<%BANNER%>

Ecology and Management of Wetland Forests Dominated by Prioria copaifera in Darien, Panama

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 E20110113_AAAABX INGEST_TIME 2011-01-13T14:31:15Z PACKAGE UFE0004366_00001
AGREEMENT_INFO ACCOUNT UF PROJECT UFDC
FILES
FILE SIZE 112030 DFID F20110113_AABJGA ORIGIN DEPOSITOR PATH grauel_w_Page_142.jp2 GLOBAL false PRESERVATION BIT MESSAGE_DIGEST ALGORITHM MD5
0e87a0adcc7c87279bce55a669a6c522
SHA-1
d311ae8f1d3c6c480d27975dd4237c865f7b7d31
106358 F20110113_AABJFL grauel_w_Page_082.jp2
e0f49f67609e5e78127d6d67be4dd28f
4255c270b5e916c34555f68de51a80e78204030f
28235 F20110113_AABJEX grauel_w_Page_177.jpg
811e4429732a3cb4c1025173e8c81d3b
b54bc69470b727e3217cdcb6826201ea0a6ab47b
7017 F20110113_AABIZR grauel_w_Page_069thm.jpg
a6d9e14fc908c49c42b43dda66d89e0b
7f63f7dd9e39877eb9d294566de432ffa2d51c1d
98619 F20110113_AABHXD grauel_w_Page_081.jpg
28ffcaf758dd08b197fe8a1bef854d0e
57b97270938979cfec4590f64e6757d79aa557d1
1053954 F20110113_AABIBV grauel_w_Page_109.tif
617cd01f4fcbaafbde1267c80afb9b01
3a381ab296f8df18ee3054d0960b645caeb08cce
34115 F20110113_AABHWP grauel_w_Page_094.pro
f17234ba6bcb88f44613913a3774c212
d9f5cc4925727813a84f1de76c5dd5aa38de6e69
675 F20110113_AABICJ grauel_w_Page_012.txt
cfd7e9d2f8b6e9edc30d680623d5feb3
e7cee1d1db7cb5f3b699f94b0bdf387bba38b66b
111471 F20110113_AABJGB grauel_w_Page_144.jp2
846a519ebf82888da0a25489efaffa2b
2710a7b98837e7316cdf9b9a771aee5aabf56e90
106824 F20110113_AABJFM grauel_w_Page_090.jp2
6f8b88c9fdaf2cfe1b02b3b907764fc5
76df13f61215e135c91c3a61a8c7ad4841cefc05
246060 F20110113_AABJEY grauel_w_Page_008.jp2
db42144da9b9e2d0257c6110769ef6f5
a784574cc4544a935c21d43eab588a5384d8b67f
8299 F20110113_AABIZS grauel_w_Page_026thm.jpg
3c9859e808eae011ce14f91f33d6a619
d4d5b0bd0fbbc42dfefa1f9f17cf1f7265dd614e
F20110113_AABHXE grauel_w_Page_144.tif
6d029a8171c42e58c52be8447f50eb97
870fc6b2fc7a8417ac75b257e9614f15aef43c65
33871 F20110113_AABIBW grauel_w_Page_132.QC.jpg
c3a4d70637d5c1e2a18d51d7c2482198
fc41d4459d6b9cfc159f42e9b918670f296262ba
110272 F20110113_AABHWQ grauel_w_Page_112.jp2
bfccfaa6f39f3994fc921cd8f9e32b8a
d68cebe8238b5b95460731353d9b8a4dddb90645
90653 F20110113_AABICK grauel_w_Page_072.jpg
5793c271765f804a56472483daaf3e64
76ae37922be11514fbc06f20bf0453949b73fdd7
865017 F20110113_AABJGC grauel_w_Page_149.jp2
b43e5d16b542245dcbbc1ace19117242
60c54e30cc5c956e77b2175e9bfa6fa70bc8f66a
28119 F20110113_AABJFN grauel_w_Page_092.jp2
492514899425861c91da500c5e9de576
506fc97c59717e5c621e63f27d41191186b107e8
1051974 F20110113_AABJEZ grauel_w_Page_010.jp2
80fd82b809a7c14f0012a00f347399cf
07458688f29ea88c1bb13c2eac5f48a9adf13ff8
8552 F20110113_AABIZT grauel_w_Page_061thm.jpg
c2bc713b1de918870b07719fae151904
cc7c96b5f39aa3e22acf353d38955ea0f307f08b
106493 F20110113_AABHXF grauel_w_Page_086.jp2
94c81f15e3fbe3f5350740a02681fe59
687b71dc50d319fce77003afaf9aac8d2986cf84
1708 F20110113_AABIBX grauel_w_Page_004.txt
c98efac02624c49a8fcdb4c21116c965
183262b29ee3c0f6758f1a5c4e8571a59627ef54
61594 F20110113_AABHWR grauel_w_Page_094.jpg
ec8fb9fecba3f4684f99ac013336fcf4
31412b203d167b2a02d016bd66b054200cc98ff8
4794 F20110113_AABIDA grauel_w_Page_126thm.jpg
0156e58171a32abfa49254ca67ea5251
2bad6197c2025a02aa0383b6afc1db836dcef90f
25528 F20110113_AABICL grauel_w_Page_111.QC.jpg
4ea2a3857c635abccc034c3aedf83b51
3c5cefe62c8e177dc5aa2fc6b3a664195c5235cd
129460 F20110113_AABJGD grauel_w_Page_159.jp2
915895e8c1a01d2096f6c6d655ecb326
b816a0decf2601641165f6310c5b960e903fb3a4
59680 F20110113_AABJFO grauel_w_Page_098.jp2
d366a9ddde3449f54a551d0f0d5e54e3
f36046b1f62a6fbcb7894cc3fc40b05c047bc677
119328 F20110113_AABIZU grauel_w_Page_166.jpg
dab7de36c5254d64df6821f8ff25957f
873cee627cbb6c7954c5fc66778e3a3a7cee0478
32235 F20110113_AABHXG grauel_w_Page_028.QC.jpg
4ca78c1ffb510dd095e01be002742815
ab1695056346eae81e8119ebb06e6fa2365a8909
8124 F20110113_AABIBY grauel_w_Page_137thm.jpg
5ff3c42ca0cfed978dcf734418f902cc
c1b50a4824966048a39bd62898c8275885ed6a20
98194 F20110113_AABHWS grauel_w_Page_047.jpg
0aeaecb36f19f1fe207990efd1211d58
8cdf8cb3e90c6e4ac2b1b54a0c905eca79586b20
19050 F20110113_AABIDB grauel_w_Page_149.pro
b6098ffe6f3b5af6e80f1ec821ceb0be
e3c1065c05180daaf151b975a93c6e9acbf679f9
100655 F20110113_AABICM grauel_w_Page_122.jpg
943197deb3dce1267990ecd2e557506f
28ec50064616cc1b2958b65bf44f090ccc2516ee
115574 F20110113_AABJGE grauel_w_Page_172.jp2
21c543a95b71aeb0ae6732340f86782e
8255292437f8a439e2291651debdbb7d86035fe9
48574 F20110113_AABJFP grauel_w_Page_100.jp2
98516577a652b32d88bbc994a64e9dc5
17fb216d58395d29785511df9bbc2153c31045c6
82387 F20110113_AABIZV grauel_w_Page_015.jp2
1ba8d2e2cdbb3c157f1e3d556e70dce2
c8eb91de68294b2f1ed9ef9a000409445e37d8a3
1054428 F20110113_AABHXH grauel_w_Page_124.tif
7100ad7b79f5652ab98bb5be665dfd89
360151258b50bc882063fc6881549ec7264fc929
105070 F20110113_AABIBZ grauel_w_Page_060.jp2
9afcca80b2ef8e72543016d8f2a4b86b
3a6f24eaab1a2876480b963caa2e586ec1a7fd31
1606 F20110113_AABHWT grauel_w_Page_097.txt
94d9f3eceebe22cc3ecabf9af53be544
629c93cb2168956cdead7fa470ce3da9370b537c
F20110113_AABIDC grauel_w_Page_161.tif
7b43a946e9df55bb4d0ec567d7478659
017aa28ff9eeee450067357c60e9efc6e762689a
F20110113_AABICN grauel_w_Page_097.tif
68c3b3ca8b394e0a91e9535022461a39
67d4e0be52ca4656ec3a74d29c6ffc4e7ad947d8
127018 F20110113_AABJGF grauel_w_Page_173.jp2
adae34f1657f5028ab89996b04ced21c
2765ed7b9a8225d59e7be49ef399336ce3b27f75
1051967 F20110113_AABJFQ grauel_w_Page_102.jp2
5bf6f818953b563488b699ebe389b5dc
6ae505031d4dbd6fa53045617b965490284633ce
110107 F20110113_AABIZW grauel_w_Page_108.jp2
65bce77dfd2167a4e59676259f82b41d
6b754b88b753b35c7f3a43574ac22428832dfb52
10490 F20110113_AABHXI grauel_w_Page_049.QC.jpg
b97ce0f45d212ea5de25efae3a386000
ebdd045f8fcc41d2bb66eae96d2a1206de23f201
30114 F20110113_AABHWU grauel_w_Page_034.QC.jpg
cca8ffde698977c5a9ec210595fb39d8
6d00f4d06bfcfa36eda46ce228b45e879e4d39f1
F20110113_AABIDD grauel_w_Page_005.tif
b19ed82a76ade13b23e58a3edbfb4f28
6002d01766a8155c7c572ce7ee568001f23e59fe
23655 F20110113_AABICO grauel_w_Page_054.jpg
1cec8e18aae2990f5c7cdf62e19a277a
16dfb5db5c2c83790b935b19ed731736c490b959
110316 F20110113_AABJFR grauel_w_Page_110.jp2
42e662b4f4a46bb72a0851a13b32a983
d4dbbfeb59e131bb761c6c1bb5c14b1aac662dbe
F20110113_AABIZX grauel_w_Page_084.tif
4542e83ebe465ae8e700528c5474cdca
8f347a7d7550ae90b59196e8614c923b85f53b75
231 F20110113_AABHXJ grauel_w_Page_147.txt
d693668f5a1f667d7b5feb6866b74c72
b9e4ea7f9d4aac872ae2abb58d5a684f78ddcdd9
22649 F20110113_AABHWV grauel_w_Page_148.QC.jpg
0d7c660f848be8cfbf3b8b3b7a8cbcad
65e1a4876cea95af14600ecd5af75112485a5962
8212 F20110113_AABICP grauel_w_Page_108thm.jpg
1e87336945bc14bcb4a8355989e23232
d8d72ac4bfc4fda18606692726474c5bbce88ca4
28217 F20110113_AABJGG grauel_w_Page_177.jp2
effd9a1f45c34cf07aba45efeda28f24
de10ec125bacd07a6aa741c5bee10b0ff0dad86d
105502 F20110113_AABJFS grauel_w_Page_113.jp2
a5ac1c01c2a0ffdf8768014c7e4bb735
9cab527f536b9fbb2ad4a1a97c3f040d6d520b4d
1809 F20110113_AABIZY grauel_w_Page_023.txt
7505f6e2cf9cc17c9d4e7ad2685d742f
9415ed0d37505c5e7c68d99ea8d831b2700d1d95
47567 F20110113_AABHXK grauel_w_Page_114.pro
32d2693beeeb4c04bbf3f5797ac7d782
8c8c8e75e911c8e00f7a16e373c17e93f2029ba8
8272 F20110113_AABHWW grauel_w_Page_121thm.jpg
ea5afd247aedcdbbe93caeee2b081a0c
9ad6bbc99230daae1c4e6409efdca165365aefd8
13564 F20110113_AABIDE grauel_w_Page_049.pro
443afd3ab8ce844fbf15dd17848707a1
76279a742c3d194204a428795e3f768ba7488471
17441 F20110113_AABICQ grauel_w_Page_124.pro
1ac93c46167a9159d8716f86c001972a
e85ac8c8b390cf6342bb841f0ccf4b29a08f785c
47077 F20110113_AABJGH grauel_w_Page_178.jp2
e14b94b8c7360b05fb9b0785de2b248c
