796 F20110115_AABYNZ muench_a_Page_028.jp2
41867954c0a500d526d7cb01ed869003
32b515e135cb860551f722838bddeb376f1e1a57
24624 F20110115_AABXRW muench_a_Page_106thm.jpg
6fc360c2affc151940333bf3e952a766
65c90ebfba34b880de5dff6ca7ba3c9da9fdafed
298972 F20110115_AABYON muench_a_Page_072.jp2
e7918f26071527acd231871f4a539307
4781c090d3df1b0cfb644a7dc6c170e9d071bbe1
212243 F20110115_AABXSL muench_a_Page_104.jpg
4dff3ebf71a00674a06e2aeecf607b0e
852c1e3623d32945e04ee82bb990b39f413b738b
F20110115_AABXRX muench_a_Page_039.tif
7422afa37ba394fce644aecdbce04f96
2617bb8dac1636200784e4149b237f59dabdf760
F20110115_AABYPC muench_a_Page_007.tif
8bb7fc976573116ab06aeb55df5d249d
83f6d3e86e11b0e7ab0884a5ae370be86f6a116e
25642 F20110115_AABXTA muench_a_Page_023thm.jpg
857b5ceb60104895eb6fa968c7c9dea7
fb36b8e63bb1d5691e818609713da0ab8ff76313
111750 F20110115_AABYOO muench_a_Page_076.jp2
18a38b4e49f51b1a440d0399caa5fa76
8718395cd2faf3fb6ad06e005fad81e0719f3458
F20110115_AABXSM muench_a_Page_013.tif
4bf7b57dbaedad1d25f52d54cbc34e66
d2f4334ce759dc031c357edfb9b14fd9c31d1901
37542 F20110115_AABXRY muench_a_Page_070thm.jpg
fb9e68f9a63db7c13aee1c6eaf82f52e
8c5d23e34b12297df8ebb4e09600cab2a64c7c94
F20110115_AABYPD muench_a_Page_020.tif
f1908d303e6a720b4f4c5d9583ce00ea
c2ebe6c3b62b63ffa138bf34bf89aa29f0943a92
F20110115_AABXTB muench_a_Page_084.tif
61f1804e9330b6277bca6362a685b206
1c5d60fc71f96ad5f6f4e765364fc08f9ed15633
1051900 F20110115_AABYOP muench_a_Page_082.jp2
b962dc6a0c3f030471d91c7e91ccbe43
7faa8713addaba60e90c014fda6af2c5dfb301a4
80119 F20110115_AABXSN muench_a_Page_057.QC.jpg
6da8b5cef182d1d47ca3abe0d48e059f
e2b5d1a8e28068795f7c04a8f1a06dc8cc128164
237311 F20110115_AABXRZ muench_a_Page_081.jpg
93a195f5468c1a2ec150c1dbb7ae8e41
b9184bf51e8927300dc2a33490be3cc31e7e4577
F20110115_AABYPE muench_a_Page_024.tif
52917f88d84f20fc27a6e5f4030edbd3
7a8af18db47cb49a44ae6ff64459b2877ae96968
112697 F20110115_AABXTC muench_a_Page_047.QC.jpg
26c37cb2f2b3e47875ec993a54c47d21
b445ae92ce7ebbf2da2e398def07b4fc1957cfc3
108327 F20110115_AABYOQ muench_a_Page_085.jp2
0c22fbfab074e6acdd25fe06aa639cb3
53897bf198cadb6a333dc5605cbaf0fe263e25c2
1786 F20110115_AABXSO muench_a_Page_101.txt
d0f1d708c78a2f545fde6b471569f564
09d591d05e11c2bac60163ca2b0bced6169c789f
F20110115_AABYPF muench_a_Page_037.tif
d22f7b94e8df2ff53b742f581cf6a05c
7d121d158cd5f05fc91ca7c199b36aaeee6484ad
1982 F20110115_AABXTD muench_a_Page_015.txt
8b900e51798bc192463acc1446b1e19d
7365390312436673f8e6e84b545ddbd626337a62
110831 F20110115_AABYOR muench_a_Page_086.jp2
d1014a5af13ed4ad485697b563d185df
d8a30f39692b1a555ab0dddd662dc4ba16b24193
22119 F20110115_AABXSP muench_a_Page_054thm.jpg
bc115aec9c15c151c1669f7bddc5ce5d
75ec975694811cc8980e0c85e8ff34559857111f
F20110115_AABYPG muench_a_Page_040.tif
c82cff09a5482c3d2455ab9f8278a534
7441b6e53b99c91b3d726dbfb8de7fa618319881
62486 F20110115_AABXTE muench_a_Page_113.pro
a26a8bd372d48424b8cc1209ec4d837e
32ff52f3bafbd01d2b895d985028e6adcd7e64f6
89472 F20110115_AABYOS muench_a_Page_090.jp2
efe9bab995dd41ede08dc6a8561a8ac9
25713c2f530d4964ee70be6e29fe2185e215a081
F20110115_AABXSQ muench_a_Page_033.tif
538363ab8a272107c9c00d7e73019f4e
3154ab3f08daf9018a0fc41893450cd7325f9bdb
F20110115_AABYPH muench_a_Page_044.tif
76bf76cd0d32bb6efc0b99c0daf5f2bd
0c0792a7122033defc3a911694641567ba5af780
45657 F20110115_AABXTF muench_a_Page_067.QC.jpg
e72602a8687744d3b6de242e66563f87
affe6dad211d330b5496b6dc0e197e0dc23b3c0f
305630 F20110115_AABYOT muench_a_Page_095.jp2
d4bebde1cb025f4071f4c49c09b76dff
4041b9ad2407ad772298fca95f264214e3e079dd
2020 F20110115_AABXSR muench_a_Page_083.txt
7cdfcb7dbe0d231a7890897cfc6a06b9
c3830d5bf5ec668177a3ca68e6555b66060a3258
F20110115_AABYPI muench_a_Page_051.tif
73e031be0f1808963f931472b913c57d
d9167d7d5a60521bd25db89bdd1aa640071f2d86
1051901 F20110115_AABXTG muench_a_Page_081.jp2
aa7628d8c33504e902478a94b4f597de
dde5b4b96054822dccc7aef4f1985d99edfc0983
325142 F20110115_AABYOU muench_a_Page_096.jp2
4b23a233eedcd4bea527fb48c46f96c4
16cb48310f1f752a7678db977be55278f8087a2b
943 F20110115_AABXSS muench_a_Page_074.txt
ed0bab5252d3bd22244c2e6a86c5e1a3
e0d52e6b27547cff78e58839354d1e84cb574513
F20110115_AABYPJ muench_a_Page_052.tif
fb5f3391ee72864133582ced8e53a054
4074e51e3dc09cf35c2ded8eb06bbb449a2166d4
102531 F20110115_AABYOV muench_a_Page_100.jp2
6f63baa217ad4e1ee91b7929aa00c976
775d24eec8d346d3f59da2e3b1f61b350779698b
80890 F20110115_AABXST muench_a_Page_103.QC.jpg
b6c59fe06101b4be158cfaa6eaba7145
cef9ad7d42a06ba42100239197cf5249714fdb66
F20110115_AABYPK muench_a_Page_053.tif
70622417af68be13fb841a1fbb3f7151
26a2bffb1ab9aced226a99836345f7b66ee887aa
75627 F20110115_AABXTH muench_a_Page_045.QC.jpg
79de9305b043d5796dde91e2c1bc5ee2
40b6e4a20281da64f180e7736a73deb45226d7ee
112331 F20110115_AABYOW muench_a_Page_106.jp2
4d1836f5189ac89fbef7220cec8244b5
23e82634761c156c7c743e79a7afd507e889a449
F20110115_AABYQA muench_a_Page_099.tif
ff51a11d25629169d188956ee2e40ea1
d87d224f82bdb9292ed0bc73329a3880163ce9ff
F20110115_AABYPL muench_a_Page_054.tif
334968fe67db865494bb607952f43c70
b1417e5ad50cdc081ea8e528c66ac7ad0ce339fe
1051980 F20110115_AABXTI muench_a_Page_049.jp2
3ad34289e80fa12443ab71514565b107
421f834167793a73054f6c989b24fc3d00f6a25d
1051959 F20110115_AABYOX muench_a_Page_111.jp2
2ca55e82d6810edea9f95fb641742839
d0daebca61fa8708271b64cbc69ab0d5fac01b5c
F20110115_AABXSU muench_a_Page_061.tif
db9c001bbfc15d20ea731f11f26af6e2
70e06bad09018dedfe796f9532b938d4473e46c1
F20110115_AABYQB muench_a_Page_100.tif
0c0690432cd82315563e0e029f48d0bf
9fea9cb65920718d49f4d8c01818b5e793955df5
F20110115_AABYPM muench_a_Page_055.tif
6f6b024c2ad7ceb083aff4202b6f9531
62ff19b26e7c63dd2aa092651081aa309257446a
F20110115_AABXTJ muench_a_Page_015.tif
191b61f0035c9c783064828b4f8c0053
04346affa234be6d6a232c345f34eba20eab00b9
35989 F20110115_AABYOY muench_a_Page_115.jp2
19a0b6f7d53465c0a5bc52871a24a808
d3d911e072045b81054952129fa8bf262a1b5eda
545311 F20110115_AABXSV muench_a_Page_073.jp2
6dd8f11f52b714c7e86a2bb6371c2251
938e9ee608fef622a8cd5cfae1c71eba9a753962
F20110115_AABYQC muench_a_Page_102.tif
cd4aa41ff2e1c4ec713234ad12210938
89c92b18369c069b25ac7a1315b778a705f693f5
F20110115_AABYPN muench_a_Page_063.tif
cb086eeb977f3628d57ef5ca961585a9
ed86bc3c538cf7f49943ace71c696ef3196fb95a
21001 F20110115_AABXTK muench_a_Page_074thm.jpg
3a2a516b63752c4d6aac83ee80446b75
ec5e743e4df8181c912d5ba678bb13895b12b5e9
59586 F20110115_AABYOZ muench_a_Page_116.jp2
ce0b3294ce6470c3652b2fba95719377
66ab67d46aed8c8f2862ef53e4f164b7f90e72ec
44511 F20110115_AABXSW muench_a_Page_074.QC.jpg
87880ed18023dad32a3b86e4c9bb9310
879ffcb0282917e344edfe3d812a582446f9c147
F20110115_AABXUA muench_a_Page_073.tif
ea2e485d535690a6dffdc477725c922f
87967c4ff4b1388485dc1486a4b51408c253b58c
F20110115_AABYPO muench_a_Page_067.tif
e24cea132c08d7e045e666a868fd4019
b847e9752ddf8aea32f8d186b19a97b1a78d477f
78650 F20110115_AABXTL muench_a_Page_085.QC.jpg
e8109b74bf73c3ce71cab976122b04de
35d6faa19614b14a45175d493573d6ebe7e1e66e
116191 F20110115_AABXSX muench_a_Page_083.jp2
e9d3238aba5bccabfe45f291d4548898
d31764f7bd5f4884d65fbe7b41eec1bbe726ec53
F20110115_AABYQD muench_a_Page_103.tif
37fdb9551a0a8c9ae556cf5a736455b0
829fb0082e8a9bf188a4e6621732679653a63bd3
48181 F20110115_AABXUB muench_a_Page_048thm.jpg
00eff46f208fa482173bc2f7a7e73542
dd64ee5431626ad57b68372bb0908a6e39c83ba0
F20110115_AABYPP muench_a_Page_069.tif
4c93ceb0ffa74414679da1fc7fcd0400
1822e11a90e4b7c202a29b550046d58c88716c76
F20110115_AABXTM muench_a_Page_027.tif
cd8ce57665eb68aad64964f0542c0339
20a57e79191af2a450430854a0107d687e27e121
122 F20110115_AABXSY muench_a_Page_058.txt
8e68fd34cbdf77d8973ace201fdc7eff
3c5083a3a4abdf4530da00072ed02ac14d2085fa
F20110115_AABYQE muench_a_Page_105.tif
22ae9d92a4fe27054b2d06576fcb7536
0233056c2d6e8c1a2bd0735cbaa039d6c785b9d7
25387 F20110115_AABXUC muench_a_Page_001.jp2
f7b74283e587ec35a9a6aab00e471f38
86890cca0990e0ca7c3fc2838a8e812454ad03d1
F20110115_AABYPQ muench_a_Page_070.tif
ea6bd9314ded9b1d1cebc5ed80db927b
94c0bd4ced442c422c92cdde6b06ed4542fe52e6
2058 F20110115_AABXTN muench_a_Page_106.txt
16a093480373b0021186aa9197f7ca67
50cb674924b28aedeba7bc9e85077dcc515adf2c
1888 F20110115_AABXSZ muench_a_Page_034.txt
bb2bac13fb013ef1c21c0b47a719e47a
072bdae0a116592ce92d193f52289d53ca52c3f8
F20110115_AABYQF muench_a_Page_107.tif
5768361e3e2ee6c7576e5ce44cf246e7
317963a1ecf32724b155f451ecda7cd66cdef25c
274082 F20110115_AABXUD muench_a_Page_041.jpg
3bb9442a4e00e702d1e46e5b5e416a52
4568e12997e03efd097a3ab27754782ad1996619
F20110115_AABYPR muench_a_Page_075.tif
3e7b55237533c4b6a6821b89e4fcd93a
3e804befb09aa9ebe70cca34871274b1342ec4e4
75304 F20110115_AABXTO muench_a_Page_019.QC.jpg
463935a3b77e947734d516d9c4f4b109
9f4ca0281d8e973ebb4dddbb7aaf6a65c771dd16
F20110115_AABYQG muench_a_Page_109.tif
1869be09712ed368cf59d2fa8ecf8e06
b48651d25b6f0f9e55351ec3fbcc15459bdf81c3
49187 F20110115_AABXUE muench_a_Page_018.pro
8c0d16224f8ced64341acc67834bee09
2242d0f11d48d1aa1173ec5dba01bcc87be478c8
F20110115_AABYPS muench_a_Page_079.tif
2ee0286f07d762048dcdd478d98049d0
e35366c1fcba2b2fed24d2e7541bd8b09ee79cfa
62868 F20110115_AABXTP muench_a_Page_110.pro
3763c14c07800ce2424b53e69c17a4a6
d5a621badebaaa68d5263ec06d36caf76922e144
F20110115_AABYQH muench_a_Page_115.tif
7c61e89bb47de8f3338ed2a548b8c6eb
0c29c62eec2bc2ecbfdd952faa8f11dcc4ef6f33
54005 F20110115_AABXUF muench_a_Page_066.QC.jpg
37a753a7d1f9ef35baae02cd6b06ed2f
121fe04fcacb61b145ed6537f839a5f4b5f3ce6d
F20110115_AABYPT muench_a_Page_081.tif
db5e385de32847fa9b8b14ed8059b060
cb61e8210d0ac11e449d3d6060e468bb51d5da0c
52964 F20110115_AABXTQ muench_a_Page_077.pro
2bce4a2422a37182679c26b3e3a99aaf
80c1723452a37285e13e849df0182908c078bebb
1200 F20110115_AABYQI muench_a_Page_002.pro
d42518a1f4ec59cf794431c29523ba8b
3c552077d8b4ec1c2b72b4b39c367d5f929d6b58
23185 F20110115_AABXUG muench_a_Page_034thm.jpg
1e227c00633f7ea260975520e5bffa80
cf35ffe4406bebcdf726a0c3a45cf6ef22487c9f
F20110115_AABYPU muench_a_Page_083.tif
5b4815ba8687d903883efe8b01035125
e7b76559adb3f5ba03b1aa88da06793a679b261d
2310 F20110115_AABXTR muench_a_Page_008.txt
5bbaf0b79735693e8370c5f8890cc39a
0a6a97f121fce26c1b52502010c7a8379f304b0e
72802 F20110115_AABYQJ muench_a_Page_005.pro
28e3ff164d041e03ee338aadf8a8606f
9f6c5d4847510137358abaad22e554cad39668fc
26194 F20110115_AABXUH muench_a_Page_076thm.jpg
8b38e89e2c01ba159fa71d65e45b8eef
4b675a6352d53a39c701990c1982b1e3d3acc241
F20110115_AABYPV muench_a_Page_085.tif
a0132b43d051687e3ee2603dfb6e26db
a7e506bce28631b448cc3a1359d718abc1a8aad6
207483 F20110115_AABXTS muench_a_Page_087.jpg
3a00896e0055c42b11a250c9804f4e01
70562824118b8b104ac8c508cede3664488c82e6
97739 F20110115_AABYQK muench_a_Page_006.pro
fc2598b8dac54df8dd5a715476224932
9cdd1a18a0c10f70cd23910b9e6cd447c999e6a1
F20110115_AABYPW muench_a_Page_090.tif
c46b9f3b22ecb67c218af9e9cbf478b1
8bfe2aec60f76bce7238ef113e2855c29b766098
91105 F20110115_AABXTT muench_a_Page_069.jpg
d15afefbe41ff4c2999542418a1cd333
d31ead6ac20d00693a8e2ae33d4c2f8020120f42
64721 F20110115_AABYQL muench_a_Page_007.pro
d01d5aa4b3c82524a4ddcb52eff53d51
99776581cab981fb03c7ef99653b7fd56b8b02e5
213581 F20110115_AABXUI muench_a_Page_086.jpg
0b6cfa5ef7b5761b8c31f24fdc8b72e2
7ce7e91c60427bfb729ef1fe333aa5d6f1850334
F20110115_AABYPX muench_a_Page_091.tif
42a40b8af6486b356dbfd74ec29bae74
e77167c771c792dff9dbd24e89912561a6ac0a34
16013 F20110115_AABXTU muench_a_Page_115.pro
29053e20769a34dfb2219a9769350e95
b9ace9cf9b5fe1c51a219aa6b8e63fd84d8982b3
17476 F20110115_AABYRA muench_a_Page_074.pro
69ee86deef7b2bb53c0b9ac668788150
c422ff1418e4e60f205dcf5e1b4a5ce3099faa54
50047 F20110115_AABYQM muench_a_Page_022.pro
5915026f2527cb7d98401e34d89a9b43
9bdc5511f88c9bd60c281d9be91ac56775ee1ab7
F20110115_AABXUJ muench_a_Page_036.tif
ee69982aec5ab150e6f720fd467a2cc3
936eb034d834912a0567c7732cc30ab399da8f74
F20110115_AABYPY muench_a_Page_092.tif
ccc6b9327eb401953c721fa9c9d7518c
89e03e8619aaf013627d48bf6a4d20152093863f
F20110115_AABXTV muench_a_Page_034.tif
adafb9d48e0119b6cabdeb123b6a1814
46439851e7a2fc7a4b49f536729bee241697d1dd
51557 F20110115_AABYRB muench_a_Page_083.pro
de0e911789ee373d8a86252d4225ec86
1b8f6375846250be27c0d7b9fc485a0baff86d45
52475 F20110115_AABYQN muench_a_Page_032.pro
8ccda1f1a8331d308aeb7edf21eca9c5
8346e7130a34213ccc7a246d16c9eec69b7a1e4f
177742 F20110115_AABXUK muench_a_Page_021.jpg
12ceeccd82a24dff63012c26f1db6580
c2ad21cb33c3d2bbd5bf847406f9938e440ad648
F20110115_AABYPZ muench_a_Page_095.tif
b13684460a3e0a031c3cffc4c25fa5f7
dd53fa8f85214c3c25709e26cf496af9f7a4bd40
2063 F20110115_AABXTW muench_a_Page_076.txt
41dc9f5a04de74871bd8aeadc5f4e98d
9b20fe93fdf1c63404f3578f4b83dffcfa9502c3
49531 F20110115_AABYRC muench_a_Page_087.pro
1399cb76caf1291346ad436a1ec6bbbe
812bbfafa4054bfff921a13b7934a2719e441110
50304 F20110115_AABYQO muench_a_Page_033.pro
13a726cca3a52d6bdb137e49b0dee94b
463bae71b8e0c74a7383a34278f82cd5cdfdcc73
51154 F20110115_AABXUL muench_a_Page_033thm.jpg
78b775f6517b401d9f8afd18dcffd131
73e39def9dc81d5a4cd4e0205599419858354aff
48561 F20110115_AABXTX muench_a_Page_050.pro
4e0e599a53157a492f0dc0b568ddbfc3
efbf942dcee6fbdca15fa3d49ff46592893cce50
52637 F20110115_AABYRD muench_a_Page_088.pro
20d3334e70c7c104bea7ab26b943b27a
e559dc102460907421bd0b2df29f38405400ce1f
114554 F20110115_AABXVA muench_a_Page_031.jp2
c4cac9efd489b8d0f3d5d4ea17c8f800
c8beeaa600b4e779e25b66700b811606e31bf267
47961 F20110115_AABYQP muench_a_Page_036.pro
774ea3ba5409470c4d95eb0c9917c03e
07658f7631ce4fea97b4baa8d83edbb2e6de2e87
25744 F20110115_AABXUM muench_a_Page_083thm.jpg
2757d98f1340345a32585a7c495b50ae
3d862390033aa21d5661746d78db9a161a8cbf94
2056 F20110115_AABXTY muench_a_Page_026.txt
5ca9ad59ad5de6749d03781b7dfb8f9a
1738b315730346e4b7fdb60dc656892f798f17a6
25508 F20110115_AABXVB muench_a_Page_112thm.jpg
ce0a7040d9108bf55ae9b83386b270b6
5b41b6e1a4987ca4ee98a304d0cdd8455c2ad2bf
50563 F20110115_AABYQQ muench_a_Page_043.pro
a32813f383a989adfe4332cc229a60fa
26f57eb623d6c3ee70bd39d8e7133b1356a06b06
22051 F20110115_AABXUN muench_a_Page_011thm.jpg
9314e82521f45e22c506619a93b4969c
63254fe2173f9596627cc5c39fa4b56394d81366
13209 F20110115_AABXTZ muench_a_Page_003.jp2
c3ae953921feb57960d8af9596c89962
8be02a4b46e5f714dba2acdede30a27da1770cb8
49817 F20110115_AABYRE muench_a_Page_089.pro
070a8ca34bd960e984bdf337a95223cf
31a43cf7813e69701750f5f06540bf134a8467c4
205237 F20110115_AABXVC muench_a_Page_085.jpg
d12035ea8fdc3b4ba8bb371d2ca13372
6e272c02ec8edd22b98b469dc7d4f14b1d513ec3
45133 F20110115_AABYQR muench_a_Page_045.pro
b44d7342bcde49d199cb40b35affef84
1bdf3917092e7b6a227679e5c7b9ff75c25127a5
1804 F20110115_AABXUO muench_a_Page_039.txt
17f00da12d1092bbc6fc8113c068e246
16706fdeb4f91d40dec2fa165cdace63fa44eb56
2881 F20110115_AABYRF muench_a_Page_091.pro
384de9c1c69459e37a6db266d5214f35
725037dc4fd9c8c62ac0945e5f16fd1cce6fcee5
110426 F20110115_AABXVD muench_a_Page_080.jp2
519a327a0a5bf436f17e07515b1dfe66
3abb89694b8d88f697edc1191e8e1c66208748b0
47094 F20110115_AABYQS muench_a_Page_052.pro
17e30cf8ceb9b7f998a5bcb8964f1825
1aeaa577718e4836e9d6a3b5b7c3dff5486273cb
1936 F20110115_AABXUP muench_a_Page_018.txt
56a0eace7a66a7ed4acf246fe720ed7a
d6483a56ef2979c5a3609f0506e094683632cbc5
20801 F20110115_AABYRG muench_a_Page_093.pro
2f5b54693ad60a242e944cd325834beb
2326413ab0d64521477d6e40d0294cdfafd17b68
8529 F20110115_AABXVE muench_a_Page_027thm.jpg
349916c22e49b5af6bf7533e43afa7dc
2aa2d3108bed0029845ddc6d249fce1949f4f97a
45487 F20110115_AABYQT muench_a_Page_054.pro
82f5b6da7f981c738ddab9fedf1796dd
9ea23d137d258ad1192e9661d015f8df1c389d14
111128 F20110115_AABXUQ muench_a_Page_057.jp2
1c2c30b1d1895b4ee3eec039cfa459eb
ca7c89f8edf7cfb5dd95de1bf681d168e5c73b3a
45080 F20110115_AABYRH muench_a_Page_101.pro
eb241977f50d184d37e6f98d9856a5f0
7be5556ee665343e056d1836a61624b4e87eadc1
12957 F20110115_AABXVF muench_a_Page_098.pro
73ebc9bd599ab42f7e80b360f044e8dd
9c21e3eaf8d3b2991a24440bc969f9a153ece119
2366 F20110115_AABYQU muench_a_Page_059.pro
46ee2682a567a305eded30c5c23d479c
05e4d522e344fbed82bc17ca736f598c021bb6bb
175 F20110115_AABXUR muench_a_Page_059.txt
fc27786f944d6620846efa7d2189e2da
6d6eba1f2ee5691e288bf5765915f7b54004753a
64075 F20110115_AABYRI muench_a_Page_112.pro
caf3ba723b833475f1ca5950f39d0e29
72d973f6a22ff85ad86db1fdf6d760eb51c3a2b5
51050 F20110115_AABXVG muench_a_Page_080.pro
cf331cc0c7ce69541da055c95acb6c23
76ab173bf1d34d4297860ca4dc10462a087940f1
19302 F20110115_AABYQV muench_a_Page_060.pro
164834747b0c541dc46a27f3097b8d5f
077a8711f798201d7e24b6b86f1377bb838ea29b
52310 F20110115_AABXUS muench_a_Page_026.pro
150dcaaf6ed5121924625cc893db2fb8
ebb3a47334399d9bcb54f10075d358b734db989d
25869 F20110115_AABYRJ muench_a_Page_116.pro
ecd6bcb3ff0542e0921f70c52be024d8
21f2e73cc570cbb99f38bf28f02f59a98429b33b
183268 F20110115_AABXVH muench_a_Page_099.jpg
614d22924f93476073c576cee0762287
f23090d73766c687528d8c51491622f68e3300a2
19904 F20110115_AABYQW muench_a_Page_062.pro
7d6e4b9624119297deb5acaa45f8440c
3ae205651772bce15ead6624c5332b9851254153
49473 F20110115_AABXUT muench_a_Page_017.pro
d566738430466a86e3deae5c448a182d
a1cdee9642709ac69369a1f77a52bd436088c51e
277 F20110115_AABYRK muench_a_Page_003.txt
cd7b36399a09e8cb0f3860a8c27154b5
82bf6badada70453af3c09b3ee541213e85f10cd
7761 F20110115_AABXVI muench_a_Page_001thm.jpg
02d8a9eb379adfd9067eeffd577ee1c7
eccdce81c4913fdf2209a1c57d7edb12e56657e3
14970 F20110115_AABXUU muench_a_Page_002.jpg
95027164a85246d97018c5d9df858510
f5a25aeb2b56b9327e48307ed85643926af4ed25
2041 F20110115_AABYSA muench_a_Page_051.txt
909a04d18cddc09943d7b57d2dec29a8
f7cc2cd5825f2325cc8013d7f26f34cef34b32a6
2618 F20110115_AABYRL muench_a_Page_007.txt
efdfe952eebc97fddf6edc20742415e9
f85c1a4af56c512aab5b1dd259a0a0401cca1de5
18521 F20110115_AABYQX muench_a_Page_064.pro
96bac3bf6acf3950b8265c506e09c14b
b6d94a5d43ccfec3d6b3d4c4d5ad1e50194d3c8c
22548 F20110115_AABXUV muench_a_Page_100thm.jpg
b92fcb0f221ae9022d2b8149fab7919a
6669c911005fc8e4a4e0056468f3ec5339c54dd1
2008 F20110115_AABYSB muench_a_Page_062.txt
f73d1fcf2238f0497b6130d0ef1cbe18
c6b4b92a8c01adf88d3c19e209c7d293a04ceccd
1716 F20110115_AABYRM muench_a_Page_021.txt
75dbf9672c55515975e01da6edffdd08
8881aebd3ff26430df8380ab9cd4daebba5d560d
103150 F20110115_AABXVJ muench_a_Page_034.jp2
7c099e11e50e94b471ee19eb59727a54
53a3b7a3798c5a7c8b10ac24dd294e3cfaa8de50
17911 F20110115_AABYQY muench_a_Page_065.pro
eabc59cf9278560bd2ad267fc7f18b2e
970262dff2dd6c7b8bba5fa0651ddf379e534f97
F20110115_AABXUW muench_a_Page_093.tif
9e7cde68cce2e0e6266b5f6a0f67ad3e
a4954ec10b8ec4e894fb1241c126188416e11323
1657 F20110115_AABYSC muench_a_Page_064.txt
cc15774b5ecad4a5994b0516bdcc0cfc
a5dee79415fb39291b1c303c60e66d58e7f8bcd5
2025 F20110115_AABYRN muench_a_Page_023.txt
b65c31c67054a8bbecc72a99e4e47c3a
5474f85318094295d6b4c55c3189407c0c9028e7
1806 F20110115_AABXVK muench_a_Page_100.txt
c9ac71a751b2d5a66db45de9e6b68ac7
1c7ae8aadc55498a55e49abbb6b41aa92a2c7d82
12570 F20110115_AABYQZ muench_a_Page_069.pro
f9dc1fca817459dc625b68cb53033bd6
440c12d3751968f763bd9324e9ce436df4ec034d
F20110115_AABXUX muench_a_Page_050.tif
4c8834c6aa007e6800b8532958bcf2be
7a76251d45b852e59aa425f40169a3cd87467a37
792 F20110115_AABYSD muench_a_Page_066.txt
87e23bf8f2f0f11f08bcc439a91e0cd8
9db8d3abf09532d9dfc9b4dfc9c8bc929e0877a9
278933 F20110115_AABXWA muench_a_Page_008.jpg
d3d0fbcb762281bf677afb6452219b4c
23f80de830c8e099f89f27f7840918fde95cca85
1856 F20110115_AABYRO muench_a_Page_025.txt
fcf9d9c71c567dd65293b8849c4929b9
43cb5ae710c3ff408a10bfbddc6fb8e6739207b4
107138 F20110115_AABXVL muench_a_Page_105.jp2
3c8c5b7a3f6f80f00cc2b8f013b873ed
e6ea901e8ef19dfd01dcfc7eb0c99cac53ff5a5a
114923 F20110115_AABXUY muench_a_Page_108.jp2
13e2fd5a05736ecfdd11214e8c71b805
99c5e4d7215d99f2c2af72852a208e97f4cf5c85
980 F20110115_AABYSE muench_a_Page_069.txt
1c224754ce86a72cc269d19b703dce34
e2f04abde66257802972778a2b48fa9d2b8df919
20536 F20110115_AABXWB muench_a_Page_099thm.jpg
66020b3512ddcbaee42033bb10445dae
b6c31a5eba7be4da08812d91a90f1fcb244bdfd9
603 F20110115_AABYRP muench_a_Page_027.txt
a7f06a11a4e35a1f0afef74cfe2e00a7
bdc9b0dab406c798aeeadff4e529b51f82a52c44
241587 F20110115_AABXVM muench_a_Page_038.jpg
8b22201fbbcfc8bcc0084c1f37db9130
76f45f666f3b4634648909f241c2f1c7846f1c3a
F20110115_AABXUZ muench_a_Page_050.jp2
44ef4f6ada73dfa5f2a6a5435697b9a3
396be18cd9d5556fb7a38351b422d5060939301c
15935 F20110115_AABXWC muench_a_Page_060thm.jpg
0c1cd1e823b7568a23595fc3bee3b8d1
edb9acd5ea1982d13b488e0d8d8189b8defc3484
1708 F20110115_AABYRQ muench_a_Page_028.txt
6f6663da233acb0b2d80543c5d53e085
968ba817a084ddd7a943b754e1e3b6846e555735
57709 F20110115_AABXVN muench_a_Page_092.QC.jpg
c5290e7422d5c6477054bd16a9bcf0c0
d6ea298db3c34508be3bf499111a6919f33cc55e
1169 F20110115_AABYSF muench_a_Page_073.txt
d7f02e44432446ee995933f272750b0d
83c18d3522e6657e78af6fcbc0f2d9fdb0903644
79339 F20110115_AABXWD muench_a_Page_055.QC.jpg
ad0409ca2bbf8e7b12e5b58505b0eb5a
ba6358b625232ec762c61c8cc0e2430d5fa69cee
2037 F20110115_AABYRR muench_a_Page_029.txt
b92a7606bf8eefacb772993e92bd91b0
56f7729d5b2b51b7baf4344953f118f109a6bcc4
108711 F20110115_AABXVO muench_a_Page_084.jp2
4a88f5e5ac8f20b73a25b413542110e4
a43fe7dc29e16ca4e53a1d301635f090b25843b8
1959 F20110115_AABYSG muench_a_Page_078.txt
851ccdc3775fbb2f55ce2d7de2a5bba6
928163c1e93ef9e3b63bdddc97e7bdc8a2c9d78f
16277 F20110115_AABXWE muench_a_Page_061thm.jpg
38925bf4f103d29cd923a51316878f6e
e00afed02bf676cceab3355768e2b1bc7b94e7ba
2062 F20110115_AABYRS muench_a_Page_031.txt
3e801d6c8e98556ae36934948b10d527
26976e255ffa2f1d24ca95fed869d94ed476a81a
620 F20110115_AABXVP muench_a_Page_068.txt
72fc930fb76e1789c7c59990a96e1a5b
22aef6603c38fd5fa73bf679ba0413ee4c9b061a
2052 F20110115_AABYSH muench_a_Page_081.txt
96e06b75f8e35893c814a5684a36acf7
e80416ec92b0958292f62ed36b215386d595c097
303697 F20110115_AABXWF muench_a_Page_071.jp2
5e6897c1322d1dd37a136caba8bf96be
1b7ab2801498d8c294803b3def4cb6e523a0aaad
1975 F20110115_AABYRT muench_a_Page_033.txt
1e84537351ad85ee255324ecec39d7f5
3769b7651a95b67d09d1bd58657f5ef05885d9d5
F20110115_AABXVQ muench_a_Page_086.tif
5c1e37a02816c3ea949d35f04d481d3e
65f07fd7946fc9defe6656f5276fab1813febeab
2921 F20110115_AABYSI muench_a_Page_082.txt
7b5cdfc2e0f76b92b3efc4a94bf10359
87ce7bbd1e974913cefa3d68b05976030ef9c949
F20110115_AABXWG muench_a_Page_068.tif
6003f62b9396cbca58bb32a43580b553
b93b3e4b5cc5618357903796ec1f4b1e3d8c692b
1891 F20110115_AABYRU muench_a_Page_036.txt
cbbe73d0e3bf1d608dc9616a69918aff
3a5410a7b56eb65d051e84bfa47d3fce70ec7877
115014 F20110115_AABXVR muench_a_Page_043.jp2
865f1eb790b44a4f8244e2e02a21d1cb
ddfb02992e365c1112950fa23fb147e3fd02eb5c
2001 F20110115_AABYSJ muench_a_Page_086.txt
7749ccacdc799c36496f9cda420631f9
908779021438db6325514b386af0c9d5cd1a9607
48962 F20110115_AABXWH muench_a_Page_019.pro
99caeb262272f6abe0969f67fb171d98
16f20b967b507c3a3044f32927fb9cc6d4aadde5
1955 F20110115_AABYRV muench_a_Page_037.txt
1e996249ebf41334c7f3a1ee2e123abd
464c9ba6adde46d7d734aa9edf56bb3dcb3a982a
63065 F20110115_AABXVS muench_a_Page_028.QC.jpg
619ef38cd2726bca6f6a3253cf584c5c
3a36abb8c2350e7815940e96c02c5d241ca7c638
1951 F20110115_AABYSK muench_a_Page_087.txt
a8af1aecdcb7943463111aac8bb5bdfb
0f93e6194c0d090350648571088819ef1a52042a
51273 F20110115_AABXWI muench_a_Page_103.pro
dc7a71b54c55210b8cd8b583357675a3
cf71dfc11861770e546185e8899242ff0cf1f277
F20110115_AABYRW muench_a_Page_038.txt
f009aafb51e3a994fe1adf54c63afdee
fbd414e86f2583de7ba27eb5e1fbd349a983000f
115784 F20110115_AABXVT muench_a_Page_007.QC.jpg
6d7091222ee8c107747a582018176e46
eeea0ef62ff569f63bb2f046f787e187fb9c24f2
1608 F20110115_AABYSL muench_a_Page_090.txt
7ae5ae1779c46cac14a3703b2d428c7b
190d67db3f2dbecfab50cb155b92cdd8f4158f1e
31327 F20110115_AABXWJ muench_a_Page_061.QC.jpg
491ca36632bb5161421af8b2958757d9
88b9a61a2245de555153693a800ac892fc6bed3e
1993 F20110115_AABYRX muench_a_Page_043.txt
accee53b03c1ba1d84246cc5c543ca71
73514bff2d3a101876c71a4bee9d1fd74d275eb3
F20110115_AABXVU muench_a_Page_030.tif
5514cbf6fd06855c4684fcc0f7dd4bf9
5c2c1a8cbb539425284d57c017cbc36cf78589e1
77132 F20110115_AABYTA muench_a_Page_014.QC.jpg
c2a085f197cffae4377e1a47c5ec5ad1
7575855fb0d4d679446d9443c4890620ebb3443a
200 F20110115_AABYSM muench_a_Page_091.txt
f0669d9d3d3dc772a4fdb0ecd360a48d
1c3f71ac693d954c979ea100525241bc850180ab
2372 F20110115_AABYRY muench_a_Page_049.txt
b591af1d9385290ffb64c652e1224c07
bf4c7f04462f4e1354389615e74bac82eba73758
12563 F20110115_AABXVV muench_a_Page_092.pro
cd30a85516efec6c08cfd7d52361d61a
bf1eacc0b4f36617afa9c1eefedb796746e7e6e0
25577 F20110115_AABYTB muench_a_Page_015thm.jpg
be55aecbd98a59479830315db4717a34
9ef922e55858b3f9bf8c3d6df7675534b9628f75
651 F20110115_AABYSN muench_a_Page_092.txt
c357c82901edefc2a6ebe2df42cc5c3f
ebf0f7a101d03cac60f4ea31f5dc57aa3360c822
51386 F20110115_AABXWK muench_a_Page_038.pro
4d7f1a7ac2dd8dc44d3e229cb54a328e
6b359e61f0c69f9cbb1458881f9ba4badb9f6048
2153 F20110115_AABYRZ muench_a_Page_050.txt
2ef1c5640e642263e91977324e459048
f1717ad633a7b31fea78643094e7b0d10a8ee4ee
24647 F20110115_AABXVW muench_a_Page_086thm.jpg
2e1259b0c735357168b780a4a25efb10
fbaa562d8cdafc34fe154f9b0f31da3b03438aea
78585 F20110115_AABYTC muench_a_Page_015.QC.jpg
182dc2da76335b2569a27bbecfae66e3
e87bb632c7582fd5a2db5f45f34cc47e83ff29b5
F20110115_AABXXA muench_a_Page_006.tif
2ed9fda24c6d415c6b28610823d6f081
7bceb2570f769b79c8aaf8db19b99dfcf413ed3e
685 F20110115_AABYSO muench_a_Page_094.txt
bec26ec0617b1441801228d2e82e13de
a480a2f14718602ab1f16b70c778d264c1d4b863
48453 F20110115_AABXWL muench_a_Page_014.pro
e67306f1e3423f678c64b7cf4ca3a133
380c49d04843f697384bac3f6f41bc7bade88615
253861 F20110115_AABXVX muench_a_Page_082.jpg
3224a399ad5300f9cb15ac22bc294eab
1eb5661a31c0c32b742e1039782975135ce2e78b
78573 F20110115_AABYTD muench_a_Page_016.QC.jpg
27849a62223021bf9bb5a7d53daef96d
6f963a114fb39077d1c82809993fa089d4174ef2
11206 F20110115_AABXXB muench_a_Page_096.pro
39552695264f5dd11b92253b6b6dcb61
7e19cc0a4eee24d366f53681df33dbdd24bafedc
577 F20110115_AABYSP muench_a_Page_096.txt
f0f24ad274db70196a72a136c09862e8
855cf704d112243c1e5f9c72d2cbf8e5479c38aa
162929 F20110115_AABXWM muench_a_Page_042.jpg
1272693b52b976da266bac85a92f1e57
50db997b1b7f6785df71b94b08f553e23738e51a
588881 F20110115_AABXVY muench_a_Page_093.jp2
cebf154f4b72139786f58632a98098d1
aade55e0b2014a402ca5656fb11e709e466c61fb
79416 F20110115_AABYTE muench_a_Page_017.QC.jpg
a3d134b4a6d888598bf91a0c2dfa8c93
61a934d3bbcaf6831478af270471c03e71133b08
114261 F20110115_AABXXC muench_a_Page_032.jp2
79e2fcdefb5726e34532c4286bfdae74
f3f011aadf1e62c36d3c20fc4cf30ef1b445fe18
516 F20110115_AABYSQ muench_a_Page_097.txt
867d99ed67bfd3f4fbbcba58181badbd
d5697ee2c6ded333145f73aa0a526df0d28be0e6
F20110115_AABXWN muench_a_Page_057.tif
a3d144790109defbf98584677e96058b
a97db004afa6982c74bf9bd20bf6f1908baa726f
326562 F20110115_AABXVZ muench_a_Page_006.jpg
4613d3358775df98920269db885c4089
3984a150c4355530aa88f84782b6606b8bd825ef
24737 F20110115_AABYTF muench_a_Page_018thm.jpg
4a8e3194d32638519f960ce0a24904b1
69a93b174d2aae7a6f6748eb350befac20e23fb5
84821 F20110115_AABXXD muench_a_Page_095.jpg
e9b73cdb00b227cd3e2e66579f5547e3
45a6bac32a08deb6d529274b222c147aae837e76
1940 F20110115_AABYSR muench_a_Page_105.txt
61d92a22fd7ec18b73c5861601eaec18
c2891e15b02e5d72fae98da82eb9336bd84286bc
70654 F20110115_AABXWO muench_a_Page_100.QC.jpg
cd4bc1e7d5b95f66a35f8a9cb8b269de
e78ee4c0400d1c8b8831daf1f7e73973c028de98
134106 F20110115_AABXXE muench_a_Page_110.jp2
c1f435163c819dc8e1424a79927e8920
9798a26ac43ca15e285e1064c885d9a9cb9dd5fc
2533 F20110115_AABYSS muench_a_Page_113.txt
5d61bb066799b96dc564778c9b1f52be
81fb96186ccd4db080bbcb1f567161df6c635963
52458 F20110115_AABXWP muench_a_Page_076.pro
48db1a3c7081f7c1bf661b7b95b6bf8d
a012a321eb55c915e904576bc957fa87d7af5348
23583 F20110115_AABYTG muench_a_Page_019thm.jpg
c787febca7ce05bdb0a3e54c53f72eac
95c4bead7e617f9279c022bc1cc10ec006a5dc05
33968 F20110115_AABXXF muench_a_Page_071thm.jpg
f3da8add4b8bed5c5329e8b775edad14
109eec1cd821ae23242e8c0eb8d5eb5bdea3212d
1075 F20110115_AABYST muench_a_Page_116.txt
e0a76ed2714e858a2486b0acba20d84e
7a7f4515ac48d647452a90bf6f406ca6c93aef6e
22998 F20110115_AABXWQ muench_a_Page_101thm.jpg
0c7791a71e424ed1db9ddadf07b4930f
d11f48ac0428973cf20876a5785aadda04539bca
20468 F20110115_AABYTH muench_a_Page_021thm.jpg
9afdc75c3fdb62fadde485e2eab19a0b
4a1e2aed629c864852e2b40914fc8f1b85fb199c
82200 F20110115_AABXXG muench_a_Page_023.QC.jpg
f985a82faf21a3b33f9825e280dd937d
a5fa4ead81cf6cd1909cf4cfd37cd1b7aa062b7e
3193 F20110115_AABYSU muench_a_Page_002thm.jpg
192e463c6bc4381dffe2fb6aa40838c5
919c4f9de891108b7675a8499012dcfa421a1f12
743 F20110115_AABXWR muench_a_Page_063.txt
c15b06bdca01e130bc051e2fb2e6fce0
831bdaa2d792015f86d40296643e0f2f9086433a
64732 F20110115_AABYTI muench_a_Page_021.QC.jpg
30bc6ae0c0c4d5da494f50ed2e7cb782
0ae0c5dab4cf1954a19258fea94a68a5b10645dc
67761 F20110115_AABXXH muench_a_Page_099.QC.jpg
da531321c91f00457f48d731422fa66d
bf22b755eb7e442ecc2810604da0ae983607b312
4665 F20110115_AABYSV muench_a_Page_003thm.jpg
cb8348b96257d3311d9b7ab230fe5b8d
15a8461f928515a2ce2f4ef7262d70ef05dadd55
25805 F20110115_AABXWS muench_a_Page_042thm.jpg
8d3112287c31df05ffbfebda43e00924
bc4ea42bd13c598f865f8d3e7dca100e3b0019ba
81203 F20110115_AABYTJ muench_a_Page_022.QC.jpg
0900500fc00ab65b62e4c3cc61e0ce0d
fad23ffdf7ff21666470feb3205a9fceb7310766
315125 F20110115_AABXXI muench_a_Page_097.jp2
bf2330e425e8a5a84377552a9ee24304
98b24200b483e81b8e849a2f5b74eec9572a3f0f
21546 F20110115_AABYSW muench_a_Page_004thm.jpg
83ff56f773a95f8807fde4e589f6df2f
7b344c870dbda618621c44464d6bbcb4035b6638
F20110115_AABXWT muench_a_Page_094.tif
1dff6ce10fe3ce56c3d5406bf8c0dabb
ac11cd896c47b613a8410395f3a752d5d49f36ab
76045 F20110115_AABYTK muench_a_Page_024.QC.jpg
5310401ce809eda321546f04f6b13c9e
b9a69dce3edba1078f0e4dd39a618cd7487808be
20102 F20110115_AABYUA muench_a_Page_075thm.jpg
77ee5090ea43d9cdfeb951038a635ce7
953ed24655316e4b261558230d62fae100338905
53347 F20110115_AABXXJ muench_a_Page_056.pro
be58bb90ad205f9fc4232875217514f1
515784327d12e1dc893a77c5834b8df984dae511
51039 F20110115_AABYSX muench_a_Page_008thm.jpg
c857475db51780ff498a42df40676b61
307751e70a66fbdd2e82f280b7188e8dc151ff58
50588 F20110115_AABXWU muench_a_Page_040.pro
320af0a6c6c7784e1dc016be5f516beb
68937b4f4bb94f216b1c32415443e2409034392a
25284 F20110115_AABYTL muench_a_Page_030thm.jpg
7854b918765ea47e5abf88760b46bfa3
d78e797e65628ebab2a59a25458d15cb699d6798
50950 F20110115_AABYUB muench_a_Page_079thm.jpg
c38a98ca4b7fc28def01d4e57c63c731
e998d333efa49f86268107bfdf7e140581b97787
35030 F20110115_AABXXK muench_a_Page_065thm.jpg
43141988776cac80ded0d820e076d484
f917585ad3c750d7019b62b96e6758dbd61ecacc
46618 F20110115_AABYSY muench_a_Page_009thm.jpg
7d4d2a158d5fbd5358ed89f33c554c49
cff122a29fac6dd3b33e5092a889a5cf3fa7cf32
1908 F20110115_AABXWV muench_a_Page_014.txt
8a2694b7ea5f72be6c1164dacc57aa67
b7fddf76d2b6ff814f6e3a730765e2540ad61e78
82677 F20110115_AABYTM muench_a_Page_032.QC.jpg
40fc5e410cf6c4aba2fa8cedc87f2246
ccd64921fe20404e6beef39aa7553291d3067c69
79077 F20110115_AABYUC muench_a_Page_080.QC.jpg
f59f282815519397c7ba2c24e99918fb
7814900a94cedf30a616ec2fde49d847c4f2c6e2
71135 F20110115_AABYSZ muench_a_Page_011.QC.jpg
e6230192cd8faa10d9b5238fa93b4bcc
e50708de4043c394d6e4ac8c8859b00b3997b1d4
114006 F20110115_AABXWW muench_a_Page_056.jp2
0013ef3a80bc947c064bd896a5fc9cdb
ea6093180ad984158ebfc960bb6a728f91cc2fa6
75192 F20110115_AABYTN muench_a_Page_036.QC.jpg
d18322145db9bac0736509e4a1bba46a
1ad46ac09c5ae15947b811d8d8f162c26f44563a
91298 F20110115_AABYUD muench_a_Page_082.QC.jpg
d7531932dbb16aab3c4a5d8d12f33526
baa993e0c715dcad0ffffb4a0bf06d28c7fe61cc
F20110115_AABXXL muench_a_Page_022.tif
9a5bab0ffb8f09717ac3176523b6be6c
f34b65c682c42b58078c3cbaa52d7a0197e04233
105794 F20110115_AABXWX muench_a_Page_014.jp2
9e6e534538f080d4a39c8170b1005d7c
ed0821883b5cb02d74ca46647c8221e5f66dac71
99030 F20110115_AABXYA muench_a_Page_048.QC.jpg
b079cabca942396ba45f1a850825faf5
2fd3f89194e5a4fa04dc7c2d8a2a27ab45a00f6a
78726 F20110115_AABYTO muench_a_Page_037.QC.jpg
8f079ae925af3d7d14b5c4edf7ccb6ad
f26ca4b47af515702af7e8225dc71d479e391758
82175 F20110115_AABYUE muench_a_Page_083.QC.jpg
9ac7a2e3d560c33bffd336cdfba823ff
b0c88a409bf17e9ef4579f36613cf1ba25f0c462
2029 F20110115_AABXXM muench_a_Page_030.txt
455a0a4ba364aa68a4ac7977daab0a5e
c7172e72ab66d0e61f18871e94b6ba4f830f02fa
68440 F20110115_AABXWY muench_a_Page_041.pro
859972e8d01948159c83d9b495e3c5a6
f2b5c1af504b2e39f577ad3fa466047f8fed88e3
F20110115_AABXYB muench_a_Page_087.tif
c4f16294f94a5dc99dd0f54f8f090728
831f07b16ab96d41773e132d6b5fd4870c689652
72035 F20110115_AABYTP muench_a_Page_039.QC.jpg
83e565f80a6ef4ab30d3815e0e8113b4
ba436468c0c202f80d982cdd999e5199d25679f9
79943 F20110115_AABYUF muench_a_Page_086.QC.jpg
65c4a1241a01766eba0ee86fe23c7916
b7c9f632fb39d6a638a38fd025d6e676593e53f1
24326 F20110115_AABYTQ muench_a_Page_040thm.jpg
b7de14c7261d8d613d14589924b9540d
873733016de889929dce671c04be65c59223a668
63404 F20110115_AABXXN muench_a_Page_060.jp2
733591a438d437c8fc72c59976231780
cff3de9c6a758f411f0de9f3428576753a1217ce
1961 F20110115_AABXWZ muench_a_Page_089.txt
244c1b45faff8bf37712680af780569b
f789282a9a17ec1881365299b37f6fdb05b6ad58
23851 F20110115_AABXYC muench_a_Page_052thm.jpg
be7004a09d84c0cea61ee9d0592f4fc0
5407113e9d95684157c83f08e7d593773b6937b3
77950 F20110115_AABYUG muench_a_Page_089.QC.jpg
a84e1ede9d1f7441b5ad2e6bfa38024d
9856174f539aaf171539da85dcc1cc5fd0620e05
80906 F20110115_AABYTR muench_a_Page_040.QC.jpg
42fb4cc949f9eee5e459c59612d9d9cc
b308a6291916b4fef0ae99f87f1e1514bb66546f
203443 F20110115_AABXXO muench_a_Page_036.jpg
ead1bbf2114690f4efb366919a287b0f
3b8582217f89b8f3ce779dfac19ef858872373e3
20195 F20110115_AABXYD muench_a_Page_107thm.jpg
613d3f14d7994ec6c2610ae835cb9f47
74d74a6118faf9420a4d81fb3e58918960203e12
107064 F20110115_AABYTS muench_a_Page_046.QC.jpg
a6fb6cd6d3ef4e5a9948929ad68e1a8e
da7d1941a6599609e27b8acbacc252302d88b2a7
2064 F20110115_AABXXP muench_a_Page_088.txt
da64d7484c92e0b1e19a27134085a897
55dcf2a7870ca55f37575ecd8c5c8cd15c1926c5
39259 F20110115_AABXYE muench_a_Page_075.pro
6b090d32d99933c15819714ede434512
214507b85fdfd12c2f87667f48f644388dd0d090
92227 F20110115_AABYUH muench_a_Page_091.QC.jpg
e8c48523d55ec0dbca93e070bb9dbebf
eef475e74c348b562f5aadb0d548a2d7b2bd040a
50870 F20110115_AABYTT muench_a_Page_049thm.jpg
f812c09cfeb90a25d9aea61dc3e686b4
cf7136da3c7c5b203f20d5a5e06514c4248faa75
103337 F20110115_AABXXQ muench_a_Page_025.jp2
a10281b45af2f7da43129ab7d99f9aef
37b7d94895351456e7697cee69de992e56a552cb
192345 F20110115_AABXYF muench_a_Page_039.jpg
3f4643335dc21ec3f5e72a0ba492f1eb
0686df98928b49c94b03dbb438ef437588704745
38980 F20110115_AABYUI muench_a_Page_093thm.jpg
5767136761d0a7b165c5567f99294ac7
c4014f15fbe05b36ac7eac8698f534245cd2a8d7
22868 F20110115_AABYTU muench_a_Page_053thm.jpg
c91979f5cc0e8e8d9894a4ac8a5a885b
b75f17f6f4b254ed1eb2616d9566dd7d046950af
2009 F20110115_AABXYG muench_a_Page_080.txt
0942527d6a3c717ef95522cefb327221
81dd590fdf0954a1c1247156da6ac3eeed10e82d
107196 F20110115_AABXXR muench_a_Page_036.jp2
38389ec740eb958d99f937379beb3263
d91338a86dc134299636bd51fc365acaa76bed07
63639 F20110115_AABYUJ muench_a_Page_093.QC.jpg
ebddb1375afae1c54c4293f97cb9b34f
2b23583642c42519881273ed060cd1a87fd120b9
25653 F20110115_AABYTV muench_a_Page_057thm.jpg
324e5243d23234e6466b3d02f899cf90
47a4be93e277e8045237f91acf8d7abf64267cd2
956 F20110115_AABXYH muench_a_Page_065.txt
31c6f2d4d5d55007876e0009a23c7893
c6b0d31bc449a215668ee81ae14d7461961a643c
57545 F20110115_AABXXS muench_a_Page_008.pro
0ca0915d4a7b0b56e0c7d41f55464830
b98b1492b1eafefb41cd47f1c0bde9256d26f45e
F20110115_AABYUK muench_a_Page_095thm.jpg
1bf6a64352b3dc54043a55788b104455
94fb438a3b18c2e351af807458262ad0466ac34c
46303 F20110115_AABYTW muench_a_Page_060.QC.jpg
9029ec8dbebbd21df9c59df75457499c
af6d9333fa4a8b1c4cf372b796b81954a415b37c
47335 F20110115_AABXYI muench_a_Page_059thm.jpg
2651e25313da9e9c1ec137d7170cb944
67498a72365b58ce8f9fa48483460dc40d20b64f
29003 F20110115_AABXXT muench_a_Page_072.QC.jpg
ed669f98b493649b92f070efc9481915
8f4e7e1a291b780a9327ef95cac7553a68953ee9
36979 F20110115_AABYUL muench_a_Page_095.QC.jpg
4f43a69d99cb3b3f9d8ffb22079a6970
f88d4f4b7671b111263a714f8303f7a9c09fc66d
52324 F20110115_AABYTX muench_a_Page_062.QC.jpg
fc84714ebbab616ee14a1fde40db27d7
a85be73c446b399f4a9e6da3451ff1086eecb2de
254125 F20110115_AABXYJ muench_a_Page_113.jpg
2e5d88a898c41f237ea0179a12871f70
2aa9ddf45de4687563d8f6fea5b2268764221654
87730 F20110115_AABXXU muench_a_Page_096.jpg
ada2548c288f55483012916573cfa4bf
09e0d575ced7728a587905452233493dc98b7ad3
19987 F20110115_AABYUM muench_a_Page_096thm.jpg
79e8981a03c740761817e591776bd32e
905a13bfe686c0912d04d53ebcbc4df96af1328c
35651 F20110115_AABYTY muench_a_Page_064thm.jpg
42869be4597ef15197712b6bbfdc4756
f0694925e74410ac9abb5ec2810b929adf7855e7
3062 F20110115_AABXYK muench_a_Page_005.txt
9ec683c5ba5cd1a0248ae31b20a3dd41
80459f71db97f6b2a358e154d79d8301b8068386
83239 F20110115_AABXXV muench_a_Page_112.QC.jpg
74ab8bdb7e020abc54dac4083aba435b
3db9065379602ee43d0cf9f8395ec66562894c26
37273 F20110115_AABYUN muench_a_Page_097.QC.jpg
4b5542e4abed970e5b0953338c7ba8cb
175803a41c1979c5ec26299e0b00ac5ec9a6d788
33959 F20110115_AABYTZ muench_a_Page_069thm.jpg
9480ebce85164b8bcf90da826ec7611d
e698cba7b1edca0da1810cd52dc7dd8314cb42e6
697 F20110115_AABXYL muench_a_Page_115.txt
f4334b766c90a95699b8b5354c9d73e2
c802bec170668a0ea79428b65c2a80f293d0ed10
57242 F20110115_AABXXW muench_a_Page_070.QC.jpg
1c740ff0d0af758af2b49f5a44ddf66d
7573cd89d878725eefa6c81592ec43f327cd431f
19153 F20110115_AABYUO muench_a_Page_098thm.jpg
875228932d075ab37808f4251f5d3757
d999ef07ef355e53e66c83ed32d1a7a93af33d7e
F20110115_AABXZA muench_a_Page_038.tif
2a591308b0881b98bd4ab4779dd42a7a
bba6555cc5c13b9cc80c615f033de652792c7a3e
25000 F20110115_AABXXX muench_a_Page_104thm.jpg
9ac2d3c55b7b87abb819173fcecf4c14
01f2fd625b0ee2542b31797b8c8a0fa1916d189f
77032 F20110115_AABYUP muench_a_Page_102.QC.jpg
4f0c898dc1e7799638cb59006ba42b16
c47a4b3d17e8a4b3ad8c718f875edb39115f6cd9
58503 F20110115_AABXZB muench_a_Page_010.QC.jpg
16bc6378af8d627a4d2524a9bc1f0e3f
11627672449391489ec2b03f0f7ba17e4a2f7d26
114566 F20110115_AABXYM muench_a_Page_077.jp2
34bf4a72101bc3103510d99bfd607fcc
2d3fdebd437be3eaab99bf4c46b1fdc2b7ba18ed
10241 F20110115_AABXXY muench_a_Page_095.pro
5992ee718adccc3093420cbca1fbc86f
db6d376be1acb83f155dbffc3194df897cfebd5d
24692 F20110115_AABYUQ muench_a_Page_105thm.jpg
96eee7368e6fd0c5d03fe5712fbc890a
782f80e968d09691d32954db9ba73b3051d25ed1
73632 F20110115_AABXZC muench_a_Page_108.QC.jpg
a7e28516ff42883c3d1b24c14b12c750
6ac6bff77e21e75d45f38eaaa992a2962d86cf55
12625 F20110115_AABXYN muench_a_Page_067.pro
873fd71584fc2d395113857ab0cf112e
5962d01531090c74a2fccdfcd57ff3e7e66c09cd
F20110115_AABXXZ muench_a_Page_114.tif
87571e69f21a02eb2daa4db7b1b39060
a2c5e7c7834006156bbdace5859d981691ca6a5f
64405 F20110115_AABYUR muench_a_Page_107.QC.jpg
834ad767cab662220466e5cb489b5044
4ddbef92e4e4a3323b79c178b4c59cb03177e383
60655 F20110115_AABXZD muench_a_Page_075.QC.jpg
7906c44d273a360b46515e7cd8e0d72c
2f9a6879842f624aa682110045aca4194f2938eb
49282 F20110115_AABXYO muench_a_Page_044thm.jpg
f95e7358ef6dec8604c39de425368e43
e181272ccc9ba289a2c2ec39d9fdda7aca9c3408
108232 F20110115_AABYUS muench_a_Page_111.QC.jpg
0ed7a8df95b709ac282e033db783ca24
acc36dbb469f1127fedb26b01dd4e9897664a929
37686 F20110115_AABXZE muench_a_Page_096.QC.jpg
c596a2915d814c668efd85885c46cf59
0d22e031abe9ee56379eb0329ac9fad751e99abc
4762 F20110115_AABXYP muench_a_Page_003.pro
cd045c2c60f3b46f5c9811604f74ab85
87049ae4582de486345eb644abc565083a981f88
44298 F20110115_AABYUT muench_a_Page_116.QC.jpg
1a35e742636ec6f577f352c3b9eb6144
ef3acb054a76c6edf7e5ec7c7a432721c2ee0a58
37951 F20110115_AABXZF muench_a_Page_010.pro
2822c61808dc4fa40917fda98af92778
c73eec14ee0d83dc5a9882e35b0174a2c5a4c5fa
25208 F20110115_AABXYQ muench_a_Page_026thm.jpg
1944042b384909ea2e22f63105faf46d
2dd263393c08f71f8bb7cdcf153499ef778e9329
134793 F20110115_AABYUU UFE0008580_00001.mets
6ce82ccddd262b52bbab3953e19442cc
fac6c2bfa9ccfcdb5d95db1cadbfcf79733634ca
51080 F20110115_AABXZG muench_a_Page_046thm.jpg
f70624e7b3077fa74fcc70703dabbc71
102d8590b296a9196ef7b86ada7d927ba0f7eeec
1750 F20110115_AABXYR muench_a_Page_099.txt
b5df3ad1c3b28e681d934ba3e960656b
6f5acc9b58e40fd8fd06c6c3c715ee49eba0fb15
1051935 F20110115_AABXZH muench_a_Page_059.jp2
fee7b824a05d2259c410d1c554655d96
421b6bb37ced2a279ce7069fe6d78aff21765fa2
25017 F20110115_AABXYS muench_a_Page_014thm.jpg
155345b33912824b2184e285d09cd013
2ce9c822c70e80057c26c24cf4572ec691e03f94
1051944 F20110115_AABXZI muench_a_Page_091.jp2
595f9e9f221043cb62d4c1a6f2c6c47a
cf21962044e69ff20bd9c8d3b403eb0855732f1c
51418 F20110115_AABXYT muench_a_Page_081.pro
56ffe5bca4632e7e75c07c04369529b7
90e755975f1916c9bd1f89539e5633a464e560d9
F20110115_AABXZJ muench_a_Page_088.tif
4e78254f6147d6611a110211ee8537c4
cd8a19eecb1eff697098b6845c1572bee5d73f79
55674 F20110115_AABXYU muench_a_Page_046.pro
478e4edd5587cb43740c4f87e73979e8
a0df1c46046b95b26c9f7411dd9d0eadf550017f
38122 F20110115_AABXZK muench_a_Page_092thm.jpg
62afef6c2617e51715b66d0950a8e810
68efc6ce5e34c477bd7e689d9c2d45c5958698fa
8860 F20110115_AABXYV muench_a_Page_097.pro
71ff45e444104d1f5d8e2c612ecab6dd
f92515396c136da89186a0a481b5861e75eb1e4e
25619 F20110115_AABXZL muench_a_Page_031thm.jpg
524780089e838f1a9e4dadec480f3cc1
1a9239cc917f5cee4623097e26a920e89f85a0e3
92421 F20110115_AABXYW muench_a_Page_099.jp2
73de6cfb1aa5626d3506f6cfdcaae725
a0546e7af292cb5a887e5debcbcf559bce73a045
51729 F20110115_AABXZM muench_a_Page_030.pro
087337ec777b41cd718df4f21f572ed9
7c9221bdca76a02b9518b6703087cbd604a18f40
170192 F20110115_AABXYX muench_a_Page_010.jpg
9a5ecb3fd603ef2d55774bc8f6105a56
7f2f1baec20c23711eec8934455f5758f3747ba8
94506 F20110115_AABXYY muench_a_Page_005.QC.jpg
3588341600f84c812b7e319cd88a15ad
36a8722e5ecdfc6939a18d3a4608e20202b131da
50370 F20110115_AABXZN muench_a_Page_013thm.jpg
1b7317d2b1eb01a33cec7404490b2d3d
8eab6fab186ad5aa318d042953a2abbcb5d5aac2
1820 F20110115_AABXYZ muench_a_Page_058.pro
ce453b6f37b616c93b5c474beb1c0f1c
36e5ff8962506c2627cdfa0f1fd9644517528965
111239 F20110115_AABXZO muench_a_Page_049.QC.jpg
1bb3b019c91246affd48fffee2037db2
865cf3359800b6fa99a1ecba5644585266ce5149
51927 F20110115_AABXZP muench_a_Page_029.pro
9d5f783a2405ac380d77caf08dff0684
247e0eafd5adbaf0994be80b992f8165ae25cdeb
14112 F20110115_AABXZQ muench_a_Page_027.pro
fe934829445bea75605e472ec6fea325
14754200f4ed00ff3d0bc3dc0760f7b1a61f9cdb
214266 F20110115_AABXZR muench_a_Page_076.jpg
8b7cce8a7df6f7255c09ab378626133a
79264d7c8b026d73820f32f2a6d3f26a50cb1738
30612 F20110115_AABXZS muench_a_Page_003.jpg
49c6eb473657efbefb1809224b92a0bf
0d84568e1420aa086aa70ac55efe0f50ad9d7974
219060 F20110115_AABXZT muench_a_Page_026.jpg
81f60baeb7c80c9fc59948b4cd421dd1
6c7cedc02aea7d827b19ea3d0011e0986a68f849
275485 F20110115_AABXZU muench_a_Page_111.jpg
82d67579d06b77ae3e109e35e474ed11
7c20acc85083e181613bb8e41248dca8519ff7b2
1203002 F20110115_AABXZV muench_a.pdf
e95963d231ea16dc3fc767a66bee238e
872cc15980b7c9f50ce4555e23c8fec6a6daf94a
19405 F20110115_AABXZW muench_a_Page_073.pro
3692ef841b07a2ff993b911cf5ad1023
9d750cf710410c5810232366f1b0950241eed107
50911 F20110115_AABXZX muench_a_Page_013.pro
84a23a3e7edac89f2b148335a01cfc64
0c78dbb3172931055b0cb223d5a51c483f56404f
2044 F20110115_AABXZY muench_a_Page_079.txt
2e1a756987536e30ba643c27a1332daa
6cf8a62b1095ef68750ea3233552660df4abc8ab
237646 F20110115_AABXZZ muench_a_Page_051.jpg
33d2d7b9deabf1577c9175b5becaa30c
95c1e3f25718cb9ecee03b7042bff9c7288ea379
99331 F20110115_AABYAA muench_a_Page_064.jpg
fa2e678dee4e85399ebd115b0e73bf71
c7e718cf7a7b244c1cbb0d0d136e77cc0208360c
1051897 F20110115_AABYAB muench_a_Page_047.jp2
5c28a409ac75eef880ddce040ec3dd55
dcae9466bbea8af544a4795860da8099e9545c35
12586 F20110115_AABYAC muench_a_Page_070.pro
3bdc6b158726d8f0b729c5e1912fa4a1
6f61e29c4eccb1813a4d1c09edb28c38294fabe4
245022 F20110115_AABYAD muench_a_Page_005.jpg
56c9dfd217637388741a9e217309c486
164a72f32612264d48550f5585dfe64710907d3b
51099 F20110115_AABYAE muench_a_Page_104.pro
5dc030364283cf08e0ad0133531502d9
5478fa406d956e571e233520119aefb812a9ab05
380165 F20110115_AABYAF muench_a_Page_062.jp2
70bfde2abaa1033cbdd4c9f50a586420
14b8d5d290b47dc14e4dbf57c5aaedc9daddea51
F20110115_AABYAG muench_a_Page_102.txt
ae0919f47b078e1cc930c0422a51d9ac
785bf0abf83fafdefd37ea719b69b9c7e39ff0be
267985 F20110115_AABYAH muench_a_Page_046.jpg
677fb99c86d9478e4dbbc47e4a2e7d30
34178e4661af6dc77c77cdf60c6c8b2c4bebd82c
687 F20110115_AABYAI muench_a_Page_070.txt
ded79c456c4903c012c10a6b8a9df463
88a89547ca250130b65c4cda195f8c26a15af9f3
1833 F20110115_AABYAJ muench_a_Page_012.txt
f063cb6afc8de8b320aee006281f47f5
d405b555c57ae26ab76a6ad1f81f4dbd6460f393
20916 F20110115_AABYAK muench_a_Page_073thm.jpg
8cb77008155644c001095946a095a7b1
206f180f9f7529d57ebbba21e6ba89109124ce67
2073 F20110115_AABYAL muench_a_Page_077.txt
8edd0160aa06717f6b76f2cfa5a7b805
4004872d099fc29a16f7300444d7b82a09c3f731
90689 F20110115_AABYBA muench_a_Page_041.QC.jpg
ed59228458938c793edc74bd7e793303
0e1c8ad1582b24d6622bd82601d559632f8798a2
1479 F20110115_AABYAM muench_a_Page_020.txt
e90e2bf1bfa256258dd55724317113b5
c9732d695f03223278bda795b2453c1427ca7975
108471 F20110115_AABYBB muench_a_Page_037.jp2
58f12c8924e96b00df2c115d3493f978
7b60ef10eb03269f944c85db4c4c66afbf79c365
25599 F20110115_AABYAN muench_a_Page_077thm.jpg
ffb8eff17f76df498e923cbc138f1467
fea47c88038110c84ad8288ecbb7dbdf86b03eab
229419 F20110115_AABYBC muench_a_Page_009.jpg
8e62fbb854f2c702d0fe8671b1af007e
ef32e697dfb2ee362fa6775da4c706ebd22a873c
35349 F20110115_AABYAO muench_a_Page_066thm.jpg
b618bbf9de1f08e2bff4135c93517b9b
79813b46e0f52e9d2700273fc39699793767f92f
2035 F20110115_AABYBD muench_a_Page_013.txt
1f6fc82f851e14137882d1f6f3e5f8ee
e8e5098041775399f88a5e249343df7705a5e6a4
81628 F20110115_AABYAP muench_a_Page_076.QC.jpg
ee20d82b9ddd201be375933c49647c85
b191afa29c2a1375e2b9b860316c899b43800f5f
51181 F20110115_AABYBE muench_a_Page_007thm.jpg
4f7f7c8a0da3da690db0705dd5791c22
33109877df74302465cdf20672f1811f984f20ff
106468 F20110115_AABYAQ muench_a_Page_051.QC.jpg
e6877fc71cf86de3f9966b7058c3507a
9b3bfdd8b5943ef4d2276b3b257d2995916957aa
23854 F20110115_AABYBF muench_a_Page_078thm.jpg
eef1ca201111adb65f985974a755b87d
78fdb061bc5a7e7a90b290c5ef9edb465c5407ce
209942 F20110115_AABYAR muench_a_Page_016.jpg
b606c146f208694160358abff1c41e83
ef02d9518148a3e939a821ecee56cdea9336c953
F20110115_AABYBG muench_a_Page_106.tif
01c221dc09fe5eede71ad8743f341b88
aad8ad03aa3490040364323991c48f73bbe3666e
95665 F20110115_AABYAS muench_a_Page_009.QC.jpg
0d41e33a82c135b501875572bd9f060e
537776f57f7e8a4c8886a7f9867560fb7401f08b
24921 F20110115_AABYBH muench_a_Page_032thm.jpg
5f9e64aa06be396e6c5768f0a5c2ffd4
098fe3ec41812fbe7eac262f6a3f3a88a6aa6d2d
1051976 F20110115_AABYBI muench_a_Page_044.jp2
e8ba6930cae4b8f65a16866e9fad260d
991229935e582926f24846e0f6178379b90d6384
72267 F20110115_AABYAT muench_a_Page_101.QC.jpg
72a44476cf39ff69e38d04bb04f993a3
eff86e06b286c9fca96eeb0d22ddaf8261add8d0
F20110115_AABYBJ muench_a_Page_004.tif
76b702621d6900ad24dcf891c810a949
6c8a350b5a2a3e7cdad5bfe874a6c4e3b8c86656
48596 F20110115_AABYAU muench_a_Page_069.QC.jpg
d894e45714f2ec66235634684adc05e7
7d97cc36b6f1e7bdbf9c7e38d80a57b98d118163
1938 F20110115_AABYBK muench_a_Page_016.txt
611fd44ac5ff616132a493aa69879681
04f49645acdf4da35670ed0e7c66427fc0d0c923
119511 F20110115_AABYAV muench_a_Page_073.jpg
af913189a6058bc119cbece3378db4e9
ca7f8ba8a66d3d508ebd22fa955a9c47741cc391
77553 F20110115_AABYBL muench_a_Page_035.QC.jpg
9098c09df88276663b6842a2c9d4e489
4b354907621c93768d70af88b6b71d305b7f96a4
78633 F20110115_AABYAW muench_a_Page_043.QC.jpg
c14ab73359e20c211caace5639aa47f9
8a17a7d4f79f1898838b195c4c7c6216cc8e647a
61779 F20110115_AABYBM muench_a_Page_114.pro
d189439507c379cfa8b007d2ef004f7c
c7a041f82d370e88985755c8f3f6f14d711f8b4c
26383 F20110115_AABYAX muench_a_Page_027.QC.jpg
2f0b611ffc776cf63008690e77f3a9da
39dbbc54bb13e5610947ba243187868982f67d9f
172533 F20110115_AABYCA muench_a_Page_028.jpg
4c08d1af75af6709c235e7a7915217c5
019b51a0e06877d514664308690c7d2aa65f6158
F20110115_AABYBN muench_a_Page_064.tif
8efbe161c9b5ea6e62fa2b38df0262f9
bad96066990877d9b3fa8e6fed422e9f91c5afa8
1590 F20110115_AABYAY muench_a_Page_107.txt
af228fd2a90c397fcb0d7061dc9ffff1
db05c0e20bc71c8102c3d8e868b15361df63906e
F20110115_AABYCB muench_a_Page_062.tif
d7cb2cf8d9ffe1340e4e690d45fe8b93
2d8144f537651cbd30c2c5c2db37a59b9a0ab755
F20110115_AABYBO muench_a_Page_019.tif
eda6d77fc23a69c5fee2e0c008ef38b0
e38d71e0e9ffc543f0fc84345e21357c074a940c
F20110115_AABYAZ muench_a_Page_095.txt
6dd1362bd36479cce4aa47aa0c48f1eb
4cc4962856324bc293b1c1362fa64e22f92dc6fd
207265 F20110115_AABYCC muench_a_Page_014.jpg
59c295b3d47c9e646d8104442d3520ad
f85947d806e89b06514b9b11ea4b660e777674ed
25146 F20110115_AABYBP muench_a_Page_035thm.jpg
3456a771eb44f1a1eb62150613dd0ebf
ed3819c192071e8ea9b636498a117696e50fd182
F20110115_AABYCD muench_a_Page_025.tif
b6822cd6ee825ad94844f01e9d60f99a
561765962fbddeeb784ee9aa55b5a5835220cc52
189784 F20110115_AABYBQ muench_a_Page_058.jpg
48ea60f966951d70f12336aed94d897d
27e41bf18ca2952eaff3dc0e38589785f63ae0a1
2005 F20110115_AABYCE muench_a_Page_104.txt
6f7f9b70633477002eb29bbbc878b105
916a69fe001d67fb6299ac34452e722e787cafea
110405 F20110115_AABYBR muench_a_Page_006.QC.jpg
5d59b940450c50f3086404a84619f3b8
9dc612dbe5d2940f3508bce6c89460e3e2cf8737
33274 F20110115_AABYCF muench_a_Page_094.QC.jpg
ebd819a33e7a64474e4ec7aabbf652e3
823d222ce6088ab37b4b4709cf3ebf048ecebe11
48839 F20110115_AABYBS muench_a_Page_078.pro
b16d2490c5bfcdd73d7c08546ff6eb8e
233d0425852b20b6959dbef843cca3cabb07132c
24965 F20110115_AABYCG muench_a_Page_114thm.jpg
4a1c7a7516b45691b12f697054a678e8
6fdbde24126b2aabe0bb2f6feeab9335a51f54ee
F20110115_AABYBT muench_a_Page_021.tif
3eb927c6baa649ce4a445eb764309330
470337f6047ac1736c1ba96e2f64262a093af1b9
50239 F20110115_AABYCH muench_a_Page_035.pro
0cd021c0bdd84bf8f68e452558d30a7c
fcf65d57d16c1fc2ea7f65244fd97c686b9061ff
83607 F20110115_AABYCI muench_a_Page_098.jpg
3b6f5431d93b291835af9a535b603556
0770626ac8d85936e780bd6ccebb0a2b33e032ba
84557 F20110115_AABYBU muench_a_Page_010.jp2
833d63ded17bdf14f49757ec74869020
358712a1f5d5e5925ffdbb779b03de6fdabd0c17
286396 F20110115_AABYCJ muench_a_Page_109.jpg
cfb9d4a32995cd059e7640d7337220fe
fbe7b8ad51d726cbb926f04ef08e075065f53000
49654 F20110115_AABYBV muench_a_Page_037.pro
2c7776cb2d957fa7b032199914be8c87
8997afd502559113ba68a084aad4c56afad90666
F20110115_AABYCK muench_a_Page_003.tif
d425de562ae2ff459058b21b742d2680
bc7738b24223dfa622de1629979992a63689012c
25318 F20110115_AABYBW muench_a_Page_037thm.jpg
093e2f91c14c21aae3aa6a22f6b9ff6f
4f7556470d538ca2eba46d814c5975d23f1001f5
45412 F20110115_AABYCL muench_a_Page_091thm.jpg
54d698ff45e0c9d8891da80f2c50c862
28baf6012b6a2bb70f8b7040c510a9c7975c0e14
49486 F20110115_AABYBX muench_a_Page_111thm.jpg
d6bfd56a0dc914001ef4fed9ff881b47
d3b804b1b4a4af4d4627716ea66741143b8991a6
247764 F20110115_AABYDA muench_a_Page_048.jpg
095d61028540bf678ebda1625c36e4f0
460af47b0a9fab031ff197269db5737b4eaaf8eb
76748 F20110115_AABYCM muench_a_Page_061.jpg
cc7a0c307937e400bcb56ed0aa3f7d67
e0f0d654a2ad486ebda645c2ab699eef877ef148
136543 F20110115_AABYBY muench_a_Page_093.jpg
49de77dde9fd44bb75c1d78ca76867ce
eb36db7e9d0709ee268ea8ce6336f36960c71506
2532 F20110115_AABYDB muench_a_Page_110.txt
d564e451d319fd5e9cb9f57db68a1fe8
3e0ae8b27c4f8314e8b6871aae9e2868b79a2c18
1051965 F20110115_AABYCN muench_a_Page_005.jp2
a3ccd437b67914ad98f3e9554530d29e
8a402e2db521164c4bd62e121a7e22afb47ed236
1051985 F20110115_AABYBZ muench_a_Page_007.jp2
abfe50312b48c113cc63e40f9f0076c4
880b31485d02ae6bd09017ca8a93b32befe2e01f
52077 F20110115_AABYDC muench_a_Page_106.pro
6dd87af3f55fb7ad7e6cb408dabe816e
26bae70cb17f2c61d71c422fb59e4aeec46011cd
152199 F20110115_AABYCO muench_a_Page_020.jpg
3a65a4e90d20bc18aae7ed804b3ef1e5
c568c59d73d84efcd20641049ba402bb37357c81
216824 F20110115_AABYDD muench_a_Page_043.jpg
d0e9d53bc7bae84b2fe871263f9bf123
88a124c4f941c1bbf635f63278fca5d48ad5fb68
241489 F20110115_AABYCP muench_a_Page_050.jpg
ea9043ad3a84e1ef1871d0cdf12c1c1f
5a550eaa3f2db4945fa28de78b138b8718d9122f
1943 F20110115_AABYDE muench_a_Page_084.txt
89db795ff84b4fdbe55166686c8ab438
5bc4734ef185e0019a52c37ae4b6dc413bd1c7d3
65812 F20110115_AABYCQ muench_a_Page_109.pro
fcef37a20eb9dc0f6ff324959abda5b6
5b1807b241c97c1b2c7304ff545e5bce86454748
87317 F20110115_AABYDF muench_a_Page_075.jp2
320aeb93909d4482b83379c60bd2d5d1
6cc7eabb483a46eac358019589bb6ddd7df856e4
78150 F20110115_AABYCR muench_a_Page_053.QC.jpg
f90d3bb987e281690abac8f6a08a036b
6b588f727eba81e6d3863ea06068c02379c250f8
47100 F20110115_AABYDG muench_a_Page_034.pro
849992444256c35825650742fd241968
535c5add3450690f6b70309f08d8fc68acea8e67
52377 F20110115_AABYCS muench_a_Page_068.QC.jpg
fdaef101096e52a654363ab564d8266f
be8f2d6355aa0d5d4a6b18bbf9dd80deb25f6d8f
25878 F20110115_AABYDH muench_a_Page_109thm.jpg
cc532859351191a7b8b886313d34d077
387b78f77c6c72477d1c55be59f51af46e7f4c5e
279361 F20110115_AABYCT muench_a_Page_114.jpg
57a17c6e2785a9aa294a0e8930fdf203
b6a078a124ac4850b94df1b3b47df47c265118a2
F20110115_AABYDI muench_a_Page_051.jp2
7e98d31f5cd2158c03aed8ec2e4d538e
92272eaec5717a9ccfd6bceff53f43195498e15a
85193 F20110115_AABYCU muench_a_Page_071.jpg
87d2a588a4f2c5978b65d848ba04aad7
0106fe9e39c67566f693247609097c7bf5383691
1051978 F20110115_AABYDJ muench_a_Page_033.jp2
4a433f913e4a018cdc8017efc7a274a3
bff60543e3a30490b47d5255631a69dac9ded376
24628 F20110115_AABYDK muench_a_Page_085thm.jpg
5de7d89becc72d0241bec99f39e7e0ad
ee0bc723ee28283bbb0ce3d469573aaa1a8520f8
87185 F20110115_AABYCV muench_a_Page_110.QC.jpg
6010854d25e5f0162a428d4cd5a018f1
ed604de3140da99135bd5365b21602774dda2d86
199440 F20110115_AABYDL muench_a_Page_025.jpg
47ca03d0e9b1953f1cc7e11bdbf20a79
78ca88adc763c6b284f0df753904ef7bcf6de302
2021 F20110115_AABYCW muench_a_Page_103.txt
729165744201c396e5f88d89025a573f
da975d06532061e57385204443fcc740e1efcf14
34691 F20110115_AABYEA muench_a_Page_067thm.jpg
6579b8d03c71743a4303f2698916ac8a
5117767a60e7a7fd383f01e8b1e58e8154ee3a69
214262 F20110115_AABYDM muench_a_Page_057.jpg
cf1169bdbbf9c5c7b9413d29c2c458a0
db004e7e2b8825bc93aa40c069b039dc61515beb
40935 F20110115_AABYCX muench_a_Page_021.pro
966c67f6fc0025a13cbdbdab2b0527e9
1a04afe79cd7ed0408052fb7af4201c0b2a6f0ce
42438 F20110115_AABYEB muench_a_Page_099.pro
41753accd0f68aa591c9bf3f0b2f17b8
ec4180df29895f3f9dcf046cea31ac517c292a21
F20110115_AABYDN muench_a_Page_032.tif
226b359cb0a082ad75c5c61d5f54553c
76468c8d108f1c9532122c80c79c221dce0a3c4d
24798 F20110115_AABYCY muench_a_Page_080thm.jpg
b109adb906e05dc81ad2cc1d38649a4a
6ef7e4e2b9668622ec3939fa8e54887663c7f4c0
229881 F20110115_AABYEC muench_a_Page_108.jpg
8fcfd894bda0ec93db664f88bc38a1cc
80b0e5c93c041e6b833d484f2f36cb2115f67704
191257 F20110115_AABYDO muench_a_Page_054.jpg
6bcdf0ae1ef2dd342046a9f7dffd846f
de91e63b12c23c894c3b1ac2dbfd0589cb8e8f4b
64558 F20110115_AABYCZ muench_a_Page_090.QC.jpg
727122a916e46716aa8d9d55f7cabd94
32028ba5264f40e745d426d95b5c34a5d8493639
5888 F20110115_AABYED muench_a_Page_002.QC.jpg
8b5dc1edb689a79803f9151f09ba02a6
6abeb4daacc271fd0efb74e3e2ee4e9971bd8c3d
F20110115_AABYDP muench_a_Page_029.tif
aed0ad9e625e0832c350f05f1b0a7151
5dcd2f76f1a8c5292dd4de38d31eb104f2c16789
1971 F20110115_AABYDQ muench_a_Page_085.txt
c3a1ec38c1918e826ccc661c6cd02427
437ab227a955f1c3e6eee4afc3f6fcab40b84a09
208020 F20110115_AABYEE muench_a_Page_037.jpg
338fba2a809be0684ab69ef71160a5fa
5418cdbb08f922ce42dd8006f1ed1a6de07e680a
2590 F20110115_AABYDR muench_a_Page_112.txt
534934ceea0b09eddbed9e6ce26cdad7
db570bdf674f66539943843d3ccadca0783fe859
31771 F20110115_AABYEF muench_a_Page_041thm.jpg
f0a1796f96b5478f70c2e878a8a34347
ae0bdf4333fd7bd5dca5af42374916ec7c877267
201482 F20110115_AABYDS muench_a_Page_053.jpg
84f195a72fe37cf151fe82d5b9750590
101b8610bc7e4cd2855420818c8476d285555f55
69407 F20110115_AABYEG muench_a_Page_058.QC.jpg
5de18471bd30f83a0b2a8b8b0973376d
37ba1b845aabc54b3ce41a9ab32383c73dce245d
200079 F20110115_AABYDT muench_a_Page_024.jpg
770925cb0eb0d7b0a9fa37354bbf07e2
9aa4258c906d84418b80ba7647a00453d3adecc4
28484 F20110115_AABYEH muench_a_Page_045thm.jpg
3da619b8172d6827dfee33bc9cbcfc6d
6472d9ab3b81f8d01578d275695603c2172fa227