e2cf43ffa8068efa419ae3fc756dd655b21b7c11
107543 F20110113_AABJFT grauel_w_Page_119.jp2
38ce2a0a38aca8923e3ef167ac273111
420a838e0ce1d0bffe67d933c596e4439072cdd7
23585 F20110113_AABIZZ grauel_w_Page_053.jpg
4ad91ec40a53ef49aec73f60dba82893
5cbca0367e541391f439a7c8ae17f3342ef4288d
129565 F20110113_AABHXL grauel_w_Page_164.jp2
5c1a20e61c2bf754665f500e464a1e54
915e52a59c8e755b413d03aed79489b42d836cbf
4751 F20110113_AABHWX grauel_w_Page_147.pro
702a945a763126e0a42f91a294faa60a
7e6a39afc46411ca9332033cafa9bca235ae6e8d
33504 F20110113_AABIDF grauel_w_Page_137.QC.jpg
c72b48e630b239d19624ba4308ab3457
10a44cc44da6d7378cb34038cb89c03d21f3ae16
99223 F20110113_AABICR grauel_w_Page_145.jpg
3eb91e9f6ccfb9d032d1dfb959be93e6
317140236b83d96cb905975d4bff3293cad7e248
F20110113_AABJGI grauel_w_Page_003.tif
25b66afc6df388458fb1991e2fe921ec
014db19e0465805b9d150aa392330a662257446e
29864 F20110113_AABJFU grauel_w_Page_123.jp2
22717ea270333f44ff22a47914239245
d2532e7c690979f27d6372cfe704c71dadb823c3
17668 F20110113_AABHXM grauel_w_Page_008.jpg
9dc3a82c50be25e4e452672ecccce70a
e4372cc7589dd271c3405d1aa201aff8c06746d0
328 F20110113_AABIDG grauel_w_Page_013.txt
07658a3fc7d10a02c46861e36a686845
9894f4e222c787e8ffe531368151af5440920870
89000 F20110113_AABHYA grauel_w_Page_084.jpg
0af094c65727584135b49954c6d09cdc
510e561d7942d75d1ebcad644297fea105e772f5
100427 F20110113_AABICS grauel_w_Page_028.jpg
a838afbd20e947fc2fd0c948deedc8d9
e0d30c7e96cb98235187d54d7cd694339c838821
25271604 F20110113_AABJGJ grauel_w_Page_007.tif
ba69fdb6df1204bb78ab0fc9b5b7220f
2d72cd8db164eba78b11f903c9a5080b4a353986
36726 F20110113_AABJFV grauel_w_Page_124.jp2
da7cbb0b76a8834792781f7805d662c4
1f27312a49728949bf1abf2b6876734de03df1a1
29913 F20110113_AABHXN grauel_w_Page_055.QC.jpg
f527ffb78d641d4862f7e6cc4af1a721
3b24ce46c570f6558c73039ebd16f46a850d2a3a
7762 F20110113_AABHWY grauel_w_Page_114thm.jpg
da0fc35a88de43de835bbe3632ef4bb7
9d3973e2a9d815be0f07a1775ac22b7054d8bc4f
39797 F20110113_AABIDH grauel_w_Page_129.jpg
0ec924a2426a9ffd5b6e16a8f8354a86
a3252d9a379df42a10a327cf501b3acc48eb62c9
103533 F20110113_AABHYB grauel_w_Page_144.jpg
1b81a740249e12e959d1f529835c9eef
aa9b1c061d870e830ce7a8d391d3d0ed6a2e78e4
7411 F20110113_AABICT grauel_w_Page_104.pro
0ecee13814051e582a3211c16b508169
4c0e5f9b5fa68f86014bda4d34ac223b6ab311ec
F20110113_AABJGK grauel_w_Page_011.tif
c039920c994bcebd8e7cf274492d4208
75f79d974a94db62d9ce2057d09f6f3322492d21
48018 F20110113_AABJFW grauel_w_Page_125.jp2
1e44b4d89800a924e1c332c14fa355ca
9a29958087c9b25300b018c177ba4ed029b7e198
48834 F20110113_AABHXO grauel_w_Page_121.pro
0dd6956c7ce5bbcf119c8f6ddef19415
f1160be2e4e25e6b0580baa706b84bf706f7ae44
F20110113_AABHWZ grauel_w_Page_138.tif
863fc660362bbd4da2d6a1cf5fac0428
a9a48412a3c7bb2abacb14fb93f78d943e88a656
1930 F20110113_AABIDI grauel_w_Page_087.txt
f0b3f565a60d09240bb41d4d62198c84
219941ab091326c2eddfb67c15221fd11cd87f52
26171 F20110113_AABHYC grauel_w_Page_074.pro
d9e37a4249146fae00b0ea5976a417e3
106db28c99870f513241440fd8209d2d3669cf99
102982 F20110113_AABICU grauel_w_Page_136.jpg
51203b58e872491952487532b8f8ce28
e1bc60eca900feda1b78bd3bdfeff598bc71e46e
F20110113_AABJHA grauel_w_Page_076.tif
cdcb982e0832766212b97702d8ed737a
b516e2f284b2233aff555493c46484f0aa26a2f6
F20110113_AABJGL grauel_w_Page_012.tif
6d18035fe6eeadfc03666b19bda2b16f
927eea2f64928be8141121e6b4268f8121df372c
42693 F20110113_AABJFX grauel_w_Page_127.jp2
e340e215162fc32615c6fa58de84897e
38bd2e30dbd94e0b23baf225816d102a7aa24380
35765 F20110113_AABHXP grauel_w_Page_169.QC.jpg
16acedf927da8fa3d8503fdb79d4ed45
f001183b587329b89e8fbba2660d14d9d0c2e724
8019 F20110113_AABIDJ grauel_w_Page_087thm.jpg
905245f1cc0172096c3d1e38b775e56f
cb4d09b4336a86b188321d25df4f412644b5821f
F20110113_AABHYD grauel_w_Page_173.tif
304d3367e516a563b471807088d2027d
7871813187962044c5a748b03d9634e5deb25723
949095 F20110113_AABICV grauel_w_Page_154.jp2
1855ac2428d3172dffef62e9bcf83639
12095fda5d4ceeddc757ce4111c02466f643470d
F20110113_AABJHB grauel_w_Page_078.tif
3a7adbde40ca4d241f0b4d9cb4d42cec
359b34b8edae8cebe0cc2e70350c3ea6ecdb7729
F20110113_AABJGM grauel_w_Page_021.tif
a72b728e5199fd58eaf9c28ae8b13d7c
3833a5982ebe04ab06d47de7bda455cec859c5f3
108065 F20110113_AABJFY grauel_w_Page_133.jp2
be1b948441cd7feb174ae0cf1b00c714
611a8a515a78a932503d41eac00e984c9f65ae11
101989 F20110113_AABHXQ grauel_w_Page_118.jpg
a961dcc5da695192a051cd85530f1bb8
b83ffb693546694233bef5cb744f5288ea7dceea
35605 F20110113_AABIDK grauel_w_Page_175.QC.jpg
0832be1f02d826bdd5f72e2959260679
120b8ffbd77cc81dcabb5d9bcc6f8434d72b4817
681 F20110113_AABHYE grauel_w_Page_049.txt
0c27e0ebfcc944c312e4fb34719e4999
fb0ca18c949703ef494dfd3b85c1f142ea3aa93d
28011 F20110113_AABICW grauel_w_Page_016.QC.jpg
82360b2e00d1ac694608e5324cd99e42
ab173adeed7cbc1d1ffd5435c79c90e261985059
F20110113_AABJHC grauel_w_Page_080.tif
95abdd8a5bba38e0d7906b73ac420303
22bc2d97ebd919ce509bdcd9d24b158d540c428a
F20110113_AABJGN grauel_w_Page_034.tif
8600dcd8260602cb0964fb4f99357117
a0f5b5f1e7943a04817b77b080eb5b64cceaea42
114716 F20110113_AABJFZ grauel_w_Page_140.jp2
7f070175add124ad9f426412801e9ff7
2d6e55eee9f269889a3b4c10d9be8168f320920f
31451 F20110113_AABHXR grauel_w_Page_019.QC.jpg
52e16d1e3025b8a2827cb85723bdfc99
b604755b8f2d720ac78de6581af0ff6654c92818
49890 F20110113_AABIEA grauel_w_Page_082.pro
0b8718a027d2614ec8b432e665d2dbec
c2c1807ba25551fb2555dcc3b074676c7096a48a
F20110113_AABIDL grauel_w_Page_036.tif
e5ca3e9325a52657668f1cc26648ee37
85bd4c58ab6f7ff5f39e53569079d8828cb7faf1
103570 F20110113_AABHYF grauel_w_Page_120.jpg
466e615ea2279baeefc178719f7343cf
d6d0e980ce46a8b850ef926fe985bc1d6c2e3d3c
7790 F20110113_AABICX grauel_w_Page_109thm.jpg
a4dcc7faa1956331e2cab861cb6bb5c8
cae1c28f9b9f5edb74bfddaae252ef47cbd9f398
F20110113_AABJHD grauel_w_Page_086.tif
50e16cf02c4ee5389301c508de5ba350
8f3cefbbe46df83179a3e68ce4c770757c1fd08d
F20110113_AABJGO grauel_w_Page_038.tif
41e343306b7986a776c28f1b832012c2
95f5e528905cd5bd69821bc031e6ec573f5c090d
F20110113_AABHXS grauel_w_Page_129.tif
bf3c02f9541b022b99e98fa5cdc0ff58
509ad337fde69743cf5dad536be263d4d646b4c6
68777 F20110113_AABIEB grauel_w_Page_148.jpg
21d05b6eaaf2a74b3bdaac0777122226
1d31adf3861a447fa93eb7792963bade0ad24a9d
50225 F20110113_AABIDM grauel_w_Page_079.pro
a9d50418700b96bb9269ad27f3158d1c
75b6d504fa3627ea8dc3b177a2663b6b75919183
62001 F20110113_AABHYG grauel_w_Page_094.jp2
0329f3a605aa3b72e1465d03f1ec70dc
2499f3b66c4f93d7a3e3c7214c4f538fc29700b4
1858 F20110113_AABICY grauel_w_Page_155thm.jpg
abcf30f716e0c39e344d93693a2c7c19
d70aa7cb4e6b97d9975fe4600920c36c82289b23
F20110113_AABJHE grauel_w_Page_094.tif
be135ff19dbf5e8a0f6e01bcfc4ed82d
6d7b3d63150f407ad000fb95c6fba3efec072641
F20110113_AABJGP grauel_w_Page_040.tif
8b2cc99c4f6b63a19dd9e779edeab25c
06bd51b0326e8b2a3f6239dc57d90a3ec139b261
F20110113_AABHXT grauel_w_Page_029.tif
d23048dd4eb6530b42013007a0497799
6d95ead4868e3e0d0fba7d8d80571176060074fb
F20110113_AABIEC grauel_w_Page_026.tif
4d76a6adc36cd75faa007df77edc22a7
a07e09055a8c505fefbb7aca9e084ef4a7155ba8
1904 F20110113_AABIDN grauel_w_Page_090.txt
5c1725c7a1ce68704962225af24539fa
329b27b3e3a852efccdf27439d29aacb34753ddd
7993 F20110113_AABHYH grauel_w_Page_030thm.jpg
7744072f5eaaba4df1e12ac3e3c56236
c3aa7f4fcd5f41b83780d41a551c83be226bf3b4
103108 F20110113_AABICZ grauel_w_Page_044.jp2
237c24665b4c1bdc5ae15a84353ae299
e4d20b4677c01b90a2f5e23fcd01692897680253
F20110113_AABJHF grauel_w_Page_105.tif
4f9603395e86079c6aa44799cc24185e
a6f1a718137da2a8f8646abbb1db07d737734713
F20110113_AABJGQ grauel_w_Page_043.tif
a3e1f652ffeab9014719061cb49eac14
c92bafc74ec34757678cb6439d73264d801318d0
48670 F20110113_AABHXU grauel_w_Page_030.pro
bfbdc3062e638aee1eed17f7801cc6d8
2d56174f2b8fa3a7a18b76fe2417b2d1d11be875
1989 F20110113_AABIED grauel_w_Page_135.txt
bd5f23cbac1ded2e5b7de6be8bd347e1
d6ad7f3c169203cba27b891144b16de13a6cb5e5
8349 F20110113_AABIDO grauel_w_Page_116thm.jpg
7f224379c7765b22f5f683f69dce4a35
370bc4b2b0803bc35bbb5266e5e2b6574e633b5c
100358 F20110113_AABHYI grauel_w_Page_119.jpg
d9a86b792c1d4abfd8672d62573fd9e3
3b4dde6336811f49dc12ac5e15dcdfb2c1a436cd
F20110113_AABJHG grauel_w_Page_108.tif
186e89acd1aaa0a3b49479c32b36b1b7
169f2f8baed02c5316efdfa3f3ed92aacd9f21fe
F20110113_AABJGR grauel_w_Page_044.tif
fe643ff8fa0bdadfd5bd99978c6f8074
eda7c0a6f934a4a1cff34cfa136e1fc6e4058d50
108967 F20110113_AABHXV grauel_w_Page_085.jp2
045692a1ec79e47187da6051c88df615
c5e7ee4ebe1edb2df5a78d19b01d495b90dbdb47
98029 F20110113_AABIEE grauel_w_Page_072.jp2
346e15e606e63deb3bf229624f1834b0
ce555344e9b12143a016c8d9a2629e11d360c315
51172 F20110113_AABIDP grauel_w_Page_132.pro
ba3af0714bfe7777dd0d5b0074b43035
5b98fa6f67edbfa9aee997b21151cc72e4dd16bd
F20110113_AABHYJ grauel_w_Page_041.tif
b87a9d6224a2f2faf9bcb93c5e625fdd
20471303405c59be66722cce6e0da4a06fd33cb5
F20110113_AABJGS grauel_w_Page_045.tif
7d5abda750bb47d1fdb7fafd376cc29d
87fdd87d436cda6e36cdc8651c8a189a705ba0c0
8168 F20110113_AABHXW grauel_w_Page_136thm.jpg
8429dd261378e07b259acebfafcdc05f
5045ed964ab338903f777e23f6117ce99e79e434
F20110113_AABIDQ grauel_w_Page_089.tif
a26cfd332444d91afe07863b14efa90c
028a20cbb97c0c0260badd855604bcb053bc1b45
7664 F20110113_AABHYK grauel_w_Page_152thm.jpg
4c51c592f4e583048cd9894e82f2c817
5bb6befedb86fcc9ba07e81d56799cc2953ac445
F20110113_AABJHH grauel_w_Page_111.tif
8612c5ec5ddb1bc4b0e3b24e92d2bc44
919e624a961af3424a33fb829559f45550a17255
F20110113_AABJGT grauel_w_Page_049.tif
cce6798cb3d27b12932b520aa7f6add3
b693b5f373f81d98b04387bd70f1cb99bd9fbda6
1051985 F20110113_AABHXX grauel_w_Page_074.jp2
53f0b9887bd08a66e1fc65c80c8e1ddc
ecbe1394265f622ed7329542db75174d187f7a42
33023 F20110113_AABIEF grauel_w_Page_012.jpg
4ad835d4754a15632390cc11e0c7347a
809ba46cedf306619977c4288081cab3cb6253c5
7909 F20110113_AABIDR grauel_w_Page_146thm.jpg
e448405e6b085769e5c658172cd1b9ca
b6e003cc2f8364bb165843607aaa35158c41f932
33009 F20110113_AABHYL grauel_w_Page_029.QC.jpg
81ca1f5b928c32afe3775fcc4c1a48f4
000ced71c36a9f523168519f4ba5e5f2761a1751
F20110113_AABJHI grauel_w_Page_117.tif
3dd33dfb10889cc439a21985a551eb8b
6818ff1ced50f2cd01061f6a5d48bac8bacf5af7
F20110113_AABJGU grauel_w_Page_051.tif
6cb4cafbfa2121d6cbda0e6e2ee79c17
b2976371ca024fcfef0694749db3d79f98e6d674
61139 F20110113_AABHXY grauel_w_Page_166.pro
32710c62eedec7ed026ad125d8623f7e
5f78e9c3a9d85be646b99ed3747ea5857d5c221d
31358 F20110113_AABIEG grauel_w_Page_009.QC.jpg
b9d3800a859c9fcf052660c9ce97b7a4
aac7044c6985f6e1f690b49b1bba9bdb8127d739
98730 F20110113_AABHZA grauel_w_Page_090.jpg
a7f60d2869b9b66800a2a0e6bd30906e
b6c689e1da98cfa5256c8ac34ff8ebd8b0f3f327
51593 F20110113_AABIDS grauel_w_Page_112.pro
ab18a8b78af50e56db3ac70c8ce88107
c5bee74ad349766d8b75aa3d7f715224668777ce
8723 F20110113_AABHYM grauel_w_Page_166thm.jpg
4f1d0ca1bf4d7b5f9bd452e441c228cc
b405acf697e7a0617e7715786b16ae096de39e28
F20110113_AABJHJ grauel_w_Page_123.tif
c990cc1d968564762886a9c7583d26c7
851e3c87b9edec80a2171fb23ae9895b587d57e9
F20110113_AABJGV grauel_w_Page_055.tif
8044a42e1c834d0e569b78757f2552f3
bdc108b18389895d63433dad4ae523d7595d5517
46842 F20110113_AABIEH grauel_w_Page_138.pro
95a55310f142296348768beb4e07efd1
93f4accf803e4a1c507ac3bd4d5475e8790c5f3f
F20110113_AABHZB grauel_w_Page_160.tif
f4d098abecb48a19cb2b0141166f5bb9
fc1dbf177232b44be559d675304adbe00dc65671
93485 F20110113_AABIDT grauel_w_Page_006.jpg
d80e4f4eb5ee55c0d501a0fd6fb88ab7
001a6ea2d2db79a1a7baef2cbfdc9a289ff8d02e
9333 F20110113_AABHYN grauel_w_Page_052.QC.jpg
eb4e13bdf92f928baa721320fb6a8bf1
335a781690ca3c748cedb296a476806f5bf4d4ef
F20110113_AABJHK grauel_w_Page_125.tif
33f9fc28176e84c19f84939470463ca6
62984601a0f095691f8b28152389ad1db2e6e8e7
F20110113_AABJGW grauel_w_Page_058.tif
af87dc21e383d68ea762efbd3cb301bb
2024114e39a3043fcfb9032728175aa64c1756d1
1963 F20110113_AABIEI grauel_w_Page_064.txt
aa13c77c716f00cb14f533c98a748ca1
2cc3892cfb2ff36e8b2ffec0aa3b8caf96175e93
35011 F20110113_AABHZC grauel_w_Page_018.QC.jpg
793d82557922e1018cca9c4d7bf3f9ef
cf7965b3a0a31cbfc052c61c153eb46b499380aa
31212 F20110113_AABIDU grauel_w_Page_042.QC.jpg
b0a6ea9e48d89dcd40ebcf40025dc4af
7a7c613599f525d4f8fa8bd7413067d37c3fabac
7589 F20110113_AABHYO grauel_w_Page_024thm.jpg
ad4b3a57091b0b59ef5a68d0cae1c67a
9f06d492848f0e6ed09ae459554bf9af5dd6786e
60481 F20110113_AABHXZ grauel_w_Page_157.pro
d5955704688e1f5b9ba3f740b46e164a
8ef97905a244cd776c18e4d2653951494aed7b4e
47246 F20110113_AABJIA grauel_w_Page_022.pro
923f70ea6b0092d417d9e0ada6f044f7
b9786befa29b4d1e5cfaa774e56d69a5c727d21a
F20110113_AABJHL grauel_w_Page_130.tif
727701263729f756a7ea37d3d896021d
6eb91ce9f4bbc4b178ff61e6111c5035c786fa1f
F20110113_AABJGX grauel_w_Page_061.tif
01698e76451d26456ed15e136b845215
7e8426b2aa11ad57f24a4590a63410cb47b9a9ab
F20110113_AABIEJ grauel_w_Page_023.tif
505b39d0f6ecc541094af9ba874db95c
feb110fda741130ad4a11e47d0d722d32979e242
29463 F20110113_AABHZD grauel_w_Page_041.QC.jpg
b28b0eaf690257e158b35043840f33ec
fb46227b62444995ed47330af3c802641bf4bc3d
F20110113_AABIDV grauel_w_Page_098.tif
344eab766e8c9df53324173ec19969d8
f5aa16db401d278182d5ec67a9b361cc1ebbb396
29834 F20110113_AABHYP grauel_w_Page_024.QC.jpg
ca5aef8e75d1ed2bf3d40b576c1433d5
a4d227f90e498d17a8661eee20ecfab9e015452a
50356 F20110113_AABJIB grauel_w_Page_027.pro
7892b46c6581b53cca5f42b21059a23f
a147a75a47b52d39dcfee7ed9adf06bfad4ec8a2
F20110113_AABJHM grauel_w_Page_133.tif
78b143d849e66b5ccd9a86c9744c5439
9a617c91442b66a569a02d33cc1f8b799a0c65db
F20110113_AABJGY grauel_w_Page_062.tif
19b475014eb7fafab48aaffaf48f6e89
ce2006386eb11874c2e467b89bfc90b6b03be12e
109244 F20110113_AABIEK grauel_w_Page_073.jp2
61187106429617a393c6954fa07b239d
c68e92ba4f6a9b0fda4274f7bb5b273fa2ce63bd
108152 F20110113_AABHZE grauel_w_Page_121.jp2
e305f6ea81623a0e5c5f4342bbf0e959
54c3cc3838dc50754c3b765385dbce664f45592c
8423998 F20110113_AABIDW grauel_w_Page_053.tif
60e427e7ba936120371ff0c49c53347b
3b48009942295b635e631a36e4ca44bb524d96a1
1882 F20110113_AABHYQ grauel_w_Page_034.txt
34c014f5eddcf15d95f03b36207c631b
25eb19b03776dba3cbc4c5588b9bbf6a98abe73e
52044 F20110113_AABJIC grauel_w_Page_043.pro
2b6e7759a4d568e1d0af56b800eb5836
7acb77bc64f9021f9ff293d0d6e0c70744d3ce7f
F20110113_AABJHN grauel_w_Page_136.tif
4bdaee3ba9ddbf4ee499f596a4a39ad3
71ccfdcf40fdec3634800a9513af81629fe453e7
F20110113_AABJGZ grauel_w_Page_069.tif
9677ba6ea79636f9766f6daf11230518
5cdd1d417579725dceade8fc88187be696a4900d
97318 F20110113_AABIEL grauel_w_Page_036.jpg
66d2c789cb6efcafda8fc5acd1dcfd53
faf97f2a7f97cfbb897206078dacd7ded5beea94
96568 F20110113_AABHZF grauel_w_Page_077.jpg
ce09e61b493d14cc45194f119af41f24
a4c9142da605ee5dd588f20ea0828ca2227f927f
47157 F20110113_AABIDX grauel_w_Page_131.pro
0358c77c63d2bac6dd5e9e8e8e2abf95
365af2e54ec12986a7de5a911cd31392929bbe11
102808 F20110113_AABHYR grauel_w_Page_057.jpg
6328953d4a8649180111b002df760fc7
1d74d364b5b4365b6b0bbbcf229069792dfc6f2b
30252 F20110113_AABIFA grauel_w_Page_023.QC.jpg
98d8beffa09f9253e337038c7c1dcc52
62fd0c72dd10866cf5b303bd8663fa756fb0015b
48889 F20110113_AABJID grauel_w_Page_047.pro
eb67a8a9eff65f1914f9c9d50a1c2405
368d6674abd0e3431cb7634368ce936a85665785
F20110113_AABJHO grauel_w_Page_137.tif
b372d229dab9aa2d8bba7d63cb048ce7
36d87919dba23d3a4debdbc01e387ec6d8012822
1728 F20110113_AABIEM grauel_w_Page_084.txt
b2328a089c996cd60ed285be2b583e13
96ef404b71543744cc91fd4c3c987fd87ad4b523