PAGE 1

AQUATIC VERTEBRATE USAGE OF LITT ORAL HABITAT PRIOR TO EXTREME HABITAT MODIFICATION IN L AKE TOHOPEKALIGA, FLORIDA By ANN MARIE MUENCH 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 2004

PAGE 2

Copyright 2004 by Ann Marie Muench

PAGE 3

This document is dedicated to my parents, Joseph F. and Mary K. Muench, whose love and support have strongly contributed to my academic, professional, and personal growth.

PAGE 4

iv ACKNOWLEDGMENTS I would like to thank my major advisor, Wiley Kitchens, for taking me on as a graduate student. He was a critical source of expert advice and encouragement, and was always accessible for consultation. I also thank my committee members, Madan Oli and Lauren Chapman, whose academic instruction and cr itical analysis of this thesis are much appreciated. My coworkers also contributed much to my education, and I am thankful. Funding for this research was provide d by the Florida Fish and Wildlife Conservation Commission (FFWCC). From this agency, Duke Hammond helped immensely with the direction of the study, and provided critical feedback on progress reports that we provided to the commission. The staff of the Kissimmee, FL, office of the FFWCC was helpful in facilitating our fi eld work at Lake Tohopekaliga. Bobbi Jo Cromwell from the Osceola County Department of Parks and Recreation allowed us to store all of the crayfish and minnow trap s on Makinson Island in Lake Toho. The field work for this study was conducte d through the time of many dedicated students and technicians from the Florida C ooperative Fish and Wildlife Research Unit. These stalwart coworkers included (in al phabetical order) Scott Berryman, Stephen Brooks, Janell Brush, Brenda Calzada, John Davis, Jamie Duberstein, Bruno Ferreira, Joey Largay, Kristianna Lindgren, Samantha Musgrave, April Norem, Derek Piotrowicz, Laura Pfenninger, Erik Powers, Vanessa Rumancik, John David Semones, Micheala Spears, Chris Tonsmeire, Paul Traylor, Zach Welch, and Christa Zweig.

PAGE 5

v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES............................................................................................................vii LIST OF FIGURES.........................................................................................................viii ABSTRACT....................................................................................................................... ..x CHAPTER 1 MAIN INTRODUCTION............................................................................................1 Lake Ecosystem............................................................................................................1 Study Area....................................................................................................................2 Research Objectives......................................................................................................8 2 DESCRIPTIONS OF FOCAL SPECIES...................................................................10 Aquatic Vertebrate Habitat.........................................................................................10 Fish Species................................................................................................................11 Centrarchids (sunfish).........................................................................................11 Exotic catfish.......................................................................................................13 Herpetofaunal Species................................................................................................13 Amphibians..........................................................................................................13 Reptiles................................................................................................................15 3 ASSEMBLAGE WITHIN THE Pontederia cordata COMMUNITY.......................17 Introduction.................................................................................................................17 Field Methods.............................................................................................................17 Trap Descriptions................................................................................................17 Whole-Lake Sampling.........................................................................................20 Analysis Methods.......................................................................................................23 Trap Comparisons...............................................................................................23 Species Richness.................................................................................................23 Assemblage Composition....................................................................................25 Influence of Temporal Gradients on Assemblage...............................................25 Proportion of Habitat Utilized by Focal Species.................................................27

PAGE 6

vi Results........................................................................................................................ .29 Trap Comparisons...............................................................................................29 Species Richness.................................................................................................31 Assemblage Composition....................................................................................32 Influence of Temporal Gradients on Assemblage...............................................34 Proportion of Habitat Utilized by Focal Species.................................................40 Discussion...................................................................................................................41 Trap Comparisons...............................................................................................41 Species Richness.................................................................................................41 Assemblage Composition....................................................................................42 Influence of Temporal Gradients on Assemblage...............................................44 Proportion of Habitat Utilized by Focal Species.................................................45 4 ASSEMBLAGE ACROSS VEGETATION COMMUNITIES.................................64 Introduction.................................................................................................................64 Field Methods.............................................................................................................64 Grid and Web Sampling......................................................................................64 Whole-Lake Sampling.........................................................................................66 Analysis Methods.......................................................................................................67 Population Estimates and Movement for Herpetofaunal Species.......................67 Capture Success for Focal Species......................................................................68 Results........................................................................................................................ .70 Population Estimates for Herpetofaunal Species................................................70 Capture success for Focal Species.......................................................................72 Discussion...................................................................................................................74 Population Estimates for Herpetofaunal Species................................................74 Capture Success for Focal Species......................................................................75 5 SUMMARY AND CONCLUSIONS.........................................................................88 Review of Aquatic Vertebrate Community Dynamics in Lake Tohopekaliga...........88 Air Temperature..................................................................................................89 Lake Stage...........................................................................................................89 Water Depth.........................................................................................................90 Vegetation Community.......................................................................................90 Population Size Estimates...................................................................................90 Lake Tohopekaliga Habitat Enhancement..................................................................91 Future Aquatic Vertebrate Monitoring Plans.............................................................95 LIST OF REFERENCES...................................................................................................97 BIOGRAPHICAL SKETCH...........................................................................................105

PAGE 7

vii LIST OF TABLES Table page 3-1 All species captured in 2002, with species codes used in subsequent figures.........30 3-2 Species capture frequencies for th e 0.6 and 1.3 cm mesh minnow traps.................31 3-3 Indicator species analysis results..............................................................................33 3-4 Stress and instability resu lts from all NMS ordinations...........................................34 3-5 Percent of variance explained (r2) by environmental variables for each axis in the vertebrate NMS with det ection/nondetection data...................................................35 3-6 Percent of variance explained (r2) for each axis by species in the vertebrate NMS with detection/nondetection data..............................................................................35 3-7 Percent of variance explained (r2) by environmental variables for each axis in the vertebrate NMS with count data...............................................................................36 3-8 Percent of variance explained (r2) for each axis by species in the vertebrate NMS with count data.........................................................................................................36 3-9 Percent of variance explained (r2) by environmental variables for each axis in the fish NMS with detection/nondetection data.............................................................37 3-10 Percent of variance explained (r2) for each axis by species in the fish NMS with detection/nondetection data......................................................................................37 3-11 Percent of variance explained (r2) by environmental variables for each axis in the fish NMS with count data.........................................................................................38 3-12 Percent of variance explained (r2) for each axis by species in the fish NMS with count data.................................................................................................................38 3-13 Percent of variance explained (r2) by environmental variables for each axis in the herpetofaunal NMS with count data.........................................................................39 3-14 Percent of variance explained (r2) for each axis by species in the herpetofaunal NMS with count data................................................................................................39 4-1 Grid sizes and population estimat es by mark recapture methods............................71

PAGE 8

viii LIST OF FIGURES Figure page 3-1 Crayfish and minnow trap in P. cordata habitat......................................................47 3-2 Locations of 2002 P. cordata sampling transects in Lake Tohopekaliga................48 3-3 2002 Vertebrate species richne ss estimates by sample date.....................................49 3-4 2002 Fish species richness estimates by sample date...............................................49 3-5 2002 Herpetofaunal species ric hness estimates by sample date..............................50 3-6 NMS ordination of sample units in vertebrate species space using detection/nondetection data......................................................................................51 3-7 NMS ordination of vertebrate species in sample unit space using detection/nondetection data......................................................................................52 3-8 NMS ordination of sample units in vert ebrate species space using count data........53 3-9 NMS ordination of vertebrate species in sample unit space using count data.........54 3-10 NMS ordination of sample units in fish species space using detection/ nondetection data......................................................................................................55 3-11 NMS ordination of fish species in sa mple unit space using detection/nondetection data........................................................................................................................... 56 3-12 NMS ordination of sample units in fish species space using count data..................57 3-13 NMS ordination of fish species in sample unit space using count data...................58 3-14 NMS ordination of sample units in herp etofaunal species space using count data..59 3-15 NMS ordination of herpetofaunal specie s in sample unit space using count data...60 3-16 Average and range of lake stage values by cluster...................................................61 3-17 Average and range of air te mperature values by cluster..........................................61 3-18 Site occupancy estimates for focal fish species in spring 2002...............................62

PAGE 9

ix 3-19 Site occupancy estimates for focal fish species in fall 2002....................................62 3-20 Site occupancy estimates for focal herpetofaunal species in spring 2002...............63 3-21 Site occupancy estimates for focal herpetofaunal species in fall 2002....................63 4-1 Locations of 2003 sampling transects in Lake Tohopekaliga..................................80 4-2 Diagram of weekly trap pl acement at specified depths............................................81 4-3 Mean maximum distances traveled with variances and maximum distances, based on results of mark-recapture sampling...........................................................81 4-4Number of trap sites sampled in each vegetation community per sample occasion.82 4-5 Salamander capture success by vegetation community............................................83 4-6 Salamander capture success by water depth............................................................83 4-7 Frog capture success by vegetation community.......................................................84 4-8 Frog capture success by water depth........................................................................84 4-9 Snake capture success by vegetation community....................................................85 4-10 Snake capture success by water depth......................................................................85 4-11 Turtle capture success by vegetation community....................................................86 4-12 Turtle capture success by water depth......................................................................86 4-13 Fish capture success by vegetation community.......................................................87 4-14 Fish capture success by water depth........................................................................87

PAGE 10

x 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 AQUATIC VERTEBRATE USAGE OF LITT ORAL HABITAT PRIOR TO EXTREME HABITAT MODIFICATION IN L AKE TOHOPEKALIGA, FLORIDA By Ann Marie Muench December 2004 Chair: Wiley Kitchens Major Department: Wildlife Ecology and Conservation Lake Tohopekaliga is a large, shallow lake in central Flor ida that is part of the Kissimmee chain of lakes. Cultural eutrophi cation and lake stabilization over the past several decades have facilitated the forma tion of a densely vege tated, often monotypic, littoral zone. Lake managers conducted an enhancement project in 2004 to improve largemouth bass ( Micropterus salmoides ) habitat. This project included an extreme water level drawdown and concurrent mechanical removal of 7.3 million cubic meters of organic sediment and vegetation from the s horeline. Following the drawdown, herbicidal treatments will keep the lake vegetation in an early state of succession in order to prolong the effects of the enhancement. Little is known about potential impacts of these procedures on wildlife, incl uding vegetation, avian, herpetofaunal, and even fish communities. This study examines the status of the reptile, amphibian and fish communities in the two years prior to the lake enhancement to provide baseline data for future assessments.

PAGE 11

xi Funnel traps were used for all sampling, allowi ng a suite of vertebrate species to be examined. In 2002, sampling was conducted in the Pontederia cordata (pickerelweed) zone of the lake. Cluster analysis, indicator species analysis, and nonmetric multidimensional scaling ordinations were us ed to examine temporal changes in the species composition of the assemblages. Envi ronmental variables such as lake stage and average air temperatures played large role s in structuring the aquatic vertebrate communities through species richness and a ssemblage composition. Fish assemblages were most correlated with air temperature, while herpetofaunal assemblages mainly showed association with lake stage. Site occupancy estimates showed that many of the herpetofaunal species are pres ent throughout the pickerelweed habitat in both the spring and fall seasons, while fish showed more fluctuation in seasonal presence. Spatial sampling took place in 2003. Sampling was conducted across vegetation communities and water depths. Both variables captured varying trends in the presence of the focal species, which included fully aquatic salamanders ( Siren spp., Amphiuma means ), water snakes (mainly Nerodia spp.), small kinosternid turtles ( Kinosternon baurii, Sternotherus odoratus ), large aquatic frogs ( Rana spp.), juvenile centrarchids (especially Micropterus salmoides ) and exotic catfish ( Hoplosternum littorale ). Attempted population density estimates for the more abundant herpetofaunal species ended in failure. Inappropriate trapping grid size and spacing for several species at one time led to poor capture proba bilities and large variances in population size estimates.

PAGE 12

1 CHAPTER 1 MAIN INTRODUCTION Lake Ecosystem The productive littoral environment in a lake system is dynamic, since the aquatic habitat has strong terrestrial influences and the terrestrial habita t has strong aquatic influences. Biological diversity is high in the ecotone due to biotic and abiotic properties that distinguish it from adjacent ecosystems, such as vegetation species, soil properties, and water chemistry (Lachavanne 1997). Water level fluctuations are the main determinants of the width of the littoral zone In lentic systems with gently sloping shorelines a wide band of macrophytes provide s patches of heterogeneous habitat for a diverse assemblage of faunal sp ecies. Animal species that have specific requirements for different life stages depend on the proximity of supralittoral (never flooded), eulittoral (occasionally flooded), and infralittoral (alway s flooded) habitats. Since the ratio of water surface to volume is much higher in the littoral zone than in the deeper pelagic region, environmental variables such as lig ht, air temperature, wind and water flow (waves) have much more critical roles in shaping the grad ient (Pieczynska 1990, Pieczynska and Zalewski 1997). Excess inputs of nutrients beyond that natu rally found in a particular lake system leads to eutrophication. This condition encour ages surplus sediment ation and vegetation growth, changing the landscape of the original littoral zone and subsequently altering the biological communities within that habitat and the lake as a whole. Eventual extinction of the lake may result from decades of nut rient pollution due to sewage discharge and

PAGE 13

2 drainage from agricultural and urban lands. Internal recyclin g of nutrients within a water body keeps it from recovering even when the inputs are reduced (Cooke et al. 1993). In order to preserve the lake for the longest ti me possible, rehabilita tion efforts are often made to counter the results of eutrophication, for example drawdowns, dredging and mechanical vegetation removal (Hasler 1947, Cooke et al. 1993). Study Area Lake Tohopekaliga (7,612 ha, 18,810 acres) is located in Osceola County, Florida, within the Upper Kissimmee Basin. This phys iographic area is known as the Osceola Plain, which is flat and has very few disti nguishing topographical ch aracteristics. The elevation in the plain ranges from 18-30 m (60-95 ft) National Vertical Geodetic Datum (NGVD), but rarely reaches maximum hei ghts (Harper 1921). Originating from prehistoric ocean bottom, the sediment mainly consists of coastal sands. Numerous shallow lakes in the region, in cluding Lake Tohopekaliga, were formed by dissolution of the carbonate-containing substrates (limes tone) in depressed areas (Schiffer 1998). Freshwater wetlands in this area include cypress sloughs, wet prairies, river swamps, floodplains, mixed forested wetlands, and marshe s. Pine flatwoods dominate the upland community (HDR Engineering, Inc. 1989). The Lake Tohopekaliga Subbasin (211.6 square km, 131.2 square miles), within the Upper Kissimmee Basin, receives wate r input from the Sh ingle Creek (184.2 square km, 118.0 square miles) and East Lake Tohopeka liga (81.7 square km, 48.4 square miles) Subbasins. Precipitation, overland flows, and to some extent groundwater from the underlying Surficial Aquifer also provides the lake with impor tant water sources. While evapotranspiration is a strong f actor in withdrawal of wate r from the lake, outflow from Lake Tohopekaliga occurs at its southernmost point, where the Sout h Port Canal connects

PAGE 14

3 it to Cypress Lake. Water from the Uppe r Kissimmee Basin flows southward through the Kissimmee Chain of Lakes, through the channelized Kissimmee River, to Lake Okeechobee, east and west coast estuaries a nd South Florida (HDR Engineering, Inc. 1989, Schiffer 1998). Human disturbance of this hydrologic syst em began in the mid-nineteenth century, with local efforts to drain wetlands. In 1882 Hamilton Disston began channelizing the watersheds in the upper basin by constructing in ter-lake canals. The ma jor results of this endeavor were lowered lake levels, drying of lake edges and interlake slough wetlands, as well as rapid transit of nutrient-laden surface waters downstream. Wetlands stretching between Lake Tohopekaliga and East Lake Toh opekaliga were strongly impacted. In the 1920’s, Shingle Creek (a major source of wa ter for Lake Tohopeka liga) was channelized, bypassing water around the swamps and marshes within that subbasin. Many landowners also dug ditches and canals to drain wetlands and improve their pastureland. In 1947, the Central and Southern Florida Flood Control Project was implemented by the U.S. Army Corps of Engineers in response to major floodi ng in the Kissimmee Ba sin. As a result of this plan, the Kissimmee River was cha nnelized, Disston’s inte r-lake canals were improved, and water control structures were built throughout the area. The goal of these actions was to use the chain of lakes for wa ter management, to provide room for water during the wet season and to stor e water during the dry season. This entailed stabilization of water levels, which historically fluctuated up to 3 m (10 ft), to a 0.6-1.2 m (2-4 ft) range. This reduction in fluctuation subs equently allowed landowners and private citizens to build on historic lake bottom and drained we tlands within the floodplain,

PAGE 15

4 further strengthening the need for tight fl ood control (Blake 1980, HDR Engineering, Inc. 1989). Lake Tohopekaliga and surrounding lakes have suffered many water quality problems due to the intense hydrologic modifica tions. The constructed canals, especially in the Shingle Creek area, allowed direct conveyance of stormwater runoff and sewage into the lakes without the be nefit of filtration through wetlands. Urban and agricultural land use continued to expand, contributing more and more overland pollution. The agricultural land in the area is mainly utilized as pasturela nd, and historically dairy farms provided significant inputs of nutrients. Although eutrophication within the lakes was rapidly increasing, water level stabilization pr evented natural fluctuations from mitigating the problem. Since the lake levels were unnaturally restricted fr om periodic flooding and drying events, thick stands of vegetation bega n invading the littoral zone, which in turn led to organic sediment buildup and decrease d water quality. As of 1988, no wastewater discharges have been permitted to the lakes. However, non-point source urban and agricultural runoff and septic tank leakage remain major contributors to eutrophication in the Kissimmee chain of lakes (HDR Engineering, Inc. 1989). As mentioned, eutrophication has caused major vegetation changes to the littoral zone of Lake Tohopekaliga. Dense monotypic expanses of aquatic vegetation began to dominate the gradually sloping shoreline, fo rmerly characterized by sandy substrate and sparse vegetation. Nuisance species such as Pontederia cordata (pickerelweed) and Typha domingensis (cattail) formed wide bands of habitat around the lake. Pontederia cordata and associated species crea ted floating mats on the lakewa rd edges of the littoral zone that rose and fell with the water level. Exotic species such as Hydrilla verticillata

PAGE 16

5 (hydrilla), Eichornia crassipes (water hyacinth), Alternanthera philoxeroides (alligator weed), and Panicum repens (torpedo grass) also benef ited from increased nutrients and high boat traffic between waterways and became a major focus of lake managers through the years. Turnover in the vegetation comm unity produced an organic muck substrate within the littoral zone. Documented faunal responses to this changing habitat have included declines in fisheries, especially s port and forage fish species, and invertebrates (HDR Engineering, Inc. 1989). In 1968, the first fish population surveys in Lake Tohopekaliga were conducted by the State of Florida Fish and Wildlife C onservation Commission (FFWCC, formerly Florida Game and Fresh Water Fish Commission, FGFWFC) with rotenone, electroshocking, and trammel nets (Wegen er 1969). Management recommendations included drawdowns every 5-7 years in order to oxidize the increasingly organic substrate and provide benefits to the growing fish population upon reflooding of littoral habitats. In 1971, Lake Tohopekaliga underwent its firs t extreme drawdown. The water stage was dropped about 2.1 m (7 ft) from high pool stage (16.8 m, 55.0 ft NGVD). Drought conditions kept water levels below 15.8 m (52.0 ft) NGVD (low pool stage) until one year after the initiation of th e drawdown. The dewatering of th e littoral zone attained the management goals, with increased acreage of desirable plant species, greater production of fish and fish-food organisms (invertebra tes) per acre, and in creased sportfishing success (Wegener and Williams 1974). Improvements made to Lake Tohopekaliga’s li ttoral zone were short lived, due to continued input of nutrients (e.g., about 53 million liters (14 million gallons) of sewage waste per day discharged into the lake (Wegener and Williams 1974)) and water level

PAGE 17

6 stabilization. An offshore berm was forming at the low-pool water li ne that was thought to be acting as a barrier to fish and inverteb rates at low water levels (Moyer et al. 1987). A second drawdown was conducted in 1979 and al so had beneficial results, although by 1986 the habitat had degraded once more to s ub-optimal fishery habitat. The organic berm in the lakeward portion of the littoral zone was increasingly becoming a management issue not able to be addre ssed by drawdowns alone (Moyer et al. 1987). When the third drawdown was performed in 1987 a pilot muck removal project was included. Along 19 km (12 miles) of shoreline, 164,830 cubic meters (225,000 cubic yards) of muck were mechanically removed fr om the organic berm. This was considered “an unprecedented large-scale restoration project to improve littoral habitat” (Moyer et al. 1993, Appendix 4, page 2). All research point ed to highly positive results from the drawdown and muck-scraping procedures. Fi shery surveys have continued since 1968, and now include roving creel surveys, blocknet/rotenone sampling, electrofishing, experimental gill nets, and shallow water sa mpling with Wegener rings. Other wildlife monitoring included snail kite ( Rostrhamus sociabilis) individual and nest counts, limited aquatic plant sampling, and littoral zone i nvertebrate community monitoring (Moyer et al. 1993). Another lake enhancement project had been planned for early 2002 in Lake Tohopekaliga, however logistical constr aints caused postponement until early 2004 (Florida Fish and Wildlife Conservation Commission 2003). The project originally included an extreme drawdown from 16.8 m (55.0 ft) NGVD to 14.9 m (49 ft) NGVD beginning in November 2003, as well as the mechanical removal of about 5.4 million cubic meters (6.8 million cubic yards) of muck and vegetation from the majority of the

PAGE 18

7 lake’s shoreline (Florida Fish and Wi ldlife Conservation Commission 2003). The subsequent estimate of the actual volume sc raped was 7.3 million cubic meters (8-million cubic yards), with 1,351 ha (3,339 acres) of s horeline habitat remove d. The entire width of the littoral zone was targeted for re moval, not just the organic berm. The Pontederia cordatadominated habitat underwent widespread elimination throughout the lake. Twenty-nine in-lake disposal islands were cr eated from much of the scraped material (Florida Fish and Wildlife Conservati on Commission 2004). On ce the water levels recover, heavy applications of herbicides will be used to keep the habitat in an early state of succession, allowing lake managers to selectively allow regr owth of desirable vegetation. Currently, the target conditions post-enhancement are undefined. The main objective of the Lake Tohopeka liga habitat enhancement project is the removal large expanses of undesira ble macrophyte stands, particularly Pontederia cordata and Typha domingensis as well as the organic substrate (muck) associated with this dense vegetation (Florida Fish a nd Wildlife Conservati on Commission 2003). Reptiles, amphibians and many juvenile fish species are known to occupy structurally complex lentic habitats and u tilize the muck and thick vege tation for foraging, cover, and also reproduction (e.g., amphibi ans). Lake enhancement techniques (both mechanical vegetation and muck removal and subsequent herbicide applications) modify these resources, changing the habitat suitability fo r aquatic vertebrates. High mortality during the scraping process and migration during the drawdown will likely also alter the community structure and dynamics. The effort to sustain high species diversity in the lake ecosystem may be important to the st ability of the system, and by examining the

PAGE 19

8 consequences of these restoration techniques managers can better eval uate their worth to wildlife and fishery species. Research Objectives While some positive responses have been documented for the fishery of Lake Tohopekaliga following past enhancement pr ojects, many wildlife guilds have been neglected. There is limited qua ntitative knowledge of vegetati on responses to mechanical removal and large-scale herbicidal treatment. It is also uncertain how wetland birds are affected. There are still unanswered questions regarding aquatic vert ebrates that utilize the thick vegetation and organic sediment, including reptiles, amphibians and fish. Herpetofaunal responses to enhancement activitie s have not been studied in the past, even though they are pervasive in the habitat. Although fishery science claims that the eutrophic littoral habitat is unsuitable for centrarchids (i.e. sport fish), conventional sampling methods may be incapable of detec ting them in highly vegetated areas (Parker 1970, Allen et al. 2003). The current study is part of a larger pr oject evaluating the wildlife response to habitat enhancement in Lake Tohopekaliga. Al so included in this project are vegetation (see Welch 2004) and avian monitoring studies. The research presented in this thesis examines the aquatic vertebrate community in the littoral zone of Lake Tohopekaliga prior to the 2003 drawdown and mechanical ve getation and muck scraping activities. The littoral zone is defined here as the ar ea occupied by emergent vegetation. However, there is particular emphasis gi ven to the pickerelweed zone due to its extensive removal during the lake enhancement. The large-scale wildlife habitat investigation will continue for at least three years after enhancement activities to examine responses to the modifications by the different guilds.

PAGE 20

9 With a large-scale habitat modification, qua ntifying the effect on a whole suite of species provides maximum information. While most of the species have common biological or ecological traits, th ey also constitute a variety of habitat requirements based on food sources, reproduction methods, and movement patterns. For this reason community metrics within the land-water ecoto ne are of main concern, as represented by species richness and commun ity composition. Species-specific site occupancy and capture frequencies also facili tate understanding of habitat utilization by focal vertebrate species. The main objectives of this research are to 1. Characterize the vertebrate faunal makeup of Lake Tohopekaliga’s littoral zone prior to the 2004 lake enhancement project, 2. Estimate parameters such as density and activity/home range for focal species in the P. cordata habitat, 3. Estimate site occupancy rates for focal species within the littoral zone, as an estimate of the proportion of the area that the species inhabits, 4. Document how temporally changing variab les including lake stage, water level fluctuation, air temperature, and rainfall sh ape the aquatic vertebrate community in the P. cordata zone, and 5. Investigate the influence of spatial vari ables such as water depth and vegetation community on the herpetofauna and fish within the landscape.

PAGE 21

10 CHAPTER 2 DESCRIPTIONS OF FOCAL SPECIES Aquatic Vertebrate Habitat Wetland communities of reptiles and amphibians show much diversity in ecological function. Often being the largest and most abundant vertebrates in this habitat, they have important places in the food we bs of lakes (Iverson 1982). Some species provide terrestrial links while others are fully aquatic and never leave the littoral zone (Joly and Morand 1997). Fish species also rely on both the littoral, pelagic, and to some extent flooded terrestrial (nurse ry) habitats as they undergo sh ifts with life stage (Werner 2002). There has been a worl dwide decline in biodivers ity, particularly seen in amphibian species. A variety of human disturbances have been identified, including climate change, habitat loss and fragmenta tion, introduced species, pollution, acid rain, and disease (Reaser 2000). Florida in particular has been severely impacted by destruction of wetlands, channelization of st reams, manipulated hydrologic cycles, and rapid human growth (HDR Engineering, In c. 1989, Pough et al. 2001). Alteration of freshwater habitats has been a problem for many aquatic species. Animals that are longlived or have delayed sexual maturity, low reproductive rates, or poor dispersal or colonization abilities are particularly vulnera ble to habitat destruction (Klemens 2000). Purposeful habitat modification should preser ve conditions necessary for aquatic animals to complete their life cycles, including a ppropriate nesting/spawning, foraging, and cover habitats.

PAGE 22

11 Species of aquatic vertebrates that ar e most at risk due to their habitat requirements are emphasized here. Most he rpetological research has been conducted on breeding populations of amphibians, large ch arismatic reptiles, or single species and guilds (but see Bancroft et al. 1983). Fishery science remains focused mainly on sportfish at the individual or population le vel (Miranda and Dibble 2002). Resident littoral zone species make up the assemblage of interest for this study and represent several different orders of animals with a variet y of life history traits. Fish guilds, such as juvenile centrarchids (especially Lepomis spp., Micropterus salmoides ) and exotic catfish ( Hoplosternum littorale ), are focused upon. Documenta tion of the presence of these species in heavily vegetated littoral habitats in Florida is very poor, probably due to inadequate sampling techniques. Reptile and amphibian species of in terest include fully aquatic salamanders ( Siren spp., Amphiuma means ), water snakes (mainly Nerodia spp.), small kinosternid turtles ( Kinosternon baurii, Sternotherus odoratus ), and large aquatic frogs ( Rana spp.). Minimal research has been conducted on the effects of lake management techniques on these herpetofaunal species. Most of th ese species and guilds have a common reliance upon vegetated wetlands for at least some part of their life cycles. They also are often preyed upon by the same species, including alligators, wading and predatory birds, large predator y fish and aquatic sn akes, and together represent many segments of the food web in the lake ecosystem. Fish Species Centrarchids (sunfish) Most species in the family Centrarchi dae in Lake Tohopekaliga are sportfish. Foremost among them in Florida lakes is the largemouth bass ( Micropterus salmoides ). This species is the primary target for be nefit by the Lake Tohopekaliga enhancement.

PAGE 23

12 Bluegills (Lepomis macrochirus) redear sunfish (Lepomis microlophus) black crappie ( Pomoxis nigromaculatus ), warmouths ( Lepomis gulosus) spotted sunfish ( Lepomis punctatus), and dollar sunfish ( Lepomis marginatus ) are also considered sportfish in Florida. Enneacanthus gloriosus (bluespotted sunfish), has a maximum total length of 80 mm, and is therefore only c onsidered a forage fish speci es (Hoyer and Canfield 1994). Most of these species depend upon the vegetate d littoral zone during juvenile stages and for spawning. Vegetated habitats provide juve nile sunfish with protection from larger predators and abundant food supplies (W erner and Hall 1988, Chapman et al. 1996, Miranda et al. 2000). The phenom enon of ontogenetic habitat shif ts is particularly well studied in bluegills. This species move s between the littoral to the pelagic zone throughout its life cycle. The lit toral zone provides nesting hab itat, as well as a preferred environment for juvenile bluegills from a pproximately 12-83 mm standard length due to size-specific predation ri sks (Werner and Hall 1988). It is claimed that these species have no access to Lake Tohopekaliga’s littoral zone due to physical and chemical barriers. The floating mats of Pontederia cordata, resulting from the eutrop hic status of the lake, are thought to form a physical barrier for centrarchids, limiting adult access to shallow water spawning sites. Even if the fish could penetrate this barrier, physicochemical char acteristics of the dense vegetation would not permit survival (Moyer et al. 1995, Allen and Tugend 2002, and Allen et al. 2003). Traditional methods of fish sampling in high-macrophyte littoral habitats in Lakes Tohopekaliga and Kissimmee, Florida, have yielded few or no centrarchid species (Moyer et al. 1993, Allen and Tugend 2002). However, since common fish sampling methods, including electrofishing and rotenone/b locknet, do not perform well in heavily

PAGE 24

13 vegetated habitats (Parker 1970, Moyer et al. 1995, Allen and Tugend 2002), many suppositions upon which lake enhancemen t projects depend are theoretical. Exotic catfish Hoplosternum littorale is an exotic species in Fl orida, originating in South America. This armored catfish was first found within the United Stat es in South Florida in 1995, and was presumably released thr ough the aquarium trade or aquaculture. Various life history and beha vioral traits, including aeria l respiration, large body size, high environmental tolerances, and nest-guard ing behaviors, are re sponsible for rapid expansion of its range in Florid a (Nico et al. 1996). This sp ecies is currently nesting in and pervasive throughout the littoral zone in Lake Tohopekaliga (personal observation). Pterygoplichthys spp. (suckermouth or sailfin catfish) has also been captured in Lake Tohopekaliga, although only on a few occasions. This species was proba bly released into Florida through the aquarium trade (Page 1994) These two species may pose significant ecological threats to native food webs and aquatic plant communities. While the Florida Fish and Wildlife Conservation Comm ission conducts yearly monitoring by electrofishing, this study is th e first known report of these sp ecies this far north in the Kissimmee chain of lakes. Herpetofaunal Species Amphibians Rana grylio (pig frog) is a highly aquatic speci es, rarely being seen on shore. They are usually associated with dense ma rsh vegetation. While leopard frogs, including Rana sphenocephala (Florida leopard frog), prefer habi tats with standing water, larger individuals can inhabit somewh at dryer on-shore habitats an d use larger home ranges, relying on plant shade, dew and soil moisture for survival (Dole 1965). Adult leopard

PAGE 25

14 frogs and their tadpoles are also noticeably absent from sandy, unvegetated shorelines (Dole 1965, Alford and Crump 1982, Bancroft et al. 1983). These two large frog species have differences in length of larval developm ent, with pig frogs taking more than a year to metamorphose and leopard frogs taking only two to three months (Bancroft et al. 1983). This, along with year-round br eeding in Florida, results in a variety of size classes of tadpoles throughout the year. Siren lacertina (Greater siren) and Amphiuma means (Two-toed amphiuma) are two of the largest sp ecies of salamanders in the wo rld (Petranka 1998). Amphiumas depend on lungs for aerial respira tion, while sirens have extern al gills as well. Although they may have lengths greater than 76 cm (Conant and Collins 1998), diminutive limbs in both species are thought to limit overland disp ersal. These salamanders burrow into organic sediment when their habitats become dry and may remain alive for up to three to five years in underground burrows without food until water comes back to the habitat (Martof 1969, Etheridge 1990). Bancroft et al. (1983) found that the density of amphiumas and greater sirens in Lake Conwa y, Florida, increased with sediment depth. They also reported that neither species i nhabited sandy, unvegetated shorelines. Sirens have compressed tails that may help to prope l them in vegetated open water as well as emergent vegetation habitats. Amphiumas on the other hand have l ong cylindrical tails and are thought to be limited to shallow water (Bancroft et al. 1983). Sirens feed mainly on mollusks, insects, crayfish and filamentous algae, as well as some other vegetation. Amphiumas eat fish, crayfish, salamanders, fr ogs and a wide variety of other species (Petranka 1998).

PAGE 26

15 Reptiles The striped mud turtle ( Kinosternon baurii ) and common musk turtle ( Sternotherus odoratus ) are both small species (maximum carapace lengths of 12.2 cm and 13.7 cm respectively) that prefer shallo w water wetlands (Conant and Collins 1998). They are both omnivorous, feeding upon animal s and some plants opportunistically. However, mud turtles are attracted to fast -moving prey while musk turtles search out more sedentary organisms as they crawl along the substrate in search of prey (Mahmoud 1968). Striped mud turtles usually occur in water greater than 60 cm deep, with lower water levels or rainfall tri ggering terrestrial activity (W ygoda 1979, Ernst et al. 1994). On the other hand, common musk turtles are highly aquatic, not leaving water unless nesting. This species seems to prefer water de pths less than 60 cm, but have been seen in up to 9 m of water (Ernst et al. 1994). Ba ncroft et al. (1983) found about 20% of all captured common musk turtles in the littora l zone, and the rest (usually larger individuals) in open water habita t. According to Mahmoud (1969), S. odoratus is found in lakes as well as riverine habitats with gravel or sandy substrates. Nerodia fasciata pictiventris (Florida water snake) is most often encountered in the shallowest regions of inhabited wetland s (Ernst and Ernst 2003). They are observed often in disturbed and white sand littoral habita ts (Bancroft et al. 1983) This species eats mainly fish until they reach a total length of 50 cm, at which point they switch to preying upon frogs (Mushinsky et al. 1982). Nerodia floridana (Florida green water snake) is the largest North American water snake, with to tal lengths approaching two meters (Conant and Collins 1998). They are inhabitants of qui et water wetlands and sometimes venture out into open water (Ernst and Ernst 2003). Bancroft et al. (1983) found them to be pervasive throughout the littora l zone, and while the dense vegetation seems to be

PAGE 27

16 preferred, the species of vege tation may not be very importa nt. Some individuals were captured up to 40 m from the edge of the lit toral zone in open water, while several terrestrial sightings occurre d during winter months. Se diment depths of 11-20 cm yielded the most individuals, and sandy beach habitats were avoided by Florida green water snakes (Bancroft et al 1983). They feed mostly upon fish, but also on frogs, salamanders, tadpoles, small turtles and invertebrates (Mushinsky and Hebrard 1977, Ernst and Ernst 2003)

PAGE 28

17 CHAPTER 3 ASSEMBLAGE WITHIN THE Pontederia cordata COMMUNITY Introduction The objective of this section is to inves tigate the temporal va riation of community composition and dynamics. The four main rese arch questions are 1) is there temporal variation in the aquatic vertebrate assemb lage, 2) does community composition change over time, 3) what environmental factors seem to be influencing the temporal variation in the assemblage and individual focal species, and 4) how are the focal species dispersed through the habitat. Key envi ronmental variables that change over the course of a year include lake stage, water leve l fluctuation, air temperature, and rainfall. Each of these will be examined for their influence on the vertebrate assemblage. The thick P. cordata (pickerelweed) habitat was the prime target for mechanical removal during the lake enhancement process and therefore was the focu s of sampling effort. This protocol was also used to select focal species (which sp ecies were present in the habitat and most detectable with the traps) and evaluate trap -sampling methods. All of this information will facilitate monitoring in the future, regard ing how, when and where to sample in order to capture the community dynamics and vari ances associated with the lake’s everchanging environment. Field Methods Trap Descriptions As previously mentioned, fishery surv eys conducted in the Kissimmee Chain of Lakes include roving creel surveys, bl ocknet/rotenone sampling, electrofishing,

PAGE 29

18 experimental gill nets, and Wegener rings (Moyer et al. 1993). These methods collect information on a variety of fish species, but spor t fish are the typical target of research. Traditional herpetofaunal sampling techniques include visual surveys and hand or dip-net collecting (Bury and Corn 1991), pitfall and funne l traps in combination with drift fences (Corn 1994), and use of seines or dredges fo r removing floating vege tation along with the animals inhabiting it (Bancroft et al. 1983). None of these methods are appropriate for the extremely thick, rooted vegeta tion in the littoral zone. Turt le traps exist, such as hoop nets and floating traps for basking turtles (Lagler 1943); however la rge turtles are not central to this research since they are not re stricted to the littoral zone. PVC pipes have also been used as passive tr aps for treefrogs (Moulton et al 1996), and audio surveys are often used for breeding ranid frogs (Zi mmerman 1994). However, a single, allencompassing technique was desired for th is community study, and the answer came from funnel traps. Recently, several research ers have noticed the be nefits of capturing aquatic organisms in thick vegetation with crayfish and minnow style funnel traps (Darby et al. 2001, Sorensen 2003, Johnson and Barich ivich 2004). Without the use of either bulky drift fences or bait, these traps have b een successful in capturing a wide variety of reptiles, amphibians, fish and some invert ebrates. Funnel traps were used for all sampling during this study. The minnow and crayfish traps were all constructed of 1.3 cm (0.5 in) mesh, dark green vinyl-coated hardware cloth (Figure 31). The crayfish traps, similar to those described by Darby et al. ( 2001), were positioned on the subs trate, or as near to the substrate as the vegetation would allow. Th ey were approximately 80 cm (30 in) tall including a “chimney” extending from the body of the trap, allowing the top to be above

PAGE 30

19 the water surface. At the base were three en try funnels leading into the trap, with each opening about 6 cm (2.5 in) in diameter, but the exact size varied slightly due to handmade construction. The modified minnow tr aps were 60 cm (24 in) long rectangular traps, which were approximately 25 cm (10 in) deep and 18 cm (7 in) high. At each end there was one entry funnel, with an opening approximately 9 cm (3.5 in) wide and 6 cm (2.5 in) tall. Floats made of Styrofoam pool toys (“Wacky Noodles”) were attached to the minnow traps to allow them to float halfwa y out of the water, wi th the funnels about even with the water surface, based on the design by Casazza et al. (2000). The funnels permitted animal access into the trap, but discouraged escape by making the exits harder to find than the entrances. By allowing the traps to remain partially above the water, the animals had access to air and mortality was re duced. Both nocturnal and diurnal species were accessible to capture since traps could be deployed without time constraint. The traps were not baited, however once an anim al was captured in the trap other animals may have been attracted to it. The dimensions of the traps restricted the assemblage of animal species captured. The traps did not confine young individuals or small species of fish, frogs, snakes and salamanders due to the 1.3 cm (0.5 in) mesh size. Also, individuals larger than the funnel diameter were excluded. To compare the di fference in species captured with 1.3 cm (0.5 in) versus 0.6 cm (0.25 in) mesh, 18 commerc ially-manufactured minnow traps, similar to the “eelpots” used in Casazza et al. (2000) were deployed at randomly assigned trap points from 11/5/2003 to 1/8/2004. These traps are cylindrical, about 60 cm (24 in) long and 23 cm (9 in) in diameter, with the funnel openings about 5 cm (2 in) in diameter. They were also fitted with floatation. Th e hardware cloth was bare metal, not vinyl-

PAGE 31

20 coated. Comparisons of species and number of captures were made between the 0.6 cm mesh minnow traps and the 1.3 cm mesh modi fied minnow traps from the same trap points during this sampling period. We exp ected to capture more species with the smaller mesh size since small species and younger individuals could escape from the larger mesh, but be retained by the 0.6 cm holes. Whole-Lake Sampling To gain information regarding temporal ha bitat utilization by the aquatic vertebrate assemblage, sampling was conducted around the pe riphery of the whole lake to maximize the inference of the results to the system. For the whol e-lake sampling, 18 sites were randomly selected from the less developed, southe rn two-thirds of the lake (Figure 3-2). At each site, a transect was established with three trap locations placed perpendicular to shore and spaced approximately 10 m (33 ft) apart, except in disturbed stretches of habitat with barriers wi thin this distance (e.g., commercial airboat trails). One crayfish and one minnow trap were placed at each tr ap location, attached to a PVC pole for extra stability. The result of this trapping arra ngement was uniform sampling effort at each transect. The trap locations were placed in the most lakeward portion of dense P. cordata when possible (mainly in the 0.6-0.9 m (2-3 ft) depth zone at 13.8 m (55 ft) NGVD). The transect sites varied in proximity to the ecotone between the open water habitat and the vegetation. Most transect s had thick stands of Typha or more diverse floating mats between the relatively monotypic sections of pickerelweed an d open water. The band of emergent macrophytes at these locations was comparatively br oad. On the other hand, at some transects the traps were relatively clos e to this ecotone due to narrowness of the pickerelweed zone at these locations, well established commercial airboat trails, or herbicide applications near the transects pr oviding large unvegetated areas. One transect

PAGE 32

21 fell in an area where the substrate had pr eviously been scraped, and the vegetation consisted mainly of Hydrilla verticillata (hydrilla) and very few emergent macrophytes. The whole-lake trap survey was c onducted year-round, pending suitable water levels (greater than approximately 16 m, 52.5 ft NGVD). Below this point, there was not enough water for the trapped animals and rode nts and birds were in advertently captured. Sampling throughout the year 2002 was as follows: January 24 – Traps were deployed to ra ndomly selected transects and sampling began. May 2 – Insufficient water levels in the pickerelweed zone caused traps to be removed and sampling suspended. June 12 – Redeployment of traps to select transects with su fficient water depth resulted in decreased trapping effort until July 24. July 24 – All traps were back in place in fixed sampling locations. December 3 – Traps were removed from pickerelweed zone due to low water associated with the attempted 2002 drawdown. When active, the traps stayed in place day and night and were typically checked once weekly. Despite efforts to keep sample s spaced seven days apart, the time interval was not always consistent due to logistical issues (e.g., air boat problems, rough weather). At each sampling occasion, two or three observers traveled to each transect in an airboat and checked the traps for their contents. All animals were brought b ack to the boat to be worked up. Reptiles and amphibians were weighed individually with Pesola spring scales and certain length measurements were taken, depending on the species. Fish were identified and grouped according to species for each trap. All individuals of each species per trap were weighed together in order to obtain the total biomass of the fish species caught. After being worked up, the animals we re released at the transects where they were captured. The types of data collected with these methods include species detection-

PAGE 33

22 nondetection, number of individuals capture d on each sample occasion, biomass, and reptile and amphibian length measurements. This sampling protocol was intended to c ontinue in the exact same locations postlake enhancement (2003) in order to compar e community traits be fore and after the modifications. However when the drawdown and muck removal was postponed for another year, it was no longer beneficial to keep sampling since there was not a before/after comparison to be made. Variab les such as water temperature and dissolved oxygen were not measured directly since this was not the initial focus of the study. Alternative environmental variables were obtai ned using Internet resources. Lake stages and rainfall were taken from the South Fl orida Water Management District’s DBHYDRO browser ( http://glades.sfwmd.gov/pls/dbhydro_pro_plsql/ ). The lake stage was the mean daily average taken from the headwater of Sta tion S61 (the water control structure in the south part of the Lake Tohopekaliga leading to Lake Cypress via the South Port Canal) in feet NGVD. Lake stage was recorded for the day of each sample occasion. Water fluctuation for one sample is the difference of the water level at that sample minus the water level at the previous sample occasion. Rainfall was also recorded at Station S61 and precipitation totals for each sample we re added up from the day of the previous sample occasion until the day before the new sample occasion. Air temperature data was gathered from the National Oceanic and Atmo spheric Administration’s National Climatic Data Center’s website ( http://www.ncdc.noaa .gov/servlets/ULCD ). The weather station location was the Orlando Inte rnational Airport (M CO) in Orlando, Florida. This is located approximately six kilometers (10 miles) from the north shore of Lake Tohopekaliga. Average temperatures were calculated for every sample occasion by

PAGE 34

23 averaging the daily average temperatures from the day of the previous sample occasion until the day before the new sample occas ion. Minimum and maximum temperatures were also recorded for each sample period. Analysis Methods Trap Comparisons To determine the utility of traps with smaller mesh si ze for this study, the species and number of captures for the 0.6 cm (0.25 in ) mesh commercial minnow traps and the 1.3 cm (0.5 in) mesh modified minnow trap s were compared. The 0.6 cm mesh traps were randomly placed at only a portion of the whole-lake trap sites, along with a 1.3 cm mesh crayfish and minnow trap. For this reason, data from both minnow traps were compared for just the trap sites with both mesh types. Species Richness Sampling was carried out with a repeated measures protocol, potentially resulting in lack of independence be tween samples. However, assuming random movement of individuals and species through the habitat over space and time, sampling over time did not result in repeated captures of the same individuals. Th is transient nature of the species and utilization of non-parametric pr ocedures for most analyses are believed remove potential bias due to re peated samples. To determin e the presence of temporal variation in the aquatic vertebrate assembla ge, total species richness was calculated for each sample occasion in 2002. Fish and he rpetofaunal species richness were also individually estimated for each sample o ccasion. Program COMDYN4 was used with detection-nondetection data to estimate richness (Hines et al. 1999), taking into account species detection probabilitie s. It uses a model (Mh) that allows each species to have a different detection probability (the probability of detecting at leas t one individual of the

PAGE 35

24 species). Since most species detection proba bilities are less than one raw count data can result in underestimations of richness. In a similar manner, the term “presence/absence data” can also be misleading since lack of detection provides no evidence of a species’ absence from the trap site. For this reas on I instead use the term “detection/nondetection data” throughout this thesis. Equal sampli ng effort is necessary for each occasion. Assumptions of this method are 1) populati on closure for species, 2) independence of captures and 3) individual species capture pr obabilities stay cons tant during sampling (Burnham and Overton 1979). However, this me thod is robust to de viations from these assumptions. Even when the assumptions are violated the model-based richness estimates are less biased than counts of species (Nichols et al. 1998). Data from seventeen of the eighteen transe cts were used in the richness analysis. The one transect that was located in the prev iously scraped habitat was removed from the analysis in order to focus solely on variations within the P. cordata habitat. Since species capture data were fairly sparse for each tr ansect per sample occas ion, the transects were randomly assigned to six groups that represent sample replicates across space. They were randomly grouped in order to remove effects of shoreline characteristics at different transects. Richness was not estimated for sa mple occasions with reduced trap effort (sample occasions 14-20). Linear regre ssions were performed using SPSS (SPSS Inc. 2001) in order to determine significant predictors of the vertebrate, fish and herpetofaunal species richness. Richness estimates from each sample occasion were used in these analyses. There were three outliers greater th an two standard deviations from the mean, which were removed for the herpetofaunal re gression analyses. Average air temperature

PAGE 36

25 (oC), lake stage (m), rainfall (cm), and water level fluctuation (cm) were used as the independent variables. Assemblage Composition Species richness estimates the number of species present, but indicates nothing about community composition. To compare the pr esence of vertebrate species over time, sample occasions were assigned to clusters us ing hierarchical cluste r analysis. This was run using PC-Ord software (McCune and Mefford 1999) with detection-nondetection data of species for each sample occasion. Sore nsen's distance measure with the flexible beta (beta=-0.25) linkage method was used. Indicator species analysis (McCune and Grace 2002) was applied to determine the most appropriate number of clusters and the best species to represent those groups. Any groups comprised of a single sample occasion were removed from the indicator sp ecies analysis. A Monte Carlo procedure was run 1000 times with randomized data to cal culate a p-value for each species, which tested the null hypothesis that their indicator values were no larger than would be expected by chance. The optimum number of groups was selected by the indicator species analysis that yielded the most speci es with statistically significant indicator values (McCune and Grace 2002). Influence of Temporal Gr adients on Assemblage Multivariate ordination was used to establish what temporally changing environmental factors were influencing th e variation in assemblage composition. Nonmetric multidimensional scaling (NMS) is an ordination technique that uses ranked distances between sample un its to reduce dimensions a nd allow description of the community in relation to environmental gradie nts. The distances represent dissimilarity between sample units in terms of species composition. This method was chosen because

PAGE 37

26 it is particularly useful for non-normal data and many sampling events with no captures (McCune and Grace 2002). Sample units, i.e. individual sample occasions, are plotted in species space using an iterat ive search for the optimal placement for the sample units. Optimal placement is determined by the maximum possible reduction in stress, which is a measure of dissimilarity between the origin al data matrix and the reduced-dimension final ordination. PC-ORD software was used for all NMS analyses (McCune and Mefford 1999). NMS was run for the entire vertebrate a ssemblage for all sample occasions using detection-nondetection and count data separa tely. Fish and herpetofaunal assemblages were then analyzed separately in the same fashion to determine if the environmental gradients affected them differently. Outliers were identified using the outlier analysis provided in PC-Ord, with the cr iteria being greater than two st andard deviations from the mean (McCune and Mefford 1999). All outliers were removed from the analyses. General relativizations by row were conducte d on the raw count data to equalize common and uncommon species and lower the coefficien t of variation (CV) of the row totals. Relativizations were followed by square root transformati ons to balance the relative importance of the species without altering their ranks. Sorensen’s distance measure was used to calculate dissimilarity matrices for the ordinations. Starting configur ations were created by random number seeds, which were generated by the time of day. Fifty runs we re conducted with the real data to find the solution with the lowest stre ss. Fifty Monte Carlo randomi zed runs were performed to select the appropriate number of dimensions that be st represent the variation in the data. Comparisons between the runs with real data and randomized data give a probability that

PAGE 38

27 final stress in the ordination could be found by chance. After the first 50 runs, the number of dimensions was determined and the fi nal NMS was rerun usi ng the random number seed from the initial ordination. From this the final stress and inst ability (fluctuation of stress per iteration) were evaluated. Measured environmental variables, includi ng lake stage, stage fluctuation, total rainfall, and average, maximum and minimum average air temperatures over the sample period, were included in the or dination graphs. The ordinations were plotted with environmental variables as biplots, indicati ng the strength of corre lations of variables with the synthetic axes. Only th e environmental variables with r2>0.2 (percent of variance represented) are shown in the ordina tion plots. Sample un its were color coded by their membership to the groups defined by the cluster analysis, representing different species composition. Proportion of Habitat Ut ilized by Focal Species Using the program PRESENCE (MacKenz ie et al. 2002), detection/nondetection data were analyzed to determine site occ upancy rates for all species with enough captures to get reasonable estimates. This method allows for numerous, representative, randomly selected sites (transects) within the much larger area of interest ( P. cordata zone) to be sampled for the presence of species. The in ference gained from sampling these sites can then be applied to the pickerelweed zone La ke Tohopekaliga. The main function of this method is to determine habitat usage for speci es with low detection probabilities (<1). Detectability is an important factor when sampling secretive aquati c organisms in thick vegetation. The program calculates (1) a “nave estimate,” which is simply the proportion of sites where the species was ca ught (considered biased low), (2) speciesspecific detection probabilities based on the capture data, and (3) the “proportion of sites

PAGE 39

28 occupied” (PSO) which is the nave estim ate corrected for detection probability (MacKenzie et al. 2002). This method assumes closure of species to changes in occupancy status over the course of sampling. However, if the sp ecies have large activity ranges and the movements are assumed to be random, the closure assumption may be relaxed (MacKenzie et al. 2002). The sample occasions were divided into two groups. The first is from the start of sampling at the beginning of February until the traps were removed at the beginning of May. The second group is fr om the beginning of August, when water levels allowed full sampling effort, to the end of sampling in November. Between these groups the lake stage became so lo w that there was no water in the P. cordata zone, which surely caused a violation of the closur e assumption for this method. This required the split of sample occasions into groups th at are assumed closed to species immigration or emigration. The first (spring) group incl udes 13 sample occasions for the herpetofauna and 11 sample occasions for the fish species, since fish were not recorded for the first sample and the last sample in the group had wa ter depths too shallow to capture fish. The second (fall) group has 16 sample occasions for all species. Parameters were estimated for each species for both groups using the single season models in PRESENCE. The data were analy zed using models with both constant and survey specific detection probabilities. Resu lts from the model with the lowest Akaike’s Information Criterion (AIC) value were reported for each species. If the AIC values were within two points of each other, the simple r, constant detection probability model was selected.