8758 F20110113_AABHZG grauel_w_Page_163thm.jpg
cc3fabc286aedc3b0292ffec0ee46932
b0a3dce543b9ab6c86cfe344be9d15e402ada12c
F20110113_AABIDY grauel_w_Page_096.tif
dc871cc82ffc516c8f13cb6918275e00
49315be8900bea43454643f2cfea8ec5108ff29c
27892 F20110113_AABHYS grauel_w_Page_069.QC.jpg
a1b02f1921f95bd1795f2d8310003627
03beb5f37cb3a21c700809724b4a8d446db0021c
14586 F20110113_AABIFB grauel_w_Page_125.QC.jpg
1169ea9eff400e7af748b66d740e4ea5
8c1d30fc0a39c42402bb9e6bb9ffc01524b54b01
37845 F20110113_AABJIE grauel_w_Page_048.pro
8aca3bc10c8dc10dba73f187c1136997
549aa22553c160e05fa45db11757e1fabb80c76d
F20110113_AABJHP grauel_w_Page_139.tif
5ec4e329eeed820e413a5c245d21c769
a28854cd292b61ff5611869ce5eebf92a4b26625
35775 F20110113_AABIEN grauel_w_Page_160.QC.jpg
5d605ee49569983531e69b45f0997b18
676f27ec0e2e21fc00b9382fc53156b042ff639a
F20110113_AABHZH grauel_w_Page_056.tif
db82eab3848c8d11fb31af84572f9706
b9e79e1313431348d38e544723b4f171d3505bd0
2471 F20110113_AABIDZ grauel_w_Page_176.txt
2e476e88fac6644e01c55d44fbfe42e1
60db0ca24a1f00bfc60991b2a2cccf9505108010
42443 F20110113_AABHYT grauel_w_Page_153.pro
a9cf07d6d9523b3391cc18740733ac0c
48291cb42f3f62efa6ccf95dfa2485be95eb8cbe
119005 F20110113_AABIFC grauel_w_Page_167.jpg
800b02419e73390e863c82b07a21bc52
60f2729d2ad8b4065929119c3a35b37dc7cfa491
12635 F20110113_AABJIF grauel_w_Page_050.pro
7bf526ec6160c5b96cf3ba47166988fa
c092b50b47a89b5dac91cc8303ed596f403073df
F20110113_AABJHQ grauel_w_Page_145.tif
aa177319643f137c0d3461f8bc0c195d
9a4793a221a75dc03824dd1cbee55699c549a9f2
2434 F20110113_AABIEO grauel_w_Page_165.txt
3999f0bd2d674738fed410e51ae4a086
d5d4c583f6c0639536d57e88d52e69fd2dca3ac8
F20110113_AABHZI grauel_w_Page_172.tif
90301d9fffa3764b16237aae40dd377c
0935c76864cbf225425eec654f7c9def7940aeba
104991 F20110113_AABHYU grauel_w_Page_035.jpg
8bda12dc6cf895d818bdf3d28bcc04d7
e1a1f98b70a80cb5d6d5fe18ec0932a82aad6939
202284 F20110113_AABIFD grauel_w_Page_054.jp2
a939530fad970f1be800f8bc768582e0
02bfeb26b61e68447f979a4f4c22e323072d5bf9
44127 F20110113_AABJIG grauel_w_Page_055.pro
23beb4921c782ec5fb195714d540f1ae
348e6aabb78743a75142f1c857dd628034ec0f15
F20110113_AABJHR grauel_w_Page_147.tif
2d8fa41e9a3ace3fdc0ead1d5d484abe
0606a2d8e06c3c05bee43dca073fa0456c3b4e9d
26320 F20110113_AABIEP grauel_w_Page_001.jp2
55acc6c3953fa180953247e724365580
1bb9a3e4b0a203edc5daaca930023a9925567045
3394 F20110113_AABHZJ grauel_w_Page_099thm.jpg
0400f78ca349c95e1155d2dae7a3ceff
ad71637ed1b9be6506072aa86aa56947922fee7c
32482 F20110113_AABHYV grauel_w_Page_119.QC.jpg
2f142575544e8cf61462fad64a6dd48d
0d86bea82eff646ba463da2e7268a8e1ab7f698d
48458 F20110113_AABIFE grauel_w_Page_028.pro
cdcadb1cf3652d12d67d07f78570f1fe
c6b1d4615be2ae0f1a20e62ffe2752bcb8de9c8b
49496 F20110113_AABJIH grauel_w_Page_056.pro
49509a9bb7e7182f14b8a4bc2940aa8e
5e4c026a37ba5ef83ad395b5a58f525aa0fbbd75
F20110113_AABJHS grauel_w_Page_148.tif
0eea350cb0ab3d3508956fa0f0c48b6f
7493de84a878a4174a1be5f679ca79af202f3ee0
52679 F20110113_AABIEQ grauel_w_Page_020.pro
4733386e3d8a45d6918ef6ee92b65234
33e8a36ac7c3965c2462337deeede4a1f8996326
51741 F20110113_AABHZK grauel_w_Page_035.pro
7b60be1b7af4f43fd13711591ee9c571
b0f59a914cce03af1e94909fa278629ab4e4e0da
2150 F20110113_AABHYW grauel_w_Page_001thm.jpg
33fe41b6f15f3d9b65c3cacee731f2c7
ce5ec41777401f22940695d843492f40cb523ca2
28972 F20110113_AABIFF grauel_w_Page_153.QC.jpg
8ef6dcc6956cbe7b173e0ab941f07fd1
6eeea38aa370140bf82a4d465ffb5834e13b9011
F20110113_AABJHT grauel_w_Page_151.tif
74d0174a69bcadc49e8eb7d3b8e580fc
0099edff8ea7a3fdc37c8d5d88165168046b7bcd
F20110113_AABIER grauel_w_Page_082.tif
e92118a0032e2dd8df5877435d733e6d
36ec2629842677edb7bc8b32e6e9ce8397eba2f3
91380 F20110113_AABHZL grauel_w_Page_041.jpg
67cc0ef70968f2b990544ae48ed6c0b7
ab9557e74572eae955ccf730384b13e3e463f46e
F20110113_AABHYX grauel_w_Page_114.tif
6bd0b70a3cbf2a835e3f8df475231a8e
c3504e9b7a82fe2727de243d676abf1e18d341f1
47391 F20110113_AABJII grauel_w_Page_060.pro
74c79580cceefdf750370e01b55c798f
166d0399985b5502437a431152c240f829ab710e
F20110113_AABJHU grauel_w_Page_154.tif
e349ea2c796ba3d2f3605092679769cb
a2b3264df920a361bc2885df07dea845058f2251
5280 F20110113_AABIES grauel_w_Page_066thm.jpg
204448625815990e938ba2a98c14834a
59e1066549c82e760f9d5c4ba5a84a7b5d7fe8ed
96430 F20110113_AABHZM grauel_w_Page_038.jpg
3ec7fb4f45fa7f7d1ec50a8d06edf8c4
4fd6ea98eb2861642e0656f47b682052edc0e77d
140738 F20110113_AABHYY grauel_w_Page_169.jp2
4724317b4bf4d1fb902ddb27fcd0fa2c
be1bc55106b42683d6332e3558463cf82ef3095b
F20110113_AABIFG grauel_w_Page_106.tif
078ae5f48a3e75b3430feeabcd581e84
69c0f9cc028ecc24e7718d7c7d8a16424de6a89b
35039 F20110113_AABJIJ grauel_w_Page_066.pro
866d19b745c22f858faf3af9445fb2e3
32aca917c46d1dcae9d4705785a15787e4a26fb3
F20110113_AABJHV grauel_w_Page_163.tif
1a052ddba79a90f60e7755141db6dfd7
69125a81ec40c0987aeab97149467f12647b02d4
113303 F20110113_AABIET grauel_w_Page_070.jp2
ae52620cde741f19b1cf91509a99c0b7
885548a6276a1d241e3982394e2848d953ff9672
43881 F20110113_AABHZN grauel_w_Page_072.pro
9fb5e033f9f234cec25a6db83ceec14b
58eebc798154f811e552b4e2d36e229d0c2b498a
F20110113_AABHYZ grauel_w_Page_157.tif
f315e55fc313ffb5006a188553a230c9
12cb3bcf876de80fe714acf528b90e394849ece9
34147 F20110113_AABIFH grauel_w_Page_026.QC.jpg
d34838ca09883f673df5875c2e59e696
3d3d38a8dcb658e7948686b5dc210ff84720b221
17418 F20110113_AABJIK grauel_w_Page_067.pro
aae92ff0a20309ce6d439b9328f9bbcf
687d4c8d1ce51c3e4da4bd544aa8c78ba8c51bc4
F20110113_AABJHW grauel_w_Page_165.tif
3ca634b15303c9d1895d70fb06d6862c
ee27df2c965d455ca97305a45e0fdf10a065cbb2
98899 F20110113_AABIEU grauel_w_Page_087.jpg
949fe278ebaee0107918310df381e35a
0f8a98074e2f661122291d3ed6336ae762869db8
44687 F20110113_AABHZO grauel_w_Page_041.pro
94b0facfbf5977e87eb6d7018c6876fa
8833ac2e6227c8fbd021df892b43bab46c48f812
105115 F20110113_AABIFI grauel_w_Page_021.jpg
d22c743584bd9fe25db2d3f4c690b19c
50054ea8d6399d356d5f415174fac71c972d3d79
42531 F20110113_AABJJA grauel_w_Page_130.pro
d5bed6d0b46885fadcc88940ab43a8e7
e78cca3830a1c5a2d989561ecc4249e4af471cef
21513 F20110113_AABJIL grauel_w_Page_068.pro
a5c9a1f4ef99cfe89957451e5c297fff
bf3c0fcc49cd0b8d629b764d842e66f7f686e2a3
F20110113_AABJHX grauel_w_Page_169.tif
5d91eef2b3754eb3f3785c335c655b5d
c19331660e950e7566e981bded174ff3378deb51
110029 F20110113_AABIEV grauel_w_Page_046.jp2
a7a49121ef81ae0cf1fb10f02e029543
27f8a2954059ad1a1d03ae8e872dcc273876466a
6858 F20110113_AABHZP grauel_w_Page_013.pro
bbdfedffef2321c13eb1b6ccec46448d
50ebc97c04e0faa120a2013548e42b06dc780a0d
102963 F20110113_AABIFJ grauel_w_Page_046.jpg
97c7443bb853e3629542864352b14ec2
1951485d4da8fc9fcff992e7e29d5a3b27390213
50494 F20110113_AABJJB grauel_w_Page_135.pro
477deb9c1eac22bd034b71eb779cfebd
5c594993737e2e1cca4da322f5217e7163cf4196
49280 F20110113_AABJIM grauel_w_Page_080.pro
5931ab44b2132dbb3d467ff77506bd99
efa93794b947d035f77aae2c2b733bebe0a55256
F20110113_AABJHY grauel_w_Page_171.tif
47faf6b1a085d9a24c9ff562209a88b9
4af096ee5227342385b43b33f68f96069adca4f4
127032 F20110113_AABIEW grauel_w_Page_165.jpg
f5e3c8724f09d73d4eee9def91a88d7e
a7ca1d748fd7c3f4953f755e7dd5d003dfebf566
5246 F20110113_AABHZQ grauel_w_Page_098thm.jpg
570cb14533e990720ed70b8f9ad2a396
f23a7ee828a9757549b6116e75835cf83f533ba5
8652 F20110113_AABIFK grauel_w_Page_159thm.jpg
81d5cdb958c42fce53f7eaccfa5f49ca
2c034fb138bcec226626bcf9fc599472abb65fb8
48388 F20110113_AABJJC grauel_w_Page_146.pro
ac1202b39218b616d8889d7830441d67
52e937abc1cd4726024a33fbd30b71711a415be6
49789 F20110113_AABJIN grauel_w_Page_089.pro
fef05272fc53a322a5b132e7864c9d79
5b85373bf3173b6d44e3cc79c09a287066d371cf
37770 F20110113_AABJHZ grauel_w_Page_014.pro
2fcd215fa109d8842b7bd6bfcc2bd862
999b74c4305659c31ec556d3b1098cd0d95f4d7c
8471 F20110113_AABHZR grauel_w_Page_107thm.jpg
9015003ece75b3211c1f2f53dc54a3e8
eeae79340fab31ea287a50953566d9f66d5195c3
F20110113_AABIGA grauel_w_Page_122.tif
af43af9001c3b61866ac307746443978
f2718162b0e4a79dac0e8b9a5b55ae2f557e1665
100239 F20110113_AABIFL grauel_w_Page_085.jpg
a78ebc5ba2d40515d32125d7d669531d
b8b9aa5f5bdd1ebff5316eb9ff3630cd8c8afe3c
F20110113_AABIEX grauel_w_Page_100.tif
aaaccd2616d6d5acaac79f47de4f4ed4
87e0740971447b9c10fe36b8d079076c539565eb
8161 F20110113_AABJJD grauel_w_Page_155.pro
0d32bebf6e66e1988229f9c6a853f1f3
f3474c39eca847d32144acb33cdf3408eb3a38d4
48479 F20110113_AABJIO grauel_w_Page_091.pro
f38de069a15031d13f1a93a43fef4954
6f2ec613f17c24b3365671496eda28f7200f8713
121902 F20110113_AABHZS grauel_w_Page_176.jpg
defd44b2f33933ccc73e1cb12effce8d
69689f9bc79b227b9af802d5aaa6b54e6175447b
7408 F20110113_AABIGB grauel_w_Page_004thm.jpg
0642fa512a40c46e4058a43178f65a47
4702bafe6f0426227493fdc3d6cb2041bd519b80
7784 F20110113_AABIFM grauel_w_Page_022thm.jpg
47821ea9f92bdaf098453adae0578e3a
6e7fbcff00e144edf4fd7ab3fcab098aef72beaf
31447 F20110113_AABIEY grauel_w_Page_131.QC.jpg
85a58ec087a41b0e4efb0c3994a23d5c
e767fe8e59405a16325a47bd6376a41c327d6262
62499 F20110113_AABJJE grauel_w_Page_159.pro
777dd92a19993b7cc82bdc9615f0720c
ddee549ddf0d24c9c3ad0a9e218adf71bf634f1f
29331 F20110113_AABJIP grauel_w_Page_099.pro
f595d48c9492c38121d55653a259e6cb
335439bd761fa58bd827967a4bb9d3deb660abd6
F20110113_AABHZT grauel_w_Page_135.tif
483f82f3dab1ce23a0f1e8559218a132
1a090345c64e5e743bf1554f4be925e0ad7458fd
7919 F20110113_AABIGC grauel_w_Page_113thm.jpg
d2ee7797cca0842f90093cd935d478a6
d85fb37de73dad7225226620f22319a2a15f8507
45708 F20110113_AABIFN grauel_w_Page_023.pro
01340bb7b98a96ba726483b2ca3cdab9
1797e2c48aefd6e3836a6a8b5d5e351c2219e4a0
6112 F20110113_AABIEZ grauel_w_Page_015thm.jpg
5e70cbd8cdd4e5d7c604318a9694e488
7e758298a4a333fe50511c0a0b6a5d2e67543aa0
60040 F20110113_AABJJF grauel_w_Page_165.pro
2ce4a2d3bace113f5a2b35defb142c3c
7d4062626aca64ca87db259a72ba0585c581c6fb
24381 F20110113_AABJIQ grauel_w_Page_100.pro
b439d28ee2b87f23f09345b016ba5816
b4f1d2aa36b76935289af7ecd6dafb5d759f114a
F20110113_AABHZU grauel_w_Page_001.tif
936e4c8df6c582844212cba33c51d5d6
beacca359ff77b4a2ab581f0642f0a083d5e262a
6465 F20110113_AABIGD grauel_w_Page_014thm.jpg
1ec03fa149cd3d40e1194380aec8c4fe
d7899fa35f0210556ac450ad5b22c7ba6a77c4ca
6359 F20110113_AABIFO grauel_w_Page_008.pro
1a0954ac6423e64f81341d703c77a821
72dd85271db451e648a9407b8435b98de9f5b0fc
62628 F20110113_AABJJG grauel_w_Page_170.pro
13907a0c0500bd0fe8c543ae4d1c282e
50f2828a5f8793cd2bae3555e6efa91f7674afda
6885 F20110113_AABJIR grauel_w_Page_102.pro
adcbebd39022b59756069fcfe534588b
3c1f12eab01d5e9bf3199ac1753be276e1020a94
111737 F20110113_AABHZV grauel_w_Page_120.jp2
f4cdf953ea4e0431a972181908103ab2
26471e3b602d1f0c3ce9654eeb469f3070b5b2a6
110209 F20110113_AABIGE grauel_w_Page_040.jp2
04c2bb40bc0093f4ff36b544ceb6ef27
822b7619c2ad0da9db28e03009b5765a5ef5758a
F20110113_AABIFP grauel_w_Page_066.tif
3ddbe0d60c38b16b5ccafe01d9d89734
c17ada2f03bbad3182813e3e0e75db8b87a39a03
55279 F20110113_AABJJH grauel_w_Page_172.pro
457d679fe55618dea47db8c3f9ff4227
c939d170afc6f55c9538ad562da1288d7e16a136
7633 F20110113_AABJIS grauel_w_Page_105.pro
629c69efee20bfde2ce93b0354923757
765d330c1dfe98cb99449666128cfbd3041ddac8
98560 F20110113_AABHZW grauel_w_Page_041.jp2
de5221e4281becd1bef0bc16a9f4d575
7fe62caaba98b113f47078ae89a4c6e7f9bc0510
47865 F20110113_AABIGF grauel_w_Page_059.pro
7a5da8296dcfbf4ffbddf4843904a1b0
2a39260865bad5953e595ce1726bb92ad45d0b19
8141 F20110113_AABIFQ grauel_w_Page_029thm.jpg
c481a63ca19086f533ae98a654baa94f
fcb1532d82167d259a20b288bfc946d0a7af33bf
433 F20110113_AABJJI grauel_w_Page_008.txt
0e4510d735a4a61acc6f151f0d7361e9
48f2e827d2946b05179e77e1ba78dab709098293
43861 F20110113_AABJIT grauel_w_Page_106.pro
38d531b06fe13489aa28cbd8c87198b5
8092e335b364b54c95631036cf29238c8c744f46
847657 F20110113_AABHZX grauel_w_Page_017.jp2
02e90a479b8a0dcc91eee5c95bf4344e
9ee5a7958d134ca85968674f020a9d0ebdc76c66
8560 F20110113_AABIGG grauel_w_Page_172thm.jpg
6b3a6bdd60fb6675e30fed2d18b8ae6f
19548df97e586e03713ff3f4d28b0b46b4953584
1459 F20110113_AABIFR grauel_w_Page_015.txt
17db01e2337d227d10eadd541f807e9f
767d4da8b27f611c558bdc47ccabd4462e686142
50740 F20110113_AABJIU grauel_w_Page_108.pro
23d39d690c9a2ac5d8b60091af701420
b84f91744294f24ef666ab7c85de5298decfbe08
1328 F20110113_AABHZY grauel_w_Page_002.pro
f0ecd25e9eda698a0ea1d56bede12bed
091d240c3d887a4ed157bdcf3ac307ae17a8ddf9
3210 F20110113_AABIFS grauel_w_Page_105thm.jpg
af15c941e8f54de00169aa391e6f74c1
2c49a482d218e6634d26fb40e543ebc7c3590004
2262 F20110113_AABJJJ grauel_w_Page_011.txt
1ac0eb8c3c83f31682f6068a4cd35763
7bcf4e54ca20f139b09e3028a59d7d4a55981e71
46739 F20110113_AABJIV grauel_w_Page_109.pro
8bbe8ce7ea5a83b5188e27d5d3050125
8f328ebf33113eec47454ecf4da65765ed32616f
49749 F20110113_AABHZZ grauel_w_Page_026.pro
e237ab70ce2450c96601b75ea7a334b3
b1e7969b6eaaedf7df692c0f95703c964af6f6cc
47013 F20110113_AABIGH grauel_w_Page_032.pro
ac2d2814c28a29581c081bae918b0b49
36daa581b4128c4ba6d01b9bba78f5a2e6ee4c07
49514 F20110113_AABIFT grauel_w_Page_097.jp2
d2ddd9499be128614c43b9d79d54b467
2e8a420a7d6d6e55a2fa8816dc11ebcda5897e70
1967 F20110113_AABJJK grauel_w_Page_025.txt
f656fb0344b9c04b9da57a18b2233669
ba8390dfae9afc11ad0588e2d8aae2d9f6c05268
50886 F20110113_AABJIW grauel_w_Page_110.pro
6274d2164f8870a62f380306b0bc0e09
3589d800f2c2d0bd3cb844c4b8d0f6d4cd4c4f0e
110815 F20110113_AABIGI grauel_w_Page_107.jp2
7d988f0b78bb9c4db6be4cb6fdd6424c
237aa6f3355b692135a8968b73c428259144a720
426 F20110113_AABIFU grauel_w_Page_051.txt
4064741b48588d73db1f297912c75c03
f4f4752c60a5a9d3e3688f1c15563bd0e9256c34
541 F20110113_AABJKA grauel_w_Page_104.txt
35f0bae1a1dc8be16cdf04c7dcfc0759
441d82520de729a5bbc3088e9c043130e9ac9661
2003 F20110113_AABJJL grauel_w_Page_026.txt
0ef690dc7863f1f5cf16c17c15ea0bde
09c85f934345ffb00ed26af6521a4966042c1349
49067 F20110113_AABJIX grauel_w_Page_113.pro
dd4b9bb60a2448b144fc81ce3340ce79
a6513ec70ada469c774821b4814546355bc211f7
96835 F20110113_AABIGJ grauel_w_Page_138.jpg
5b554b0dd2748279967b078de6c90c97
ae87f6a73c7798147046c40c6ffe6ee91219720c
96788 F20110113_AABIFV grauel_w_Page_130.jp2
c671da7434db82d111b14d6356d9f9a7
340c846ac11f03f42f0300d763629d3f2e5a3726
487 F20110113_AABJKB grauel_w_Page_105.txt
07d3a627a28acb3808de9a3f10574942
48b381d959ca698f32a9ff352540e13cbd7a5b85
507 F20110113_AABJJM grauel_w_Page_052.txt
25c50e80bf2a9326ab5028ac95c0e1c3
610e68dcfa5c12d2bb7f5d8023b2bf57d3f58a76
50399 F20110113_AABJIY grauel_w_Page_118.pro
ba85368dc07da6fb4ce2d79ed3e4d4f1
fa120fe1b02dae866bcc94ff8e7ab97d81c5dcfc
32658 F20110113_AABIGK grauel_w_Page_148.pro
e99c9a6d2d2ec531a8f2647f76de21ec
e763ee3aceff7bdd00eea0f3def6f86d88bb414b
654 F20110113_AABIFW grauel_w_Page_054.txt
5e97d1b1de41c28da43865fe0a3bacdb
71a34a4334eeda39f80a930212010822e7c3f071
1830 F20110113_AABJKC grauel_w_Page_106.txt
288b5b9db46b901aa23c396e36524bba
3188b764235e452559e9a5d4dad7cbe2d97d5b9c
F20110113_AABJJN grauel_w_Page_053.txt
0aaf7bdd6200d5601d2b2e4659854471
3b6dbd4f47b6f7680590cb468c4e44cbb33cc469
23217 F20110113_AABJIZ grauel_w_Page_125.pro
273b109c702ff3ddab58c019a39b3779
b84c7da94331880e8cb354fe81cd6d3ffad8222a
8416 F20110113_AABIGL grauel_w_Page_037thm.jpg
e9ff1587f59bb8e8d2c517e126bcdc21
d55972f5a409b9208103d406a0522fe8d373212c
7847 F20110113_AABIFX grauel_w_Page_023thm.jpg
b74623a5bd73ac8d31c036e68414bee9
2da726fc00a8d258c599f03c476854c76db86405
7975 F20110113_AABIHA grauel_w_Page_042thm.jpg