PAGE 40

29 Results Trap Comparisons All reptile, amphibian and fish species captured during the 2002 whole-lake sampling, along with species codes used in the figures ar e listed in Table 3-1. Due to restrictive funnel sizes, most fish species (especially centrarchids) were represented by juvenile life stages, except small species such as mollie s and killifish. On the other hand, adult individuals characterized the majority of the reptile and amphibian species, since most young individuals could escape through the me sh. Table 3-2 shows the species and number of captures for the two types of minnow traps. Eleven vertebrate species were captured with the 0.6 cm (0.25 in) mesh minnow traps, while 19 species were captured in the 1.3 cm (0.5 in) mesh minnow traps at the sa me sample locations. Three species were unique to the 0.6 cm mesh traps on th ese occasions: black swamp snake ( Seminatrix pygaea ), flagfish ( Jordanella floridae ), and mosquitofish ( Gambusia spp). Only three reptile or amphibian species were captured: black swamp snake, Florida leopard frog ( Rana sphenocephala ), and pig frog ( Rana grylio ). More tadpoles were captured with the 0.6 cm mesh (n=31) than with the 1.3 cm mesh (n=2). Nine species were unique to the 1.3 cm mesh traps: striped mud turtle ( Kinosternon baurii ), striped crayfish snake ( Regina alleni ), Florida water snake ( Nerodia fasciata pictiventris ), Florida green water snake ( Nerodia floridana ), siren ( Siren spp), redfin pickerel ( Esox americanus ), armored catfish ( H. littorale ), dollar sunfish ( Lepomis marginatus ), spotted sunfish ( Lepomis punctatus ), largemouth bass ( Micropterus salmoides ), and redear sunfish ( Lepomis microlophus ). Seven combined reptile and amphibi an species were caught with these traps. Species common to both traps were leopard frog, pig frog, bluegill ( Lepomis macrochirus ), bluespotted sunfish ( Enneacanthus gloriosus ), golden topminnow

PAGE 41

30 Table 3-1. All species captured in 2002, with species codes used in subsequent figures. Fish Species Scientific Name Family Species Code Armored catfish Hoplosternum littorale Callichthyidae HOPLI Black crappie Pomoxis nigromaculatus Centrarchidae POMNI Bluegill Lepomis macrochirus Centrarchidae LEPMAC Bluespotted sunfish Enneacanthus gloriosus Centrarchidae ENNGL Bowfin Amia calva Amiidae AMICA Chain pickerel Esox niger Esocidae ESONI Chubsucker Erimyzon spp. Catostomidae ERIMY Dollar sunfish Lepomis marginatus Centrarchidae LEPMAR Flagfish Jordanella floridae Cyprinodontidae JORFL Gar Lepisosteus spp. Lepisosteidae LEPIS Golden shiner Notemigonus crysoleucas Cyprinidae NOTCR Golden topminnow Fundulus chrysotus Fundulidae FUNCH Largemouth bass Micropterus salmoides Centrarchidae MICSA Pterygoplichthys Pterygoplichthys spp. Loricariidae PTERY Redear sunfish Lepomis microlophus Centrarchidae LEPMI Redfin pickerel Esox americanus Esocidae ESOAM Sailfin molly Poecilia latipinna Poeciliidae POELA Seminole killifish Fundulus seminolis Fundulidae FUNSE Spotted sunfish Lepomis punctatus Centrarchidae LEPPU Warmouth Lepomis gulosus Centrarchidae LEPGU Herpetofaunal Species Scientific Name Family Species Code Amphiuma Amphiuma means Amphiumidae AMPME Cottonmouth Agkistrodon piscivorous conanti Viperidae AGKPICO Fl. banded water snake Nerodia fasciata pictiventris Colubridae NERFAPI Fl. green water snake Nerodia floridana Colubridae NERFL Fl. snapping turtle Chelydra serpentina osceola Chelydridae CHESEOS Fl. softshell turtle Apalone ferox Trionychidae APAFE Leopard frog Rana sphenocephala Ranidae RANSP Mud snake Farancia abacura abacura Colubridae FARABAB Peninsula cooter Pseudemys floridana peninsularis Emydidae PSEFLPE Pig frog Rana grylio Ranidae RANGR Siren Siren spp. Sirenidae SIREN Stinkpot Sternotherus odoratus Kinosternidae STEOD Striped crayfish snake Regina alleni Colubridae REGAL Striped mud turtle Kinosternon baurii Kinosternidae KINBA Tadpole-leopard frog Rana sphenocephala Ranidae TADRANSP Tadpole-pig frog Rana grylio Ranidae TADRANGR

PAGE 42

31 Table 3-2. Species capture frequencies for the 0.6 and 1.3 cm mesh minnow traps. Vertebrate Species 0.6 cm mesh 1.3 cm mesh Armored catfish 0 2 Black swamp snake 1 0 Bluegill 2 1 Bluespotted sunfish 18 31 Dollar sunfish 0 2 Flagfish 147 0 Florida green water snake 0 4 Florida water snake 0 1 Gambusia 75 0 Golden topminnow 17 1 Largemouth bass 0 1 Florida leopard frog 4 2 Pig frog 2 7 Redear sunfish 0 1 Redfin pickerel 0 1 Sailfin molly 56 5 Siren 0 1 Spotted sunfish 0 2 Striped crayfish snake 0 1 Striped mud turtle 0 1 Tadpoles 31 2 Warmouth 2 6 ( Fundulus chrysotus ), sailfin molly ( Poecilia latipinna ), and warmouth ( Lepomis gulosus ). Species Richness Vertebrate species richness (Figure 3-3) was negatively correlated with average air temperature (r2=0.330, df=1, p=0.001) and rainfall (r2=0.173, df=1, p=0.028). Average air temperature was the only si gnificant predictor of richne ss for the fish (Figure 3-4), (r2=0.316, df=1, p=0.002), with higher species richness estimates occurring with lower air temperatures. The estimated richness of the herpetofaunal assemblage (Figure 3-5) was negatively correlated with lake stage (r2=0.327, df=1, p=0.003), rainfall (r2=0.166, df=1, p=0.043), and water level fluctuation (r2=0.211, df=1, p=0.021).

PAGE 43

32 Assemblage Composition Six clusters were chosen to represent th e 34 sample occasions. Thirteen indicator species were determined with p<0.05 (Table 33). Indicator values are given for these species, with 100 representing perfect indi cation of that group based on relative abundances and frequency of occurrence. A zer o indicates complete absence of a species from a particular group. Since these cryptic sp ecies are quite mobile (with respect to the traps) and dependent upon detection for quantif ication, indicator values are relatively low compared to vegetation studies where virtua lly all species are de tectable. Group 1 is identified by several species, including Amia calva (bowfin), Fundulus chrysotus (golden topminnow), Lepomis macrochirus (bluegill), Lepomis marginatus (dollar sunfish), Nerodia fasciata pictiventris (Florida water snake), Rana sphenocephala (Florida leopard frog), and R. sphenocephala and Rana grylio (pig frog) tadpoles. The second group consisted only of sample occasion #13 (an extremely low water sample), and was therefore omitted. Hoplosternum littorale (armored catfish) and Regina alleni (striped crayfish snake) are the indicator species for Group 3. There were no significant indicator species for Group 4, but Amphiuma means (amphiuma) and Lepisosteus spp. (gar) show the highest indicator values with 28 and 22 respectively. Sternotherus odoratus (common musk turtle) was the sole indicator species for Group 5. The two pickerel species, Esox americanus (redfin) and Esox niger (chain), were the only two species indicating Group 6. Some species were captured duri ng every sample occasion, including Enneacanthus gloriosus (bluespotted sunfish), Lepomis gulosus (warmouth), and Siren spp. (siren). Hoplosternum littorale and Kinosternon baurii (striped mud turtle) were found on almost every sample occasion. It is unclear why the armored catfish is an

PAGE 44

33 indicator species for Group 3, when it has an in dicator value of 22 for all groups but one. Also, the bluegill had indicator values of 47 for both Group 1 and Group 6, although it was assigned to Group 1 with p=0.028. Table 3-3. Indicator species analysis results. Significant indicator species are highlighted (p<0.05) and displayed with associated indicator values and the clusters to which the species was assigned. Species Code Cluster Max Indicator Value Mean Standard Deviation Probability AGKPICO 1 25 15 7.4 0.211 AMICA 1 48.5 18 9.86 0.017 AMPME 1 27.9 24.8 4.25 0.234 APAFE 5 9.1 15.3 7.72 1 CHESEOS 4 12.5 15.3 7.49 0.696 ENNGL 1 20 20 0.63 1 ERIMY 6 27.9 23.1 7.1 0.201 ESOAM 6 40.9 22.6 7.6 0.022 ESONI 6 43.3 18.1 10.46 0.022 FARABAB 6 21.3 18.2 10.31 0.29 FUNCH 1 43.8 21.3 8.26 0.04 HOPLI 3 22.2 21.3 0.82 0.039 JORFL 1 34.1 16.8 9.59 0.077 KINBA 1 20.5 20.6 0.73 0.672 LEPGU 1 20 20 0.63 1 LEPIS 4 22.3 24.9 3.41 0.76 LEPMAC 1 47.1 21.9 8.23 0.028 LEPMAR 1 36.8 23.8 7.01 0.05 LEPMI 6 33.8 22.3 8.45 0.1 LEPPU 5 12 22.2 8.8 0.976 MICSA 6 28.8 23 7.01 0.178 NERFAPI 1 40.8 22.7 7.03 0.018 NERFL 3 21.6 21.3 0.83 0.295 NONE 1 31.7 19.9 9.88 0.107 NOTCR 6 28.6 16.5 8.92 0.059 POELA 1 28.8 24.6 4.2 0.115 POMNI 1 25 14.9 7.44 0.196 PSEFLPE 5 9.1 15.1 7.52 1 PTERY 5 18.7 16.9 9.76 0.179 RANGR 1 22.4 23.4 1.3 0.92 RANSP 1 100 17.4 9.61 0.001 REGAL 3 72.6 17.8 10.18 0.002 SIREN 1 20 20 0.63 1 STEOD 5 29.8 24.4 2.19 0.001 TADRANGR 1 34.6 24.4 5.45 0.009 TADRANSP 1 75 17 9.16 0.001

PAGE 45

34 Influence of Temporal Gr adients on Assemblage Stable three-dimensional ordinations were produced with all NMS analyses except the herpetofaunal analysis w ith detection/nondetection data. The final solutions were based on the criteria of stress being redu ced by at least 5% with each additional dimension. The final stress values were lowe r with the real data than was found by the Monte Carlo randomized runs (p<0.05), which indicates that there was real structure found in the data. The final stress values fo r all ordinations (except for herpetofaunal detection/nondetection) were between 12 and 18 (Table 3-4), which are common values for ecological data and depict a fair portrayal of the da ta (McCune and Grace 2002). Table 3-4. Stress and instability results from all NMS ordinations Assemblage Data Type Final Stress Instability Iterations Vertebrate Detection/ nondetection17.65 0.00254 500 Vertebrate Counts 13.27 0.00049 69 Fish Detection/nondetection15.31 0.00095 29 Fish Counts 12.93 0.00016 49 Herpetofauna Counts 14.08 0.00045 38 For the NMS with vertebrate detecti on/nondetection data, Axes 1 and 3 best explained 61% of the variance found in th e assemblage composition. Environmental variables with r2>0.2 were shown as biplots on the pl ots and include la ke stage and air temperature measures (Figure 3-6, Table 3-5) Axis 1 was correlated with lake stage (r2=0.349). Axis 3 was most correlated with bo th lake stage and average air temperature, (r2=0.354 and 0.259 respectively). Sixt een species were correlated with either Axis 1 or 3 with r2>0.2 (Figure 3-7, Table 3-6), 10 of them being indicator species.

PAGE 46

35 Table 3-5. Percent of variance explained (r2) by environmental variables for each axis in the vertebrate NMS with detection/n ondetection data. Variables with r2 > 0.2 are highlighted. Axis 1 2 3 Variable r2 r2 r2 Stage(m) 0.349 0.021 0.345 Fluc(cm) 0.039 0.009 0.183 Rain(cm) 0.037 0.006 0.029 Max(C) 0.038 0.362 0.127 Min(C) 0.002 0.255 0.229 Ave(C) 0.009 0.375 0.259 Table 3-6. Percent of variance explained (r2) for each axis by species in the vertebrate NMS with detection/nondetection da ta. Indicator species with r2 > 0.2 are highlighted in blue, while all other species with r2 > 0.2 are highlighted in yellow. Axis 1 2 3 Axis 1 2 3 Species r2 r2 r2 Species r2 r2 r2 AGKPICO 0.073 0.002 0.094 LEPMI 0.035 0.56 0.018 AMICA 0.027 0.077 0.327 LEPPU 0.064 0.027 0 AMPME 0.002 0.171 0.01 MICSA 0.291 0.227 0.242 APAFE 0.035 0.004 0.001 NERFAPI 0.227 0.167 0.257 CHESEOS 0.021 0 0.063 NERFL 0.003 0 0 ENNGL 0.381 0.019 0.002 NONE 0.057 0.004 0.102 ERIMY 0.031 0.536 0.007 NOTCR 0.011 0.05 0.017 ESOAM 0.033 0.186 0.353 POELA 0.188 0.018 0.248 ESONI 0.126 0.166 0.089 POMNI 0.007 0.011 0.067 FARABAB 0.049 0 0.21 PSEFLPE 0.005 0.085 0.001 FUNCH 0.019 0.419 0.547 PTERY 0.002 0.081 0.038 HOPLI 0.376 0.003 0.155 RANGR 0.149 0.035 0 JORFL 0.087 0.047 0.15 RANSP 0.218 0.02 0.331 KINBA 0 0.07 0.016 REGAL 0.01 0.117 0.037 LEPGU 0.381 0.019 0.002 SIREN n/a n/a n/a LEPIS 0.36 0.007 0.083 STEOD 0.021 0.006 0.018 LEPMAC 0.002 0.311 0.53 TADRANGR0.016 0.127 0.311 LEPMAR 0.019 0.04 0.422 TADRANSP0.07 0.039 0.287 With vertebrate assemblage count da ta, Axes 1 and 3 had the highest r2. Together these axes represent 63% of the variance in the species composition. Axis 2 also had an r2>0.2, and was best correlated with air temperature measures. As with the detection/nondetection analysis lake stage and air temper ature measures showed the

PAGE 47

36 highest correlation with these axes (Figure 3-8, Table 3-7). Air temperature is correlated with Axis 1, with maximum air temperature having the highest r2 of 0.353. Axis 3 was most highly correlated with lake stage (r2=0.458) and average air temperature (r2=0.375). Seventeen species had an r2>0.2 for at least one of the axes nine of them being indicator species (Figure 3-9, Table 3-8). Table 3-7. Percent of variance explained (r2) by environmental variables for each axis in the vertebrate NMS with count data. Variables with r2 > 0.2 are highlighted. Axis 1 2 3 Variable r2 r2 r2 Stage(m) 0.146 0.010 0.458 Fluc(cm) 0.013 0.006 0.108 Rain(cm) 0.001 0.001 0.044 Max(C) 0.353 0.299 0.232 Min(C) 0.261 0.346 0.281 Ave(C) 0.298 0.379 0.375 Table 3-8. Percent of variance explained (r2) for each axis by species in the vertebrate NMS with count data. Indicator species with r2 > 0.2 are highlighted in blue, while all other species with r2 > 0.2 are highlighted in yellow. Axis 1 2 3 Axis 1 2 3 Species r2 r2 r2 Species r2 r2 r2 AGKPICO 0.16 0.005 0.095 LEPMI 0.241 0.331 0.139 AMICA 0.059 0.089 0.192 LEPPU 0.06 0.02 0.018 AMPME 0.183 0.124 0.211 MICSA 0.13 0.184 0.048 APAFE 0.044 0.021 0.017 NERFAPI 0.166 0.001 0.373 CHESEOS 0 0.002 0.087 NERFL 0.068 0.024 0.004 ENNGL 0.809 0.342 0.108 NONE 0.031 0.011 0.164 ERIMY 0.217 0.024 0.092 NOTCR 0.124 0.136 0.022 ESOAM 0.014 0.455 0.09 POELA 0.133 0.023 0.341 ESONI 0.051 0.108 0.044 POMNI 0.022 0.005 0.087 FARABAB 0.041 0.028 0.018 PSEFLPE 0.065 0 0.038 FUNCH 0.002 0.291 0.575 PTERY 0.063 0.005 0.115 HOPLI 0.05 0.285 0.653 RANGR 0.125 0.157 0 JORFL 0.186 0.04 0.172 RANSP 0.092 0.038 0.345 KINBA 0.43 0.003 0.005 REGAL 0.039 0.039 0.005 LEPGU 0 0.104 0.013 SIREN 0.428 0.46 0.172 LEPIS 0.005 0.158 0.288 STEOD 0.268 0.182 0.288 LEPMAC 0.079 0.423 0.386 TADRANGR 0.147 0.159 0.008 LEPMAR 0.023 0.182 0.4 TADRANSP 0.137 0.065 0.24

PAGE 48

37 The fish assemblage alone with detection/ nondetection data resulted in the first two axes representing 63% of the variance expl ained. Average temperature was most highly correlated with Axis 1 (r2=0.291), and also with Axis 2 (r2=0.368), while lake stage was correlated with Axis 2 (r2=0.275), (Figure 3-10, Table 3-9). For these two axes, there were a total of 11 species with an r2>0.2, with six indicator sp ecies (Figure 3-11, Table 310). Table 3-9. Percent of variance explained (r2) by environmental variables for each axis in the fish NMS with detection/nondet ection data. Variables with r2 > 0.2 are highlighted. Axis 1 2 3 Variable r2 r2 r2 Stage(m) 0.042 0.275 0.105 Fluc(cm) 0.008 0.044 0.062 Rain(cm) 0.001 0.031 0.018 Max(C) 0.250 0.177 0.115 Min(C) 0.205 0.330 0.106 Ave(C) 0.291 0.368 0.083 Table 3-10. Percent of variance explained (r2) for each axis by species in the fish NMS with detection/nondetection data Indicator species with r2 > 0.2 are highlighted in blue, while all other species with r2 > 0.2 are highlighted in yellow. Axis 1 2 3 Axis 1 2 3 Species r2 r2 r2 Species r2 r2 r2 AMICA 0.152 0.296 0.12 LEPMAC 0.361 0.646 0 ENNGL n/a n/a n/a LEPMAR 0.406 0.229 0.168 ERIMY 0.149 0.089 0.223 LEPMI 0.118 0.205 0.436 ESOAM 0.287 0.223 0.083 LEPPU 0.002 0.009 0.021 ESONI 0.195 0.066 0.013 MICSA 0.706 0.049 0.001 FUNCH 0.486 0.572 0.003 NOTCR 0.072 0.026 0.018 HOPLI 0.058 0.251 0.149 POELA 0.007 0.263 0.264 JORFL 0.038 0.181 0.137 POMNI 0.034 0.009 0.004 LEPGU n/a n/a n/a PTERY 0.253 0.003 0.046 LEPIS 0.021 0.339 0.004 For the fish ordination with count data, Axes 2 and 3 represent the most variation in the assemblage data with a cumulative r2 of 0.76. Again, air temperature and stage are

PAGE 49

38 the environmental variables most correlated wi th these axes (Figure 3-12, Table 3-11). Axis 2 was most highly correlated with maximum air temperature (r2=0.380), while lake stage (r2=0.287) and average air temperature (r2=0.333) are most correlated with Axis 3. Fourteen of the fish species had an r2>0.2 for these two axes, with six of them being indicator species (Fi gure 3-13, Table 3-12). Table 3-11. Percent of variance explained (r2) by environmental variables for each axis in the fish NMS with count data. Variables with r2 > 0.2 are highlighted. Axis 1 2 3 Variable r2 r2 r2 Stage(m) 0.070 0.005 0.287 Fluc(cm) 0.008 0.001 0.064 Rain(cm) 0.003 0.000 0.024 Max(C) 0.242 0.380 0.189 Min(C) 0.346 0.303 0.243 Ave(C) 0.319 0.354 0.333 Table 3-12. Percent of variance explained (r2) for each axis by species in the fish NMS with count data. Indicator species with r2 > 0.2 are highlighted in blue, while all other species with r2 > 0.2 are highlighted in yellow. Axis 1 2 3 Axis 1 2 3 Species r2 r2 r2 Species r2 r2 r2 AMICA 0 0.008 0.445 LEPIS 0.359 0.044 0.07 ENNGL 0.171 0.57 0.239 LEPMAC 0.085 0.308 0.503 ERIMY 0.221 0.367 0.135 LEPMAR 0.033 0.185 0.624 ESOAM 0.159 0.165 0.372 LEPMI 0.227 0.359 0.164 ESONI 0.002 0.252 0.156 LEPPU 0.074 0.274 0.006 FUNCH 0.113 0.106 0.757 MICSA 0 0.64 0.294 HOPLI 0.108 0.03 0.194 NOTCR 0 0.152 0.058 JORFL 0.004 0.046 0.303 POELA 0.17 0 0.423 LEPGU 0.191 0.337 0.223 POMNI 0.054 0.077 0.016 The herpetofaunal assemblage detection/ nondetection data were analyzed with NMS, but results yielded only a one-dimensi onal solution. Stress on the final run was 51.36, which represents an unacceptable amount of variation from the original data set. Values of stress greater than 20 indicate that the solution may be misleading, while ordinations with stress values over 40 represent very little of the structure in the original

PAGE 50

39 data matrix (McCune and Grace 2002). Due to these outcomes, NMS was considered unsuccessful for the reptile and amphibian a ssemblage with detection/nondetection alone. Count data for the reptiles and amphibians did yield a successful ordination. The first and third axes explain a total of 69% of the variation in the assemblage composition. Lake stage is highly corr elated with Axis 1 (r2=0.267) and Axis 3 (r2=0.389), and water level fluctuation is correlated with Axis 3 (r2=0.204), (Figure 3-14, Table 3-13). Eight species were correlated w ith Axes 1 and 3 with r2>0.2, six of them being indicator species (Figure 3-15, Table 3-14). Table 3-13. Percent of variance explained (r2) by environmental variables for each axis in the herpetofaunal NMS with count data. Variables with r2 > 0.2 are highlighted. Axis 1 2 3 Variable r2 r2 r2 Stage(m) 0.267 0.081 0.389 Fluc(cm) 0.006 0.002 0.204 Rain(cm) 0.000 0.001 0.057 Max(C) 0.052 0.020 0.031 Min(C) 0.022 0.034 0.091 Ave(C) 0.058 0.016 0.073 Table 3-14. Percent of variance explained (r2) for each axis by species in the herpetofaunal NMS with count data. Indicator species with r2 > 0.2 are highlighted in blue, while all other species with r2 > 0.2 are highlighted in yellow. Axis 1 2 3 Axis 1 2 3 Species r2 r2 r2 Species r2 r2 r2 AGKPICO 0.041 0.188 0.012 PSEFLPE 0.042 0.079 0.04 AMPME 0.014 0.291 0.094 RANGR 0.025 0.229 0.229 APAFE 0.087 0 0.017 RANSP 0.185 0.061 0.263 CHESEOS 0.154 0.018 0.036 REGAL 0.384 0.055 0.109 FARABAB 0.004 0.024 0.409 SIREN 0.018 0.031 0.047 KINBA 0.083 0.215 0.351 STEOD 0.437 0.091 0.033 NERFAPI 0.275 0.023 0.498 TADRANGR 0.002 0.095 0.311 NERFL 0.097 0.258 0.03 TADRANSP 0.107 0.275 0.057

PAGE 51

40 To determine the separation of the sample occasion clusters along lake stage and average air temperature gradients means and ranges of each are shown in Figures 3-16 and 3-17 respectively. Clusters one and two have lake stage ranges lower and nonoverlapping with clusters four, five and six. Stage values for cluster 3 span much of the ranges for every other group. Air temperature va lues are very similar for clusters one and six, both being below and non-overlapping with cl usters two, three and four. Cluster five overlaps with most of the ra nges of all other groups. Proportion of Habitat Ut ilized by Focal Species Figures 3-18 and 3-19 show the occupancy rate estimates for the fish species. Several species exhibited a decrease in site occupancy from the spring to the fall season, most pronounced in M. salmoides, L. macrochirus and L. microlophus. Lepomis microlophus, L. gulosus and E. gloriosus occurred at all transects during the spring. Lepisosteidae spp., H. littorale and L. gulosus were present in all transects in the fall. Lepomis punctatus had invalid occupancy estimates during the spring due to low detection probabilities, w ith the same problem for L. macrochirus and L. microlophus in the fall. In the spring, the only fish species th at used survey-specific detection probability models for occupancy estimation were E. gloriosus and L. macrochirus, which indicates that detection of these two species varied between sample occasions. All species were modeled with time-constant detectio n probabilities in the fall, except E. gloriosus, P. latipinna and Lepisosteidae spp. Estimates of the proportion of sites occ upied for the eight focal herpetofaunal species are presented in Figures 3-20 and 3-24. Site occ upancy was similar in the spring and fall for most species. There were not enough data to make valid estimates for A. means in the spring and for R. sphenocephala and N. fasciata pictiventris in the fall. All

PAGE 52

41 species were modeled with the time constant detection probability except N. floridana and R. sphenocephala in the spring. With the exception of R. sphenocephala and A. means most species were estimated to be in 90-1 00% of all transects at some point in the year. Discussion Trap Comparisons The 1.3 cm (0.5 in) mesh traps were bette r at capturing the sp ecies of interest, including reptiles, amphibia ns, centrarchids and exotic catfish. Although it had been assumed that many species would be under-rep resented with the larger mesh, the few unique species caught in the 0.6 cm (0.25 in) mesh traps were not of particular interest to this study. The fish species included mosquito fish and flagfish, which are not suspected of being impacted by the removal of the vege tation from the littoral zone. The one black swamp snake was actually stuck about halfwa y through the mesh, indicating that even 0.6 cm mesh may not be small enough to capture th is species representa tively. The size and shape of the traps and openings may also have affected capture proba bilities of different species. Also, the bare metal hardware cloth may be more visible to the animals than the dark green vinyl-coated hardwa re cloth of the larger mesh traps. In comparing practicality of sampling with each type of trap, the 0.6 cm commercially manufactured traps were much less durable and prone to breaking during use, as well as being more expensive ($16.16 each for 0.6 cm vs. $11.00 each for 1.3 cm). Given these factors, the 1.3 cm mesh traps were more useful than the 0.6 cm for sampling in this habitat. Species Richness Ectothermic fish communities respond to water temperature due to its effect on metabolism and growth rates, especially fo r juveniles (Holt 2002) This relationship

PAGE 53

42 determines the length of time that species will benefit from residing in a given habitat. Water temperature is closely associated with air temperature due to the shallow aquatic habitat. Therefore, it is not surprising th at the average air temperature is negatively correlated with fish species richness. In the summer season, water temperature may exceed tolerance limits for sensitive fish such as select species of sunfish, causing them to leave the habitat and resulting in fewer fish species. Temperature also indirectly determines the amount of dissolved oxygen in the water; however this variable was not measured in the field. Lake stage, rainfall, and water level fluc tuations were signifi cant predictors of herpetofaunal richness, all of which have a re lationship to water dept h at the fixed trap locations. The gradually sloping contour of Lake Tohopekaliga causes small increases in lake stage to flood broad expanses of previ ously dry habitat, allo wing species to enter into new habitat for spawning or foraging. A lternatively, moderate drops in lake stage may cause a rapid decrease in the area of th e littoral zone inundated with water, causing species to emigrate or burrow, or else risk being trapped by unsuitable conditions. Receding water levels may bring in more terr estrial species into th e previously aquatic habitat, such as cottonmouths, Florida water snakes, and Florida leopard frogs. Alternatively, deeper water in the habitat may restrict species that prefer shallow water and promote more aquatic species su ch as the common musk turtle. Assemblage Composition Since animal species move throughout the habitat and are detected with probabilities less than one, and animal assemb lages tend to be transient in nature, the groups determined by cluster analysis are not very discrete with respect to species

PAGE 54

43 composition. This resulted in fairly low indicator values, but with several being significant nonetheless. The four sample occasions in Group 1 had the lowest lake stage values and were tied with Group 6 for the lowest average air temperatures. The Florida water snake and Florida leopard frog are the herp etofaunal indicator species fo r this group, being the more terrestrial of the focal rep tile and amphibian species. Bowfins, golden topminnows, bluegill and dollar sunfis h are the fish indicator species. Bowfins are one of the most tolerant freshwater fish speci es and are often called "mud fish" (Boschhung et al. 1995), so it is not unreasonable that this species w ould occupy this shallow vegetated habitat. Golden topminnows prefer to occupy la kes with abundant vegetation (Hoyer and Canfield 1994), and therefore may be more tolerant of dense vegetation communities than other species. It is not as easy to explain why the two sunfish species were indicators for these shallow water depths. They may have been stranded by receding waters and captured more easily in trap s with puddles of water remaining. Group 2 only had one sample occasion attributed to it. This sample was the last one in late April before the traps were rem oved due to lack of water in the habitat. Although not included in the indicator species analysis, this sample included no fish species. Only three sample occasions were cl ustered into Group 3. During these hot summer samples, the lake stage was rapidly ri sing, and trap effort was reduced pending appropriate water depths at th e trap sites. The striped crayfish snake was the main indicator species for this gr oup. Being a specialist predator on crayfish (Godley 1980),

PAGE 55

44 prey availability in recently flooded habitats may have been the driving factor for their indicator values on these occasions. Eight samples fell into Group 4, which had no indicator species attributed to it. These sample occasions occurred in early fall with the highest lake stages and average air temperatures. Group 5 occurred mainly in the summer months (eight samples), but also contained two samples in February. Water depths were moderately high. Air temperatures were low in February and high in summer, spanni ng a wide range of temperatures. The sole indicator species for this group was the comm on musk turtle. This species is known for being highly aquatic, leaving the water onl y to nest (Wygoda 1979, Gibbons et al. 1983). It also is active for the widest temperatur e ranges of any other kinosternid species in North America, being able to retreat to deeper waters to buffer the effects of air temperature extremes (Mahmoud 1969, Ernst 1986). The last cluster, Group 6, included seven samples, with six occurring February through April and the other one in November. Low air temperatures and moderately high but dropping lake stages characterize these sample occasions. The two Esox spp. (pike) are the indicators for this group. These speci es breed from February to March in the south, spawning in densely vegetated habitats less than 50 cm deep (Billard 1996). This may explain their presence in the habita t during these environmental conditions. Influence of Temporal Gr adients on Assemblage Detection/nondetection data were used for ordinations, and are generally recommended when comparing habitat distribu tions of species (Hayek 1994), and when sample unit heterogeneity is large (McCune and Grace 2002). Counts were also used to compare results obtained by the two types of data, but although agreement between the

PAGE 56

45 two provides more support, lack of detection probabilities make count data less valuable. For the vertebrate and fish ordinations, result s were similar for both types of data. The herpetofaunal ordination was unsuccessful with detection/ nondetection data, and therefore counts were used solely. Average air temperature and lake stage cam e out as the most important variables correlated with the axes representing varia tion in species composition in the vertebrate and fish assemblages. As mentioned before temperature influences growth rates for young fish, as well as the amount of dissolved ox ygen in the water. Both of these factors limits the time that fish are able to occupy a habitat. Physical access to heavily vegetated habitat is also limited by water depth, which is controlled by lake stage. This determines the volume of water the animals have to move through, as well as the effect of vegetation density in the water column. However, for herpetofaunal species alone air temperature is not associated with variation in the species composition. In th is case, lake stage is most important, with water level fluctuation also s howing a correlation with one of the axes. Lake stage probably dictates movements of sp ecies that do not show site fidelity, in response to habitat requirements and prey ava ilability. For species that are not known to move long distances, for example sirens a nd amphiumas, low water levels trigger burrowing activities (Aresco 2001), ther eby reducing captu re opportunities. Proportion of Habitat Ut ilized by Focal Species Due to the large-scale removal of picker elweed from the littoral zone of Lake Tohopekaliga during enhancement activities, it was important to investigate the spatial distribution of species in this habitat. Site occupancy analys es were used to estimate the proportion of this habitat type that was used throughout the year by various fish, reptile and amphibian species. While some species ar e temporally and spatially pervasive in the

PAGE 57

46 habitat (e.g., warmouths, blues potted sunfish, Florida green wa ter snakes, sirens, striped mud turtles, and pig frogs), others seem to use the Pontederia cordata zone intermittently. Of the fish species, sailfin mollies, spotted sunfish, chubsuckers, and largemouth bass were found in a moderate propo rtion of transects ( 30-70%) in both the spring and fall. Bluegill and redear sunfis h were both in a high proportion of sites (>80%) in the spring, but were found in less than 20% of the transects in the fall. This trend may be due to juvenile fish using the littoral zone for foraging and predator avoidance during the spring when suitable physicochemical conditions permit survival (Werner and Hall 1979, Crowder and Cooper 1982, Werner and Hall 1988, Chapman et al. 1996). Unmeasured environmental charact eristics such as low dissolved oxygen may have kept the sunfish out of the thick vegetation after the summer low-water spell (Miranda and Hodges 2000). Gars and armore d catfish went from about 65-80% of the sites in the spring to 100% occ upancy in the fall. These two species are far more tolerant of harsh environmental conditions than most sunfish due in part to their capacity for aerial respiration (Boschung et al. 1995, Braune r et al. 1995). For the armored catfish, the greater presence in the fall may be due to the breeding season and sufficiently high water levels for nesting (Mol 1993). Most reptile and amphibian species occupi ed a similar proportion of sites in both the spring and fall. Florida leopard frogs a nd Florida water snakes were only captured when water levels were very low, which restri cted reasonable estimates of site occupancy to the spring season. Since the heavily vege tated littoral zone is known as prime habitat for several of these species due to life histor y requirements, it is not surprising to find most of the focal species in such a high proportion of the s ites (>70% occupancy).

PAGE 58

47 Figure 3-1. Crayfish and minnow trap in P. cordata habitat.

PAGE 59

48 Figure 3-2. Locations of 2002 P. cordata sampling transects in Lake Tohopekaliga

PAGE 60

49 Date 1/1/02 2/1/02 3/1/02 4/1/02 5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02 1/1/03 Species Richness with 95% CI 0 5 10 15 20 25 30 35 40 45 50 55 Figure 3-3. 2002 Vertebrate sp ecies richness estimates by sample date, with points representing richness for the time between the last sample occasion and the sample date. Date 1/1/02 2/1/02 3/1/02 4/1/02 5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02 1/1/03 Species Richness with 95% CI 0 5 10 15 20 25 30 35 40 Figure 3-4. 2002 Fish species ri chness estimates by sample date, with points representing richness for the time between the last sample occasion and the sample date.

PAGE 61

50 Date 1/1/02 2/1/02 3/1/02 4/1/02 5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02 1/1/03 Species Richness with 95% CI 0 10 20 30 40 Figure 3-5. 2002 Herpetofaunal species richne ss estimates by sample date, with points representing richness for the time between the last sample occasion and the sample date.

PAGE 62

51 10 11 12 13 15 16 17 18 19 2 20 21 22 23 24 25 26 27 28 29 3 30 31 32 33 34 35 36 4 5 6 7 8 9 Stage(m) A ve(C) A xis 1Axis 3 Cluster 1 2 3 4 5 6 Figure 3-6. NMS ordination of sample uni ts in vertebrate species space using detection/nondetection data. Points re present sample occasions and distances between points show the relative diffe rences in species composition. The length of each line is proportional to th e strength of the correlation between the environmental gradient and the synthetic axes.

PAGE 63

52 Figure 3-7. NMS ordination of vertebrate species in sample unit space using detection/nondetection data. Points represent average species positions with respect to sample units. The length of each line is proportional to the strength of the correlation between the environmental gradient and the synthetic axes. AGKPICO AMICA AMPME APAFE CHESEOS ENNGL ERIMY ESOAM ESONI FARABAB FUNCH HOPLI JORFL KINBA LEPGU LEPIS LEPMAC LEPMAR LEPMI LEPPU MICSA NERFAPI NERFL NONE NOTCR POELA POMNI PSEFLPE PTERY RANGR RANSP REGAL SIREN STEOD TADRANGR TADRANSP Stage(m) Ave(C) Axis 1Axis 3

PAGE 64

53 Figure 3-8. NMS ordination of sample units in vertebrate species space using count data. Points represent sample occasions and distances between points show the relative differences in species comp osition. The length of each line is proportional to the strength of the correlation between the environmental gradient and the synthetic axes. 10 11 12 15 16 17 18 19 2 20 21 22 23 24 25 26 27 28 29 3 30 31 32 33 34 35 36 4 5 6 7 8 9 Stage(m) Ave(C) Axis 1Axis 3 Cluster 1 3 4 5 6

PAGE 65

54 Figure 3-9. NMS ordination of ve rtebrate species in sample unit space using count data. Points represent average species positions with respect to sample units. The length of each line is proportional to th e strength of the correlation between the environmental gradient and the synthetic axes. AGKPICO AMICA AMPME APAFE CHESEOS ENNGL ERIMY ESOAM ESONI FARABAB FUNCH HOPLI JORFL KINBA LEPGU LEPIS LEPMAC LEPMAR LEPMI LEPPU MICSA NERFAPI NERFL NONE NOTCR POELA POMNI PSEFLPE PTERY RANGR RANSP REGAL SIREN STEOD TADRANGR TADRANSP Stage(m) Ave(C) Axis 1Axis 3

PAGE 66

55 Figure 3-10. NMS ordination of sample units in fish species space using detection/nondetection data. Points re present sample occasions and distances between points show the relative diffe rences in species composition. The length of each line is proportional to th e strength of the correlation between the environmental gradient and the synthetic axes. Ave(C) 10 11 12 15 16 17 18 19 2 20 21 22 23 24 25 26 27 28 29 3 30 31 32 33 34 35 36 4 5 6 7 8 9 Stage(m) Axis 1Axis 2 Cluster 1 3 4 5 6

PAGE 67

56 Figure 3-11. NMS ordination of fish species in sample unit space using detection/nondetection data. Points represent average species positions with respect to sample units. The length of each line is proportional to the strength of the correlation between the environmental gradient and the synthetic axes. AMICA ENNGL ERIMY ESOAM ESONI FUNCH HOPLI JORFL LEPGU LEPIS LEPMAC LEPMAR LEPMI LEPPU MICSA NOTCR POELA POMNI PTERY Stage(m) Ave(C) Axis 1Axis 2

PAGE 68

57 Figure 3-12. NMS ordination of sample units in fish species space using count data. Points represent sample occasions and distances between points show the relative differences in species comp osition. The length of each line is proportional to the strength of the correlation between the environmental gradient and the synthetic axes. 10 11 12 15 16 17 18 19 2 20 21 22 23 24 25 26 27 28 29 3 30 31 32 33 34 35 36 4 5 6 7 8 9 Stage(m) Ave(C) Axis 2Axis 3 Cluster 1 3 4 5 6

PAGE 69

58 Figure 3-13. NMS ordination of fish species in sample unit space using count data. Points represent average species positions with respect to sample units. The length of each line is proportional to th e strength of the correlation between the environmental gradient and the synthetic axes. AMICA ENNGL ERIMY ESOAM ESONI FUNCH HOPLI JORFL LEPGU LEPIS LEPMAC LEPMAR LEPMI LEPPU MICSA NOTCR POELA POMNI Stage(m) Ave(C) Axis 2Axis 3

PAGE 70

59 Figure 3-14. NMS ordination of sample units in herpetofaunal species space using count data. Points represent sample occasions and distances between points show the relative differences in species composition. The length of each line is proportional to the strength of the correlation between the environmental gradient and the synthetic axes. 10 11 12 13 15 16 17 18 19 2 20 21 22 23 24 25 26 27 28 29 3 30 31 32 33 34 35 36 4 5 6 7 8 9 Stage(m) Fluct(cm) Axis 1Axis 3 Cluster 1 2 3 4 5 6

PAGE 71

60 Figure 3-15. NMS ordination of herpetofauna l species in sample unit space using count data. Points represent average species pos itions with respect to sample units. The length of each line is proportional to the strength of the correlation between the environmental grad ient and the synthetic axes. AGKPICO AMPME APAFE CHESEOS FARABAB KINBA NERFAPI NERFL PSEFLPE RANGR RANSP REGAL SIREN STEOD TADRANGR TADRANSP Stage(m) Fluct(cm) Axis 1Axis 3 Cluster 1 2 3 4 5 6

PAGE 72

61 Cluster 123456 Lake Stage (m) 16.0 16.1 16.2 16.3 16.4 16.5 16.6 16.7 16.8 Figure 3-16. Average and range of lake stage values by cluster. Cluster 123456 Average Air Temperature (OC) 12 14 16 18 20 22 24 26 28 30 Figure 3-17. Average and range of air temperature values by cluster.

PAGE 73

62 FISH SPECIES POELALEPPUERIMYLEPISMICSAHOPLILEPMACLEPMILEPGUENNGL PROPORTION -0.1 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1 NAIVE ESTIMATE PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR Figure 3-18. Site occupancy estimates for focal fish species in spring 2002. FISH SPECIES POELALEPPUERIMYLEPISMICSAHO PLILEPMACLEPMILEPGUENNGL PROPORTION -0.1 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1 NAIVE ESTIMATE PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR Figure 3-19. Site occupancy estimate s for focal fish species in fall 2002.

PAGE 74

63 HERPETOFAUNAL SPECIES RANSPSTEODAMPMERANGRNERFAPIKINBASIRENNERFL PROPORTION -0.1 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1 NAIVE ESTIMATE PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR Figure 3-20. Site occupancy estimates for focal herpetofaunal sp ecies in spring 2002. HERPETOFAUNAL SPECIES RANSPSTEODAMPMERANGRNERFAPIKINBASIRENNERFL PROPORTION -0.1 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 1.1 NAIVE ESTIMATE PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR Figure 3-21. Site occupancy estimates fo r focal herpetofauna l species in fall 2002.