e369735bc3166feb4e613a499305f013
3eb3c8a534394bd4433b9eb7a85db3ea85e0aa1a
1883 F20110113_AABJKD grauel_w_Page_114.txt
b5edf246e6e8ecf39ef8a74cbf537b91
71bd6807f03ded67c27deeee08abfd257db53c39
1889 F20110113_AABJJO grauel_w_Page_059.txt
80cb22e0ff304d9891f1fb4dbfd3d4b7
6b90b4a3d4efbdb6ddb56b87f57db2dbbd3d0649
91149 F20110113_AABIGM grauel_w_Page_024.jpg
e1a24bd05ade928dbd999f4ee3ba1d55
dd6510fbe8f5c29dfd8d4cad3488b6cdf07c7e04
8532 F20110113_AABIFY grauel_w_Page_140thm.jpg
0f74d896d00229023c0639efd89bc54a
c2dd6ff29a656358f1312ac0415187851f6f796c
104418 F20110113_AABIHB grauel_w_Page_132.jpg
1a8c8988988d5960745b8e918fdab3e2
77c993442a0e86457f98f393be4a1c47240dccff
1964 F20110113_AABJKE grauel_w_Page_116.txt
a2f9b4a0c28a369c57a7252d74250722
308e17c75c496b4878a462f1232debe23fc68686
1966 F20110113_AABJJP grauel_w_Page_071.txt
c1f59cf00b6eb249441c58e54c111730
51d2f6dea6f2f667ae6d348e806a0722d57ff641
F20110113_AABIGN grauel_w_Page_113.tif
1746c4b3e33bd2d3af1c3773a2270dc2
bc6f399a110fc6f082c1d49f7a0af54b3c9e39b9
94091 F20110113_AABIFZ grauel_w_Page_114.jpg
862be089a547918f899a76749e3ed5c4
d8e2583928d52a519cbb80eff7ab397c6ae4e348
8426 F20110113_AABIHC grauel_w_Page_062thm.jpg
6431c3d5c031c818dccf5474c4c623e7
a5a5c6e27b6a86b8f258b7adb82ba12171e33fb2
1976 F20110113_AABJKF grauel_w_Page_122.txt
a1e58ae439531b74ace973ab105bbd98
1f00b95c44af210fdac81ed921963522b41ee5d7
1040 F20110113_AABJJQ grauel_w_Page_074.txt
d7704c216aeb587218226926527aa063
d364b175239015665fb778f11773952cf34766af
F20110113_AABIGO grauel_w_Page_121.tif
af1c7d7a709e5d020bc2a386704cd933
e4d5fe8f6b10d234bee28284467cdcebc5f01faa
25975 F20110113_AABIHD grauel_w_Page_001.jpg
da8c511aada1a9b6876c8d9d7b13a121
23f64acaf986344fd8145ff58affeec9076be6ea
1970 F20110113_AABJKG grauel_w_Page_137.txt
179aaf2f998e9a316fb4c945ef30817d
d6b90f2262614f41d34bceb52c92ae1f57fc9752
1979 F20110113_AABJJR grauel_w_Page_080.txt
94418593dba5f5a9e259fb1de0391368
bab8f32e3615669f65ed716ea8c4a881a53d5b18
1983 F20110113_AABIGP grauel_w_Page_031.txt
284b2b3f7294d99ff54a565f9f404604
74576703dae352d9a281e2cc180d328f9cc4936f
2016 F20110113_AABIHE grauel_w_Page_120.txt
5f3c62a3918605377f0b12134563e7a1
98bf2134be52a704d20d26d49f2573956693cbc3
1874 F20110113_AABJKH grauel_w_Page_138.txt
67dbd534af17e2e0b8f5da135c6accdf
db76559f08646fb5916e599cc9d7756b4d45018b
1780 F20110113_AABJJS grauel_w_Page_083.txt
ac09a67833c26e81fe3ddb559798347e
6d19b5bd48567ca7cbb8fe5784c1f0ac496fb59f
107992 F20110113_AABIGQ grauel_w_Page_081.jp2
7948c718347c4f357e364f2983a389b9
a6948b7dd2bd33aae60c15ac1b84d85490be9ec1
51410 F20110113_AABIHF grauel_w_Page_120.pro
7cc87bbe5c4fb6c59f72aaec35a7ef4d
eb3dc73603dd851265387c4bf9a271ae3f2ebe86
1402 F20110113_AABJKI grauel_w_Page_148.txt
b628943c103a37905c7df4c9cd6ccf09
20f560b743bffedd1c7ae2c4fa4d14cafaaddff5
2007 F20110113_AABJJT grauel_w_Page_088.txt
35787cfae852a01f73e686d6d9b534c8
48348e5af108ecdf6ecb6815505fbe989f37499c
2021 F20110113_AABIGR grauel_w_Page_063.txt
da51664d365159f77d5750134582430d
882d82dc0de50d1e7d0c2423ab5a32ab49b02748
5268 F20110113_AABIHG grauel_w_Page_050thm.jpg
3f11177de696d6b0434fc08e0be30231
2ca992ec7430585e278c0b66da29ffb5ae9a660b
1999 F20110113_AABJKJ grauel_w_Page_151.txt
225f6c5d5193bf2a849eaa889780d119
1395b68fcafeb8dd1585a10f44725feffc0fd924
1925 F20110113_AABJJU grauel_w_Page_091.txt
48c4684d79da0ec77eb0596cfb0f6e2c
7a31369ef8625f68dad30f251336e7872389c573
24852 F20110113_AABIGS grauel_w_Page_015.QC.jpg
03526a710c18acc9844d9c4e13727cfa
1b46d72b0fbaebbc784e2f2119352a73009b469c
103224 F20110113_AABIHH grauel_w_Page_038.jp2
99dabc64dadae83605c65500b222b62a
c5f01f6dd8815a4d2741eacf73728c5b717e198c
498 F20110113_AABJJV grauel_w_Page_092.txt
80080dab96bbcbe5094f6c96b738c108
9adaf0db9fd79c40b74cf6edbbd02724f8366c10
944 F20110113_AABIGT grauel_w_Page_003thm.jpg
e9d3f01c4917d14e454c619451d3be80
325bc48f57a42562bbaa0b94c3e2b0800467fb34
2073 F20110113_AABJKK grauel_w_Page_152.txt
acb34e8d909ebf7378314f0a47c1c952
7b126ae293c6cf647d939761e3f06bd473621de6
1351 F20110113_AABJJW grauel_w_Page_098.txt
b8bc22ec4066ce77cc73f01bdf05c603
4f05ca6016d5b4c8f0d7ffbe1784ee82f7f2ccfa
1745 F20110113_AABIGU grauel_w_Page_069.txt
a1f485021621b504690b7c51454724e8
f45833d7682cd362f264fa12a01cf5b74acb70c8
F20110113_AABIHI grauel_w_Page_075.tif
54e0f466cf47adbad57c80921d4a1771
be3c580864d60a334729d1ddec2701134f7abdac
32773 F20110113_AABJLA grauel_w_Page_107.QC.jpg
9805f8b9f299c9a98d633b97896f9a18
0d109810200bf0a38736a6c10099fae12330b2b8
2412 F20110113_AABJKL grauel_w_Page_158.txt
8e74ae4f5d5c60186d8857a2c31d1850
c18e3ab038bedb0dc43a123b56f83c465546a471
1848 F20110113_AABJJX grauel_w_Page_099.txt
9fbf8f5748d29ae5b60cf291ad0bc950
68e769fad3725ba5af72bf49d747b36c1d74f49e
F20110113_AABIGV grauel_w_Page_170thm.jpg
e963532fab7ce0a666d182831597c551
ee90fe07a90a2f8b723ec49c360639d840dc3adc
106741 F20110113_AABIHJ grauel_w_Page_045.jp2
93b09f84e89b9d00015b0284d5b2d80a
a5be953c063dc71cf4a15be3a41a0fced3e6c765
8526 F20110113_AABJLB grauel_w_Page_135thm.jpg
f10d10fd2353e45d97ef59723572b56b
d4c2bd6600abab141aff6d9d589e8de128e49f5f
2483 F20110113_AABJKM grauel_w_Page_160.txt
11d4803fa6f60673a8e8969e7fbba74a
92dcdb4f6548c37e811d24be70e2bf1d0eab3ea3
1673 F20110113_AABJJY grauel_w_Page_101.txt
444d4148e2080fde83ac72312e3ff7ef
43dd2aaf087f65917d7791da190bba5c2c956d64
3276 F20110113_AABIGW grauel_w_Page_033thm.jpg
82dab108521225b47b7d7377bd4f3529
49271e590e7fea12df8a89d17862bc46d31d54c4
8424 F20110113_AABIHK grauel_w_Page_118thm.jpg
d35eeee34cff17b935bf076fb9b45a7b
c87e279ec51db95471d92c6dae8d324baa560b88
7971 F20110113_AABJLC grauel_w_Page_085thm.jpg
50efcb15c34d97612e5f4f46b5642176
49d9159c06905cb7203299cbbf01ce521dc77536
2605 F20110113_AABJKN grauel_w_Page_161.txt
f2456036aa6709eb5ca9949af91a34f8
41d1a41e6bb9b01408f37709b7869ae830acb6ab
F20110113_AABJJZ grauel_w_Page_102.txt
032dac92ba10fad35ba89209c42a793c
72444354ed310a0c4527b96ef9403b66a3953d5c
51873 F20110113_AABIGX grauel_w_Page_134.pro
9f767cb1a4d1442149a2fe5259563624
9d754d9958ccf460c58be8d78002ab9a9e18f445
25426 F20110113_AABIIA grauel_w_Page_098.pro
aa193fd9e4841493ff8fd1eb5f90bd79
423d8344cbe6d5493f9f65fd0b7c1e28fa6faa1b
110702 F20110113_AABIHL grauel_w_Page_043.jp2
4f851b99d383b700a94b2b26448eb605
c76ae7148cf0856957a71232f8314ab7e6e510e1
7755 F20110113_AABJLD grauel_w_Page_138thm.jpg
52da7551bbc545aa3af20fe6fab3c3a1
3bc24d5462565abc4023dadf388a1b3964e6b919
2495 F20110113_AABJKO grauel_w_Page_171.txt
9c9aaa285576b613043c54dc048a251f
11f4e93fa13c39596d8afc43da0996992f3a1e15
F20110113_AABIGY grauel_w_Page_063.tif
03e8c1bec8f1cde4e8a46e85a6de9ac5
df81c0023946e3c91b80440f0d24153bf7a7bb0d
94829 F20110113_AABIIB grauel_w_Page_042.jpg
16bc74d363d2488f0f7bc26a6a2c38cb
a498a70dcb6675f49e72d5f71703298a4b92279c
101789 F20110113_AABIHM grauel_w_Page_079.jpg
836d05ab6fe840980dfd3cb2a9672bfe
4476fbe6199604060190ecec17fcb1badf4b3a76
29315 F20110113_AABJLE grauel_w_Page_150.QC.jpg
310a4c363d246546af7e7cd10cf5225c
24546e001dd2967e6bdb950a036e5c250b25e867
2241 F20110113_AABJKP grauel_w_Page_172.txt
5c20b4aad4fdd465e8e0ae2900f8cc79
2a83bc0171c5ba383895e503e4aa26b43da1cc1d
36839 F20110113_AABIGZ grauel_w_Page_078.pro
5caf5e077d5a0da7fd0411e5eb99393f
4ce2da3e3296232c23d1dd96c38536fec08b847f
7912 F20110113_AABIIC grauel_w_Page_127.pro
24be54b93a852e358ab99774b7097762
7151b9f85483fefa1b37618c5823bf7da3c23b53
29402 F20110113_AABIHN grauel_w_Page_104.jpg
e20d74cfbdeee05d0ff10da2330d4def
b3b7fc66f6ff962920d711d5c41723e195e0ae27
8291 F20110113_AABJLF grauel_w_Page_081thm.jpg
8b0082759b5812f0d73eb6758148d67f
c03d5736482ac6e95b13da2ef7a0453e311faa02
2624 F20110113_AABJKQ grauel_w_Page_174.txt
25fc982c6bcf8ccf5c4e41948eedf07d