PAGE 75

64 CHAPTER 4 ASSEMBLAGE ACROSS VEGETATION COMMUNITIES Introduction The objective of this section is to inves tigate the influence of vegetation type and water depth on spatial variati on in the aquatic vertebrate community. The three main research questions are 1) are we able to estimate population parameters such as abundance and density for the focal species us ing trapping grids or webs, 2) is there spatial variation in the aquatic vertebra te assemblage, and 3) do the vegetation communities or water depths influence the spatial variation for the individual focal species? Field Methods Grid and Web Sampling A pilot mark-recapture protocol was employed in the summer of 2002 in order to estimate activity ranges, abundances and densi ties of species of in terest. Trap points were arranged in a square grid pattern (W hite et al. 1982). In order to have uniform sampling effort within the grids, one minnow and one crayfish trap was placed at each point. All newly captured reptiles and amphi bians were weighed and measured, then tagged with Passive In tegrated Transponders (PIT tags), which are small microchips inserted under the skin to permit individual identification when scanned. Each animal was released at the cap ture location after being worked up. Traps were checked daily for PIT tagged individuals, and all new individua ls were measured and tagged during each sampling event. In order to satisfy the assu mption of closure for analysis and minimize

PAGE 76

65 temporal variation in detecti on probabilities, the grids were sampled for 5-7 days. This technique was used only for the most abunda nt species, since dete ction probabilities would be too small to make accurate estimat es of any others. These species included A. means Siren spp., N. floridana and K. baurii The first grid consisted of 49 trap points, each three m (9.8 ft) apart, in a seven by seven grid ("GRID1"), and sampled for seven consecutive days in July. Next, 100 trap points were placed in a 10 by 10 grid ("GR ID2A"), five meters (16.4 ft) apart, and sampled for six consecutive days in August. The same grid was then sampled for five non-consecutive days ("GRID2B"), using frozen sardines as bait, until it was decided that the bait was logistically imp ractical. The last design wa s 100 trap points ("GRID3"), spaced three meters (9.8 ft) apart, in a 10 by 10 grid. This was sampled for five times over seven days, due to logistic problems. A ttempts to sample several times within a day were also made, but were discontinued imme diately due to low numbers of captures. In addition to these grids, a trapping web was attempted in September 2002. The web is a variation of point-transect distance sampling, typically used for small terrestrial animals. It is designed to have lines of tr aps radiating out from a center point, forming a gradient of sampling effort and detection probab ilities. Data from each concentric ring is grouped according to distance intervals (Anderson et al. 1983). This web consisted of eight radiating arms of 12 trap points each, placed at threemeter (9.8 ft) intervals, for a total of 96 trap points. The benefit of this me thod is that it uses onl y the initial captures, so the recapture rate is irrelevant. The a ssumptions of this method are that all animals near the center of the web are captured, the size of the web is large relative to the movements of the animals, distances are measured accurately from the center, and

PAGE 77

66 individual captures are i ndependent events. Sampling continues daily until no new animals are caught at the center of the we b, indicating 100% dete ction at the center (Anderson et al. 1983) Since it was evident that the assumptions were not met in this web, sampling was discontinued after six sample occasions. Whole-Lake Sampling After the postponement of the fall 2002 drawdown, we began sampling again with modifications to the 2002 temporal sampling protocol. The goal was to investigate differences in vertebrate habitat usage of vegetation communities beyond the P. cordata zone, as well as varying water depths. Ei ghteen new locations were randomly chosen for transects (Figure 4-1), since th e sampling of 2002 had disturbed the habitat in some of the previous locations. Sampling sites in Goblet ’s Cove were included and none were placed in the disturbed stretch of shoreline in the southern part of the lake. Transects were placed at least 200 m (656 ft) apart. At each transect, there were four trap points, each with a minnow and crayfish trap. However, instead of placing the trap sites at fixed locations in the habitat as in 2002, they were placed at fixed depths and moved with the water level. When the water was rising or remaining stationary, each transect had four trap points located at 15, 30, 45, and 60 cm ( 6, 12, 18, 24 in) deep (Figure 4-2). During falling water levels, the trap points were placed at 30, 45, 60, and 75 cm (12, 18, 24, and 30 in) deep, except when falling lake stages were not predicted. The traps were still checked once weekly, and at each sampling occasion, the traps were moved to the appropriate depth. This result ed in trapping animals at sp ecific depth ranges over the course of the week, with similar water depths between sample occasions. For example if a trap point was located at the 30 cm (12 in) depth and the water level rose several centimeters during the week, the trap was cons idered to be sampling the 30-45 cm (12-18

PAGE 78

67 in) water depth. For falling wa ter levels, the 30 cm (12 in) traps were sampling the 15-30 cm (6-12 in) depth. Percent cover of vegeta tion species was also estimated for a 2 m (6.5 ft) radius around every trap site on each sampling occasion. Continuous sampling was conducted from 1/30/2003 to 1/5/2004, during which time traps were located up in the shallow grassy habitat at high water levels (characterized mainly by Luziola fluitans and Panicum repens ), through the thick emergent habitat (with Pontederia cordata and Typha domingensis ), down to more open water zones (with Hydrilla verticillata and floating leaf species Nuphar luteum and Nymphaea odorata ) at lower water levels. Besides the added environmental variables, this new protocol also allowed us to samp le year-round, instead of having to remove the traps at moderately low water levels. Sampling ended on 1/5/2004 at about 15.5 m (50.8 ft) NGVD, with lake stage droppi ng due to the 2003 drawdown. Analysis Methods Population Estimates and Movement for Herpetofaunal Species Trapping grids are known to exhibit “edge eff ects” due to animals near the edges of the grid moving in and out of the sample d area. To account for this phenomenon a boundary strip is typically estimated and added to the grid area to estimate an effective sampling area. Wilson and Anderson (1985) propose using the mean maximum distance moved (MMDM) by animals recaptured at least once to estimate the activity range of the species. Although lacking a solid theoretica l explanation, this method works well in simulations. The alternative method, nested grid design, requires a large data set for estimation of density (Williams et al. 2002). Since our data were fairly sparse with recaptures, we used the MMDM method to calc ulate the effective areas of the trapping grids for each species.

PAGE 79

68 Low numbers of recaptures were attained fo r each grid; so in or der to calculate the MMDM for each species movement distances were pooled from all grids and the trapping web. To calculate the diagonal distances within the trapping grids, the Pythagorean theorem was used: [a2+b2=c2], where sides a and b are sides of known lengths. For diagonal distan ces in the trapping web, [a2=b2+c2-2bc(cosA)] was used, where A is the degree measure between side s b and c of known length (Larson et al. 1994). For each species, MMDM and its variance were calculated using formulas from Wilson and Anderson (1985). The widths of th e boundary strips were estimated as half the MMDM for each species. The effective grid areas and associated variances were then calculated for each grid per species (Wilson and Anderson 1985). Program MARK (White and Burnham 1999) was used to estimate population sizes for each species per trapping grid. Estimates were obtained using models M(o), (constant capture probabilities), and M( t), (time dependent capture pr obabilities). The Akaike’s Information Criteria (AIC) were compared to determine which model best fit the small data set. Density was then calculated for each grid per species by dividing the population size by the effective sampling area (Wilson and Anderson 1985). Capture Success for Focal Species Basic analyses were conducted on the 2003 data to look for trends in the data associated with the main sampling variables involved in this protocol. All trap points were divided into groups according to vegetation community (see results section below for descriptions) and water depth at each tr ap location. Ten species were examined, two each of salamanders ( Siren spp ., A. means ), snakes (both Nerodia spp.), turtles ( K. baurii and S. odoratus ), frogs (both Rana spp.) and fish ( H. littorale and M. salmoides ). The reptile and amphibian pairs represent species with similar life history traits but which

PAGE 80

69 have slightly different habitat requirements or preferences. They were also the most frequently captured reptile and amphibian sp ecies in this study. The two fish species were chosen to characterize oppos ite ends of the spectrum of habitat selection. While the armored catfish is a generalist species with great tolerance for low dissolved oxygen, high temperatures, thick vegetation and other extr eme environmental variables (e.g., Nico and Fuller 1999), largemouth bass and other sunfishe s are thought to be highly intolerant of these same habitat characteristic s (e.g., Allen and Tugend 2002). Capture success was calculated as the num ber of captures per species divided by the total number of trap points for the particular variable of interest. This usually resulted in a very low frequency, due to the large num ber of trap points and low detectability of species. An arcsine squareroot transformation was applied to all success values, in order to spread the ends of the scale, whil e improving normality for the proportion data (McCune and Grace 2002). This allowed the relative values to s how up more clearly while reducing the effect of la rge sample units. The assumpti on of equal detectability of the different species between habitats or wa ter depths may be violated, but detection probabilities cannot be calculated for this particular analysis. However, uniform sampling methods were used over space (trans ects) and time (sampling occasions) to reduce variability in detection. For this sampling protocol, dependence of trap placement upon water depth (i.e. lake stage) resulted in unequal sampling for vegetation communities. In addition, the vast number of trap sites (n= 3,426) and spar se nature of the data made multivariate analyses virtually impossible. For example, dividing data into groups (either subjectively or with cluster analysis) depending on sa mple occasions, water depth or vegetation

PAGE 81

70 communities would neglect important differences in the other variables and/or result in groups of vastly unequal numbers of trap poi nts. Attempted NMS analyses of all trap sites (ungrouped) yielded no re sults due to the great number of zeroes in the matrices. The data were not even appr opriate for most univariate analyses. For example, chisquare analyses of capture success would indicate whether there were significant differences in the counts of focal species between vegetation type s or water depths, however the large sample sizes invariably lead to significant differences. Repeatedmeasures analysis of variance was considered to test the differences between water depths over time, however the data were t oo sparse to divide the counts between both sample occasions and water depths. Species richness was also inestimable because of the frequent nondetection of sp ecies and unequal sample si zes. As a result of the complicated nature of the data, the analyses were largely descriptive in nature. These descriptions of habitat usage rely mainly on comparisons of capt ure success across two categorical environmental variables: water depth and vegetation community. Results Population Estimates for Herpetofaunal Species Each species showed different movement distances over the sampling grids and web. Figure 4-3 shows the mean maximum distances traveled and variances, along with the associated widths and variances of the boundary strips for each species. Maximum distances traveled for A. means Siren spp., N. floridana and K. baurii were 18 m (59 ft), 24 m (79 ft), 34 m (112 ft), and 59 m (194 ft) respectively. These m ovements are fairly large relative to the sizes of the grids, with 72 m (236 ft) being the absolute maximum distance between any two traps during all samp ling. This indicates that the assumption of closure was violated. Table 4-1 contains th e percent increase in the size of each grid

PAGE 82

71 when the boundary widths for each species were added to the sizes of the grids. While the effective sampling areas are fairly acceptable for the amphiumas and sirens (Wilson and Anderson 1985), Florida green water snak es and striped mud turtles add excessive area to the original sampling areas. After recogn izing the fact that closure was violated in these grids, population sizes and densities we re estimated with unreliable accuracy, but a best attempt was made given the data. Estimates of population size and variances are shown in Table 4-1. Several times the capture history data were so sparse that estimates could not be calculated for some grids and species. The estimates generated with sufficient data often have large variances due to low recapture probabilities. The null model of no variation in detection Table 4-1. Grid sizes a nd population estimates by mark recapture methods. Parameter AMPME SIREN NERFL KINBA GRID1 Actual size of grid (ha) 0.0324 0.0324 0.0324 0.0324 Est. effective sampling area and variance (ha)0.078 (0.40)0.08 (0.93) 0.12 (1.79)0.16 (9.17) Percent of original grid (%) 239 264 376 493 Est. population size and variance (# indivs.) n/a n/a 16 (184) 7 (30) Density estimate (#/ha) n/a n/a 131 44 GRID2A Actual size of gr id (ha) 0.2025 0.2025 0.2025 0.2025 Est. effective sampling area and variance (ha)0.30 (1.66)0.32 (3.71) 0.39 (6.08)0.46 (27.83) Percent of original grid (%) 149 157 192 225 Est. population size and variance (# indivs.) n/a n/a n/a n/a Density estimate (#/ha) n/a n/a n/a n/a GRID2B Actual size of gr id (ha) 0.2025 0.2025 0.2025 0.2025 Est. effective sampling area and variance (ha)0.30 (1.66)0.32 (3.71) 0.39 (6.08)0.46 (27.83) Percent of original grid (%) 149 157 192 225 Est. population size and variance (# indivs.) 14 (37) n/a 26 (99) 9 (5) Density estimate (#/ha) 49 n/a 68 20 GRID3 Actual size of grid (ha) 0.0729 0.0729 0.0729 0.0729 Est. effective sampling area and variance (ha)0.14 (0.72)0.15 (1.65) 0.19 (2.94)0.24 (14.27) Percent of original grid (%) 187 201 267 332 Est. population size and variance (# indivs.) 19 (117) n/a 25 (24) 26 (525) Density estimate (#/ha) 142 n/a 133 108

PAGE 83

72 probability was always selected over the tim e-varying capture probability model. Density estimates based on these abunda nces are also included in Table 4-1. Capture success for Focal Species Figure 4-4 shows the types of vegeta tion communities that were sampled throughout 2003. Each sampling occasion corresponds to one weekly sample, which includes 72 trap points (4 trap points for each of 18 transects). O ccasionally there were less than 72 samples for a given sample occa sion, usually because data recording for a sample was inadvertently neglected, traps went unchecked due to dangerous weather, or traps were missing. Each trap point was s ubjectively categorized in the field into different vegetation communities, based on the dominant species present. The “Grass” community is the closest shor eward, and is characterized by Panicum repens, Luziola fluitans, Juncus effusus, and Eleocharis spp. Lakeward from this is the “Rooted-HE”, which refers to the herbaceous emergent species, especially Pontederia cordata and Typha domingensis The community termed “G/HE” is the border of the grass and rooted herbaceous emergent zones, which ha d enough samples to be a separate category. “Floating-HE” is the floating mat community, consisting of P. cordata, Bidens spp., Ludwigia leptocarpa and a variety of other species. Out past the herbaceous emergent communities are the deeper “Outward” communities. These include floating-leaf emergents ( Nelumbo lutea, Nymphaea odorata, and Nuphar luteum ), submersed plants ( Hydrilla verticillata ), and deep emergents ( Paspalidium geminatum ). Trap points that were on the borders between distinct ve getation communities, or were part of communities with too few samples to have its own category, were classified as “Mixed”. Since transects were randomly chosen and the trap points were moved with the water level, there was no way to collect equal num bers of samples from each community.

PAGE 84

73 Figures 4-5 and 4-6 show results fr om capture success comparisons for salamanders. Both species tend to be capture d more frequently in the outer herbaceous emergent vegetation communities and community edges, as well as greater water depths. Pig frogs seemed to show a preference fo r grassy habitats and community edges, decreasing towards more outward vegetation communities, while leopard frogs did not show much of a pattern (Figure 4-7). Howe ver, much stronger trends appeared with water depth (Figure 4-8). Both species, espe cially the leopard frogs, showed an inverse relationship to water depth. The two snake species showed divergent trends in capture rate. In Figure 4-9, Florida green water snakes ( Nerodia floridana ) appeared more in the rooted pickerelweed communities, while the Florida water snakes ( Nerodia fasciata pictiventris ) did not have quite as strong a tendency to be in specific habitats. Water depth seemed to be more important in Florida water snake occurrence (Figure 4-10), with most being trapped in shallow water and decreasing steadily with dept h. The Florida green water snake did not have such a trend. Figures 4-11 and 4-12 show that common musk turtles were captured more frequently in vegetation habitats furt hest from shore, as well as deeper water depths. Striped mud turtles on the other ha nd did not have strong trends, but peak in rooted pickerelweed habitats and intermediate water depths. Largemouth bass did not show a strong affi nity for any certain habitat, however they were captured slightly more frequently in rooted herbaceous emergent and borderline communities (Figure 4-13). They also appeared most in intermediate water depths, (Figure 4-14), e.g.45-60 cm (18-24 in) deep. Bass ca ptured in the traps were juvenile fish, with total lengths in 2003 ra nging from 5.1-14.0 cm (2-5.5 in), (n=60).

PAGE 85

74 The armored catfish were found mostly in bo rder vegetation communities (Figure 4-13). They also were positively correlate d with water depth (Figure 4-14). Discussion Population Estimates for Herpetofaunal Species The small activity ranges estimated for the amphiumas and sirens were similar to what have been found in other studies (Gehlbach and Kennedy 1978, Sorensen 2004). Maximum distances were higher in this case, (18 m (59 ft) vs. 5 m (16 ft) for amphiumas and 24 m (79 ft) vs. 10 m (33 ft) for greater sirens (Sorensen 2004)). These estimates suggest that lack of movement in these an imals make them susceptible to mortality during muck removal operations. The sizes of the grids were proba bly not large enough to make reasonable activity range estimates fo r the Florida green water snake or striped mud turtle. Bancroft et al (1983) documented a Florida green water snake moving 223 m (731 ft) in less than two hours. As another example, Nerodia taxispilota (brown water snakes) have been documented moving distances greater than 1 km ( 0.62 miles) (Mills et al. 1995). Mahmoud (1969) found maximum distan ces for several species of kinosternid turtles, including 525.5 m (1,723.6 ft) for S. odoratus 435.3 m (1,427.8 ft) for Kinosternon flavescens (yellow mud turtle), 408.4 m (1,340 ft) for Kinosternon subrubrum (Mississippi mud turtle ), and 93.9 m (308 ft) for Sternotherus carinatus (Mississippi musk turtle). Due to low recapture rates and large movements of individuals, poor density estimates were attained with these protocols. Even when the simplest models were utilized to estimate population sizes (the estimated number of animals with no account of area sampled), variances were unacceptably high. Even if the estimates had been reasonable, the study would have been limite d to a small number of species, a narrow

PAGE 86

75 window of opportunity when the pickerelweed zone was completely inundated with water, and non-random locations which po ssessed a wide enough band of habitat to contain the grids within a rela tively homogeneous habitat. Therefore, the mark-recapture grids will not be utilized for post-enhancement sampling. Capture Success for Focal Species Vegetation communities offer different tradeoffs to animal species that inhabit them. Variations in predator efficiency, pr ey type and abundance, or abiotic properties associated with dissimilar macrophyte types strongly determine their use to aquatic vertebrates (Miranda et al. 2000) Physical properties of plan t species such as branching, leaf shape and number, plant biomass and pos ition throughout the water column affect animals’ ability to maneuver in the habitat, as well as phys ical and chemical properties such as dissolved oxygen, nutrient levels, water temperature, light pe netration and current (Chick and McIvor 1994). Welch (2004) c onducted a thorough ecol ogical investigation into the vegetation communities of Lake T ohopekaliga prior to the 2004 enhancement. One finding was that the soils associated with the intermediate littoral zone depths and Pontederia cordata communities were highly organic and low in bulk density. On the other hand, the shallow grassy communities and the various deeper water communities had soils higher in bulk density, and theref ore were sandier in composition. Substrate alone may provide benefits or detriments to animal species, dependi ng on their specific life history traits. Water depth at a given location has st rong influences on vertebrate species distribution and habitat use. It is the main determinant of the boundaries of the littoral ecotone, limiting emergent aquatic vegetati on growth to the limits of the water fluctuation. The gradual slope of the shoreline causes sma ll increases or decreases in

PAGE 87

76 lake stage to flood or dry out broad expanses ha bitat, altering its use to different species. Water depth also establishes th e volume of water that aqua tic organisms have to move through, and can provide enough space for the pr esence of a thermocline (Miranda et al. 2000). The aquatic salamander species are known for burrowing in the organic sediment associated with dense vegetation as refuge from predators and drought (Etheridge 1990, Conant and Collins 1998). In fact many of these large salamanders have been uncovered during muck removal operations around Flor ida, even when there was water still covering deeper areas of the lake (Aresco 2001) The findings of this study indicate the same pattern, with highest capture success occurring in densely vegetated communities. As mentioned previously, these communities are also most associated with low bulk density and high organic composition of the so ils (Welch 2004). This indicates that not only may the dense vegetation provide ample fo rage and cover for these creatures, but also the organic sediment (muck) is prefe rred for burrowing. While deeper water depths yielded more sirens and amphiumas, it is possi ble that they are si mply more active in deeper water, increasing de tection probabilities. Higher capture success in shallow water si tes was expected for leopard frogs, since all individuals captured in 2002 occurred in Ap ril and November when water levels were low. This species is known to travel rela tively far from aquatic habitats, given proper cover and shade from terrestrial vegetati on, depending on soil moisture and dew to prevent desiccation (Dole 1965, Conant a nd Collins 1998). Leopard frogs are particularly dense in herbaceous vegetation ar ound lakes, with plenty of protection and food sources in the grasses (Kilby 1936). On the other hand, pig fr ogs are highly aquatic,

PAGE 88

77 often being associated with emergent and floating vegetation (Conant and Collins 1998). Pig frogs were captured the most in the “m ixed” vegetation community. This category was mainly represented by floating vegetati on (water hyacinth) and borders between communities, (mainly between the floating mats of emergent vegetation and submersed vegetation). The dominance of pig frogs found in this community may indicate that border communities provide a tradeoff between predator avoidance and prey availability. Decreases in capture success with increased water depth and relative distance from shore indicate that water depth was very influentia l in determining the presence of both species of ranid frogs. The leopard frog in particular has a very strong decreasing trend with water depth, which is consistent with its more terrestrial nature. Water depth was also an important factor in the Florida water snakes’ habitat preference, which was expected since in th e 2002 sampling most i ndividuals were caught in April and November, both during relativel y low water periods. Water depth does not seem to have as strong an influence on Florid a green water snakes. To illustrate this difference, Seigel et al. (1995) found that during a three-year dr ought in Ellenton Bay, South Carolina, many N. fasciata left the habitat only seven days after it dried out, while N. floridana never left in large numbers. While the abundance of snakes was generally lower, it was five years before N. floridana was captured after the drought. As discussed previously, water snakes typically show little site fidelity and are capable of long-range movements (Bancroft et al. 1983, Mills et al 1995). Several of the species show ontogenetic niche shifts with ag e and size, often changing diet and habitat preference at a certain size (Mushinsky et al. 1982, Mushinsky and Miller 1993). Fl orida water snakes in particular are known for feeding on fish when young, and then at 50 cm (20 in) snout-

PAGE 89

78 vent length they begin to feed almost ex clusively on frogs (Mushi nsky et al. 1982). The traps used in this study mainly catch adult sn akes, so prey (i.e. a nuran) availability in shallow habitats may result in this species preference for shallower water depths. Alternatively, Florida green water snakes ar e not so specialized, being caught with and regurgitating a variety of fish, frogs, and ev en large sirens. While both species were captured more frequently in the emergent vegetation communities, the water depth seems to influence the presence of these species the most. The turtle species have different life hi story traits that may explain observed differences in capture success. For exampl e, common musk turtles are highly aquatic, rarely leaving the wate r except to nest. When water le vels drop, at least in ponds, they follow the water down and then burrow into th e sediment to avoid desiccation (Wygoda 1979, Gibbons et al. 1983). Ma hmoud (1969) suggests that Sternotherus spp are more dependent upon water depth than Kinosternon spp. The former appears to prefer water depths greater than 30 cm and have been found in up to seven meters of water. As shown in this study, there is a sharp increase in capture success associated with both lakeward vegetation communities and deeper water depths Alternatively, striped mud turtles are much more terrestrial, usually dispersing over land during drought or heavy rainfall to find alternative habitats (B ennet 1972, Wygoda 1979). On th e other hand, Gibbons et al. (1983) found that Kinosternon subrubrum experienced no increased emigration due to drought in Ellenton Bay, South Carolina, since it is a fairly terrestrial species and is not negatively affected by dry conditions. Maximu m activity of striped mud turtles occurred in 15 cm in Oklahoma (Mahmoud 1969). In this study, peak captures occur in rooted

PAGE 90

79 emergent vegetation and intermediate water dept hs. All species of ki nosternid turtles are thought to prefer vegetated habitats to unvegetated ones (Mahmoud 1969). The captures of young largemouth bass in a ll parts of the littoral habitat were contrary to common fisherie s doctrine. One would expect them to occur almost exclusively in the open water/submersed habi tats, due to physicochemical requirements, as well as the physical barrier of the organic berm formed by the floating vegetation mats (Moyer et al. 1995, Allen and Tugend 2002, Allen et al. 2003). However, relatively high water levels evidently allow young bass and ot her centrarchid species to enter grassy habitats, as well as inhabit the pickerelweed zone. Mira nda et al. (2000) described vegetated aquatic habitats as a mosaic of microhabitats within larger seemingly inhospitable macrophyte stands. Although from a human’s pe rspective the habitat may seem uniformly unsuitable, fish can move bot h horizontally and vertically to find pockets of suitable physical (e.g., temperature) and chemical (e.g., dissolved oxygen) water conditions for survival. Perhaps this explai ns the bass’ ability to move through this landscape relatively unscathed. The armo red catfish on the other hand, have a high tolerance for poor water quality due to their ab ility to breathe air (B rauner et al. 1995). They were still found at deeper water de pths, but his was likely due to breeding requirements in the littoral zone for greater than 0.3 m water depths (Hostache and Mol 1998).

PAGE 91

80 60 cm 45 cm 30 cm 15 cm 60 cm 45 cm 30 cm 15 cm Figure 4-1. Locations of 2003 sampli ng transects in Lake Tohopekaliga.

PAGE 92

81 15 cm 30 cm 45 cm Crayfish traps are positioned on the substrate Minnow traps float on surface of water 60 cm 15 cm 30 cm 45 cm Crayfish traps are positioned on the substrate Minnow traps float on surface of water 60 cm 15 cm 30 cm 45 cm Crayfish traps are positioned on the substrate Minnow traps float on surface of water 60 cm Figure 4-2. Diagram of weekly trap placement at specified depths. Distance (m) 0510152025303540455055606570 Amphiuma Siren Fl.green water snake Striped mud turtle Maximum mean distance moved and variance Maximum distances moved per species Figure 4-3. Mean maximum distances travel ed with variances and maximum distances, based on results of mark-recapture sampling.

PAGE 93

82 Sample Occasion 024681012141618202224262830323436384042444648 Number of Each Vegetation Community 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 GRASS (n=799) G/HE (n=230) ROOTED-HE (n=1174) FLOATING-HE (n=714) MIXED (n=162) OUTWARD (n=347) Figure 4-4. Number of trap sites sample d in each vegetation community per sample occasion.

PAGE 94

83 Vegetation Community GG/HER-HEF-HEMO Arcsine Squareroot Transformed (#captures/# trap points) 0.00 0.05 0.10 0.15 0.20 0.25 0.30 0.35 SIREN AMPHIUMA Figure 4-5. Salamander capture success by vegetation community. Water Depth (cm) 0 15 30 45 60 Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.05 0.10 0.15 0.20 SIREN AMPHIUMA Figure 4-6. Salamander capture success by water depth.

PAGE 95

84 Vegetation Community GG/HER-HEF-HEMO Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.05 0.10 0.15 0.20 0.25 PIG FROG SOUTHERN LEOPARD FROG Figure 4-7. Frog capture su ccess by vegetation community. Water Depths (cm) 0 15 30 45 60 Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 PIG FROG SOUTHERN LEOPARD FROG Figure 4-8. Frog capture success by water depth.

PAGE 96

85 Vegetation Communities GG/HER-HEF-HEMO Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.05 0.10 0.15 0.20 0.25 FLORIDA GREEN WATER SNAKE FLORIDA WATER SNAKE Figure 4-9. Snake capture success by vegetation community. Water Depth (cm) 0 15 30 45 60 Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 0.16 0.18 FLORIDA GREEN WATER SNAKE FLORIDA WATER SNAKE Figure 4-10. Snake capture success by water depth.

PAGE 97

86 Vegetation Communities GG/HER-HEF-HEMO Arcsine Squareroot Transformed (# capture/ #trap points) 0.00 0.05 0.10 0.15 0.20 0.25 0.30 STRIPED MUD TURTLE COMMON MUSK TURTLE Figure 4-11. Turtle capture success by vegetation community. Water Depths (cm) 0 15 30 45 60 Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 STRIPED MUD TURTLE COMMON MUSK TURTLE Figure 4-12. Turtle captu re success by water depth.

PAGE 98

87 Vegetation Communities GG/HER-HEF-HEMO Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.05 0.10 0.15 0.20 LARGEMOUTH BASS ARMORED CATFISH Figure 4-13. Fish capture su ccess by vegetation community. Water Depths (cm) 0 15 30 45 60 Arcsine Squareroot Transformed (# captures/ # trap points) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 LARGEMOUTH BASS ARMORED CATFISH Figure 4-14. Fish capture success by water depth.

PAGE 99

88 CHAPTER 5 SUMMARY AND CONCLUSIONS Review of Aquatic Vertebrate Communi ty Dynamics in Lake Tohopekaliga This study has documented the conditi ons of the fish and herpetofaunal communities in the littoral zone of Lake Tohopekaliga prior to extreme habitat modifications performed in 2004. The littoral zone in this eutrophic lake defined here includes the entire vegetated shoreline, fr om the grassy vegetation community on the shore that is occasionally inundated with water to the lakeward band of emergent vegetation that is always floode d. Animals captured in this ha bitat represent species that are dependent in some way upon its unique ch aracteristics for their survival, including still water, cover from predators or light, nest ing substrate, organic sediment in which to burrow or forage, and appropriate prey. Perh aps the most useful information from this study so far has been the species list, which includes juvenile centrarchids and exotic catfish. While it is no surprise to find most of the herpetofaunal species in this type of landscape, the quantity and quality of fish cap tured was not expected. Previous research indicates that sunfish cannot and do not inhabit heavily vegetated littoral habitats, while the two species of exotic catfish captured have not been documented at all this far north in Florida. Species such as warmouths, bl uespotted sunfish, Florida green water snakes, sirens, striped mud turtles, and pig frogs were found in all parts of the habitat throughout the year, indicating that they may be threatened by the habitat alterations more than other species.

PAGE 100

89 Research efforts in 2002 were focused on the Pontederia cordata zone only, which was the species most considered a nuisance by lake managers. The widespread removal of this species threatens to disrupt the wildli fe community structure in a large area of the lake, making research of this specific ve getation community a n ecessary component of the study. In 2003, sampling was conducted ac ross vegetation communities within the littoral zone in an attempt to understand how different fo cal species are distributed throughout the habitat. The following secti ons will provide a revi ew of the observed effects of these environmental variables on species richness, community composition and distribution of focal species. Air Temperature Average air temperatures over the course of each sample occasion accounted for much variation in the aquatic vertebrate co mmunity. Both fish and total vertebrate richness were negatively correlated with average air temperature over time. NMS analyses also indicated that this variable explained large percentages of variance in the species composition of the fish and vertebra te communities. Vertebrate assemblages represented by Regina alleni and Esox spp. were associated w ith high air temperatures, while groups with Nerodia fasciata pictiventris and Rana sphenocephala as indicator species were found at lowe r air temperatures. Lake Stage Only herpetofaunal species richness was found to decline with increased lake stages. This variable expl ained much of the variation in fish, herpetofaunal, and combined vertebrate assemblages. In periods of high lake stages, Sternotherus odoratus and Esox spp. were the indicator species fo r sample occasion clusters. N. fasciata

PAGE 101

90 pictiventris and R. sphenocephala characterized assemblages occurring with low lake stages. Water Depth Water depths from 0-75 cm were samp led for aquatic vertebrates. Sirens, amphiumas, armored catfish and especially common musk turtles showed increasing capture success with water depth. On the ot her hand, pig frogs, leopard frogs and Florida water snakes showed strong decreases in capture success as depth increased. The main drawback to this sampling protocol was th e limiting size of the cr ayfish traps, with restricted sampling beyond 75 cm deep. Species occurrence beyond this depth would have provided valuable information that would be relevant to potenti al responses to the lake drawdown and scraping activities. Vegetation Community Both salamander species tended to be captu red mostly in the emergent vegetation, but probably had a stronger association with the decreased bulk density in the soils of these vegetation communities. Pig frogs were found to occupy the more shoreward vegetations disproportionately. Florida green water snakes were more often found in the rooted grasses and pickerelweed than in th e outermost communities. While striped mud turtles peaked in the rooted emergent vege tation zone, common musk turtle increased in frequency in the more lakeward communities. Population Size Estimates Due to unsuitable sizes and trap spacing of the grids and web, large movements and low recapture rates of the focal species resu lted in poor population size estimates. Large variances indicate the unreliability of the result s. This attempt illustrates the difficulty of using one method to sample several different species that have di ssimilar life history

PAGE 102

91 traits. Even if reasonable estimates had result ed from these pilot studies, the limitation of the protocol to only four species and such a specific habitat ( P. cordata zone while inundated with water) narrows the focus of the study beyond much use to lake managers. Lake Tohopekaliga Habitat Enhancement The fall 2003 drawdown has been successfu lly implemented and an estimated 7.3 million cubic meters of muck and vegetation have been removed (scraped) from the littoral zone. Bulldozers and dump trucks re moved a total area of 1,351 ha of habitat (Florida Fish and Wildlife Conservation Commissi on 2004). Repercussions or benefits to the various wildlife guilds inhabitin g the lake have yet to be determined. As noted in this study, there are multiple species that utili ze the heavily vegetated littoral habitat throughout the year, whether by preference or necessity. While there remain intact stretches of habitat that have not been scra ped, the majority of the shoreline has been radically altered. The main reason for the extreme nature of this project was to attain maximum benefits to the largemouth bass population for the longest period of time possible. High costs associated with the enhancement procedures prohibit frequent drawdowns and scraping efforts. Due to these factors, managers feel pressure to remove the most vegetation and muck as possible at one time in order to produce long-term benefits (Allen and Tugend 2002). If it were not for monetary restrictions, perhaps alternatives such as limiting removal of mu ck and vegetation to a smaller portion of the shoreline, or using mechanical removal as a method of increasing the patchiness of the vegetated area would be prefe rred. In these cases, adult bass would still have access to shallow spawning sites, while species requi ring more complex habitats would not be displaced.

PAGE 103

92 Direct mortality will doubtless cause change s in the abundance of several species, most notably for the sirens and amphiuma s. During just a few hours of bulldozer activity, several dozen sirens and amphiumas and one pig frog were unearthed from beneath the vegetation (personal observation) The machinery also crushed one large snapping turtle in the dry pickerelweed z one. Scraping was conducted from the grassy vegetation communities out into the floati ng leaf/submersed vegetation communities, removing virtually all of the emergent vegeta tion across the ecotonal gradient. The depth of sediment removal was established by white sand substrate and absence of all root structures. Sediment and ve getation was piled into large windrows and subsequently loaded into dump trucks and deposited in uplan d disposal areas or in -lake spoil islands. Although most salamanders that were aestiva ting beneath the soil surface seemed to survive the initial pass with the equipment, most probably were crushed by the weight of the debris or physically removed from the lake altogether. Bulldozer operators reported numerous small turtles being uncovered, in addition to the “muck eels” (i.e., large salamanders). In agreemen t with this study, Aresco ( 2001) also reported aestivating sirens and amphiumas uncovered by bulldozer operations in Lake Jackson, Leon County, Florida, even though open-water ha bitat was still present past the vegetated zone. Due to the terrestrial nature of K. baurii, N. fascia ta pictiventris, and R. sphenocephala, and the highly aquatic characteristics of S. odoratus these species may not have been in the littoral zone of Lake Tohope kaliga during the drawdown and may have experienced less mortality associated with the bulldozers. In addition to mortality due to enhancem ent operations, ensuing lack of suitable habitat will further alter vertebrate communities. A suite of herbicides is being utilized to

PAGE 104

93 stop natural vegetation succession so that managers may selectively allow regrowth of specific species at subjectively desirable (i.e. more sparse) de nsities. As a result, at any point in the former littoral zone, the ha bitat will have underg one several physical alterations. Complete absence of emergent vegetation and or ganic sediment will prohibit recolonization by many of the herpetofaunal species examin ed in this study. For example, species such as sirens, amphiumas, striped mud turtles, and Florida green water snakes will likely avoid the enhanced shorel ines for these reasons. Juvenile sunfish may also limit their use of this habitat due to lack of prey and cover. Bancroft et al. (1983) found that Florida green water snakes, greater sirens, southe rn leopard frogs, striped mud turtles and common musk turtles were nega tively impacted by vegetation loss resulting from grass carp introductions and shoreline development in Lake Conway, Florida. Alternatively, other species such as amphiumas and Florida water snakes did not seem to have adverse reactions to th e resulting loss of vegetation and increase in sandy beach shoreline. In addition to the obvious lack of muck and vegetation (except for the exotic hydrilla), several more subtle changes will have taken place. For any given lake stage, any affected location will have deeper water due to the removal of the sediment and root structure. As seen in this study, increased water depths are associated with lower herpetofaunal species richness. In particular aquatic frogs and Flor ida water snakes may be less inclined to inhabit these deeper habitats, while the co mmon musk turtle may increase presence. Pelagic fish species may replace juvenile or more littoral zone fish species. Increased wave action and water cu rrent will result from vegetation removal in the shoreline habitat. In these large lakes, strong wi nds and thunderstorms blowing

PAGE 105

94 across the open water often cause large wa ves and whitecaps. Light penetration and water temperature will also increase due to vegetation loss. This may influence fish species richness, which was show n to decrease with increased air temperatures in this study. Fish species with higher environmenta l tolerances such as H. littorale, Esox spp., and Lepisosteus spp. may increase in abundance relative to Lepomis spp. Most of these factors will not only impact ad ult habitat preference for severa l native species of fish and herpetofauna, but will certai nly have strong effects on nes ting potential for fish and amphibians, several species of which require still water and/or emergent vegetation stems. Even though this expansive vegetated habitat is a result of anthropogenic eutrophication in central Florid a lakes, it may provide an a lternative habitat for species affected by decades of extensive wetland de struction. Wetland isolation resulting from human modifications reduces co rridor travel between habita ts and abundances for species such as sirens that rely on wetlands fo r survival (Snodgrass et al. 1999). Through the previous century, channelization of natura l streams, wetland drainage to improve pastureland, and water level stabilization within the Upper Kissimmee Basin have eliminated and fragmented many habitats used by wetland animal species. As a consequence of these landscape manipulations the lush, vegetated, lake littoral zones may provide refuge for displaced aquatic verteb rates in this area, regardless of the fact that the eutrophic lake edges are not natural features. There is also the concern that the extr eme techniques used during the enhancement project were unnecessary and based upon false suppositions. Following a similar drawdown and muck-scraping project in 1995 in Lake Kissimmee, Flor ida, Allen et al.

PAGE 106

95 (2003) found no increase in harvestable largem outh bass or angler catch rate in the following six years compared to pre-enhancemen t. Due to the lack of response from the adult bass population, the rese archers concluded that ma nagers should view lake enhancement as a way to improve recreational (i .e. fishing boat) access in the lake rather than setting the stated goal of improving bass fishing. Mi randa and Dibble (2002) stated the need for fishery scientists to focus at al l organizational levels to properly manage for bass. Relying on just the individual and population response is not enough to understand and manipulate the behavior of the species or manage an ecosystem. Ecological interactions at the community and ecosystem levels must also be considered before single-species management techniques are impos ed on a habitat, since there are important interactions between other sp ecies and the abiotic environment that may influence the success of the species. Future Aquatic Vertebrate Monitoring Plans While this study has provided baseline information on species presence and identified the influence of temporally and spatially changing environmental variables on the vertebrate communities, it is only the be ginning of the necessary research. The continuation of the current project on Lake Tohopekaliga will provide missing information needed to determine the changes in the wildlife communities. In late 2004, pending suitable water depths in the littoral z one, traps will be redeployed at the original transects sampled in 2002. The fixed-location trapping protocol will be implemented and will continue indefinitely. Comparisons will be made in site occupancy and temporal community utilization of this habitat, be tween the preand post-enhancement littoral zone. Although the spatial sampling protoc ol (used in 2003) provided information on habitat use beyond the pickerelweed zone, it will not be continued. The dependence of

PAGE 107

96 trap locations on lake stage results in little investigator control over sampling effort in different vegetation communities, and therefor e is limited in its use as a long-term sampling technique. In addition to whole-lake sampling, three study areas have been defined which each contained four 400 m long sections that underwent different treatments, including scraping and herbicide application, only scrapi ng, only herbicide application and control. Since the mark-recapture grids were unsucce ssful in attaining reliable population estimates, an alternative sampling protocol will be used in these study plots. Fixed transects similar to the whole-lake transects will be laid out in the 12 plots. Sampling will take place in the spring and fall seasons when lake stages will allow sufficient water in the habitat. In both seasons traps will be checked for 3 months, yielding 12 weekly sampling occasions. Captured individuals of the focal herpetofaunal species will be PIT tagged to determine movement patterns within and among treatment plots. With this protocol repeated samples in the same locations will allow estimation of detection probabilities, facilitating estimation of site occupancy and community diversity measures. Assemblage composition and association with measured environmen tal variables will also be analyzed. Comparisons of the aquatic vertebrate communities between treatment plots will reveal effects of the vegetation and sediment remo val by herbicidal treatments and mechanical removal.

PAGE 108

97 LIST OF REFERENCES Alford, R. A., and M. L. Crump. 1982. Habitat partitioning among size classes of larval Southern leopard frogs, Rana utricularia Copeia 1982:367-373. Allen, M. S., and K. I. Tugend. 2002. Eff ects of a large-scale habitat enhancement project on habitat quality for age-0 largem outh bass at Lake Kissimmee, Florida. Pages 265-276 in D. P. Philipp and M. S. Ri dgeway. Black bass: ecology, conservation, and management. Americ an Fisheries Society, Symposium 31, Bethesda, Maryland, USA. Allen, M. S., K. I. Tugend, and M. J. Mann. 2003. Largemouth bass abundance and angler catch rates following a habitat enhancement project at Lake Kissimmee, Florida. North American Journa l of Fisheries Management 23:845-855. Anderson, D. R., K. P. Burnham, G. C. White and D. L. Otis. 1983. Density estimation of small-mammal populations using a trapping web and distance sampling methods. Ecology 64:674-680. Aresco, M. J. 2001. Siren lacertina (greater siren). Aestivation chamber. Herpetological Review 32:32-33. Bancroft, G. T., S. J. Godley, D. T. Gross, N. N. Rojas, and D. A. Sutphen. 1983. Largescale operations management test of use of the white amur for control of problem aquatic plants; the herpetof auna of Lake Conway: spec ies accounts: miscellaneous papers A-85-3, U.S. Army Engineer Waterways Experiment Station, CE, Vicksburg, MS. Bennet, D. H. 1972. Notes on the terre strial wintering of mud turtles ( Kinosternon subrubrum ). Herpetologica 28:245-247. Billard, R. 1996. Reproduction of pike: ga metogenesis, gamete biology and early development. Pages 13-43 in J. F. Craig, editor. Pike : biology and exploitation. Chapman & Hall, London, England. Blake, N. M. 1980. Land into water water into land, a history of water management in Florida. University Presses of Fl orida, Tallahassee, Florida, USA. Boschung, H. T. Jr., J. C. Williams, D. W. Gotsha ll, D. K. Caldwell, and M. C. Caldwell. 1995. National Audubon Society field guide to North American fishes, whales, and dolphins. Alfred A. Knopf, Inc., New York, USA.

PAGE 109

98 Brauner, C. J., C. L. Ballantyne, D. J. Randall, and A. L. Val. 1995. Air breathing in the armoured catfish ( Hoplosternum littorale ) as an adaptation to hypoxic, acidic, and hydrogen sulphide rich wa ters. Canadian Journal of Zoology 73:739-744. Burnham, K. P., and W. S. Overton. 1979. Robust estimation of population size when capture probabilities vary among animals. Ecology 60:927-936. Bury, R. B., and P. S. Corn. 1991. Sampli ng methods for amphibians in streams in the Pacific Northwest. Gen. Tech. Rep. PNW-GTR-257. Portland, Oregon: U.S. Department of Agriculture, Forest Servi ce, Pacific Northwest Research Station. Casazza, M. L., G. D. White, and C. J. Gr egory. 2000. A funnel trap modification for surface collection of aquatic amphibians and reptiles. Herpetological Review 31:91-92. Chapman, L. J., C. A. Chapman, and M. Ch andler. 1996. Wetland ecotones as refugia for endangered fishes. Biol ogical Conservation 78:263-270. Chick, J. H., and C. C. McIvor. 1994. Pa tterns in abundance and composition of fishes among beds of different macrophytes: view ing a littoral zone as a landscape. Canadian Journal of Fisheries and Aquatic Sciences 51: 2873-2882. Conant, R., and J. T. Collins. 1998. A field guide to reptiles and amphibians of eastern and central North America, third edi tion, expanded. Houghton Mifflin Company, New York, New York, USA. Cooke, G. D., E. B. Welch, S. A. Peterson, and P. R. Newroth. 1993. Restoration and management of lakes and reservoirs, second edition. Lewis Publishers, Boca Raton, Florida, USA. Corn, P. S. 1994. Straight-line drift fences and pitfall traps. Pages 109-117 in W. R. Heyer, M. A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M. S. Foster, editors. Measuring and monitoring biol ogical diversity: standard methods for amphibians. Smithsonian Institution, Washington D.C., USA. Crowder, L. B., and W. E. Cooper. 1982. Habitat structural complexity and the interaction between bluegill a nd their prey. Ecology 63:1802-1813. Darby, P. C., P. L. Valentine-Darby, H. F. Percival, and W. M. Kitchens. 2001. Collecting Florida applesnails ( Pomacea paludosa ) from wetland habitats using funnel traps. Wetlands 21:308-311. Dole, J. W. 1965. Spatial relations in natural populations of the leopard frog, Rana pipiens Schreber, in northern Michigan. The American Midland Naturalist 74: 464-478. Ernst, C. H. 1986. Ecology of the turtle, Sternotherus odoratus in Southeastern Pennsylvania. Journal of Herpetology 20:341-352.

PAGE 110

99 Ernst, C. H., and E. Ernst. 2003. Snakes of the United States and Canada. Smithsonian Institute Press, Washington DC, USA. Ernst, C. H., J. E. Lovich, and R. W. Bar bour. 1994. Turtles of the United States and Canada. Smithsonian Institute Press, Washington DC, USA. Etheridge, K. 1990. The energetics of estivating sirenid salamanders ( Siren lacertina and Pseudobranchus striatus ). Herpetologica 46:407-414. Florida Fish and Wildlife Conservation Commission. 2003. 2004 Lake Tohopekaliga habitat enhancement project, a fishery mana gement program. Freshwater Fisheries Division, Kissimmee, Florida, USA. Florida Fish and Wildlife Conservation Commission. 2004. Kissi mmee Chain of Lakes highlights, August 13, 2004. Aquatic Ha bitat Conservation and Restoration Section, Kissimmee, Florida, USA. Gehlbach, F. R., and S. E. Kennedy. 1978. Population ecology of a highly productive aquatic salamander ( Siren intermedia ). Southwestern Naturalist 23: 423-430. Gibbons, J. W., J. L. Greene, and J. D. Congdon. 1983. Drought-related responses of aquatic turtle populations. J ournal of Herpetology 17:242-246. Godley, J. S. 1980. Foraging ecolo gy of the striped swamp snake, Regina alleni in Southern Florida. Ecol ogical Monographs 50:411-436. Godley, J. S. 1983. Observations on the courtship, nests and young of Siren intermedia in southern Florida. The Amer ican Midland Naturalist 110:215-219. Harper, R. M. 1921. Geography of cent ral Florida. Florida State Geological Survey, 13th Annual Report. Hasler, A. D. 1947. Eutrophication of lakes by domestic drainage. Ecology 28:383–395 Hayek, L. C. 1994. Analysis of amphi bian biodiversity data. Pages 207-269 in W. R. Heyer, M. A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M. S. Foster, editors. Measuring and monitoring biol ogical diversity: standard methods for amphibians. Smithsonian Institution, Washington D.C., USA. HDR Engineering, Inc. 1989. Technical report for the deve lopment of a surface water improvement and management plan for Lake Tohopekaliga/East Lake Tohopekaliga, Final Report. South Florida Water Management District, West Palm Beach, Florida, USA. Contract No. 88-475-0961. Hines, J. E., T. Boulinier, J. D. Nichols, J. R. Saur, and K. H. Pollock. 1999. COMDYN: software to study the dynamics of animal communities using a capture-recapture approach. Bird Study 46 Supplement, 209-217.