9a61e391cfa4996f385837550814e813a303eb38
14750 F20110113_AABIID grauel_w_Page_126.QC.jpg
5a19c37e250902cd1aa38e733081a2b7
3329baa38fa801c7a7cc68e18f18d966e0c5cb18
9687 F20110113_AABIHO grauel_w_Page_003.jp2
2682e02a53adb02a43086cd6fafdb4ba
145636c8213b6cb7d121c30e6775c94f1caa4cc2
32953 F20110113_AABJLG grauel_w_Page_152.QC.jpg
6cc7de349fce666db68809fb31cd9653
69e7246f6b01e037e562fff83ad05319a366c48e
8237 F20110113_AABJKR grauel_w_Page_077thm.jpg
29002a57c78cb9250a51e7ceb35f1e27
8157883bfa2c9e2b28da40fa5d0eb90ab839a19f
345742 F20110113_AABIIE grauel_w_Page_050.jp2
ac875b6c27bf38f606569f37ff473dac
f2b215ca95afcf1c876ae64fb369e30ed7a15343
27120 F20110113_AABIHP grauel_w_Page_154.QC.jpg
afb20c8ce628590965f5a8965a15ff1e
13482c501d3c6c2c74fbdd511c61e825ab85a0b3
8057 F20110113_AABJLH grauel_w_Page_086thm.jpg
2f2cf41b2e661a785db9e74c1336717c
60d2639e6c454f587aafa9e4671385c58176a6a5
32864 F20110113_AABJKS grauel_w_Page_158.QC.jpg
f6d3955414a8dfd384220859ccaa328c
74b692ac1ad8cfa2e06f4880439f4983eff142f8
47280 F20110113_AABIIF grauel_w_Page_125.jpg
50fa9d57370a3550a1fe04dc66afc3e9
0ee48f694a9d7e3fb1dc51a178da9173620fd6ca
353 F20110113_AABIHQ grauel_w_Page_127.txt
57bb13091209c1c638d57db187660dd5
b82ca79b7f253de6af38278c8dbda231f899d660
3829 F20110113_AABJLI grauel_w_Page_125thm.jpg
85a700f2216d20f3c566d0a3119977cf
78e6c27e1046e1ba46ca4d52923fa016ca91995a
32834 F20110113_AABJKT grauel_w_Page_031.QC.jpg
2a58de6027b0b24238fd3c5c50736713
8e1e3a717a1748d94d51debcc9486d6693efe558
66580 F20110113_AABIIG grauel_w_Page_169.pro
962d03dc0a5a97d25c0072c923cb5376
3b666327b5fccc4e2abb35fbc4308957f3c7a698
30224 F20110113_AABIHR grauel_w_Page_095.pro
c6ed8cf8e10e8b42b2ed335ae4ffaa30
5cedffc7c7d0dc75357c8dacdfef413904639a39
24740 F20110113_AABJLJ grauel_w_Page_096.QC.jpg
9b35312541bf87073967a4e1018d5406
b6ed9420a7ff4443b1b03749d0b59b069c651ccc
31210 F20110113_AABJKU grauel_w_Page_036.QC.jpg
d70969e154706d0843a190f533e3417c
2e1a4e32c37f10886d97d37bd14773ef4265bd5f
29474 F20110113_AABIIH grauel_w_Page_128.jp2
ae7490f439710ef35cd75b36d818481f
f2aff9d0c87d41a914b0e3823fa0367808e12863
102277 F20110113_AABIHS grauel_w_Page_137.jpg
33adac05baaa05bb7247327ad393984e
4e55545943db3d0b691199c94fad45980f45dc48
10113 F20110113_AABJLK grauel_w_Page_012.QC.jpg
e722e6de363a36ed9db8822e3e6b49bd
da52cc6b5c2eb2d6b4082c0466aacea3f95d6c09
29351 F20110113_AABJKV grauel_w_Page_106.QC.jpg
9abeb48f985db11573b7655e909a9921
b1d83be26cc10d62ab3c9df3533aefb0f041b73f
65043 F20110113_AABIII grauel_w_Page_174.pro
b7b15f1bc469319dc8851c17935f4e04
d1062a12b7ffee2c814e906a8024cd19e8583415
6931 F20110113_AABIHT grauel_w_Page_153thm.jpg
93844b6406efdc172d29096e57a8bb73
988a95e8f382fb59279cbb4a92686c5d3ce06455
7918 F20110113_AABJKW grauel_w_Page_044thm.jpg
4e4ddba3710210e30d4b7c4edaac05e4
0abaa2e500f397daf73c9c17d4b5c3bf2bc67f40
98577 F20110113_AABIHU grauel_w_Page_086.jpg
babd0f70db4227fb12507bbffd284d24
32ffd9d5110d07b10f8b03888fbf61f90fd9dac2
33095 F20110113_AABJMA grauel_w_Page_080.QC.jpg
c3ac2dd66b403c237da3cf64a403b5ef
73af54eaf8d3dba8ca8153a3b92478b0c43843f5
6364 F20110113_AABJLL grauel_w_Page_048thm.jpg
e2a56d14e40337d22ee099bea01ed13e
3bfac9c0f29f9cde84ea8f707e31d8f682fe8de5
30037 F20110113_AABJKX grauel_w_Page_025.QC.jpg
6851296ab9595050dccd3c3ffd196919
7f115592f5ff30e6687e2e2f44cf73aa01c26da8
2050 F20110113_AABIIJ grauel_w_Page_046.txt
8aebf50758d0a3ee311b56ec786d8a36
65db49403fbf84271620600903049dd5fd1fa2b5
10044 F20110113_AABIHV grauel_w_Page_128.QC.jpg
19b9cb5ca19ce4487bb9304749a428e2
4a97687576a13eda8ddaa06feaa1a75c41bde185
29687 F20110113_AABJMB grauel_w_Page_004.QC.jpg
569ff3dff06639222b5c7dd4dfc1003d
0edb544797622c3af296e7180c5d07dd8ecd2311
4684 F20110113_AABJLM grauel_w_Page_147.QC.jpg
d9b7f31d377825c251ef0e47d27b48bd
0f782c6f5d9d8e4bc0ec1690e0d09bd0f164e9aa
7560 F20110113_AABJKY grauel_w_Page_034thm.jpg
d2a71b3529dd9f18af198726771e4904
0d2027d18f5d745234859f596c9a77275c4bd5ce
47970 F20110113_AABIIK grauel_w_Page_090.pro
b7b96d902e43eb6c3dad1857132a99f0
ddff37da62116137081a66b590ec649be1af5e56
97085 F20110113_AABIHW grauel_w_Page_022.jpg
89444df31533b0888ce5a8944d5738af
f8b3cc13cfb36e0571c3389aa0d14d4a9c0c6a98
3201 F20110113_AABJMC grauel_w_Page_104thm.jpg
022c9fa159573f42067dbf8608ffb9a8
f8117fe28a8b0fd39d3cbc1a6f63bd220a9c5792
637 F20110113_AABJLN grauel_w_Page_002thm.jpg
ec24cd5b8da0038fb329c9999f32a410
978d3472fc009a4b66bdbf953f977bfad6055e54
33632 F20110113_AABJKZ grauel_w_Page_063.QC.jpg
6918854343cc08f5979a965a7cb81fd6
fb950e780ffef9b9728b94cb07568cc499f974eb
32253 F20110113_AABIJA grauel_w_Page_113.QC.jpg
37ed3008946f419c1a8d7870aeeb839f
a05c5b158500b78617d2996f97cab95a86d151aa
106337 F20110113_AABIIL grauel_w_Page_117.jp2
2bed27d0f4698ea7cbe7c4e84a9d4c02
484064b4c6635c2694c83b6d0d78eeb6dcdfa1cc
34101 F20110113_AABIHX grauel_w_Page_166.QC.jpg
af9ea19b4d33879e8bbde13a93790beb
aa15fd884bab5e5f171c87418259442bb2b18e13
2974 F20110113_AABJMD grauel_w_Page_052thm.jpg
f9f43ed8400a0f42355c167c1ade5f29
ce82201cbb6e7618c2621c8c1374953d4d24d13c
31422 F20110113_AABJLO grauel_w_Page_039.QC.jpg
caa8a3e90b507a194fb6935b2f6eb106
e0bf01026699b0d42d5ac17310f5f7de6a1244dd
108858 F20110113_AABIJB grauel_w_Page_029.jp2
e41587af30b45938b0c3e9d88c1ea76b
1ca419a3bcc1c99393df84db1c03a7e95bbc372d
1887 F20110113_AABIIM grauel_w_Page_042.txt
6a33faa90945082f600a056333b7ba9a
e766e2a1a81e18fd41b9c71d12c7fceae209b52a
16520 F20110113_AABIHY grauel_w_Page_100.QC.jpg
ed2a118f6470d98fc71fc0138b579625
c1b93ff00eddfa35af2a7156f86b44a1a6f5852c
7226 F20110113_AABJME grauel_w_Page_130thm.jpg
14e0393a805d11cc5df5b8488c6d8eec
cdae2e839a01e5c858aaa3c4ec3f41cd0453b596
7462 F20110113_AABJLP grauel_w_Page_150thm.jpg
c370e028a7095870a2306e12604a807e
b7370803fe622de617a3d66b182a59a6b9cdb0b0
91680 F20110113_AABIJC grauel_w_Page_055.jpg
e6b03dfee79b7fb11b0ebcd20340eb5c
351c3075ae9c030fa91baadf3dc82b9c6e393b63
2069 F20110113_AABIIN grauel_w_Page_134.txt
4dae796bbcbde2c60f10566b3b57b6d7
10187d6178a6e10e69529a8a88ab2870d5a5427b
792 F20110113_AABIHZ grauel_w_Page_050.txt
83c5d233921d1b7bb36c219cb16a0c7d
0276f0cbb314a7a7547e47186660aa302419c08e
7622 F20110113_AABJMF grauel_w_Page_177.QC.jpg
284a82f4abdffd31782943f57178086e
5bb71d82fad150cf5b7361a19109503f2fbdec9e
7511 F20110113_AABJLQ grauel_w_Page_084thm.jpg
a4ea12ad1900ea24696c7d11a8caf6fe
fcf479a311cd7ba9ad7b6e106a93345ffc4d842e
110098 F20110113_AABIJD grauel_w_Page_135.jp2
f95f119c94f95ee3226b3eed8284309f
667de4f0473ca8cf7b2073cd78278aaaaf81c6b8
2025 F20110113_AABIIO grauel_w_Page_142.txt
c5abe8589da4d370a73fefe177c70b36
9f9516fbc71f7cf81bb9d6d42893196f9813dc94
8649 F20110113_AABJMG grauel_w_Page_040thm.jpg
cbdb18a19d9e0d8463006d2007f88396
c28c370248b75276db42c431d5c95593ca352a66
33602 F20110113_AABJLR grauel_w_Page_035.QC.jpg
5f2203c7674f096a10b5e6f1981d368c
1da90948195cad1a95bd03e210dc5698b4a9252d
2484 F20110113_AABIJE grauel_w_Page_166.txt
02048db580c83350e9c675950fb636af
06b28a655993d5500d3156cf2616a36057c354e8
31415 F20110113_AABIIP grauel_w_Page_081.QC.jpg
c9ebf66cc6df2dee00a5e7c1ef46104b
85767c594840cf6660c5e08fbe7f698da29e9a79
16212 F20110113_AABJMH grauel_w_Page_067.QC.jpg
bc2bc05bbea85cbe3935007b338ca7a3
2d6b21a02178f2af94d5aeaf916d0c6051d0651e
7866 F20110113_AABJLS grauel_w_Page_009thm.jpg
93353018f47c319990249683990996e5
80d73e12217b45585d72706387848e190c52c276
8118 F20110113_AABIJF grauel_w_Page_080thm.jpg
3b649ffcabf3415be9faf2ab13d298a7
cc3a076801f191addf7f17767242e8d48d0bed26
F20110113_AABIIQ grauel_w_Page_027.tif
3608cf533d2fa78f51b6cb4202d057e8
f1492f2444d0db551602123956d7131e76278789
35988 F20110113_AABJMI grauel_w_Page_170.QC.jpg
2e69f77bb533c96d11fdbae03feb5145