PAGE 111

100 Holt, G. J. 2002. Human impacts. Pages 222-242 in L. A. Fuiman and R. G. Werner, editors. Fishery science: the unique cont ributions of early life stages. Blackwell Science, Ltd., Osney Mead, Oxford, UK. Hostache, G., and J. H. Mol. 1998. Reproduc tive biology of the neotropical armoured catfish Hoplosternum littorale (Siluriformes-Callichthyida e): a synthesis stressing the role of the floating bubble nest Aquatic Living Resources 11:173-185. Hoyer, M. V., and D. E. Canfield, Jr. 1994. Handbook of common freshwater fish in Florida lakes. University of Fl orida, Gainesville, Florida, USA. Iverson, J. B. 1982. Biomass in turtle populations: a neglected subject. Oecologia 55:69-76. Johnson, S. A., and W. J. Barichivich. 2004. A simple technique for trapping Siren lacertina Amphiuma means and other aquatic vertebrate s. Journal of Freshwater Ecology 19:263-269. Joly, P., and A. Morand. 1997. Amphibian di versity in land-water ecotones. Pages 161182 in Lachavanne, J. B., and R. Juge, editors Biodiversity in land-inland water ecotones Man and the Biosphere Series. Volume 18. UNESCO, Paris, France, and Parthenon Publishing, Carnforth, England. Kilby, J. D. 1936. A biological analysis of the food and feeding habits of Rana sphenocephala (Cope) and Hyla cinerea (Schneider). M.S. Thesis. University of Florida, Gainesville, Florida, USA. Klemens, M. W. 2000. Turtle Conservation. Smithsonian Institution Press, Washington, DC, USA. Lachavanne, J. B. 1997. Why study biodiversit y in land-inland water ecotones? Pages 1-45 in Lachavanne, J.B., and R. Juge, edito rs. Biodiversity in Land-Inland Water Ecotones Man and the Biosphere Series. Volume 18. UNESCO, Paris, France, and Parthenon Publishing, Carnforth, England. Lagler, K. F. 1943. Methods of collecti ng freshwater turtles. Copeia 1943:21-25. Larson, R. E., R. P. Hostetler, and B. H. Edwards. 1994. Calculus with analytic geometry, fifth edition. D.C. Heath and Company, Massachusetts, USA. MacKenzie, D. I., J. D. Nichols, G. B. L achman, S. Droege, J. A. Royle, and C. A. Langtimm. 2002. Estimating site occupanc y rates when detec tion probabilities are less than one. Ecology 83:2248-2255. Mahmoud, I. Y. 1968. Feeding behavior in ki nosternid turtles. Herpetologica 24: 300305.

PAGE 112

101 Mahmoud, I. Y. 1969. Comparative ecology of the kinosternid tur tles of Oklahoma. The Southwestern Naturalist 14:31-66. Martof, B. S. 1969. Prolonged inanition in Siren lacertina Copeia 1969:285-288. McCune, B. and J. B. Grace. 2002. Analysis of ecological communities. MjM Software Design, Gleneden Beach, Oregon, USA. McCune, B., and M. J. Mefford. 1999. Multivariate analysis of ecological data, version 4.20. MjM Software, Gleneden Beach, Oregon, USA. Mills, M. S., C. J. Hudson, and H. J. Berna. 1995. Spatial ecology and movements of the brown water snake ( Nerodia taxispilota ). Herpetologica 51:412-423. Miranda, L. E., and E. D. Dibble. 2002. An ecological foundation for black bass management. Pages 433-453 in D. P. Philipp and M. S. Ridgeway. Black bass: ecology, conservation, and management. Am erican Fisheries Society, Symposium 31, Bethesda, Maryland, USA. Miranda, L. E., and K. B. Hodges. 2000. Role of aquatic vegetation coverage on hypoxia and sunfish abundance in bays of a eutrophic reservoir. Hydrobiologia 427:51-57. Miranda, L. E., M. P. Driscoll, and M. S. Allen. 2000. Transient physiochemical microhabitats facilitate fish survival in inhospitable aquatic plant stands. Freshwater Biology 44:617-628. Mol, J. H. 1993. Structure and function of floating bubble nests of three armoured catfishes (Callichthyidae) in relation to the aquatic environment. Pages 167-197 i n P. E. Ouboter, editor. Freshwater Ec osystems of Suriname. Monographiae Biologicae 70. Klewer, Do rdrecht, the Netherlands. Moulton, C. A., W. J. Fleming, and B. R. Nerney. 1996. The use of PVC pipes to capture hylid frogs. Herpet ological Review 27:186-187. Moyer, E. J., M. W. Hulon, R. S. Butler, D. C. Arwood, C. Michael, and C. A. Harris. 1987. Kissimmee Chain of Lakes studi es, completion report for Lake Tohopekaliga investigations. State of Florida Game and Fresh Water Fish Commission, Tallahassee, Florida, USA. Moyer, E. J., M. W. Hulon, J. Buntz, R. W. Hujik, J. J. Sweatman, C. S. Michael, D. C. Arwood, and A. C. Jasent. 1993. Comp letion report for Kissimmee Chain of Lakes studies (1987-1992). State of Florida Game and Fresh Water Fish Commission, Tallahassee, Florida, USA. Moyer, E. J., M. W. Hulon, J. J. Sweatman, R. S. Butler, and V. P. Williams. 1995. Fishery responses to habitat restoration in Lake Tohopekaliga, Florida. North American Journal of Fish eries Management 15:591-595.

PAGE 113

102 Mushinsky, H. R., and J. J. Hebrard. 1977. Food partitioning by five species of water snakes in Louisiana. Herpetologica 33:162-166. Mushinsky, H. R., J. J. Hebrard, and D. S. Vodopich. 1982. Ontogeny of water snake foraging ecology. Ecology 63:1624-1629. Mushinsky, H. R., and D. E. Miller. 1993. Predation on water snakes: ontogenetic and interspecific considerations. Copeia 1993:660-665. Nichols, J. D., T. Boulinier, J. E. Hines, K. H. Pollock, and J. R. Saur. 1998. Estimating rates of local species extinction, colonizat ion, and turnover in animal communities. Ecological Appli cations 8:1213-1225. Nico, L. G., S. J. Walsh, and R. H. Robins 1996. An introduced population of the South American Callichthyid catfish Hoplosternum littorale in the Indian River Lagoon system, Florida. Flor ida Scientist 59:189-200. Nico, L. G., and P. L. Fuller. 1999. Spatial a nd temporal patterns of nonindigenous fish introductions in the United States. Fisheries 24:16-27. Page, L. M. 1994. Identification of sailfin catfishes introduced to Florida. Florida Scientist 57:171-172. Parker, J. O., Jr. 1970. Surfacing of dead fish following application of rotenone. Transactions of the American Fisheries Society 99:805-807. Petranka, J. W. 1998. Salamanders of th e United States and Canada. Smithsonian Institute Press, Washington, DC, USA Pieczynska, E. 1990. Lentic a quatic-terrestrial ecotones: th eir structure, functions and importance. Pages 103-140 in : R. J. Naiman and H. Decamps, editors. The ecology and management of aquatic-terrest rial ecotones. The Parthenon Publishing Group, Paris, France. Pieczynska, E, and M. Zalewski. 1997. Ha bitat complexity in land-inland water ecotones. Pages 61-108 in : Lachavanne, J. B., and R. J uge, editors. Biodiversity in land-inland water ecotones Man and the Biosphere Series. Volume 18. UNESCO, Paris, France, and Parthenon P ublishing, Carnforth, England. Pough, F. H., R. M. Andrews, J. E. Cadle, M. L. Crump, A. H. Savitzky, and K. D. Wells. 2001. Herpetology, second edition. Prentice-Hall, Inc., Upper Saddle River, New Jersey, USA. Reaser, J. K. 2000. Amphibian declines: an issue overview. Federal Taskforce on Amphibian Declines and Deformities, Washington, DC, USA. Schiffer, D. M. 1998. Hydrology of central Florida lakes: a primer. U.S. Geological Survey Circular 1137, Denver, Colorado, USA.

PAGE 114

103 Seigel, R. A., J. W. Gibbons, and T. K. Lynch. 1995. Temporal changes in reptile populations: effects of a severe drought on aquatic snakes. Herpetologica 51: 424434. Snodgrass, J. W., J. W. Ackerman, A. L. Br yan, Jr., and J. Burger. 1999. Influence of hydroperiod, isolation, and heterospecif ics on the distribution of aquatic salamanders ( Siren and Amphiuma ) among depression wetlands. Copeia 1999:107113. Sorensen, K. 2003. Trapping success and population analysis of Siren lacertina and Amphiuma means M.S. Thesis. University of Flor ida, Gainesville, Florida, USA. Sorensen, K. 2004. Population characteristics of Siren lacertina and Amphiuma means in north Florida. Southe astern Naturalist 3:249-258. SPSS Inc. 2001. SPSS 11.0 for Windows. Chicago, Illinois, USA. Wegener, W. 1969. Lake Tohopekaliga fisher y investigation. Florida Game and Fresh Water Fish Commission, Tallaha ssee, Florida, USA. Wegener, W., and V. Williams. 1974. Extreme lake drawdown, a working fish management technique. Dingell-Johnson Federal Aid Project F-29-R. Florida Game and Fresh Water Fish Commissi on, Tallahassee, Florida, USA. Welch, Z. C. 2004. Littoral vegetation of Lake Tohopekaliga: community descriptions prior to a large-scale fisheries habita t-enhancement project. M.S. Thesis. University of Florida, Gainesville, Florida, USA. Werner, R. G. 2002. Habitat requirements. Pages 161-182 in L. A. Fuiman and R. G. Werner, editors. Fishery science: the uni que contributions of early life stages. Blackwell Science, Ltd., Osney Mead, Oxford, UK. Werner, E. E., and D. J. Hall. 1979. Fo raging efficiency and habitat switching in competing sunfishes. Ecology 60:256-264. Werner, E. E., and D. J. Hall. 1988. Ontoge netic habitat shifts in bluegill: the foraging rate-predation risk tradeoff. Ecology 69: 1352-1366. White, G. C., D. R. Anderson, K. P. Burnham, and D. L. Otis. 1982. Capture-recapture and removal methods for sampling clos ed populations. Los Alamos National Laboratory Report LA-8787-NERP, Los Alamos, New Mexico, USA. White, G. C., and K. P. Burnham. 1999. Program MARK: Survival estimation from populations of marked animals. Bird Study 46 Supplement, 120-138. Williams, B. K., J. D. Nichols, and M. J. Conroy. 2002. Analysis and management of animal populations. Academic Pre ss, San Diego, California, USA.

PAGE 115

104 Wilson, K. R., and D. R. Anderson. 1985. Eval uation of two density estimators of small mammal population size. J ournal of Mammalogy 66:13-21. Wygoda, M. L. 1979. Terrestrial activity of striped mud turtles, Kinosternon baurii (Reptilia, Testudines, Kinos ternidae) in West-central Florida. Journal of Herpetology 13:469-480. Zimmerman, B. L. 1994. Audio-st rip transects. Pages 92-97 i n W. R. Heyer, M. A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M. S. Foster, editors. Measuring and monitoring biological diversity: standard methods for amphibians. Smithsonian Institution, Washington D.C, USA.

PAGE 116

105 BIOGRAPHICAL SKETCH Ann Marie Muench was born in Louisvil le, KY, in 1977. Through middle and high school she also lived in Cumberland, MD, and Jacksonville Beach, FL. After high school, she attended the Univer sity of North Florida in Jacksonville, FL, where in 1998 she received a Bachelor of Science degree in biology. After graduation she worked as a Metals Analyst at Environmental Conserva tion Laboratories, Inc., an environmental testing laboratory in Jacksonv ille, FL. Ann Marie decided to pursue her career goal of catching reptiles and amphibian s, and applied to the Depart ment of Wildlife Ecology and Conservation at the University of Florida fo r graduate school. For the next three years she examined the ecology of the aquatic ve rtebrate community in Lake Tohopekaliga, FL, as a graduate assistant with the Florida Cooperative Fish and Wildlife Research Unit. She received her Master of Science degr ee in December, 2004, and subsequently moved to Black Mountain, NC.


Permanent Link: http://ufdc.ufl.edu/UFE0008580/00001

Material Information

Title: Aquatic Vertebrate Usage of Littoral Habitat Prior to Extreme Habitat Modification in Lake Tohopekaliga, Florida
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: UFE0008580:00001

Permanent Link: http://ufdc.ufl.edu/UFE0008580/00001

Material Information

Title: Aquatic Vertebrate Usage of Littoral Habitat Prior to Extreme Habitat Modification in Lake Tohopekaliga, Florida
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: UFE0008580:00001


This item has the following downloads:


Full Text












AQUATIC VERTEBRATE USAGE OF LITTORAL HABITAT PRIOR TO EXTREME
HABITAT MODIFICATION IN LAKE TOHOPEKALIGA, FLORIDA















By

ANN MARIE MUENCH


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


2004

































Copyright 2004

by

Ann Marie Muench


































This document is dedicated to my parents, Joseph F. and Mary K. Muench, whose love
and support have strongly contributed to my academic, professional, and personal
growth.















ACKNOWLEDGMENTS

I would like to thank my major advisor, Wiley Kitchens, for taking me on as a

graduate student. He was a critical source of expert advice and encouragement, and was

always accessible for consultation. I also thank my committee members, Madan Oli and

Lauren Chapman, whose academic instruction and critical analysis of this thesis are much

appreciated. My coworkers also contributed much to my education, and I am thankful.

Funding for this research was provided by the Florida Fish and Wildlife

Conservation Commission (FFWCC). From this agency, Duke Hammond helped

immensely with the direction of the study, and provided critical feedback on progress

reports that we provided to the commission. The staff of the Kissimmee, FL, office of

the FFWCC was helpful in facilitating our field work at Lake Tohopekaliga. Bobbi Jo

Cromwell from the Osceola County Department of Parks and Recreation allowed us to

store all of the crayfish and minnow traps on Makinson Island in Lake Toho.

The field work for this study was conducted through the time of many dedicated

students and technicians from the Florida Cooperative Fish and Wildlife Research Unit.

These stalwart coworkers included (in alphabetical order) Scott Berryman, Stephen

Brooks, Janell Brush, Brenda Calzada, John Davis, Jamie Duberstein, Bruno Ferreira,

Joey Largay, Kristianna Lindgren, Samantha Musgrave, April Norem, Derek Piotrowicz,

Laura Pfenninger, Erik Powers, Vanessa Rumancik, John David Semones, Micheala

Spears, Chris Tonsmeire, Paul Traylor, Zach Welch, and Christa Zweig.
















TABLE OF CONTENTS

page

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

LIST OF TABLES ............. ................... ......... .... ... ............... .. vii

LIST O F FIG U RE S ......................................................... ......... .. ............. viii

A B ST R A C T .......... ..... ...................................................................................... x

CHAPTER

1 M A IN IN TR OD U CTION .................................................. .............................. .

L ake E cosy stem .....................................................................................................1
Stu dy A rea ......................................................................... . 2
R research Objectives.............................................. 8

2 DESCRIPTIONS OF FOCAL SPECIES .................................. ...............10

A quatic V ertebrate H habitat ................................................................................... 10
Fish Species ................................. ........................................... 11
C entrarch id s (su n fish ) ................................................................................... 1 1
E x o tic c catfish ................................................................ ............................... 1 3
H erpetofaunal Species ............................................................13
A m phibians ............................................. 13
R reptiles .................................................................................... ................... 15

3 AS SEMBLAGE WITHIN THE Pontederia cordata COMMUNITY ................. 17

In tro du ctio n ...............17..............................................
Field Methods ......... ......... ......... .. ...............17
T rap D description s .............................................................17
W hole-Lake Sam pling .............. ........ ................ ............... ............... 20
A analysis M methods ..................... .................. ......................... 23
Trap C om prisons ...................... ............ .......................................... 23
Species R richness .................................. ...........................................23
Assemblage Composition ............................ ........ ............25
Influence of Temporal Gradients on Assemblage .......... ..................... 25
Proportion of Habitat Utilized by Focal Species ........................... ............. 27


v









R e su lts ...........................................................................................2 9
T rap C om prison s ......................... ............................ .. ......... .... ............29
S p e cie s R ich n e ss ........................................................................................... 3 1
A ssem blage Com position.................................... ......................... ............ ... 32
Influence of Temporal Gradients on Assemblage.............................................34
Proportion of Habitat Utilized by Focal Species....................................40
Discussion ..................................................41
T rap C om p arison s .............................. .... ...................... .. ........ .... ............4 1
S p e cie s R ich n e ss ........................................................................................... 4 1
A ssem blage C om position ................................................................................ 42
Influence of Temporal Gradients on Assemblage.................. .............. 44
Proportion of Habitat Utilized by Focal Species............................. ..............45

4 ASSEMBLAGE ACROSS VEGETATION COMMUNITIES ..............................64

In tro d u ctio n ............. ........... ... .................. ................. ................ 6 4
F ield M ethods ....................................................... 64
G rid and W eb Sam pling ............................................. ............................. 64
W hole-Lake Sam pling....................... ......................... ................. .. ............. 66
A analysis M ethods ............... .......... .. .......... .. ...... .. ................. .... ... ........... 67
Population Estimates and Movement for Herpetofaunal Species .....................67
Capture Success for Focal Species ......... .............. ................. ... .............. 68
R e su lts ....................... .. .. .............. .. .......................... ................ 7 0
Population Estimates for Herpetofaunal Species .............................................70
Capture success for Focal Species.................................... ....................... 72
D iscu ssion .................. ....... .... ....... ...... ......................... 74
Population Estimates for Herpetofaunal Species .........................................74
Capture Success for Focal Species .............. ..................... .............. 75

5 SUMMARY AND CONCLUSIONS................... .. .................. ...............88

Review of Aquatic Vertebrate Community Dynamics in Lake Tohopekaliga ...........88
A ir T em perature ........................................ ................. .... ..... .. 89
L ak e S tag e ...................................................... ................ 8 9
W ater D epth ............ ...... ....... ..... .............. .. ......... ..... ........... 90
V egetation C om m unity ............................................... ............................ 90
Population Size E stim ates ........................................... ............................ 90
Lake Tohopekaliga Habitat Enhancement.................... ............ ............... 91
Future Aquatic Vertebrate M monitoring Plans .................................. ............... 95

L IST O F R E FE R E N C E S ...................... ........................................................ .......... 97

BIOGRAPH ICAL SKETCH ....................... .......... ......... .................................. 105















LIST OF TABLES


Table page

3-1 All species captured in 2002, with species codes used in subsequent figures.........30

3-2 Species capture frequencies for the 0.6 and 1.3 cm mesh minnow traps ................31

3-3 Indicator species analysis results........................................ .......................... 33

3-4 Stress and instability results from all NMS ordinations.............................34

3-5 Percent of variance explained (r2) by environmental variables for each axis in the
vertebrate NMS with detection/nondetection data............... ....... ............... 35

3-6 Percent of variance explained (r2) for each axis by species in the vertebrate NMS
w ith detection/nondetection data ......................................................... ............... 35

3-7 Percent of variance explained (r2) by environmental variables for each axis in the
vertebrate N M S w ith count data........................................ ........................... 36

3-8 Percent of variance explained (r2) for each axis by species in the vertebrate NMS
w ith count data. .......................................................................36

3-9 Percent of variance explained (r2) by environmental variables for each axis in the
fish NM S with detection/nondetection data.......................................................37

3-10 Percent of variance explained (r2) for each axis by species in the fish NMS with
detection/nondetection data.......................................................... ................37

3-11 Percent of variance explained (r2) by environmental variables for each axis in the
fish N M S w ith count data............................................... .............................. 38

3-12 Percent of variance explained (r2) for each axis by species in the fish NMS with
c o u n t d ata .......................................................................... 3 8

3-13 Percent of variance explained (r2) by environmental variables for each axis in the
herpetofaunal NM S with count data.................................... ......................... 39

3-14 Percent of variance explained (r2) for each axis by species in the herpetofaunal
NMS with count data..................... ................... ............ 39

4-1 Grid sizes and population estimates by mark recapture methods. .........................71
















LIST OF FIGURES


Figure page

3-1 Crayfish and minnow trap in P. cordata habitat. .............................................. 47

3-2 Locations of 2002 P. cordata sampling transects in Lake Tohopekaliga ...............48

3-3 2002 Vertebrate species richness estimates by sample date..................................49

3-4 2002 Fish species richness estimates by sample date......................................... 49

3-5 2002 Herpetofaunal species richness estimates by sample date ............................ 50

3-6 NMS ordination of sample units in vertebrate species space using
detection/nondetection data.......................................................... ............... 51

3-7 NMS ordination of vertebrate species in sample unit space using
detection/nondetection data.......................................................... ................52

3-8 NMS ordination of sample units in vertebrate species space using count data........53

3-9 NMS ordination of vertebrate species in sample unit space using count data .........54

3-10 NMS ordination of sample units in fish species space using detection/
n on d election d ata ............ ........................................................................... .. ....... .. 5 5

3-11 NMS ordination of fish species in sample unit space using detection/nondetection
d a ta .............................................................................. 5 6

3-12 NMS ordination of sample units in fish species space using count data..................57

3-13 NMS ordination of fish species in sample unit space using count data .................58

3-14 NMS ordination of sample units in herpetofaunal species space using count data..59

3-15 NMS ordination of herpetofaunal species in sample unit space using count data ...60

3-16 Average and range of lake stage values by cluster................................................61

3-17 Average and range of air temperature values by cluster. .......................................61

3-18 Site occupancy estimates for focal fish species in spring 2002. ...........................62









3-19 Site occupancy estimates for focal fish species in fall 2002. ..................................62

3-20 Site occupancy estimates for focal herpetofaunal species in spring 2002. ..............63

3-21 Site occupancy estimates for focal herpetofaunal species in fall 2002 ..................63

4-1 Locations of 2003 sampling transects in Lake Tohopekaliga. ..............................80

4-2 Diagram of weekly trap placement at specified depths........................................81

4-3 Mean maximum distances traveled with variances and maximum distances,
based on results of mark-recapture sampling. .......................................................81

4-4 Number of trap sites sampled in each vegetation community per sample occasion. 82

4-5 Salamander capture success by vegetation community..............................83

4-6 Salamander capture success by water depth. ................................... ..................... 83

4-7 Frog capture success by vegetation community........................................... 84

4-8 Frog capture success by water depth ..................................................................... 84

4-9 Snake capture success by vegetation community. ................................................85

4-10 Snake capture success by water depth............................................. ...............85

4-11 Turtle capture success by vegetation community. ................................................86

4-12 Turtle capture success by water depth................................... ....................... 86

4-13 Fish capture success by vegetation community. ................... ................... .......... 87

4-14 Fish capture success by water depth. ............................................ ............... 87















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

AQUATIC VERTEBRATE USAGE OF LITTORAL HABITAT PRIOR TO EXTREME
HABITAT MODIFICATION IN LAKE TOHOPEKALIGA, FLORIDA
By

Ann Marie Muench

December 2004

Chair: Wiley Kitchens
Major Department: Wildlife Ecology and Conservation

Lake Tohopekaliga is a large, shallow lake in central Florida that is part of the

Kissimmee chain of lakes. Cultural eutrophication and lake stabilization over the past

several decades have facilitated the formation of a densely vegetated, often monotypic,

littoral zone. Lake managers conducted an enhancement project in 2004 to improve

largemouth bass (Micropterus salmoides) habitat. This project included an extreme water

level drawdown and concurrent mechanical removal of 7.3 million cubic meters of

organic sediment and vegetation from the shoreline. Following the drawdown, herbicidal

treatments will keep the lake vegetation in an early state of succession in order to prolong

the effects of the enhancement. Little is known about potential impacts of these

procedures on wildlife, including vegetation, avian, herpetofaunal, and even fish

communities. This study examines the status of the reptile, amphibian and fish

communities in the two years prior to the lake enhancement to provide baseline data for

future assessments.









Funnel traps were used for all sampling, allowing a suite of vertebrate species to be

examined. In 2002, sampling was conducted in the Pontederia cordata pickerelweedd)

zone of the lake. Cluster analysis, indicator species analysis, and nonmetric

multidimensional scaling ordinations were used to examine temporal changes in the

species composition of the assemblages. Environmental variables such as lake stage and

average air temperatures played large roles in structuring the aquatic vertebrate

communities through species richness and assemblage composition. Fish assemblages

were most correlated with air temperature, while herpetofaunal assemblages mainly

showed association with lake stage. Site occupancy estimates showed that many of the

herpetofaunal species are present throughout the pickerelweed habitat in both the spring

and fall seasons, while fish showed more fluctuation in seasonal presence.

Spatial sampling took place in 2003. Sampling was conducted across vegetation

communities and water depths. Both variables captured varying trends in the presence of

the focal species, which included fully aquatic salamanders (Siren spp., Amphiuma

means), water snakes (mainly Nerodia spp.), small kinostemid turtles (Kinosternon

baurii, S.i ntu,,hei ,1n odoratus), large aquatic frogs (Rana spp.), juvenile centrarchids

(especially Micropterus salmoides) and exotic catfish (Hoplosternum littorale).

Attempted population density estimates for the more abundant herpetofaunal species

ended in failure. Inappropriate trapping grid size and spacing for several species at one

time led to poor capture probabilities and large variances in population size estimates.














CHAPTER 1
MAIN INTRODUCTION

Lake Ecosystem

The productive littoral environment in a lake system is dynamic, since the aquatic

habitat has strong terrestrial influences and the terrestrial habitat has strong aquatic

influences. Biological diversity is high in the ecotone due to biotic and abiotic properties

that distinguish it from adjacent ecosystems, such as vegetation species, soil properties,

and water chemistry (Lachavanne 1997). Water level fluctuations are the main

determinants of the width of the littoral zone. In lentic systems with gently sloping

shorelines a wide band of macrophytes provides patches of heterogeneous habitat for a

diverse assemblage of faunal species. Animal species that have specific requirements for

different life stages depend on the proximity of supralittoral (never flooded), eulittoral

(occasionally flooded), and infralittoral (always flooded) habitats. Since the ratio of

water surface to volume is much higher in the littoral zone than in the deeper pelagic

region, environmental variables such as light, air temperature, wind and water flow

(waves) have much more critical roles in shaping the gradient (Pieczynska 1990,

Pieczynska and Zalewski 1997).

Excess inputs of nutrients beyond that naturally found in a particular lake system

leads to eutrophication. This condition encourages surplus sedimentation and vegetation

growth, changing the landscape of the original littoral zone and subsequently altering the

biological communities within that habitat and the lake as a whole. Eventual extinction

of the lake may result from decades of nutrient pollution due to sewage discharge and









drainage from agricultural and urban lands. Internal recycling of nutrients within a water

body keeps it from recovering even when the inputs are reduced (Cooke et al. 1993). In

order to preserve the lake for the longest time possible, rehabilitation efforts are often

made to counter the results of eutrophication, for example drawdowns, dredging and

mechanical vegetation removal (Hasler 1947, Cooke et al. 1993).

Study Area

Lake Tohopekaliga (7,612 ha, 18,810 acres) is located in Osceola County, Florida,

within the Upper Kissimmee Basin. This physiographic area is known as the Osceola

Plain, which is flat and has very few distinguishing topographical characteristics. The

elevation in the plain ranges from 18-30 m (60-95 ft) National Vertical Geodetic Datum

(NGVD), but rarely reaches maximum heights (Harper 1921). Originating from

prehistoric ocean bottom, the sediment mainly consists of coastal sands. Numerous

shallow lakes in the region, including Lake Tohopekaliga, were formed by dissolution of

the carbonate-containing substrates (limestone) in depressed areas (Schiffer 1998).

Freshwater wetlands in this area include cypress sloughs, wet prairies, river swamps,

floodplains, mixed forested wetlands, and marshes. Pine flatwoods dominate the upland

community (HDR Engineering, Inc. 1989).

The Lake Tohopekaliga Subbasin (211.6 square km, 131.2 square miles), within the

Upper Kissimmee Basin, receives water input from the Shingle Creek (184.2 square km,

118.0 square miles) and East Lake Tohopekaliga (81.7 square km, 48.4 square miles)

Subbasins. Precipitation, overland flows, and to some extent groundwater from the

underlying Surficial Aquifer also provides the lake with important water sources. While

evapotranspiration is a strong factor in withdrawal of water from the lake, outflow from

Lake Tohopekaliga occurs at its southernmost point, where the South Port Canal connects









it to Cypress Lake. Water from the Upper Kissimmee Basin flows southward through the

Kissimmee Chain of Lakes, through the channelized Kissimmee River, to Lake

Okeechobee, east and west coast estuaries and South Florida (HDR Engineering, Inc.

1989, Schiffer 1998).

Human disturbance of this hydrologic system began in the mid-nineteenth century,

with local efforts to drain wetlands. In 1882 Hamilton Disston began channelizing the

watersheds in the upper basin by constructing inter-lake canals. The major results of this

endeavor were lowered lake levels, drying of lake edges and inter-lake slough wetlands,

as well as rapid transit of nutrient-laden surface waters downstream. Wetlands stretching

between Lake Tohopekaliga and East Lake Tohopekaliga were strongly impacted. In the

1920's, Shingle Creek (a major source of water for Lake Tohopekaliga) was channelized,

bypassing water around the swamps and marshes within that subbasin. Many landowners

also dug ditches and canals to drain wetlands and improve their pastureland. In 1947, the

Central and Southern Florida Flood Control Project was implemented by the U.S. Army

Corps of Engineers in response to major flooding in the Kissimmee Basin. As a result of

this plan, the Kissimmee River was channelized, Disston's inter-lake canals were

improved, and water control structures were built throughout the area. The goal of these

actions was to use the chain of lakes for water management, to provide room for water

during the wet season and to store water during the dry season. This entailed stabilization

of water levels, which historically fluctuated up to 3 m (10 ft), to a 0.6-1.2 m (2-4 ft)

range. This reduction in fluctuation subsequently allowed landowners and private

citizens to build on historic lake bottom and drained wetlands within the floodplain,









further strengthening the need for tight flood control (Blake 1980, HDR Engineering, Inc.

1989).

Lake Tohopekaliga and surrounding lakes have suffered many water quality

problems due to the intense hydrologic modifications. The constructed canals, especially

in the Shingle Creek area, allowed direct conveyance of stormwater runoff and sewage

into the lakes without the benefit of filtration through wetlands. Urban and agricultural

land use continued to expand, contributing more and more overland pollution. The

agricultural land in the area is mainly utilized as pastureland, and historically dairy farms

provided significant inputs of nutrients. Although eutrophication within the lakes was

rapidly increasing, water level stabilization prevented natural fluctuations from mitigating

the problem. Since the lake levels were unnaturally restricted from periodic flooding and

drying events, thick stands of vegetation began invading the littoral zone, which in turn

led to organic sediment buildup and decreased water quality. As of 1988, no wastewater

discharges have been permitted to the lakes. However, non-point source urban and

agricultural runoff and septic tank leakage remain major contributors to eutrophication in

the Kissimmee chain of lakes (HDR Engineering, Inc. 1989).

As mentioned, eutrophication has caused major vegetation changes to the littoral

zone of Lake Tohopekaliga. Dense monotypic expanses of aquatic vegetation began to

dominate the gradually sloping shoreline, formerly characterized by sandy substrate and

sparse vegetation. Nuisance species such as Pontederia cordata pickerelweedd) and

Typha domingensis (cattail) formed wide bands of habitat around the lake. Pontederia

cordata and associated species created floating mats on the lakeward edges of the littoral

zone that rose and fell with the water level. Exotic species such as Hydrilla verticillata









(hydrilla), Eichornia crassipes (water hyacinth), Alternanthera philoxeroides (alligator

weed), and Panicum repens (torpedo grass) also benefited from increased nutrients and

high boat traffic between waterways and became a major focus of lake managers through

the years. Turnover in the vegetation community produced an organic muck substrate

within the littoral zone. Documented faunal responses to this changing habitat have

included declines in fisheries, especially sport and forage fish species, and invertebrates

(HDR Engineering, Inc. 1989).

In 1968, the first fish population surveys in Lake Tohopekaliga were conducted by

the State of Florida Fish and Wildlife Conservation Commission (FFWCC, formerly

Florida Game and Fresh Water Fish Commission, FGFWFC) with rotenone,

electroshocking, and trammel nets (Wegener 1969). Management recommendations

included drawdowns every 5-7 years in order to oxidize the increasingly organic substrate

and provide benefits to the growing fish population upon reflooding of littoral habitats.

In 1971, Lake Tohopekaliga underwent its first extreme drawdown. The water stage was

dropped about 2.1 m (7 ft) from high pool stage (16.8 m, 55.0 ft NGVD). Drought

conditions kept water levels below 15.8 m (52.0 ft) NGVD (low pool stage) until one

year after the initiation of the drawdown. The dewatering of the littoral zone attained the

management goals, with increased acreage of desirable plant species, greater production

of fish and fish-food organisms (invertebrates) per acre, and increased sportfishing

success (Wegener and Williams 1974).

Improvements made to Lake Tohopekaliga's littoral zone were short lived, due to

continued input of nutrients (e.g., about 53 million liters (14 million gallons) of sewage

waste per day discharged into the lake (Wegener and Williams 1974)) and water level









stabilization. An offshore berm was forming at the low-pool water line that was thought

to be acting as a barrier to fish and invertebrates at low water levels (Moyer et al. 1987).

A second drawdown was conducted in 1979 and also had beneficial results, although by

1986 the habitat had degraded once more to sub-optimal fishery habitat. The organic

berm in the lakeward portion of the littoral zone was increasingly becoming a

management issue not able to be addressed by drawdowns alone (Moyer et al. 1987).

When the third drawdown was performed in 1987 a pilot muck removal project was

included. Along 19 km (12 miles) of shoreline, 164,830 cubic meters (225,000 cubic

yards) of muck were mechanically removed from the organic berm. This was considered

"an unprecedented large-scale restoration project to improve littoral habitat" (Moyer et al.

1993, Appendix 4, page 2). All research pointed to highly positive results from the

drawdown and muck-scraping procedures. Fishery surveys have continued since 1968,

and now include roving creel surveys, blocknet/rotenone sampling, electrofishing,

experimental gill nets, and shallow water sampling with Wegener rings. Other wildlife

monitoring included snail kite (Rostrhamus sociabilis) individual and nest counts, limited

aquatic plant sampling, and littoral zone invertebrate community monitoring (Moyer et

al. 1993).

Another lake enhancement project had been planned for early 2002 in Lake

Tohopekaliga, however logistical constraints caused postponement until early 2004

(Florida Fish and Wildlife Conservation Commission 2003). The project originally

included an extreme drawdown from 16.8 m (55.0 ft) NGVD to 14.9 m (49 ft) NGVD

beginning in November 2003, as well as the mechanical removal of about 5.4 million

cubic meters (6.8 million cubic yards) of muck and vegetation from the majority of the









lake's shoreline (Florida Fish and Wildlife Conservation Commission 2003). The

subsequent estimate of the actual volume scraped was 7.3 million cubic meters (8-million

cubic yards), with 1,351 ha (3,339 acres) of shoreline habitat removed. The entire width

of the littoral zone was targeted for removal, not just the organic berm. The Pontederia

cordata-dominated habitat underwent widespread elimination throughout the lake.

Twenty-nine in-lake disposal islands were created from much of the scraped material

(Florida Fish and Wildlife Conservation Commission 2004). Once the water levels

recover, heavy applications of herbicides will be used to keep the habitat in an early state

of succession, allowing lake managers to selectively allow regrowth of desirable

vegetation. Currently, the target conditions post-enhancement are undefined.

The main objective of the Lake Tohopekaliga habitat enhancement project is the

removal large expanses of undesirable macrophyte stands, particularly Pontederia

cordata and Typha domingensis, as well as the organic substrate (muck) associated with

this dense vegetation (Florida Fish and Wildlife Conservation Commission 2003).

Reptiles, amphibians and many juvenile fish species are known to occupy structurally

complex lentic habitats and utilize the muck and thick vegetation for foraging, cover, and

also reproduction (e.g., amphibians). Lake enhancement techniques (both mechanical

vegetation and muck removal and subsequent herbicide applications) modify these

resources, changing the habitat suitability for aquatic vertebrates. High mortality during

the scraping process and migration during the drawdown will likely also alter the

community structure and dynamics. The effort to sustain high species diversity in the

lake ecosystem may be important to the stability of the system, and by examining the









consequences of these restoration techniques managers can better evaluate their worth to

wildlife and fishery species.

Research Objectives

While some positive responses have been documented for the fishery of Lake

Tohopekaliga following past enhancement projects, many wildlife guilds have been

neglected. There is limited quantitative knowledge of vegetation responses to mechanical

removal and large-scale herbicidal treatment. It is also uncertain how wetland birds are

affected. There are still unanswered questions regarding aquatic vertebrates that utilize

the thick vegetation and organic sediment, including reptiles, amphibians and fish.

Herpetofaunal responses to enhancement activities have not been studied in the past, even

though they are pervasive in the habitat. Although fishery science claims that the

eutrophic littoral habitat is unsuitable for centrarchids (i.e. sport fish), conventional

sampling methods may be incapable of detecting them in highly vegetated areas (Parker

1970, Allen et al. 2003).

The current study is part of a larger project evaluating the wildlife response to

habitat enhancement in Lake Tohopekaliga. Also included in this project are vegetation

(see Welch 2004) and avian monitoring studies. The research presented in this thesis

examines the aquatic vertebrate community in the littoral zone of Lake Tohopekaliga

prior to the 2003 drawdown and mechanical vegetation and muck scraping activities.

The littoral zone is defined here as the area occupied by emergent vegetation. However,

there is particular emphasis given to the pickerelweed zone due to its extensive removal

during the lake enhancement. The large-scale wildlife habitat investigation will continue

for at least three years after enhancement activities to examine responses to the

modifications by the different guilds.









With a large-scale habitat modification, quantifying the effect on a whole suite of

species provides maximum information. While most of the species have common

biological or ecological traits, they also constitute a variety of habitat requirements based

on food sources, reproduction methods, and movement patterns. For this reason

community metrics within the land-water ecotone are of main concern, as represented by

species richness and community composition. Species-specific site occupancy and

capture frequencies also facilitate understanding of habitat utilization by focal vertebrate

species. The main objectives of this research are to

1. Characterize the vertebrate faunal makeup of Lake Tohopekaliga's littoral zone
prior to the 2004 lake enhancement project,

2. Estimate parameters such as density and activity/home range for focal species in
the P. cordata habitat,

3. Estimate site occupancy rates for focal species within the littoral zone, as an
estimate of the proportion of the area that the species inhabits,

4. Document how temporally changing variables including lake stage, water level
fluctuation, air temperature, and rainfall shape the aquatic vertebrate community in
the P. cordata zone, and

5. Investigate the influence of spatial variables such as water depth and vegetation
community on the herpetofauna and fish within the landscape.














CHAPTER 2
DESCRIPTIONS OF FOCAL SPECIES

Aquatic Vertebrate Habitat

Wetland communities of reptiles and amphibians show much diversity in

ecological function. Often being the largest and most abundant vertebrates in this habitat,

they have important places in the food webs of lakes (Iverson 1982). Some species

provide terrestrial links while others are fully aquatic and never leave the littoral zone

(Joly and Morand 1997). Fish species also rely on both the littoral, pelagic, and to some

extent flooded terrestrial (nursery) habitats as they undergo shifts with life stage (Werner

2002). There has been a worldwide decline in biodiversity, particularly seen in

amphibian species. A variety of human disturbances have been identified, including

climate change, habitat loss and fragmentation, introduced species, pollution, acid rain,

and disease (Reaser 2000). Florida in particular has been severely impacted by

destruction of wetlands, channelization of streams, manipulated hydrologic cycles, and

rapid human growth (HDR Engineering, Inc. 1989, Pough et al. 2001). Alteration of

freshwater habitats has been a problem for many aquatic species. Animals that are long-

lived or have delayed sexual maturity, low reproductive rates, or poor dispersal or

colonization abilities are particularly vulnerable to habitat destruction (Klemens 2000).

Purposeful habitat modification should preserve conditions necessary for aquatic animals

to complete their life cycles, including appropriate nesting/spawning, foraging, and cover

habitats.









Species of aquatic vertebrates that are most at risk due to their habitat

requirements are emphasized here. Most herpetological research has been conducted on

breeding populations of amphibians, large charismatic reptiles, or single species and

guilds (but see Bancroft et al. 1983). Fishery science remains focused mainly on

sportfish at the individual or population level (Miranda and Dibble 2002). Resident

littoral zone species make up the assemblage of interest for this study and represent

several different orders of animals with a variety of life history traits. Fish guilds, such as

juvenile centrarchids (especially Lepomis spp., Micropterus salmoides) and exotic catfish

(Hoplosternum littorale), are focused upon. Documentation of the presence of these

species in heavily vegetated littoral habitats in Florida is very poor, probably due to

inadequate sampling techniques. Reptile and amphibian species of interest include fully

aquatic salamanders (Siren spp., Amphiuma means), water snakes (mainly Nerodia spp.),

small kinosternid turtles (Kinosternon baurii, Si.e inlheil %// odoratus), and large aquatic

frogs (Rana spp.). Minimal research has been conducted on the effects of lake

management techniques on these herpetofaunal species. Most of these species and guilds

have a common reliance upon vegetated wetlands for at least some part of their life

cycles. They also are often preyed upon by the same species, including alligators,

wading and predatory birds, large predatory fish and aquatic snakes, and together

represent many segments of the food web in the lake ecosystem.

Fish Species

Centrarchids (sunfish)

Most species in the family Centrarchidae in Lake Tohopekaliga are sportfish.

Foremost among them in Florida lakes is the largemouth bass (Micropterus salmoides).

This species is the primary target for benefit by the Lake Tohopekaliga enhancement.









Bluegills (Lepomis macrochirus), redear sunfish (Lepomis microlophus), black crappie

(Pomoxis nigromaculatus), warmouths (Lepomis gulosus), spotted sunfish (Lepomis

punctatus), and dollar sunfish (Lepomis marginatus) are also considered sportfish in

Florida. Enneacanthus glorious, (bluespotted sunfish), has a maximum total length of

80 mm, and is therefore only considered a forage fish species (Hoyer and Canfield 1994).

Most of these species depend upon the vegetated littoral zone during juvenile stages and

for spawning. Vegetated habitats provide juvenile sunfish with protection from larger

predators and abundant food supplies (Werner and Hall 1988, Chapman et al. 1996,

Miranda et al. 2000). The phenomenon of ontogenetic habitat shifts is particularly well

studied in bluegills. This species moves between the littoral to the pelagic zone

throughout its life cycle. The littoral zone provides nesting habitat, as well as a preferred

environment for juvenile bluegills from approximately 12-83 mm standard length due to

size-specific predation risks (Werner and Hall 1988).

It is claimed that these species have no access to Lake Tohopekaliga's littoral

zone due to physical and chemical barriers. The floating mats ofPontederia cordata,

resulting from the eutrophic status of the lake, are thought to form a physical barrier for

centrarchids, limiting adult access to shallow water spawning sites. Even if the fish could

penetrate this barrier, physicochemical characteristics of the dense vegetation would not

permit survival (Moyer et al. 1995, Allen and Tugend 2002, and Allen et al. 2003).

Traditional methods of fish sampling in high-macrophyte littoral habitats in Lakes

Tohopekaliga and Kissimmee, Florida, have yielded few or no centrarchid species

(Moyer et al. 1993, Allen and Tugend 2002). However, since common fish sampling

methods, including electrofishing and rotenone/blocknet, do not perform well in heavily









vegetated habitats (Parker 1970, Moyer et al. 1995, Allen and Tugend 2002), many

suppositions upon which lake enhancement projects depend are theoretical.

Exotic catfish

Hoplosternum littorale is an exotic species in Florida, originating in South

America. This armored catfish was first found within the United States in South Florida

in 1995, and was presumably released through the aquarium trade or aquaculture.

Various life history and behavioral traits, including aerial respiration, large body size,

high environmental tolerances, and nest-guarding behaviors, are responsible for rapid

expansion of its range in Florida (Nico et al. 1996). This species is currently nesting in

and pervasive throughout the littoral zone in Lake Tohopekaliga (personal observation).

Ptei gqpliI hIiy\ spp. (suckermouth or sailfin catfish) has also been captured in Lake

Tohopekaliga, although only on a few occasions. This species was probably released into

Florida through the aquarium trade (Page 1994). These two species may pose significant

ecological threats to native food webs and aquatic plant communities. While the Florida

Fish and Wildlife Conservation Commission conducts yearly monitoring by

electrofishing, this study is the first known report of these species this far north in the

Kissimmee chain of lakes.

Herpetofaunal Species

Amphibians

Rana grylio (pig frog) is a highly aquatic species, rarely being seen on shore.

They are usually associated with dense marsh vegetation. While leopard frogs, including

Rana sphenocephala (Florida leopard frog), prefer habitats with standing water, larger

individuals can inhabit somewhat dryer on-shore habitats and use larger home ranges,

relying on plant shade, dew and soil moisture for survival (Dole 1965). Adult leopard









frogs and their tadpoles are also noticeably absent from sandy, unvegetated shorelines

(Dole 1965, Alford and Crump 1982, Bancroft et al. 1983). These two large frog species

have differences in length of larval development, with pig frogs taking more than a year

to metamorphose and leopard frogs taking only two to three months (Bancroft et al.

1983). This, along with year-round breeding in Florida, results in a variety of size classes

of tadpoles throughout the year.

Siren lacertina (Greater siren) and Amphiuma means (Two-toed amphiuma) are

two of the largest species of salamanders in the world (Petranka 1998). Amphiumas

depend on lungs for aerial respiration, while sirens have external gills as well. Although

they may have lengths greater than 76 cm (Conant and Collins 1998), diminutive limbs in

both species are thought to limit overland dispersal. These salamanders burrow into

organic sediment when their habitats become dry and may remain alive for up to three to

five years in underground burrows without food until water comes back to the habitat

(Martof 1969, Etheridge 1990). Bancroft et al. (1983) found that the density of

amphiumas and greater sirens in Lake Conway, Florida, increased with sediment depth.

They also reported that neither species inhabited sandy, unvegetated shorelines. Sirens

have compressed tails that may help to propel them in vegetated open water as well as

emergent vegetation habitats. Amphiumas on the other hand have long cylindrical tails

and are thought to be limited to shallow water (Bancroft et al. 1983). Sirens feed mainly

on mollusks, insects, crayfish and filamentous algae, as well as some other vegetation.

Amphiumas eat fish, crayfish, salamanders, frogs and a wide variety of other species

(Petranka 1998).









Reptiles

The striped mud turtle (Kinosternon baurii) and common musk turtle

(.Siie nithe/i % odoratus) are both small species (maximum carapace lengths of 12.2 cm

and 13.7 cm respectively) that prefer shallow water wetlands (Conant and Collins 1998).

They are both omnivorous, feeding upon animals and some plants opportunistically.

However, mud turtles are attracted to fast-moving prey while musk turtles search out

more sedentary organisms as they crawl along the substrate in search of prey (Mahmoud

1968). Striped mud turtles usually occur in water greater than 60 cm deep, with lower

water levels or rainfall triggering terrestrial activity (Wygoda 1979, Ernst et al. 1994).

On the other hand, common musk turtles are highly aquatic, not leaving water unless

nesting. This species seems to prefer water depths less than 60 cm, but have been seen in

up to 9 m of water (Ernst et al. 1994). Bancroft et al. (1983) found about 20% of all

captured common musk turtles in the littoral zone, and the rest (usually larger

individuals) in open water habitat. According to Mahmoud (1969), S. odoratus is found

in lakes as well as riverine habitats with gravel or sandy substrates.

Nerodiafasciatapictiventris (Florida water snake) is most often encountered in

the shallowest regions of inhabited wetlands (Ernst and Ernst 2003). They are observed

often in disturbed and white sand littoral habitats (Bancroft et al. 1983). This species eats

mainly fish until they reach a total length of 50 cm, at which point they switch to preying

upon frogs (Mushinsky et al. 1982). Nerodiafloridana (Florida green water snake) is the

largest North American water snake, with total lengths approaching two meters (Conant

and Collins 1998). They are inhabitants of quiet water wetlands and sometimes venture

out into open water (Ernst and Ernst 2003). Bancroft et al. (1983) found them to be

pervasive throughout the littoral zone, and while the dense vegetation seems to be






16


preferred, the species of vegetation may not be very important. Some individuals were

captured up to 40 m from the edge of the littoral zone in open water, while several

terrestrial sightings occurred during winter months. Sediment depths of 11-20 cm

yielded the most individuals, and sandy beach habitats were avoided by Florida green

water snakes (Bancroft et al. 1983). They feed mostly upon fish, but also on frogs,

salamanders, tadpoles, small turtles and invertebrates (Mushinsky and Hebrard 1977,

Ernst and Ernst 2003)














CHAPTER 3
ASSEMBLAGE WITHIN THE Pontederia cordata COMMUNITY

Introduction

The objective of this section is to investigate the temporal variation of community

composition and dynamics. The four main research questions are 1) is there temporal

variation in the aquatic vertebrate assemblage, 2) does community composition change

over time, 3) what environmental factors seem to be influencing the temporal variation in

the assemblage and individual focal species, and 4) how are the focal species dispersed

through the habitat. Key environmental variables that change over the course of a year

include lake stage, water level fluctuation, air temperature, and rainfall. Each of these

will be examined for their influence on the vertebrate assemblage. The thick P. cordata

pickerelweedd) habitat was the prime target for mechanical removal during the lake

enhancement process and therefore was the focus of sampling effort. This protocol was

also used to select focal species (which species were present in the habitat and most

detectable with the traps) and evaluate trap-sampling methods. All of this information

will facilitate monitoring in the future, regarding how, when and where to sample in order

to capture the community dynamics and variances associated with the lake's ever-

changing environment.

Field Methods

Trap Descriptions

As previously mentioned, fishery surveys conducted in the Kissimmee Chain of

Lakes include roving creel surveys, blocknet/rotenone sampling, electrofishing,









experimental gill nets, and Wegener rings (Moyer et al. 1993). These methods collect

information on a variety of fish species, but sport fish are the typical target of research.

Traditional herpetofaunal sampling techniques include visual surveys and hand or dip-net

collecting (Bury and Corn 1991), pitfall and funnel traps in combination with drift fences

(Corn 1994), and use of seines or dredges for removing floating vegetation along with the

animals inhabiting it (Bancroft et al. 1983). None of these methods are appropriate for

the extremely thick, rooted vegetation in the littoral zone. Turtle traps exist, such as hoop

nets and floating traps for basking turtles (Lagler 1943); however large turtles are not

central to this research since they are not restricted to the littoral zone. PVC pipes have

also been used as passive traps for treefrogs (Moulton et al. 1996), and audio surveys are

often used for breeding ranid frogs (Zimmerman 1994). However, a single, all-

encompassing technique was desired for this community study, and the answer came

from funnel traps. Recently, several researchers have noticed the benefits of capturing

aquatic organisms in thick vegetation with crayfish and minnow style funnel traps (Darby

et al. 2001, Sorensen 2003, Johnson and Barichivich 2004). Without the use of either

bulky drift fences or bait, these traps have been successful in capturing a wide variety of

reptiles, amphibians, fish and some invertebrates. Funnel traps were used for all

sampling during this study.

The minnow and crayfish traps were all constructed of 1.3 cm (0.5 in) mesh, dark

green vinyl-coated hardware cloth (Figure 3-1). The crayfish traps, similar to those

described by Darby et al. (2001), were positioned on the substrate, or as near to the

substrate as the vegetation would allow. They were approximately 80 cm (30 in) tall

including a "chimney" extending from the body of the trap, allowing the top to be above









the water surface. At the base were three entry funnels leading into the trap, with each

opening about 6 cm (2.5 in) in diameter, but the exact size varied slightly due to

handmade construction. The modified minnow traps were 60 cm (24 in) long rectangular

traps, which were approximately 25 cm (10 in) deep and 18 cm (7 in) high. At each end

there was one entry funnel, with an opening approximately 9 cm (3.5 in) wide and 6 cm

(2.5 in) tall. Floats made of Styrofoam pool toys ("Wacky Noodles") were attached to

the minnow traps to allow them to float halfway out of the water, with the funnels about

even with the water surface, based on the design by Casazza et al. (2000). The funnels

permitted animal access into the trap, but discouraged escape by making the exits harder

to find than the entrances. By allowing the traps to remain partially above the water, the

animals had access to air and mortality was reduced. Both nocturnal and diurnal species

were accessible to capture since traps could be deployed without time constraint. The

traps were not baited, however once an animal was captured in the trap other animals

may have been attracted to it.

The dimensions of the traps restricted the assemblage of animal species captured.

The traps did not confine young individuals or small species of fish, frogs, snakes and

salamanders due to the 1.3 cm (0.5 in) mesh size. Also, individuals larger than the funnel

diameter were excluded. To compare the difference in species captured with 1.3 cm (0.5

in) versus 0.6 cm (0.25 in) mesh, 18 commercially-manufactured minnow traps, similar

to the "eelpots" used in Casazza et al. (2000), were deployed at randomly assigned trap

points from 11/5/2003 to 1/8/2004. These traps are cylindrical, about 60 cm (24 in) long

and 23 cm (9 in) in diameter, with the funnel openings about 5 cm (2 in) in diameter.

They were also fitted with floatation. The hardware cloth was bare metal, not vinyl-









coated. Comparisons of species and number of captures were made between the 0.6 cm

mesh minnow traps and the 1.3 cm mesh modified minnow traps from the same trap

points during this sampling period. We expected to capture more species with the

smaller mesh size since small species and younger individuals could escape from the

larger mesh, but be retained by the 0.6 cm holes.

Whole-Lake Sampling

To gain information regarding temporal habitat utilization by the aquatic vertebrate

assemblage, sampling was conducted around the periphery of the whole lake to maximize

the inference of the results to the system. For the whole-lake sampling, 18 sites were

randomly selected from the less developed, southern two-thirds of the lake (Figure 3-2).

At each site, a transect was established with three trap locations placed perpendicular to

shore and spaced approximately 10 m (33 ft) apart, except in disturbed stretches of

habitat with barriers within this distance (e.g., commercial airboat trails). One crayfish

and one minnow trap were placed at each trap location, attached to a PVC pole for extra

stability. The result of this trapping arrangement was uniform sampling effort at each

transect. The trap locations were placed in the most lakeward portion of dense P. cordata

when possible (mainly in the 0.6-0.9 m (2-3 ft) depth zone at 13.8 m (55 ft) NGVD). The

transect sites varied in proximity to the ecotone between the open water habitat and the

vegetation. Most transects had thick stands of Typha or more diverse floating mats

between the relatively monotypic sections of pickerelweed and open water. The band of

emergent macrophytes at these locations was comparatively broad. On the other hand, at

some transects the traps were relatively close to this ecotone due to narrowness of the

pickerelweed zone at these locations, well established commercial airboat trails, or

herbicide applications near the transects providing large unvegetated areas. One transect









fell in an area where the substrate had previously been scraped, and the vegetation

consisted mainly ofHydrilla verticillata (hydrilla) and very few emergent macrophytes.

The whole-lake trap survey was conducted year-round, pending suitable water

levels (greater than approximately 16 m, 52.5 ft NGVD). Below this point, there was not

enough water for the trapped animals and rodents and birds were inadvertently captured.

Sampling throughout the year 2002 was as follows:

* January 24 Traps were deployed to randomly selected transects and sampling
began.

* May 2 Insufficient water levels in the pickerelweed zone caused traps to be
removed and sampling suspended.

* June 12 Redeployment of traps to select transects with sufficient water depth
resulted in decreased trapping effort until July 24.

* July 24 All traps were back in place in fixed sampling locations.

* December 3 Traps were removed from pickerelweed zone due to low water
associated with the attempted 2002 drawdown.

When active, the traps stayed in place day and night and were typically checked

once weekly. Despite efforts to keep samples spaced seven days apart, the time interval

was not always consistent due to logistical issues (e.g., airboat problems, rough weather).

At each sampling occasion, two or three observers traveled to each transect in an airboat

and checked the traps for their contents. All animals were brought back to the boat to be

worked up. Reptiles and amphibians were weighed individually with Pesola spring

scales and certain length measurements were taken, depending on the species. Fish were

identified and grouped according to species for each trap. All individuals of each species

per trap were weighed together in order to obtain the total biomass of the fish species

caught. After being worked up, the animals were released at the transects where they

were captured. The types of data collected with these methods include species detection-









nondetection, number of individuals captured on each sample occasion, biomass, and

reptile and amphibian length measurements.

This sampling protocol was intended to continue in the exact same locations post-

lake enhancement (2003) in order to compare community traits before and after the

modifications. However when the drawdown and muck removal was postponed for

another year, it was no longer beneficial to keep sampling since there was not a

before/after comparison to be made. Variables such as water temperature and dissolved

oxygen were not measured directly since this was not the initial focus of the study.

Alternative environmental variables were obtained using Internet resources. Lake stages

and rainfall were taken from the South Florida Water Management District's DBHYDRO

browser (http://glades.sfwmd.gov/pls/dbhydro_pro_plsql/). The lake stage was the mean

daily average taken from the headwater of Station S61 (the water control structure in the

south part of the Lake Tohopekaliga leading to Lake Cypress via the South Port Canal) in

feet NGVD. Lake stage was recorded for the day of each sample occasion. Water

fluctuation for one sample is the difference of the water level at that sample minus the

water level at the previous sample occasion. Rainfall was also recorded at Station S61

and precipitation totals for each sample were added up from the day of the previous

sample occasion until the day before the new sample occasion. Air temperature data was

gathered from the National Oceanic and Atmospheric Administration's National Climatic

Data Center's website (http://www.ncdc.noaa.gov/servlets/ULCD). The weather station

location was the Orlando International Airport (MCO) in Orlando, Florida. This is

located approximately six kilometers (10 miles) from the north shore of Lake

Tohopekaliga. Average temperatures were calculated for every sample occasion by









averaging the daily average temperatures from the day of the previous sample occasion

until the day before the new sample occasion. Minimum and maximum temperatures

were also recorded for each sample period.

Analysis Methods

Trap Comparisons

To determine the utility of traps with smaller mesh size for this study, the species

and number of captures for the 0.6 cm (0.25 in) mesh commercial minnow traps and the

1.3 cm (0.5 in) mesh modified minnow traps were compared. The 0.6 cm mesh traps

were randomly placed at only a portion of the whole-lake trap sites, along with a 1.3 cm

mesh crayfish and minnow trap. For this reason, data from both minnow traps were

compared for just the trap sites with both mesh types.

Species Richness

Sampling was carried out with a repeated measures protocol, potentially resulting

in lack of independence between samples. However, assuming random movement of

individuals and species through the habitat over space and time, sampling over time did

not result in repeated captures of the same individuals. This transient nature of the

species and utilization of non-parametric procedures for most analyses are believed

remove potential bias due to repeated samples. To determine the presence of temporal

variation in the aquatic vertebrate assemblage, total species richness was calculated for

each sample occasion in 2002. Fish and herpetofaunal species richness were also

individually estimated for each sample occasion. Program COMDYN4 was used with

detection-nondetection data to estimate richness (Hines et al. 1999), taking into account

species detection probabilities. It uses a model (Mh) that allows each species to have a

different detection probability (the probability of detecting at least one individual of the









species). Since most species detection probabilities are less than one, raw count data can

result in underestimations of richness. In a similar manner, the term "presence/absence

data" can also be misleading since lack of detection provides no evidence of a species'

absence from the trap site. For this reason I instead use the term "detection/nondetection

data" throughout this thesis. Equal sampling effort is necessary for each occasion.

Assumptions of this method are 1) population closure for species, 2) independence of

captures and 3) individual species capture probabilities stay constant during sampling

(Burnham and Overton 1979). However, this method is robust to deviations from these

assumptions. Even when the assumptions are violated the model-based richness

estimates are less biased than counts of species (Nichols et al. 1998).

Data from seventeen of the eighteen transects were used in the richness analysis.

The one transect that was located in the previously scraped habitat was removed from the

analysis in order to focus solely on variations within the P. cordata habitat. Since species

capture data were fairly sparse for each transect per sample occasion, the transects were

randomly assigned to six groups that represent sample replicates across space. They were

randomly grouped in order to remove effects of shoreline characteristics at different

transects. Richness was not estimated for sample occasions with reduced trap effort

(sample occasions 14-20). Linear regressions were performed using SPSS (SPSS Inc.

2001) in order to determine significant predictors of the vertebrate, fish and herpetofaunal

species richness. Richness estimates from each sample occasion were used in these

analyses. There were three outliers greater than two standard deviations from the mean,

which were removed for the herpetofaunal regression analyses. Average air temperature









(C), lake stage (m), rainfall (cm), and water level fluctuation (cm) were used as the

independent variables.

Assemblage Composition

Species richness estimates the number of species present, but indicates nothing

about community composition. To compare the presence of vertebrate species over time,

sample occasions were assigned to clusters using hierarchical cluster analysis. This was

run using PC-Ord software (McCune and Mefford 1999) with detection-nondetection

data of species for each sample occasion. Sorensen's distance measure with the flexible

beta (beta=-0.25) linkage method was used. Indicator species analysis (McCune and

Grace 2002) was applied to determine the most appropriate number of clusters and the

best species to represent those groups. Any groups comprised of a single sample

occasion were removed from the indicator species analysis. A Monte Carlo procedure

was run 1000 times with randomized data to calculate a p-value for each species, which

tested the null hypothesis that their indicator values were no larger than would be

expected by chance. The optimum number of groups was selected by the indicator

species analysis that yielded the most species with statistically significant indicator

values (McCune and Grace 2002).

Influence of Temporal Gradients on Assemblage

Multivariate ordination was used to establish what temporally changing

environmental factors were influencing the variation in assemblage composition.

Nonmetric multidimensional scaling (NMS) is an ordination technique that uses ranked

distances between sample units to reduce dimensions and allow description of the

community in relation to environmental gradients. The distances represent dissimilarity

between sample units in terms of species composition. This method was chosen because









it is particularly useful for non-normal data and many sampling events with no captures

(McCune and Grace 2002). Sample units, i.e. individual sample occasions, are plotted in

species space using an iterative search for the optimal placement for the sample units.

Optimal placement is determined by the maximum possible reduction in stress, which is a

measure of dissimilarity between the original data matrix and the reduced-dimension

final ordination. PC-ORD software was used for all NMS analyses (McCune and

Mefford 1999).

NMS was run for the entire vertebrate assemblage for all sample occasions using

detection-nondetection and count data separately. Fish and herpetofaunal assemblages

were then analyzed separately in the same fashion to determine if the environmental

gradients affected them differently. Outliers were identified using the outlier analysis

provided in PC-Ord, with the criteria being greater than two standard deviations from the

mean (McCune and Mefford 1999). All outliers were removed from the analyses.

General relativizations by row were conducted on the raw count data to equalize common

and uncommon species and lower the coefficient of variation (CV) of the row totals.

Relativizations were followed by square root transformations to balance the relative

importance of the species without altering their ranks.

Sorensen's distance measure was used to calculate dissimilarity matrices for the

ordinations. Starting configurations were created by random number seeds, which were

generated by the time of day. Fifty runs were conducted with the real data to find the

solution with the lowest stress. Fifty Monte Carlo randomized runs were performed to

select the appropriate number of dimensions that best represent the variation in the data.

Comparisons between the runs with real data and randomized data give a probability that









final stress in the ordination could be found by chance. After the first 50 runs, the number

of dimensions was determined and the final NMS was rerun using the random number

seed from the initial ordination. From this the final stress and instability (fluctuation of

stress per iteration) were evaluated.

Measured environmental variables, including lake stage, stage fluctuation, total

rainfall, and average, maximum and minimum average air temperatures over the sample

period, were included in the ordination graphs. The ordinations were plotted with

environmental variables as biplots, indicating the strength of correlations of variables

with the synthetic axes. Only the environmental variables with r2>0.2 (percent of

variance represented) are shown in the ordination plots. Sample units were color coded

by their membership to the groups defined by the cluster analysis, representing different

species composition.

Proportion of Habitat Utilized by Focal Species

Using the program PRESENCE (MacKenzie et al. 2002), detection/nondetection

data were analyzed to determine site occupancy rates for all species with enough captures

to get reasonable estimates. This method allows for numerous, representative, randomly

selected sites transectss) within the much larger area of interest (P. cordata zone) to be

sampled for the presence of species. The inference gained from sampling these sites can

then be applied to the pickerelweed zone Lake Tohopekaliga. The main function of this

method is to determine habitat usage for species with low detection probabilities (<1).

Detectability is an important factor when sampling secretive aquatic organisms in thick

vegetation. The program calculates (1) a "naive estimate," which is simply the

proportion of sites where the species was caught (considered biased low), (2) species-

specific detection probabilities based on the capture data, and (3) the "proportion of sites









occupied" (PSO) which is the naive estimate corrected for detection probability

(MacKenzie et al. 2002).

This method assumes closure of species to changes in occupancy status over the

course of sampling. However, if the species have large activity ranges and the

movements are assumed to be random, the closure assumption may be relaxed

(MacKenzie et al. 2002). The sample occasions were divided into two groups. The first

is from the start of sampling at the beginning of February until the traps were removed at

the beginning of May. The second group is from the beginning of August, when water

levels allowed full sampling effort, to the end of sampling in November. Between these

groups the lake stage became so low that there was no water in the P. cordata zone,

which surely caused a violation of the closure assumption for this method. This required

the split of sample occasions into groups that are assumed closed to species immigration

or emigration. The first (spring) group includes 13 sample occasions for the herpetofauna

and 11 sample occasions for the fish species, since fish were not recorded for the first

sample and the last sample in the group had water depths too shallow to capture fish. The

second (fall) group has 16 sample occasions for all species.

Parameters were estimated for each species for both groups using the single season

models in PRESENCE. The data were analyzed using models with both constant and

survey specific detection probabilities. Results from the model with the lowest Akaike's

Information Criterion (AIC) value were reported for each species. If the AIC values were

within two points of each other, the simpler, constant detection probability model was

selected.









Results

Trap Comparisons

All reptile, amphibian and fish species captured during the 2002 whole-lake sampling,

along with species codes used in the figures are listed in Table 3-1. Due to restrictive

funnel sizes, most fish species (especially centrarchids) were represented by juvenile life

stages, except small species such as mollies and killifish. On the other hand, adult

individuals characterized the majority of the reptile and amphibian species, since most

young individuals could escape through the mesh. Table 3-2 shows the species and

number of captures for the two types of minnow traps. Eleven vertebrate species were

captured with the 0.6 cm (0.25 in) mesh minnow traps, while 19 species were captured in

the 1.3 cm (0.5 in) mesh minnow traps at the same sample locations. Three species were

unique to the 0.6 cm mesh traps on these occasions: black swamp snake (Seminatrix

pygaea), flagfish (Jordanellafloridae), and mosquitofish (Gambusia spp). Only three

reptile or amphibian species were captured: black swamp snake, Florida leopard frog

(Rana sphenocephala), and pig frog (Rana grylio). More tadpoles were captured with the

0.6 cm mesh (n=31) than with the 1.3 cm mesh (n=2). Nine species were unique to the

1.3 cm mesh traps: striped mud turtle (Kinosternon baurii), striped crayfish snake

(Regina alleni), Florida water snake (Nerodiafasciatapictiventris), Florida green water

snake (Nerodiafloridana), siren (Siren spp), redfin pickerel (Esox americanus), armored

catfish (H. littorale), dollar sunfish (Lepomis marginatus), spotted sunfish (Lepomis

punctatus), largemouth bass (Micropterus salmoides), and redear sunfish (Lepomis

microlophus). Seven combined reptile and amphibian species were caught with these

traps. Species common to both traps were leopard frog, pig frog, bluegill (Lepomis

macrochirus), bluespotted sunfish (Enneacanthus gloriouss, golden topminnow










Table 3-1. All species captured in 2002, with species codes used in subsequent figures.
Fish Species Scientific Name Family Species Code
Armored catfish Hoplosternum littorale Callichthyidae HOPLI
Black crappie Pomoxis nigromaculatus Centrarchidae POMNI
Bluegill Lepomis macrochirus Centrarchidae LEPMAC
Bluespotted sunfish Enneacanthus glorious Centrarchidae ENNGL
Bowfin Amia calva Amiidae AMICA
Chain pickerel Esox niger Esocidae ESONI
Chubsucker Erimyzon spp. Catostomidae ERIMY
Dollar sunfish Lepomis marginatus Centrarchidae LEPMAR
Flagfish Jordanella floridae Cyprinodontidae JORFL
Gar Lepisosteus spp. Lepisosteidae LEPIS
Golden shiner Notemigonus crysoleucas Cyprinidae NOTCR
Golden topminnow Fundulus chrysotus Fundulidae FUNCH
Largemouth bass Micropterus salmoides Centrarchidae MICSA
Pterygoplichthys Pterygoplichthys spp. Loricariidae PTERY
Redear sunfish Lepomis microlophus Centrarchidae LEPMI
Redfin pickerel Esox americanus Esocidae ESOAM
Sailfin molly Poecilia latipinna Poeciliidae POELA
Seminole killifish Fundulus seminolis Fundulidae FUNSE
Spotted sunfish Lepomis punctatus Centrarchidae LEPPU
Warmouth Lepomis gulosus Centrarchidae LEPGU
Herpetofaunal Species Scientific Name Family Species Code
Amphiuma Amphiuma means Amphiumidae AMPME
Cottonmouth Agkistrodon piscivorous conanti Viperidae AGKPICO
Fl. banded water snake Nerodia fasciata pictiventris Colubridae NERFAPI
Fl. green water snake Nerodia floridana Colubridae NERFL
Fl. snapping turtle Chelydra serpentina osceola Chelydridae CHESEOS
Fl. softshell turtle Apalone ferox Trionychidae APAFE
Leopard frog Rana sphenocephala Ranidae RANSP
Mud snake Farancia abacura abacura Colubridae FARABAB
Peninsula cooter Pseudemys floridana peninsularis Emydidae PSEFLPE
Pig frog Rana grylio Ranidae RANGR
Siren Siren spp. Sirenidae SIREN
Stinkpot Sternotherus odoratus Kinosternidae STEOD
Striped crayfish snake Regina alleni Colubridae REGAL
Striped mud turtle Kinosternon baurii Kinosternidae KINBA
Tadpole-leopard frog Rana sphenocephala Ranidae TADRANSP
Tadpole-pig frog Rana grylio Ranidae TADRANGR










Table 3-2. Species capture frequencies for the 0.6 and 1.3 cm mesh minnow traps.
Vertebrate Species 0.6 cm mesh 1.3 cm mesh
Armored catfish 0 2
Black swamp snake 1 0
Bluegill 2 1
Bluespotted sunfish 18 31
Dollar sunfish 0 2
Flagfish 147 0
Florida green water snake 0 4
Florida water snake 0 1
Gambusia 75 0
Golden topminnow 17 1
Largemouth bass 0 1
Florida leopard frog 4 2
Pig frog 2 7
Redearsunfish 0 1
Redfin pickerel 0 1
Sailfin molly 56 5
Siren 0 1
Spotted sunfish 0 2
Striped crayfish snake 0 1
Striped mud turtle 0 1
Tadpoles 31 2
Warmouth 2 6


(Fundulus chrysotus), sailfin molly (Poecilia latipinna), and warmouth (Lepomis

gulosus).

Species Richness

Vertebrate species richness (Figure 3-3) was negatively correlated with average air

temperature (r2=0.330, df=l, p=0.001) and rainfall (r2=0.173, df=l, p=0.028). Average

air temperature was the only significant predictor of richness for the fish (Figure 3-4),

(r2=0.316, df=l, p=0.002), with higher species richness estimates occurring with lower

air temperatures. The estimated richness of the herpetofaunal assemblage (Figure 3-5)

was negatively correlated with lake stage (r2=0.327, df=l, p=0.003), rainfall (r2=0.166,

df=l, p=0.043), and water level fluctuation (r2=0.211, df=l, p=0.021).









Assemblage Composition

Six clusters were chosen to represent the 34 sample occasions. Thirteen indicator

species were determined with p<0.05 (Table 3-3). Indicator values are given for these

species, with 100 representing perfect indication of that group based on relative

abundances and frequency of occurrence. A zero indicates complete absence of a species

from a particular group. Since these cryptic species are quite mobile (with respect to the

traps) and dependent upon detection for quantification, indicator values are relatively low

compared to vegetation studies where virtually all species are detectable. Group 1 is

identified by several species, including Amia calva (bowfin), Fundulus chrysotus (golden

topminnow), Lepomis macrochirus (bluegill), Lepomis marginatus (dollar sunfish),

Nerodiafasciata pictiventris (Florida water snake), Rana sphenocephala (Florida leopard

frog), and R. sphenocephala and Rana grylio (pig frog) tadpoles. The second group

consisted only of sample occasion #13 (an extremely low water sample), and was

therefore omitted. Hoplosternum littorale (armored catfish) and Regina alleni (striped

crayfish snake) are the indicator species for Group 3. There were no significant indicator

species for Group 4, but Amphiuma means amphiumaa) and Lepisosteus spp. (gar) show

the highest indicator values with 28 and 22 respectively. Si. intihel i/ odoratus

(common musk turtle) was the sole indicator species for Group 5. The two pickerel

species, Esox americanus (redfin) and Esox niger (chain), were the only two species

indicating Group 6.

Some species were captured during every sample occasion, including

Enneacanthus glorious (bluespotted sunfish), Lepomis gulosus (warmouth), and Siren

spp. (siren). Hoplosternum littorale and Kinosternon baurii (striped mud turtle) were

found on almost every sample occasion. It is unclear why the armored catfish is an










indicator species for Group 3, when it has an indicator value of 22 for all groups but one.

Also, the bluegill had indicator values of 47 for both Group 1 and Group 6, although it

was assigned to Group 1 with p=0.028.

Table 3-3. Indicator species analysis results. Significant indicator species are
highlighted (p<0.05) and displayed with associated indicator values and the
clusters to which the species was assigned.
Species Code Cluster Max Indicator Value Mean Standard Deviation Probability
AGKPICO 1 25 15 7.4 0.211
AMICA 1 48.5 18 9.86 0.017
AMPME 1 27.9 24.8 4.25 0.234
APAFE 5 9.1 15.3 7.72 1
CHESEOS 4 12.5 15.3 7.49 0.696
ENNGL 1 20 20 0.63 1
ERIMY 6 27.9 23.1 7.1 0.201
ESOAM 6 40.9 22.6 7.6 0.022
ESONI 6 43.3 18.1 10.46 0.022
FARABAB 6 21.3 18.2 10.31 0.29
FUNCH 1 43.8 21.3 8.26 0.04
HOPLI 3 22.2 21.3 0.82 0.039
JORFL 1 34.1 16.8 9.59 0.077
KINBA 1 20.5 20.6 0.73 0.672
LEPGU 1 20 20 0.63 1
LEPIS 4 22.3 24.9 3.41 0.76
LEPMAC 1 47.1 21.9 8.23 0.028
LEPMAR 1 36.8 23.8 7.01 0.05
LEPMI 6 33.8 22.3 8.45 0.1
LEPPU 5 12 22.2 8.8 0.976
MICSA 6 28.8 23 7.01 0.178
NERFAPI 1 40.8 22.7 7.03 0.018
NERFL 3 21.6 21.3 0.83 0.295
NONE 1 31.7 19.9 9.88 0.107
NOTCR 6 28.6 16.5 8.92 0.059
POELA 1 28.8 24.6 4.2 0.115
POMNI 1 25 14.9 7.44 0.196
PSEFLPE 5 9.1 15.1 7.52 1
PTERY 5 18.7 16.9 9.76 0.179
RANGR 1 22.4 23.4 1.3 0.92
RANSP 1 100 17.4 9.61 0.001
REGAL 3 72.6 17.8 10.18 0.002
SIREN 1 20 20 0.63 1
STEOD 5 29.8 24.4 2.19 0.001
TADRANGR 1 34.6 24.4 5.45 0.009
TADRANSP 1 75 17 9.16 0.001









Influence of Temporal Gradients on Assemblage

Stable three-dimensional ordinations were produced with all NMS analyses except

the herpetofaunal analysis with detection/nondetection data. The final solutions were

based on the criteria of stress being reduced by at least 5% with each additional

dimension. The final stress values were lower with the real data than was found by the

Monte Carlo randomized runs (p<0.05), which indicates that there was real structure

found in the data. The final stress values for all ordinations (except for herpetofaunal

detection/nondetection) were between 12 and 18 (Table 3-4), which are common values

for ecological data and depict a fair portrayal of the data (McCune and Grace 2002).

Table 3-4. Stress and instability results from all NMS ordinations
Assemblage Data Type Final Stress Instability Iterations
Vertebrate Detection/nondetection 17.65 0.00254 500
Vertebrate Counts 13.27 0.00049 69
Fish Detection/nondetection 15.31 0.00095 29
Fish Counts 12.93 0.00016 49
Herpetofauna Counts 14.08 0.00045 38


For the NMS with vertebrate detection/nondetection data, Axes 1 and 3 best

explained 61% of the variance found in the assemblage composition. Environmental

variables with r2>0.2 were shown as biplots on the plots and include lake stage and air

temperature measures (Figure 3-6, Table 3-5). Axis 1 was correlated with lake stage

(r2=0.349). Axis 3 was most correlated with both lake stage and average air temperature,

(r2=0.354 and 0.259 respectively). Sixteen species were correlated with either Axis 1 or

3 with r2>0.2 (Figure 3-7, Table 3-6), 10 of them being indicator species.










Table 3-5. Percent of variance explained (r2) by environmental variables for each axis in
the vertebrate NMS with detection/nondetection data. Variables with r2 > 0.2
are highlighted.
Axis 1 2 3
Variable r2 r2 r2
Stage (m) 0.349 0.021 0.345
Fluc (cm) 0.039 0.009 0.183
Rain (cm) 0.037 0.006 0.029
Max (C) 0.038 0.362 0.127
Min (C) 0.002 0.255 0.229
Ave (C) 0.009 0.375 0.259

Table 3-6. Percent of variance explained (r2) for each axis by species in the vertebrate
NMS with detection/nondetection data. Indicator species with r2 > 0.2 are
highlighted in blue, while all other species with r2 > 0.2 are highlighted in
yellow.
Axis 1 2 3 Axis 1 2 3
Species r2 r2 r2 Species r2 r2
AGKPICO 0.073 0.002 0.094 LEPMI 0.035 0.56 0.018
AMICA 0.027 0.077 0.327 LEPPU 0.064 0.027 0
AMPME 0.002 0.171 0.01 MICSA 0.291 0.227 0.242
APAFE 0.035 0.004 0.001 NERFAPI 0.227 0.167 0.257
CHESEOS 0.021 0 0.063 NERFL 0.003 0 0
ENNGL 0.381 0.019 0.002 NONE 0.057 0.004 0.102
ERIMY 0.031 0.536 0.007 NOTCR 0.011 0.05 0.017
ESOAM 0.033 0.186 0.353 POELA 0.188 0.018 0.248
ESONI 0.126 0.166 0.089 POMNI 0.007 0.011 0.067
FARABAB 0.049 0 0.21 PSEFLPE 0.005 0.085 0.001
FUNCH 0.019 0.419 0.547 PTERY 0.002 0.081 0.038
HOPLI 0.376 0.003 0.155 RANGR 0.149 0.035 0
JORFL 0.087 0.047 0.15 RANSP 0.218 0.02 0.331
KINBA 0 0.07 0.016 REGAL 0.01 0.117 0.037
LEPGU 0.381 0.019 0.002 SIREN n/a n/a n/a
LEPIS 0.36 0.007 0.083 STEOD 0.021 0.006 0.018
LEPMAC 0.002 0.311 0.53 TADRANGR 0.016 0.127 0.311
LEPMAR 0.019 0.04 0.422 TADRANSP 0.07 0.039 0.287


With vertebrate assemblage count data, Axes 1 and 3 had the highest r2. Together

these axes represent 63% of the variance in the species composition. Axis 2 also had an

r2>0.2, and was best correlated with air temperature measures. As with the

detection/nondetection analysis, lake stage and air temperature measures showed the










highest correlation with these axes (Figure 3-8, Table 3-7). Air temperature is correlated

with Axis 1, with maximum air temperature having the highest r2 of 0.353. Axis 3 was

most highly correlated with lake stage (r2=0.458) and average air temperature (r2=0.375).

Seventeen species had an r2>0.2 for at least one of the axes, nine of them being indicator

species (Figure 3-9, Table 3-8).


Table 3-7.


Percent of variance explained (r2) by environmental variables for each axis in
e ht vertebrate NMS with count data Variab d


Axis 1 2 3
Variable r r
Stage (m) 0.146 0.010 0.458
Fluc (cm) 0.013 0.006 0.108
Rain (cm) 0.001 0.001 0.044
Max (C) 0.353 0.299 0.232
Min (C) 0.261 0.346 0.281
Ave (C) 0.298 0.379 0.375


--.--- ----- ..--- -. -.----


Table 3-8.


Percent of variance explained (r2) FOr each axis by species in the vertebrate
NMS with count data. Indicator species with r2 > 0.2 are highlighted in blue,
while all other species with / > 02 are highlighted in yellow


Axis 1 2 3 Axis 1 2 3
Species r2 r2 r2 Species r2 r2
AGKPICO 0.16 0.005 0.095 LEPMI 0.241 0.331 0.139
AMICA 0.059 0.089 0.192 LEPPU 0.06 0.02 0.018
AMPME 0.183 0.124 0.211 MICSA 0.13 0.184 0.048
APAFE 0.044 0.021 0.017 NERFAPI 0.166 0.001 0.373
CHESEOS 0 0.002 0.087 NERFL 0.068 0.024 0.004
ENNGL 0.809 0.342 0.108 NONE 0.031 0.011 0.164
ERIMY 0.217 0.024 0.092 NOTCR 0.124 0.136 0.022
ESOAM 0.014 0.455 0.09 POELA 0.133 0.023 0.341
ESONI 0.051 0.108 0.044 POMNI 0.022 0.005 0.087
FARABAB 0.041 0.028 0.018 PSEFLPE 0.065 0 0.038
FUNCH 0.002 0.291 0.575 PTERY 0.063 0.005 0.115
HOPLI 0.05 0.285 0.653 RANGR 0.125 0.157 0
JORFL 0.186 0.04 0.172 RANSP 0.092 0.038 0.345
KINBA 0.43 0.003 0.005 REGAL 0.039 0.039 0.005
LEPGU 0 0.104 0.013 SIREN 0.428 0.46 0.172
LEPIS 0.005 0.158 0.288 STEOD 0.268 0.182 0.288
LEPMAC 0.079 0.423 0.386 TADRANGR 0.147 0.159 0.008
LEPMAR 0.023 0.182 0.4 TADRANSP 0.137 0.065 0.24










The fish assemblage alone with detection/nondetection data resulted in the first two

axes representing 63% of the variance explained. Average temperature was most highly

correlated with Axis 1 (r2=0.291), and also with Axis 2 (r2=0.368), while lake stage was

correlated with Axis 2 (r2=0.275), (Figure 3-10, Table 3-9). For these two axes, there

were a total of 11 species with an r2>0.2, with six indicator species (Figure 3-11, Table 3-

10).

Table 3-9. Percent of variance explained (r2) by environmental variables for each axis in
the fish NMS with detection/nondetection data. Variables with r2 > 0.2 are
highlighted.
Axis 1 2 3
Variable r2 r
Stage (m) 0.042 0.275 0.105
Fluc (cm) 0.008 0.044 0.062
Rain (cm) 0.001 0.031 0.018
Max(C) 0.250 0.177 0.115
Min(C) 0.205 0.330 0.106
Ave (C) 0.291 0.368 0.083

Table 3-10. Percent of variance explained (r2) for each axis by species in the fish NMS
with detection/nondetection data. Indicator species with r2 > 0.2 are
highlighted in blue, while all other species with r2 > 0.2 are highlighted in
yellow.
Axis 1 2 3 Axis 1 2 3
Species r2 r2 r2 Species r2 r2
AMICA 0.152 0.296 0.12 LEPMAC 0.361 0.646 0
ENNGL n/a n/a n/a LEPMAR 0.406 0.229 0.168
ERIMY 0.149 0.089 0.223 LEPMI 0.118 0.205 0.436
ESOAM 0.287 0.223 0.083 LEPPU 0.002 0.009 0.021
ESONI 0.195 0.066 0.013 MICSA 0.706 0.049 0.001
FUNCH 0.486 0.572 0.003 NOTCR 0.072 0.026 0.018
HOPLI 0.058 0.251 0.149 POELA 0.007 0.263 0.264
JORFL 0.038 0.181 0.137 POMNI 0.034 0.009 0.004
LEPGU n/a n/a n/a PTERY 0.253 0.003 0.046
LEPIS 0.021 0.339 0.004


For the fish ordination with count data, Axes 2 and 3 represent the most variation in

the assemblage data with a cumulative r2 of 0.76. Again, air temperature and stage are










the environmental variables most correlated with these axes (Figure 3-12, Table 3-11).

Axis 2 was most highly correlated with maximum air temperature (r2=0.380), while lake

stage (r2=0.287) and average air temperature (r2=0.333) are most correlated with Axis 3.

Fourteen of the fish species had an r2>0.2 for these two axes, with six of them being

indicator species (Figure 3-13, Table 3-12).

Table 3-11. Percent of variance explained (r2) by environmental variables for each axis
in the fish NMS with count data. Variables with r2 > 0.2 are highlighted.
Axis 1 2 3
Variable
Stage (m) 0.070 0.005 0.287
Fluc (cm) 0.008 0.001 0.064
Rain (cm) 0.003 0.000 0.024
Max (C) 0.242 0.380 0.189
Min (C) 0.346 0.303 0.243
Ave (C) 0.319 0.354 0.333


Table 3-


12. Percent of variance explained (r2) for each axis by species in the fish NMS
with count data. Indicator species with r2 > 0.2 are highlighted in blue, while
,211 .I, 24t \ t. t, 1 t 11 T


a ot er 1 ispec1es witl / L u.z. arie 1111111t 111d n yIe w.
Axis 1 2 3 Axis 1 2 3
Species Species
AMICA 0 0.008 0.445 LEPIS 0.359 0.044 0.07
ENNGL 0.171 0.57 0.239 LEPMAC 0.085 0.308 0.503
ERIMY 0.221 0.367 0.135 LEPMAR 0.033 0.185 0.624
ESOAM 0.159 0.165 0.372 LEPMI 0.227 0.359 0.164
ESONI 0.002 0.252 0.156 LEPPU 0.074 0.274 0.006
FUNCH 0.113 0.106 0.757 MICSA 0 0.64 0.294
HOPLI 0.108 0.03 0.194 NOTCR 0 0.152 0.058
JORFL 0.004 0.046 0.303 POELA 0.17 0 0.423
LEPGU 0.191 0.337 0.223 POMNI 0.054 0.077 0.016


The herpetofaunal assemblage detection/nondetection data were analyzed with

NMS, but results yielded only a one-dimensional solution. Stress on the final run was

51.36, which represents an unacceptable amount of variation from the original data set.

Values of stress greater than 20 indicate that the solution may be misleading, while

ordinations with stress values over 40 represent very little of the structure in the original










data matrix (McCune and Grace 2002). Due to these outcomes, NMS was considered

unsuccessful for the reptile and amphibian assemblage with detection/nondetection alone.

Count data for the reptiles and amphibians did yield a successful ordination. The

first and third axes explain a total of 69% of the variation in the assemblage composition.

Lake stage is highly correlated with Axis 1 (r2=0.267) and Axis 3 (r2=0.389), and water

level fluctuation is correlated with Axis 3 (r2=0.204), (Figure 3-14, Table 3-13). Eight

species were correlated with Axes 1 and 3 with r2>0.2, six of them being indicator

species (Figure 3-15, Table 3-14).

Table 3-13. Percent of variance explained (r2) by environmental variables for each axis
in the herpetofaunal NMS with count data. Variables with r2 > 0.2 are
highlighted.
Axis 1 2 3
Variable r2 r2 r2
Stage (m) 0.267 0.081 0.389
Fluc (cm) 0.006 0.002 0.204
Rain (cm) 0.000 0.001 0.057
Max (C) 0.052 0.020 0.031
Min (C) 0.022 0.034 0.091
Ave(C) 0.058 0.016 0.073

Table 3-14. Percent of variance explained (r2) for each axis by species in the
herpetofaunal NMS with count data. Indicator species with r2 > 0.2 are
highlighted in blue, while all other species with r2 > 0.2 are highlighted in
yellow.
Axis 1 2 3 Axis 1 2 3
Species r2 r2 r2 Species r2 r2
AGKPICO 0.041 0.188 0.012 PSEFLPE 0.042 0.079 0.04
AMPME 0.014 0.291 0.094 RANGR 0.025 0.229 0.229
APAFE 0.087 0 0.017 RANSP 0.185 0.061 0.263
CHESEOS 0.154 0.018 0.036 REGAL 0.384 0.055 0.109
FARABAB 0.004 0.024 0.409 SIREN 0.018 0.031 0.047
KINBA 0.083 0.215 0.351 STEOD 0.437 0.091 0.033
NERFAPI 0.275 0.023 0.498 TADRANGR 0.002 0.095 0.311
NERFL 0.097 0.258 0.03 TADRANSP 0.107 0.275 0.057









To determine the separation of the sample occasion clusters along lake stage and

average air temperature gradients means and ranges of each are shown in Figures 3-16

and 3-17 respectively. Clusters one and two have lake stage ranges lower and non-

overlapping with clusters four, five and six. Stage values for cluster 3 span much of the

ranges for every other group. Air temperature values are very similar for clusters one and

six, both being below and non-overlapping with clusters two, three and four. Cluster five

overlaps with most of the ranges of all other groups.

Proportion of Habitat Utilized by Focal Species

Figures 3-18 and 3-19 show the occupancy rate estimates for the fish species.

Several species exhibited a decrease in site occupancy from the spring to the fall season,

most pronounced in M. salmoides, L. macrochirus and L. microlophus. Lepomis

microlophus, L. gulosus and E. glorious occurred at all transects during the spring.

Lepisosteidae spp., H. littorale and L. gulosus were present in all transects in the fall.

Lepomispunctatus had invalid occupancy estimates during the spring due to low

detection probabilities, with the same problem for L. macrochirus and L. microlophus in

the fall. In the spring, the only fish species that used survey-specific detection probability

models for occupancy estimation were E. glorious and L. macrochirus, which indicates

that detection of these two species varied between sample occasions. All species were

modeled with time-constant detection probabilities in the fall, except E. glorious, P.

latipinna and Lepisosteidae spp.

Estimates of the proportion of sites occupied for the eight focal herpetofaunal

species are presented in Figures 3-20 and 3-24. Site occupancy was similar in the spring

and fall for most species. There were not enough data to make valid estimates for A.

means in the spring and for R. sphenocephala and N. fasciatapictiventris in the fall. All









species were modeled with the time constant detection probability except N. floridana

and R. sphenocephala in the spring. With the exception ofR. sphenocephala and A.

means, most species were estimated to be in 90-100% of all transects at some point in the

year.

Discussion

Trap Comparisons

The 1.3 cm (0.5 in) mesh traps were better at capturing the species of interest,

including reptiles, amphibians, centrarchids and exotic catfish. Although it had been

assumed that many species would be under-represented with the larger mesh, the few

unique species caught in the 0.6 cm (0.25 in) mesh traps were not of particular interest to

this study. The fish species included mosquitofish and flagfish, which are not suspected

of being impacted by the removal of the vegetation from the littoral zone. The one black

swamp snake was actually stuck about halfway through the mesh, indicating that even 0.6

cm mesh may not be small enough to capture this species representatively. The size and

shape of the traps and openings may also have affected capture probabilities of different

species. Also, the bare metal hardware cloth may be more visible to the animals than the

dark green vinyl-coated hardware cloth of the larger mesh traps. In comparing

practicality of sampling with each type of trap, the 0.6 cm commercially manufactured

traps were much less durable and prone to breaking during use, as well as being more

expensive ($16.16 each for 0.6 cm vs. $11.00 each for 1.3 cm). Given these factors, the

1.3 cm mesh traps were more useful than the 0.6 cm for sampling in this habitat.

Species Richness

Ectothermic fish communities respond to water temperature due to its effect on

metabolism and growth rates, especially for juveniles (Holt 2002). This relationship









determines the length of time that species will benefit from residing in a given habitat.

Water temperature is closely associated with air temperature due to the shallow aquatic

habitat. Therefore, it is not surprising that the average air temperature is negatively

correlated with fish species richness. In the summer season, water temperature may

exceed tolerance limits for sensitive fish such as select species of sunfish, causing them

to leave the habitat and resulting in fewer fish species. Temperature also indirectly

determines the amount of dissolved oxygen in the water; however this variable was not

measured in the field.

Lake stage, rainfall, and water level fluctuations were significant predictors of

herpetofaunal richness, all of which have a relationship to water depth at the fixed trap

locations. The gradually sloping contour of Lake Tohopekaliga causes small increases in

lake stage to flood broad expanses of previously dry habitat, allowing species to enter

into new habitat for spawning or foraging. Alternatively, moderate drops in lake stage

may cause a rapid decrease in the area of the littoral zone inundated with water, causing

species to emigrate or burrow, or else risk being trapped by unsuitable conditions.

Receding water levels may bring in more terrestrial species into the previously aquatic

habitat, such as cottonmouths, Florida water snakes, and Florida leopard frogs.

Alternatively, deeper water in the habitat may restrict species that prefer shallow water

and promote more aquatic species such as the common musk turtle.

Assemblage Composition

Since animal species move throughout the habitat and are detected with

probabilities less than one, and animal assemblages tend to be transient in nature, the

groups determined by cluster analysis are not very discrete with respect to species









composition. This resulted in fairly low indicator values, but with several being

significant nonetheless.

The four sample occasions in Group 1 had the lowest lake stage values and were

tied with Group 6 for the lowest average air temperatures. The Florida water snake and

Florida leopard frog are the herpetofaunal indicator species for this group, being the more

terrestrial of the focal reptile and amphibian species. Bowfins, golden topminnows,

bluegill and dollar sunfish are the fish indicator species. Bowfins are one of the most

tolerant freshwater fish species and are often called mudfishh" (Boschhung et al. 1995),

so it is not unreasonable that this species would occupy this shallow vegetated habitat.

Golden topminnows prefer to occupy lakes with abundant vegetation (Hoyer and

Canfield 1994), and therefore may be more tolerant of dense vegetation communities

than other species. It is not as easy to explain why the two sunfish species were

indicators for these shallow water depths. They may have been stranded by receding

waters and captured more easily in traps with puddles of water remaining.

Group 2 only had one sample occasion attributed to it. This sample was the last

one in late April before the traps were removed due to lack of water in the habitat.

Although not included in the indicator species analysis, this sample included no fish

species.

Only three sample occasions were clustered into Group 3. During these hot

summer samples, the lake stage was rapidly rising, and trap effort was reduced pending

appropriate water depths at the trap sites. The striped crayfish snake was the main

indicator species for this group. Being a specialist predator on crayfish (Godley 1980),









prey availability in recently flooded habitats may have been the driving factor for their

indicator values on these occasions.

Eight samples fell into Group 4, which had no indicator species attributed to it.

These sample occasions occurred in early fall, with the highest lake stages and average

air temperatures.

Group 5 occurred mainly in the summer months (eight samples), but also contained

two samples in February. Water depths were moderately high. Air temperatures were

low in February and high in summer, spanning a wide range of temperatures. The sole

indicator species for this group was the common musk turtle. This species is known for

being highly aquatic, leaving the water only to nest (Wygoda 1979, Gibbons et al. 1983).

It also is active for the widest temperature ranges of any other kinosternid species in

North America, being able to retreat to deeper waters to buffer the effects of air

temperature extremes (Mahmoud 1969, Ernst 1986).

The last cluster, Group 6, included seven samples, with six occurring February

through April and the other one in November. Low air temperatures and moderately high

but dropping lake stages characterize these sample occasions. The two Esox spp. (pike)

are the indicators for this group. These species breed from February to March in the

south, spawning in densely vegetated habitats less than 50 cm deep (Billard 1996). This

may explain their presence in the habitat during these environmental conditions.

Influence of Temporal Gradients on Assemblage

Detection/nondetection data were used for ordinations, and are generally

recommended when comparing habitat distributions of species (Hayek 1994), and when

sample unit heterogeneity is large (McCune and Grace 2002). Counts were also used to

compare results obtained by the two types of data, but although agreement between the









two provides more support, lack of detection probabilities make count data less valuable.

For the vertebrate and fish ordinations, results were similar for both types of data. The

herpetofaunal ordination was unsuccessful with detection/ nondetection data, and

therefore counts were used solely.

Average air temperature and lake stage came out as the most important variables

correlated with the axes representing variation in species composition in the vertebrate

and fish assemblages. As mentioned before, temperature influences growth rates for

young fish, as well as the amount of dissolved oxygen in the water. Both of these factors

limits the time that fish are able to occupy a habitat. Physical access to heavily vegetated

habitat is also limited by water depth, which is controlled by lake stage. This determines

the volume of water the animals have to move through, as well as the effect of vegetation

density in the water column. However, for herpetofaunal species alone air temperature is

not associated with variation in the species composition. In this case, lake stage is most

important, with water level fluctuation also showing a correlation with one of the axes.

Lake stage probably dictates movements of species that do not show site fidelity, in

response to habitat requirements and prey availability. For species that are not known to

move long distances, for example sirens and amphiumas, low water levels trigger

burrowing activities (Aresco 2001), thereby reducing capture opportunities.

Proportion of Habitat Utilized by Focal Species

Due to the large-scale removal of pickerelweed from the littoral zone of Lake

Tohopekaliga during enhancement activities, it was important to investigate the spatial

distribution of species in this habitat. Site occupancy analyses were used to estimate the

proportion of this habitat type that was used throughout the year by various fish, reptile

and amphibian species. While some species are temporally and spatially pervasive in the









habitat (e.g., warmouths, bluespotted sunfish, Florida green water snakes, sirens, striped

mud turtles, and pig frogs), others seem to use the Pontederia cordata zone

intermittently. Of the fish species, sailfin mollies, spotted sunfish, chubsuckers, and

largemouth bass were found in a moderate proportion of transects (30-70%) in both the

spring and fall. Bluegill and redear sunfish were both in a high proportion of sites

(>80%) in the spring, but were found in less than 20% of the transects in the fall. This

trend may be due to juvenile fish using the littoral zone for foraging and predator

avoidance during the spring when suitable physicochemical conditions permit survival

(Werner and Hall 1979, Crowder and Cooper 1982, Werner and Hall 1988, Chapman et

al. 1996). Unmeasured environmental characteristics such as low dissolved oxygen may

have kept the sunfish out of the thick vegetation after the summer low-water spell

(Miranda and Hodges 2000). Gars and armored catfish went from about 65-80% of the

sites in the spring to 100% occupancy in the fall. These two species are far more tolerant

of harsh environmental conditions than most sunfish due in part to their capacity for

aerial respiration (Boschung et al. 1995, Brauner et al. 1995). For the armored catfish,

the greater presence in the fall may be due to the breeding season and sufficiently high

water levels for nesting (Mol 1993).

Most reptile and amphibian species occupied a similar proportion of sites in both

the spring and fall. Florida leopard frogs and Florida water snakes were only captured

when water levels were very low, which restricted reasonable estimates of site occupancy

to the spring season. Since the heavily vegetated littoral zone is known as prime habitat

for several of these species due to life history requirements, it is not surprising to find

most of the focal species in such a high proportion of the sites (>70% occupancy).
































Figure 3-1. Crayfish and minnow trap in P. cordata habitat.
















































Figure 3-2. Locations of 2002 P. cordata sampling transects in Lake Tohopekaliga

























2102 3102 4102
2/1/02 3/1/02 4/1/02


5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02


Date


Figure 3-3. 2002 Vertebrate species richness estimates by sample date, with points
representing richness for the time between the last sample occasion and the
sample date.


2/1/02 3/1/02 4/1/02


5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02 1/1/03


Date
Figure 3-4. 2002 Fish species richness estimates by sample date, with points representing
richness for the time between the last sample occasion and the sample date.


III
0 ** .. .


*. ; .. .**'"+" -7 "."
: : : 0 ~+.,""


0o 1
1/1/02


1/1/03


0 +
1/1/02







50



40





S30




20





10
S. .............................
.






0
0 i i i i i i ii

1/1/02 2/1/02 3/1/02 4/1/02 5/1/02 6/1/02 7/1/02 8/1/02 9/1/02 10/1/02 11/1/02 12/1/02 1/1/03

Date


Figure 3-5. 2002 Herpetofaunal species richness estimates by sample date, with points
representing richness for the time between the last sample occasion and the
sample date.








51



18 Cluster

A1
@2
30 31 3
33
4
S5
15 34 28 6
V 2V Stage(m)
CO 2 Ave(C) 16
.) 2 V 26 2 25 3 1
X V
< 3
32
21
13 27
S V
24 0
v, 6 6




35 7
o0 36
10
12 11 A
A A

Axis 1


Figure 3-6. NMS ordination of sample units in vertebrate species space using
detection/nondetection data. Points represent sample occasions and distances
between points show the relative differences in species composition. The
length of each line is proportional to the strength of the correlation between
the environmental gradient and the synthetic axes.













CHESEOS
+


PTERY
+


POELA
+4-


NERFAPI NONE
++


JORFL


Stage(m)


APAFE
+


TADRANGR
+ NOTCR MICSA
+ +
LEPM ESOAM ESONI
W +


FARABAB
+ LEPMACFUNCH
++


AMICA


RANSP
+


AGKPICO
+


POMNI
+


TADRANSP


Axis 1



Figure 3-7. NMS ordination of vertebrate species in sample unit space using
detection/nondetection data. Points represent average species positions with
respect to sample units. The length of each line is proportional to the strength
of the correlation between the environmental gradient and the synthetic axes.








53




23
31 Cluster
30 A
3
4
17 Stage(m) 29 V1 5
Ave(C) 32 6
18 V 27
18 2
26 V
V 22
28 33 V
19 2 20
CO 16 V 24
34
S25 V




335
V 5



35
4 6 80 9

7 10
0A
15
36
11 12


Axis 1


Figure 3-8. NMS ordination of sample units in vertebrate species space using count data.
Points represent sample occasions and distances between points show the
relative differences in species composition. The length of each line is
proportional to the strength of the correlation between the environmental
gradient and the synthetic axes.














CHESEOS
+


Stage(m)


ENN
REGAL LI


PTERY
+


Ave(C)

LEPISHOPLI STEOD
\-r ++
S AMPME

KINBA
GL -lfI+ TADRANGR
EPPU +


+ -'1ICSA
ERIMY' ESOAM
LEPMI 'ESONI +
+ +


LtIVI/KI
+
LEPMAC
+


FUNCH


POMNI
+


FARABAB
+
POELA
+
NONE NERFAPI
+ +
AMICA


JORFL
+


RANSP TADRANSP
+


Axis 1



Figure 3-9. NMS ordination of vertebrate species in sample unit space using count data.
Points represent average species positions with respect to sample units. The
length of each line is proportional to the strength of the correlation between
the environmental gradient and the synthetic axes.


PSEFLPE
+


APAFE
+


NOTCR
+


AGKPICO
+













23
V19
17


3
v 16


22
2729Ave(C)33
_427Stage(m 34
34
20

2
31 28
V


15 32
V


7 64
0 00
A

8


30 I


Cluster
S1
3
4
S5
0 6


Axis 1


Figure 3-10. NMS ordination of sample units in fish species space using
detection/nondetection data. Points represent sample occasions and distances
between points show the relative differences in species composition. The
length of each line is proportional to the strength of the correlation between
the environmental gradient and the synthetic axes.


























i ERIMY
.A + POELA
x
< LEPMAR
ESOAM
POMNI ESONI + LEPMI
+ +
NOTCR
+





FUNCH
LEPMAC
+
AMICA JORFL
+ +

Axis 1




Figure 3-11. NMS ordination of fish species in sample unit space using
detection/nondetection data. Points represent average species positions with
respect to sample units. The length of each line is proportional to the strength
of the correlation between the environmental gradient and the synthetic axes.















9
o0
7
10
A
6
8 4 5
36
A


33 Ave(C)
26 34
V V


Cluster
S1
3
4
S5
0 6


Axis 2


Figure 3-12. NMS ordination of sample units in fish species space using count data.
Points represent sample occasions and distances between points show the
relative differences in species composition. The length of each line is
proportional to the strength of the correlation between the environmental
gradient and the synthetic axes.

























0)
*.U ESOAM POMNI
< + +

+ +

ERIMY
ENNGL-
+
LEPGU
LEPPU



Stage(m)

Ave(C) HOPLI EPIS
++

Axis 2



Figure 3-13. NMS ordination offish species in sample unit space using count data.
Points represent average species positions with respect to sample units. The
length of each line is proportional to the strength of the correlation between
the environmental gradient and the synthetic axes.




























28 34
'V


5
23
V Fluct(cm)
25
Stage(m) 3

32
33 30 V


Cluster
A 1
S2
3
4
S5
0 6


Axis 1


Figure 3-14. NMS ordination of sample units in herpetofaunal species space using count
data. Points represent sample occasions and distances between points show
the relative differences in species composition. The length of each line is
proportional to the strength of the correlation between the environmental
gradient and the synthetic axes.









60





"NP Cluster
A 1
*2
FARABAB 2
3
PSEFLPE 4
S5
06
NERFAPI
+ LADRANSP
APAFE +

+

+
A GPICo

TADRANGR



+
"RANGR


++
SIREN 4




Fluct(cm)
Stage(m)



REGAL




+
CHESEOS

Axis 1



Figure 3-15. NMS ordination of herpetofaunal species in sample unit space using count

data. Points represent average species positions with respect to sample units.

The length of each line is proportional to the strength of the correlation

between the environmental gradient and the synthetic axes.












































Figure 3-16.





30

28

o 26
0

24

22
E
I- 20

g) 18
03

> 16

14

12


1 2 3 4 5 6

Cluster


Average and range of lake stage values by cluster.











0


1 2 3 4 5 6


Cluster


Figure 3-17. Average and range of air temperature values by cluster.








62




1.1

1.0 0 0 0

0.9

0.8

Z 0.7

0.6
F--
o 0.5

o 0.4

0.3
*
0.2

0.1

0.0 -

-0 .1 ....
POELA LEPPU ERIMY LEPIS MICSA HOPLI LEPMAC LEPMI LEPGU ENNGL

FISH SPECIES

NAIVE ESTIMATE
O PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR



Figure 3-18. Site occupancy estimates for focal fish species in spring 2002.



1.1

1.0 -0 0 0

0.9

0.8

0.7 -

0.6

0 0.5 -

S0.4 -

S0.3

0.2 -

0.1 -

0.0 0 0

-0.1
POELA LEPPU ERIMY LEPIS MICSA HOPLI LEPMAC LEPMI LEPGU ENNGL

FISH SPECIES

NAIVE ESTIMATE
O PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR



Figure 3-19. Site occupancy estimates for focal fish species in fall 2002.

















1.0 0 0 0 0

0.9

0.8



0.6 { .






0.2
0.1
of-
O 0.5
0 0.4

0.3

0.2

0.1

0.0 -

-0.1 .
RANSP STEOD AMPME RANGR NERFAPI KINBA SIREN NERFL

HERPETOFAUNAL SPECIES

NAIVE ESTIMATE
O PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR



Figure 3-20. Site occupancy estimates for focal herpetofaunal species in spring 2002.


0 0
o o


RANSP STEOD AMPME RANGR NERFAPI KINBA SIREN NERFL

HERPETOFAUNAL SPECIES

* NAIVE ESTIMATE
O PROPORTION OF SITES OCCUPIED WITH STANDARD ERROR


Site occupancy estimates for focal herpetofaunal species in fall 2002.


Figure 3-21.














CHAPTER 4
ASSEMBLAGE ACROSS VEGETATION COMMUNITIES

Introduction

The objective of this section is to investigate the influence of vegetation type and

water depth on spatial variation in the aquatic vertebrate community. The three main

research questions are 1) are we able to estimate population parameters such as

abundance and density for the focal species using trapping grids or webs, 2) is there

spatial variation in the aquatic vertebrate assemblage, and 3) do the vegetation

communities or water depths influence the spatial variation for the individual focal

species?

Field Methods

Grid and Web Sampling

A pilot mark-recapture protocol was employed in the summer of 2002 in order to

estimate activity ranges, abundances and densities of species of interest. Trap points

were arranged in a square grid pattern (White et al. 1982). In order to have uniform

sampling effort within the grids, one minnow and one crayfish trap was placed at each

point. All newly captured reptiles and amphibians were weighed and measured, then

tagged with Passive Integrated Transponders (PIT tags), which are small microchips

inserted under the skin to permit individual identification when scanned. Each animal

was released at the capture location after being worked up. Traps were checked daily for

PIT tagged individuals, and all new individuals were measured and tagged during each

sampling event. In order to satisfy the assumption of closure for analysis and minimize









temporal variation in detection probabilities, the grids were sampled for 5-7 days. This

technique was used only for the most abundant species, since detection probabilities

would be too small to make accurate estimates of any others. These species included A.

means, Siren spp., N. floridana, and K. baurii.

The first grid consisted of 49 trap points, each three m (9.8 ft) apart, in a seven by

seven grid ("GRID1"), and sampled for seven consecutive days in July. Next, 100 trap

points were placed in a 10 by 10 grid ("GRID2A"), five meters (16.4 ft) apart, and

sampled for six consecutive days in August. The same grid was then sampled for five

non-consecutive days ("GRID2B"), using frozen sardines as bait, until it was decided that

the bait was logistically impractical. The last design was 100 trap points ("GRID3"),

spaced three meters (9.8 ft) apart, in a 10 by 10 grid. This was sampled for five times

over seven days, due to logistic problems. Attempts to sample several times within a day

were also made, but were discontinued immediately due to low numbers of captures.

In addition to these grids, a trapping web was attempted in September 2002. The

web is a variation of point-transect distance sampling, typically used for small terrestrial

animals. It is designed to have lines of traps radiating out from a center point, forming a

gradient of sampling effort and detection probabilities. Data from each concentric ring is

grouped according to distance intervals (Anderson et al. 1983). This web consisted of

eight radiating arms of 12 trap points each, placed at three-meter (9.8 ft) intervals, for a

total of 96 trap points. The benefit of this method is that it uses only the initial captures,

so the recapture rate is irrelevant. The assumptions of this method are that all animals

near the center of the web are captured, the size of the web is large relative to the

movements of the animals, distances are measured accurately from the center, and









individual captures are independent events. Sampling continues daily until no new

animals are caught at the center of the web, indicating 100% detection at the center

(Anderson et al. 1983) Since it was evident that the assumptions were not met in this

web, sampling was discontinued after six sample occasions.

Whole-Lake Sampling

After the postponement of the fall 2002 drawdown, we began sampling again with

modifications to the 2002 temporal sampling protocol. The goal was to investigate

differences in vertebrate habitat usage of vegetation communities beyond the P. cordata

zone, as well as varying water depths. Eighteen new locations were randomly chosen for

transects (Figure 4-1), since the sampling of 2002 had disturbed the habitat in some of the

previous locations. Sampling sites in Goblet's Cove were included and none were placed

in the disturbed stretch of shoreline in the southern part of the lake. Transects were

placed at least 200 m (656 ft) apart. At each transect, there were four trap points, each

with a minnow and crayfish trap. However, instead of placing the trap sites at fixed

locations in the habitat as in 2002, they were placed at fixed depths and moved with the

water level. When the water was rising or remaining stationary, each transect had four

trap points located at 15, 30, 45, and 60 cm (6, 12, 18, 24 in) deep (Figure 4-2). During

falling water levels, the trap points were placed at 30, 45, 60, and 75 cm (12, 18, 24, and

30 in) deep, except when falling lake stages were not predicted. The traps were still

checked once weekly, and at each sampling occasion, the traps were moved to the

appropriate depth. This resulted in trapping animals at specific depth ranges over the

course of the week, with similar water depths between sample occasions. For example if

a trap point was located at the 30 cm (12 in) depth and the water level rose several

centimeters during the week, the trap was considered to be sampling the 30-45 cm (12-18









in) water depth. For falling water levels, the 30 cm (12 in) traps were sampling the 15-30

cm (6-12 in) depth. Percent cover of vegetation species was also estimated for a 2 m (6.5

ft) radius around every trap site on each sampling occasion.

Continuous sampling was conducted from 1/30/2003 to 1/5/2004, during which

time traps were located up in the shallow grassy habitat at high water levels

(characterized mainly by Luziolafluitans and Panicum repens), through the thick

emergent habitat (with Pontederia cordata and Typha domingensis), down to more open

water zones (with Hydrilla verticillata and floating leaf species Nuphar luteum and

Nymphaea odorata) at lower water levels. Besides the added environmental variables,

this new protocol also allowed us to sample year-round, instead of having to remove the

traps at moderately low water levels. Sampling ended on 1/5/2004 at about 15.5 m (50.8

ft) NGVD, with lake stage dropping due to the 2003 drawdown.

Analysis Methods

Population Estimates and Movement for Herpetofaunal Species

Trapping grids are known to exhibit "edge effects" due to animals near the edges of

the grid moving in and out of the sampled area. To account for this phenomenon a

boundary strip is typically estimated and added to the grid area to estimate an effective

sampling area. Wilson and Anderson (1985) propose using the mean maximum distance

moved (MMDM) by animals recaptured at least once to estimate the activity range of the

species. Although lacking a solid theoretical explanation, this method works well in

simulations. The alternative method, nested grid design, requires a large data set for

estimation of density (Williams et al. 2002). Since our data were fairly sparse with

recaptures, we used the MMDM method to calculate the effective areas of the trapping

grids for each species.









Low numbers of recaptures were attained for each grid; so in order to calculate the

MMDM for each species movement distances were pooled from all grids and the

trapping web. To calculate the diagonal distances within the trapping grids, the

Pythagorean theorem was used: [a2+b2=c2], where sides a and b are sides of known

lengths. For diagonal distances in the trapping web, [a2=b2+c2-2bc(cosA)] was used,

where A is the degree measure between sides b and c of known length (Larson et al.

1994). For each species, MMDM and its variance were calculated using formulas from

Wilson and Anderson (1985). The widths of the boundary strips were estimated as half

the MMDM for each species. The effective grid areas and associated variances were then

calculated for each grid per species (Wilson and Anderson 1985).

Program MARK (White and Burnham 1999) was used to estimate population sizes

for each species per trapping grid. Estimates were obtained using models M(o), (constant

capture probabilities), and M(t), (time dependent capture probabilities). The Akaike's

Information Criteria (AIC) were compared to determine which model best fit the small

data set. Density was then calculated for each grid per species by dividing the population

size by the effective sampling area (Wilson and Anderson 1985).

Capture Success for Focal Species

Basic analyses were conducted on the 2003 data to look for trends in the data

associated with the main sampling variables involved in this protocol. All trap points

were divided into groups according to vegetation community (see results section below

for descriptions) and water depth at each trap location. Ten species were examined, two

each of salamanders (Siren spp., A. means), snakes (both Nerodia spp.), turtles (K. baurii

and S. odoratus), frogs (both Rana spp.) and fish (H. littorale and M. salmoides). The

reptile and amphibian pairs represent species with similar life history traits but which









have slightly different habitat requirements or preferences. They were also the most

frequently captured reptile and amphibian species in this study. The two fish species

were chosen to characterize opposite ends of the spectrum of habitat selection. While the

armored catfish is a generalist species with great tolerance for low dissolved oxygen, high

temperatures, thick vegetation and other extreme environmental variables (e.g., Nico and

Fuller 1999), largemouth bass and other sunfishes are thought to be highly intolerant of

these same habitat characteristics (e.g., Allen and Tugend 2002).

Capture success was calculated as the number of captures per species divided by

the total number of trap points for the particular variable of interest. This usually resulted

in a very low frequency, due to the large number of trap points and low detectability of

species. An arcsine squareroot transformation was applied to all success values, in order

to spread the ends of the scale, while improving normality for the proportion data

(McCune and Grace 2002). This allowed the relative values to show up more clearly

while reducing the effect of large sample units. The assumption of equal detectability of

the different species between habitats or water depths may be violated, but detection

probabilities cannot be calculated for this particular analysis. However, uniform

sampling methods were used over space transectss) and time (sampling occasions) to

reduce variability in detection.

For this sampling protocol, dependence of trap placement upon water depth (i.e.

lake stage) resulted in unequal sampling for vegetation communities. In addition, the

vast number of trap sites (n= 3,426) and sparse nature of the data made multivariate

analyses virtually impossible. For example, dividing data into groups (either subjectively

or with cluster analysis) depending on sample occasions, water depth or vegetation









communities would neglect important differences in the other variables and/or result in

groups of vastly unequal numbers of trap points. Attempted NMS analyses of all trap

sites (ungrouped) yielded no results due to the great number of zeroes in the matrices.

The data were not even appropriate for most univariate analyses. For example, chi-

square analyses of capture success would indicate whether there were significant

differences in the counts of focal species between vegetation types or water depths,

however the large sample sizes invariably lead to significant differences. Repeated-

measures analysis of variance was considered to test the differences between water

depths over time, however the data were too sparse to divide the counts between both

sample occasions and water depths. Species richness was also inestimable because of the

frequent nondetection of species and unequal sample sizes. As a result of the

complicated nature of the data, the analyses were largely descriptive in nature. These

descriptions of habitat usage rely mainly on comparisons of capture success across two

categorical environmental variables: water depth and vegetation community.

Results

Population Estimates for Herpetofaunal Species

Each species showed different movement distances over the sampling grids and

web. Figure 4-3 shows the mean maximum distances traveled and variances, along with

the associated widths and variances of the boundary strips for each species. Maximum

distances traveled for A. means, Siren spp., N. floridana, and K. baurii were 18 m (59 ft),

24 m (79 ft), 34 m (112 ft), and 59 m (194 ft) respectively. These movements are fairly

large relative to the sizes of the grids, with 72 m (236 ft) being the absolute maximum

distance between any two traps during all sampling. This indicates that the assumption of

closure was violated. Table 4-1 contains the percent increase in the size of each grid










when the boundary widths for each species were added to the sizes of the grids. While

the effective sampling areas are fairly acceptable for the amphiumas and sirens (Wilson

and Anderson 1985), Florida green water snakes and striped mud turtles add excessive

area to the original sampling areas. After recognizing the fact that closure was violated in

these grids, population sizes and densities were estimated with unreliable accuracy, but a

best attempt was made given the data.

Estimates of population size and variances are shown in Table 4-1. Several times

the capture history data were so sparse that estimates could not be calculated for some

grids and species. The estimates generated with sufficient data often have large variances

due to low recapture probabilities. The null model of no variation in detection

Table 4-1. Grid sizes and population estimates by mark recapture methods.
Parameter AMPME SIREN NERFL KINBA
GRID1 Actual size of grid (ha) 0.0324 0.0324 0.0324 0.0324
Est. effective sampling area and variance (ha) 0.078 (0.40) 0.08 (0.93) 0.12 (1.79) 0.16 (9.17)
Percent of original grid (%) 239 264 376 493
Est. population size and variance (# indivs.) n/a n/a 16(184) 7 (30)
Density estimate (#/ha) n/a n/a 131 44
GRID2A Actual size of grid (ha) 0.2025 0.2025 0.2025 0.2025
Est. effective sampling area and variance (ha) 0.30 (1.66) 0.32 (3.71) 0.39 (6.08) 0.46 (27.83)
Percent of original grid (%) 149 157 192 225
Est. population size and variance (# indivs.) n/a n/a n/a n/a
Density estimate (#/ha) n/a n/a n/a n/a
GRID2B Actual size of grid (ha) 0.2025 0.2025 0.2025 0.2025
Est. effective sampling area and variance (ha) 0.30 (1.66) 0.32 (3.71) 0.39 (6.08) 0.46 (27.83)
Percent of original grid (%) 149 157 192 225
Est. population size and variance (# indivs.) 14 (37) n/a 26 (99) 9 (5)
Density estimate (#/ha) 49 n/a 68 20
GRID3 Actual size of grid (ha) 0.0729 0.0729 0.0729 0.0729
Est. effective sampling area and variance (ha) 0.14 (0.72) 0.15 (1.65) 0.19 (2.94) 0.24 (14.27)
Percent of original grid (%) 187 201 267 332
Est. population size and variance (# indivs.) 19(117) n/a 25(24) 26(525)
Density estimate (#/ha) 142 n/a 133 108









probability was always selected over the time-varying capture probability model.

Density estimates based on these abundances are also included in Table 4-1.

Capture success for Focal Species

Figure 4-4 shows the types of vegetation communities that were sampled

throughout 2003. Each sampling occasion corresponds to one weekly sample, which

includes 72 trap points (4 trap points for each of 18 transects). Occasionally there were

less than 72 samples for a given sample occasion, usually because data recording for a

sample was inadvertently neglected, traps went unchecked due to dangerous weather, or

traps were missing. Each trap point was subjectively categorized in the field into

different vegetation communities, based on the dominant species present. The "Grass"

community is the closest shoreward, and is characterized by Panicum repens, Luziola

fluitans, Juncus effusus, and Eleocharis spp. Lakeward from this is the "Rooted-HE",

which refers to the herbaceous emergent species, especially Pontederia cordata and

Typha domingensis. The community termed "G/HE" is the border of the grass and

rooted herbaceous emergent zones, which had enough samples to be a separate category.

"Floating-HE" is the floating mat community, consisting ofP. cordata, Bidens spp.,

Ludwigia leptocarpa and a variety of other species. Out past the herbaceous emergent

communities are the deeper "Outward" communities. These include floating-leaf

emergents (Nelumbo lutea, Nymphaea odorata, andNuphar luteum), submersed plants

(Hydrilla verticillata), and deep emergents (Paspalidium geminatum). Trap points that

were on the borders between distinct vegetation communities, or were part of

communities with too few samples to have its own category, were classified as "Mixed".

Since transects were randomly chosen and the trap points were moved with the water

level, there was no way to collect equal numbers of samples from each community.









Figures 4-5 and 4-6 show results from capture success comparisons for

salamanders. Both species tend to be captured more frequently in the outer herbaceous

emergent vegetation communities and community edges, as well as greater water depths.

Pig frogs seemed to show a preference for grassy habitats and community edges,

decreasing towards more outward vegetation communities, while leopard frogs did not

show much of a pattern (Figure 4-7). However, much stronger trends appeared with

water depth (Figure 4-8). Both species, especially the leopard frogs, showed an inverse

relationship to water depth.

The two snake species showed divergent trends in capture rate. In Figure 4-9,

Florida green water snakes (Nerodiafloridana) appeared more in the rooted pickerelweed

communities, while the Florida water snakes (Nerodiafasciatapictiventris) did not have

quite as strong a tendency to be in specific habitats. Water depth seemed to be more

important in Florida water snake occurrence (Figure 4-10), with most being trapped in

shallow water and decreasing steadily with depth. The Florida green water snake did not

have such a trend. Figures 4-11 and 4-12 show that common musk turtles were captured

more frequently in vegetation habitats furthest from shore, as well as deeper water

depths. Striped mud turtles on the other hand did not have strong trends, but peak in

rooted pickerelweed habitats and intermediate water depths.

Largemouth bass did not show a strong affinity for any certain habitat, however

they were captured slightly more frequently in rooted herbaceous emergent and

borderline communities (Figure 4-13). They also appeared most in intermediate water

depths, (Figure 4-14), e.g.45-60 cm (18-24 in) deep. Bass captured in the traps were

juvenile fish, with total lengths in 2003 ranging from 5.1-14.0 cm (2-5.5 in), (n=60).









The armored catfish were found mostly in border vegetation communities (Figure 4-13).

They also were positively correlated with water depth (Figure 4-14).

Discussion

Population Estimates for Herpetofaunal Species

The small activity ranges estimated for the amphiumas and sirens were similar to

what have been found in other studies (Gehlbach and Kennedy 1978, Sorensen 2004).

Maximum distances were higher in this case, (18 m (59 ft) vs. 5 m (16 ft) for amphiumas

and 24 m (79 ft) vs. 10 m (33 ft) for greater sirens (Sorensen 2004)). These estimates

suggest that lack of movement in these animals make them susceptible to mortality

during muck removal operations. The sizes of the grids were probably not large enough

to make reasonable activity range estimates for the Florida green water snake or striped

mud turtle. Bancroft et al. (1983) documented a Florida green water snake moving 223

m (731 ft) in less than two hours. As another example, Nerodia taxispilota (brown water

snakes) have been documented moving distances greater than 1 km (0.62 miles) (Mills et

al. 1995). Mahmoud (1969) found maximum distances for several species of kinosternid

turtles, including 525.5 m (1,723.6 ft)for S. odoratus, 435.3 m (1,427.8 ft) for

Kinosternonflavescens (yellow mud turtle), 408.4 m (1,340 ft) for Kinosternon

subrubrum (Mississippi mud turtle), and 93.9 m (308 ft) for Si. ,i/hel ii'n carinatus

(Mississippi musk turtle).

Due to low recapture rates and large movements of individuals, poor density

estimates were attained with these protocols. Even when the simplest models were

utilized to estimate population sizes (the estimated number of animals with no account of

area sampled), variances were unacceptably high. Even if the estimates had been

reasonable, the study would have been limited to a small number of species, a narrow









window of opportunity when the pickerelweed zone was completely inundated with

water, and non-random locations which possessed a wide enough band of habitat to

contain the grids within a relatively homogeneous habitat. Therefore, the mark-recapture

grids will not be utilized for post-enhancement sampling.

Capture Success for Focal Species

Vegetation communities offer different tradeoffs to animal species that inhabit

them. Variations in predator efficiency, prey type and abundance, or abiotic properties

associated with dissimilar macrophyte types strongly determine their use to aquatic

vertebrates (Miranda et al. 2000). Physical properties of plant species such as branching,

leaf shape and number, plant biomass and position throughout the water column affect

animals' ability to maneuver in the habitat, as well as physical and chemical properties

such as dissolved oxygen, nutrient levels, water temperature, light penetration and current

(Chick and Mclvor 1994). Welch (2004) conducted a thorough ecological investigation

into the vegetation communities of Lake Tohopekaliga prior to the 2004 enhancement.

One finding was that the soils associated with the intermediate littoral zone depths and

Pontederia cordata communities were highly organic and low in bulk density. On the

other hand, the shallow grassy communities and the various deeper water communities

had soils higher in bulk density, and therefore were sandier in composition. Substrate

alone may provide benefits or detriments to animal species, depending on their specific

life history traits.

Water depth at a given location has strong influences on vertebrate species

distribution and habitat use. It is the main determinant of the boundaries of the littoral

ecotone, limiting emergent aquatic vegetation growth to the limits of the water

fluctuation. The gradual slope of the shoreline causes small increases or decreases in









lake stage to flood or dry out broad expanses habitat, altering its use to different species.

Water depth also establishes the volume of water that aquatic organisms have to move

through, and can provide enough space for the presence of a thermocline (Miranda et al.

2000).

The aquatic salamander species are known for burrowing in the organic sediment

associated with dense vegetation as refuge from predators and drought (Etheridge 1990,

Conant and Collins 1998). In fact many of these large salamanders have been uncovered

during muck removal operations around Florida, even when there was water still

covering deeper areas of the lake (Aresco 2001). The findings of this study indicate the

same pattern, with highest capture success occurring in densely vegetated communities.

As mentioned previously, these communities are also most associated with low bulk

density and high organic composition of the soils (Welch 2004). This indicates that not

only may the dense vegetation provide ample forage and cover for these creatures, but

also the organic sediment (muck) is preferred for burrowing. While deeper water depths

yielded more sirens and amphiumas, it is possible that they are simply more active in

deeper water, increasing detection probabilities.

Higher capture success in shallow water sites was expected for leopard frogs, since

all individuals captured in 2002 occurred in April and November when water levels were

low. This species is known to travel relatively far from aquatic habitats, given proper

cover and shade from terrestrial vegetation, depending on soil moisture and dew to

prevent desiccation (Dole 1965, Conant and Collins 1998). Leopard frogs are

particularly dense in herbaceous vegetation around lakes, with plenty of protection and

food sources in the grasses (Kilby 1936). On the other hand, pig frogs are highly aquatic,









often being associated with emergent and floating vegetation (Conant and Collins 1998).

Pig frogs were captured the most in the "mixed" vegetation community. This category

was mainly represented by floating vegetation (water hyacinth) and borders between

communities, (mainly between the floating mats of emergent vegetation and submersed

vegetation). The dominance of pig frogs found in this community may indicate that

border communities provide a tradeoff between predator avoidance and prey availability.

Decreases in capture success with increased water depth and relative distance from shore

indicate that water depth was very influential in determining the presence of both species

of ranid frogs. The leopard frog in particular has a very strong decreasing trend with

water depth, which is consistent with its more terrestrial nature.

Water depth was also an important factor in the Florida water snakes' habitat

preference, which was expected since in the 2002 sampling most individuals were caught

in April and November, both during relatively low water periods. Water depth does not

seem to have as strong an influence on Florida green water snakes. To illustrate this

difference, Seigel et al. (1995) found that during a three-year drought in Ellenton Bay,

South Carolina, many N. fasciata left the habitat only seven days after it dried out, while

N. floridana never left in large numbers. While the abundance of snakes was generally

lower, it was five years before N. floridana was captured after the drought. As discussed

previously, water snakes typically show little site fidelity and are capable of long-range

movements (Bancroft et al. 1983, Mills et al. 1995). Several of the species show

ontogenetic niche shifts with age and size, often changing diet and habitat preference at a

certain size (Mushinsky et al. 1982, Mushinsky and Miller 1993). Florida water snakes

in particular are known for feeding on fish when young, and then at 50 cm (20 in) snout-









vent length they begin to feed almost exclusively on frogs (Mushinsky et al. 1982). The

traps used in this study mainly catch adult snakes, so prey (i.e. anuran) availability in

shallow habitats may result in this species' preference for shallower water depths.

Alternatively, Florida green water snakes are not so specialized, being caught with and

regurgitating a variety of fish, frogs, and even large sirens. While both species were

captured more frequently in the emergent vegetation communities, the water depth seems

to influence the presence of these species the most.

The turtle species have different life history traits that may explain observed

differences in capture success. For example, common musk turtles are highly aquatic,

rarely leaving the water except to nest. When water levels drop, at least in ponds, they

follow the water down and then burrow into the sediment to avoid desiccation (Wygoda

1979, Gibbons et al. 1983). Mahmoud (1969) suggests that Scite ii,,I/ it'l spp are more

dependent upon water depth than Kinosternon spp. The former appears to prefer water

depths greater than 30 cm and have been found in up to seven meters of water. As shown

in this study, there is a sharp increase in capture success associated with both lakeward

vegetation communities and deeper water depths. Alternatively, striped mud turtles are

much more terrestrial, usually dispersing over land during drought or heavy rainfall to

find alternative habitats (Bennet 1972, Wygoda 1979). On the other hand, Gibbons et al.

(1983) found that Kinosternon subrubrum experienced no increased emigration due to

drought in Ellenton Bay, South Carolina, since it is a fairly terrestrial species and is not

negatively affected by dry conditions. Maximum activity of striped mud turtles occurred

in 15 cm in Oklahoma (Mahmoud 1969). In this study, peak captures occur in rooted









emergent vegetation and intermediate water depths. All species of kinosternid turtles are

thought to prefer vegetated habitats to unvegetated ones (Mahmoud 1969).

The captures of young largemouth bass in all parts of the littoral habitat were

contrary to common fisheries doctrine. One would expect them to occur almost

exclusively in the open water/submersed habitats, due to physicochemical requirements,

as well as the physical barrier of the organic berm formed by the floating vegetation mats

(Moyer et al. 1995, Allen and Tugend 2002, Allen et al. 2003). However, relatively high

water levels evidently allow young bass and other centrarchid species to enter grassy

habitats, as well as inhabit the pickerelweed zone. Miranda et al. (2000) described

vegetated aquatic habitats as a mosaic of microhabitats within larger seemingly

inhospitable macrophyte stands. Although from a human's perspective the habitat may

seem uniformly unsuitable, fish can move both horizontally and vertically to find pockets

of suitable physical (e.g., temperature) and chemical (e.g., dissolved oxygen) water

conditions for survival. Perhaps this explains the bass' ability to move through this

landscape relatively unscathed. The armored catfish on the other hand, have a high

tolerance for poor water quality due to their ability to breathe air (Brauner et al. 1995).

They were still found at deeper water depths, but his was likely due to breeding

requirements in the littoral zone for greater than 0.3 m water depths (Hostache and Mol

1998).









































Figure 4-1. Locations of 2003 sampling transects in Lake Tohopekaliga.


60 cm
45 On
30 cm
15 cm


































Figure 4-2. Diagram of weekly trap placement at specified depths.


Striped mud turtle



Fl.green water snake



Siren



Amphiuma


-


-- U

- U


0 5 10 15 20 25 30 35 40 45 50 55 60 65 70

Distance (m)

W Maximum mean distance moved and variance
U Maximum distances moved per species


Figure 4-3. Mean maximum distances traveled with variances and maximum distances,
based on results of mark-recapture sampling.







82




75
> 70 -
C 65
E 60 II GRASS
0 55 (n=799)
L) G/HE
c 50 -
o (n=230)
45 ROOTED-HE
40 (n=1174)
S35 I FLOATING-HE
> (n=714)
= 30
U30 MIXED
L 25 (n=162)
20 OUTWARD
(n=347)
15-
E 10
S5-

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48
Sample Occasion


Figure 4-4. Number of trap sites sampled in each vegetation community per sample
occasion.
















O 0.35
Q_

S0.30


-. 0.25


0 0.20


U- 0.15
c
1-
5 0.10
0

0.05
CO
0-



. 0.00
U)


Figure 4-5. Salamander capture success by vegetation community.




o





P 0.15 -





E 0.10





S0.05


Co
09














E 0.10 -
E 0 15 30 45 60


Water Depth (cm)
I SIREN
0-










I AMPHIUMA



Figure 4-6. Salamander capture success by water depth.


G G/HE R-HE F-HE M O

Vegetation Community

I SIREN
W AMPHIUMA















0
1?

.. 0.25
Q_



. 0.20




S0.15

E
0
S0.10
cr



0.05


0)
c 0.00
o G G/HE R-HE F-HE M O

Vegetation Community

S PIG FROG
I SOUTHERN LEOPARD FROG



Figure 4-7. Frog capture success by vegetation community.


o 0 15 30 45 60

Water Depths (cm)

SPIG FROG
I SOUTHERN LEOPARD FROG



Figure 4-8. Frog capture success by water depth.
















. 0.25








0-
"| 0.20


a-
o
- 0.15
03
E
0
S0.10
-0




2 0.05


"II
U)
. 0.00
o G G/HE R-HE F-HE M O

Vegetation Communities

SFLORIDA GREEN WATER SNAKE
II FLORIDA WATER SNAKE



Figure 4-9. Snake capture success by vegetation community.


o 0 15 30 45

Water Depth (cm)

SFLORIDA GREEN WATER SNAKE
I FLORIDA WATER SNAKE



Figure 4-10. Snake capture success by water depth.



































2 G G/HE R-HE F-HE M O
<
Vegetation Communities
SSTRIPED MUD TURTLE
II COMMON MUSK TURTLE


Figure 4-11. Turtle capture success by vegetation community.


o 0 15 30 45

Water Depths (cm)

SSTRIPED MUD TURTLE
I COMMON MUSK TURTLE


Figure 4-12. Turtle capture success by water depth.















1?

o. 0.20
0-




P 0.15
03



0
a)


E 0.10

U)
r-

0.05





c 0.00
U)
Co


Figure 4-13. Fish capture success by vegetation community.


0 15 30 45 60

Water Depths (cm)

LARGEMOUTH BASS
I ARMORED CATFISH


Figure 4-14. Fish capture success by water depth.


G G/HE R-HE F-HE M O

Vegetation Communities

LARGEMOUTH BASS
II ARMORED CATFISH


.o
. 0.12
0-


S0.10
I)


0-
E 0.08
C-)
0





0.04
o
1-
E 0.06

UI)

0.04
1-
o

S0.02


0)
r 0.00
Co
<














CHAPTER 5
SUMMARY AND CONCLUSIONS

Review of Aquatic Vertebrate Community Dynamics in Lake Tohopekaliga

This study has documented the conditions of the fish and herpetofaunal

communities in the littoral zone of Lake Tohopekaliga prior to extreme habitat

modifications performed in 2004. The littoral zone in this eutrophic lake defined here

includes the entire vegetated shoreline, from the grassy vegetation community on the

shore that is occasionally inundated with water to the lakeward band of emergent

vegetation that is always flooded. Animals captured in this habitat represent species that

are dependent in some way upon its unique characteristics for their survival, including

still water, cover from predators or light, nesting substrate, organic sediment in which to

burrow or forage, and appropriate prey. Perhaps the most useful information from this

study so far has been the species list, which includes juvenile centrarchids and exotic

catfish. While it is no surprise to find most of the herpetofaunal species in this type of

landscape, the quantity and quality of fish captured was not expected. Previous research

indicates that sunfish cannot and do not inhabit heavily vegetated littoral habitats, while

the two species of exotic catfish captured have not been documented at all this far north

in Florida. Species such as warmouths, bluespotted sunfish, Florida green water snakes,

sirens, striped mud turtles, and pig frogs were found in all parts of the habitat throughout

the year, indicating that they may be threatened by the habitat alterations more than other

species.









Research efforts in 2002 were focused on the Pontederia cordata zone only, which

was the species most considered a nuisance by lake managers. The widespread removal

of this species threatens to disrupt the wildlife community structure in a large area of the

lake, making research of this specific vegetation community a necessary component of

the study. In 2003, sampling was conducted across vegetation communities within the

littoral zone in an attempt to understand how different focal species are distributed

throughout the habitat. The following sections will provide a review of the observed

effects of these environmental variables on species richness, community composition and

distribution of focal species.

Air Temperature

Average air temperatures over the course of each sample occasion accounted for

much variation in the aquatic vertebrate community. Both fish and total vertebrate

richness were negatively correlated with average air temperature over time. NMS

analyses also indicated that this variable explained large percentages of variance in the

species composition of the fish and vertebrate communities. Vertebrate assemblages

represented by Regina alleni and Esox spp. were associated with high air temperatures,

while groups with Nerodiafasciata pictiventris and Rana sphenocephala as indicator

species were found at lower air temperatures.

Lake Stage

Only herpetofaunal species richness was found to decline with increased lake

stages. This variable explained much of the variation in fish, herpetofaunal, and

combined vertebrate assemblages. In periods of high lake stages, .Sicil n, iih' odoratus

and Esox spp. were the indicator species for sample occasion clusters. N. fasciata