ee7cef685389a52270926af77df9d2bdd2573363
13370 F20110113_AABJLT grauel_w_Page_065.QC.jpg
5c81c382a2b43f1bc5384a28a0ec14ab
29bad8cf2e6158362f85db4dfdb90acc0fd41586
2754 F20110113_AABIJG grauel_w_Page_054thm.jpg
bf57a0832544fd53c741fab2232883dd
035dc6839894986a778c3f5085af0bac98d94482
109245 F20110113_AABIIR grauel_w_Page_118.jp2
36604cea924397ac7fbb61a8880463e5
b5780f99092663aef0d3ab86a8c4b4744981ea7e
33130 F20110113_AABJMJ grauel_w_Page_046.QC.jpg
aa2a9b5f99562f2b453f9d6ebeadf3fd
47c4b66eecc40d9bb10663ffcaa8e0a716e4ff5d
8812 F20110113_AABJLU grauel_w_Page_164thm.jpg
50f2a475bc60351a18ac017692893462
47884b44b75b632a0724076924d0ca8a3d13d500
33939 F20110113_AABIJH grauel_w_Page_173.QC.jpg
d425faf4cc69d196690bccc52a41d820
b022c674354a5abab1447dc72cb208b956b6e36d
F20110113_AABIIS grauel_w_Page_037.tif
c98abcc2149fb91b1e2b074d4cec6cfe
8ec5719ff8d4473ce6423fe2d30cd62d94d48885
8305 F20110113_AABJMK grauel_w_Page_057thm.jpg
7562f4da4a24b105e0817674904fa6d3
a6bc8cdbe1af8a0c789ba3c9fdcbd229f628b0fa
8578 F20110113_AABJLV grauel_w_Page_035thm.jpg
71ddb04a928a6e2fae31240d1d6bcc3a
eeae59dbd391bdfd9ebf4892e087b1adbd12afb0
1051969 F20110113_AABIJI grauel_w_Page_006.jp2
bee5176c1b5ca38e66ea5f9f4e55e202
15b7e7bd80e299756ed357251e88c6378653d216
51830 F20110113_AABIIT grauel_w_Page_058.pro
2df533123a79685dab1464b6d089c89f
632aa799de96967b561bc3fc4a58c49432e51a03
8278 F20110113_AABJML grauel_w_Page_119thm.jpg
2119fabdef44fbd4e6a5b2cbde263dff
756e4d6db4c446b565bd47b447381f74032f92bb
25139 F20110113_AABJLW grauel_w_Page_102.QC.jpg
51c473446dadf8f16a2c817ad603a74a
a2d5f0149b2123ed48eb5bd850a73e0e2f3ebb88
33954 F20110113_AABIJJ grauel_w_Page_157.QC.jpg
feb20fee6431c6e68a3421608f3131d5
c23e12f71ae0365c7860f2dc6489a8403431e1d0
7503 F20110113_AABIIU grauel_w_Page_156thm.jpg
c7d3b8ec597f9804a7f3276d99e3279e
278d124061f22c866ebeab19ed46451581d9910f
29329 F20110113_AABJNA grauel_w_Page_072.QC.jpg
6cc6ba2dd66a759cb90c8102ae014f00
f72b60cc450febb5729efba606611ce23bb80d5e
1924 F20110113_AABJLX grauel_w_Page_177thm.jpg
3b858bfdb463123a637a345d8ae48b08
43dbef868567e4ea4cd96345baf7249693862d70
F20110113_AABIIV grauel_w_Page_074.tif
2d3222bd02325d5e630306c1bf24c067
9652f3c616b14d31360886454e68ec22059fcf0c
23012 F20110113_AABJNB grauel_w_Page_093.QC.jpg
70920c2f3db1d459932dcd190991bcb1
f0ccb341a42f2db8c20734e73c0c3a39e4d15cb8
8178 F20110113_AABJMM grauel_w_Page_139thm.jpg
4d34c0f2259cfef915dc74e2a955eea1
88555b95ef570269990ea2dfa0aed6dc567f575a
8254 F20110113_AABJLY grauel_w_Page_053.QC.jpg
673d0cd02e779694fc22a29d6c128a2e
92ec8a123f0e242021904d417fc6addea6087795
8049 F20110113_AABIJK grauel_w_Page_028thm.jpg
b8cf726e7b92c0027dfad4a3aae83b81
eb73ee794066301bb77b684b5999063343147604
F20110113_AABIIW grauel_w_Page_153.tif
512437012633d33026595de9d894e68c
970c7e8e19ded15212b1c6452a8814b9952cce3c
8413 F20110113_AABJNC grauel_w_Page_018thm.jpg
994fa2764dfbb9999beb84673525997b
6178519278fc3c699c1914d49f9639106b4fd523
31622 F20110113_AABJMN grauel_w_Page_146.QC.jpg
bebcb9e40a71ec9a735cfd6f92558405
1afc4aa4c08999c931ba95c76387a873ec433e3d
7222 F20110113_AABJLZ grauel_w_Page_076thm.jpg
fd0ceba7302e9f58c56416647d48f82e
7b8eb1eb5f53f4d54007f62a4c3dab52dc6ec631
1781 F20110113_AABIJL grauel_w_Page_072.txt
39528766014dafd94249b5ff863d6a4f
2f63c50023de9a9e7b4eaf06774fbc7a2a309f21
3510 F20110113_AABIIX grauel_w_Page_005thm.jpg
c0411c4cdf0f1f0ef1a4f48d5fb9a46d
b065a2bd5831c6bddb10170d4404037ba391c0fc
33078 F20110113_AABIKA grauel_w_Page_118.QC.jpg
4772a50049478a9e6017317c868bf2c1
e4ab4bb23ec6dc0c46606bc6740a4e222b165b20
18502 F20110113_AABJND grauel_w_Page_098.QC.jpg
7f88e38d01c2fa381234aafbe4596032
1555d35703d2b9477bcdfa5971d85f9cd86b7d1c
6317 F20110113_AABJMO grauel_w_Page_148thm.jpg
88931a26683329b5a1a1d4e02470a0e4
a9ec7e4950b4cb2fd45c66b594085fe66a54e894
19238 F20110113_AABIJM grauel_w_Page_155.jpg
b46de755660c0d085f9f8048453e8180
60f582112e5909c4cebd9772325f2118bf439612
1888 F20110113_AABIIY grauel_w_Page_145.txt
b1fc57f7f7bf4fc4eb2f56dd240e78c4
053a2bef41eff874b600d9ddc618af20d7e76c21
51192 F20110113_AABIKB grauel_w_Page_063.pro
9c820ca3ef5b8ff36c1cc62deacbad55
6d7933ff762bb27c40843e5c49f856f5bc8ff453
31973 F20110113_AABJNE grauel_w_Page_047.QC.jpg
aa31675f90433d1bba9634575b368a14
bdb5fe6aeb31fc948c13ba7a91bdd79dc631f562
8000 F20110113_AABJMP grauel_w_Page_019thm.jpg
bfe29db13072db456a38c8d8dd6ff48f
5cff88fe5030469d89e8131a315a5f052551a8b1
2448 F20110113_AABIJN grauel_w_Page_157.txt
b9c26b8c50075fafdd991e8bcd80d668
b2f76169856ca604c24ceed9a6112d87e5a40869
48298 F20110113_AABIIZ grauel_w_Page_086.pro
877ce85c4dde2db1c4bafb994565a4e2
4400411eda804e143cfb1639462dcda2454510b3
15425 F20110113_AABIKC grauel_w_Page_127.QC.jpg
854798d3ef8f5b58354ebd77528fc759
eae0ee6469a49f61e5b2c3df42ac2c952477314a
8408 F20110113_AABJNF grauel_w_Page_157thm.jpg
ac14284c7eb1a974441d94d8d6dfc75a
04c2d0230bf6fd970eaa657d27e69aac80814024
8170 F20110113_AABJMQ grauel_w_Page_036thm.jpg
949d59b180235a843388610e5383cf22
12f9b0825c5ed114c724deacc1f395d126679cb5
F20110113_AABIJO grauel_w_Page_013.tif
50fd7b6cc388e16a6d873b24a2507908
679724ab60c46aac229d8e650a71e563642ab7af
8397 F20110113_AABIKD grauel_w_Page_063thm.jpg
b3fe78fb30fec26c298673690db2b2db
3d7dc586c0f3a9da83dea02b7cc12540d4eaf11f
6645 F20110113_AABJNG grauel_w_Page_051.QC.jpg
08aae30dc8a5a53aa5cdbecdc07a98f5
3ce2f734f1602632bf414fef1e0d53a08bd3b7fc
36513 F20110113_AABJMR grauel_w_Page_168.QC.jpg
5a0001cf7b7b3aeff0753cd5524aee4b
95b4e9fa812adfddea7cfd3ba4dadd4bf78eddd2
2058 F20110113_AABIJP grauel_w_Page_144.txt
ab1953a41d9cf148f0c69be4409a47e7
edb0bcc4024150d39b0336a456fd1bb1e8779c6f
8063 F20110113_AABIKE grauel_w_Page_090thm.jpg
74a109c53a2fd5d5a2b076752f76b506
358eb71e0715ffa853b9a8f2fb6a412928bca1a9
265645 F20110113_AABJNH UFE0004366_00001.xml FULL
39f88c6ea3e34cfd94b93b7d9062edeb
be6a4cc26d5b96e3d5bd93570999ac74f710fd04
7653 F20110113_AABJMS grauel_w_Page_032thm.jpg
6cff46ff2b6db1026d2ada2a3cf31607
9685559c2caf47573be8f92fb3f0ec88bb5a251d
51078 F20110113_AABIJQ grauel_w_Page_144.pro
859a04f26127f0463ed03f047e1cef19
f3ad9d29240c4da9e108676808fb90b6ac5cc78b
F20110113_AABIKF grauel_w_Page_140.tif
be42f80a0236a05ed40409139defd07d
423013f780fe915c15ebcd1eaf300c9fd1f6d060
2552 F20110113_AABJNI grauel_w_Page_053thm.jpg
abb6cecbfcdfdef03486239ce7f1caeb
31154ba93d1865de84f95a49843c62d119230183
34028 F20110113_AABJMT grauel_w_Page_078.QC.jpg
2a9d85a0dccbd1eb392484acd84dbef4
fca1dbe1b3b42a3970df38bd44cebc31112af7cc
8702 F20110113_AABIJR grauel_w_Page_092.QC.jpg
736bccabd67bdd988961d7c63a382e6e
e73a838c37a2a9a3829ffd6a05e407108fd0bc3f
F20110113_AABIKG grauel_w_Page_121.txt
15f311241c3026923ce803bd7a0f3b99
64ebb59e365d7d6a118bbf500ce8a1bbfe877634
3577 F20110113_AABJNJ grauel_w_Page_065thm.jpg
894115082e2b8d504e57eeee9f8cb239
63a54ff725e5d83be11845ad41f00479a40351b7
32068 F20110113_AABJMU grauel_w_Page_045.QC.jpg
2c8231d4f7371249db8262a63e761c60
7b13bea54f5481c949d18f96f2f9b3fabb1b2f75
1911 F20110113_AABIJS grauel_w_Page_028.txt
17cdfb2225b4bbfd2bc5f687775d745c
71dea826ef5ef068885f2c789a180dbfefae87b4
67080 F20110113_AABIKH grauel_w_Page_066.jpg
8f512ba1afacd31d6b14692fc17439df
7a469ebaf8cee975b52deee9bc384d6cfee7c5a1
8219 F20110113_AABJNK grauel_w_Page_088thm.jpg
57d5dc0b8a21ad42b85226fd51095fa5
4d329c0765417a43071d9c1a8b6dac4e9847062f
8500 F20110113_AABJMV grauel_w_Page_173thm.jpg
6fa6868c75638c29837818135f2f057a
b19c44915415232c6a95caa5b7432218c8593a03
2132 F20110113_AABIJT grauel_w_Page_020.txt
07da19c49ff1ec01e8aef11ff3e23f69
3aff5cd7b66fd3641c7d605bd5c4adb0ee92b3ee
4846 F20110113_AABIKI grauel_w_Page_127thm.jpg
843464f1602391af34470900061df183
661c5f294e445e2a809de138a7afa06ca6d2de7b
33033 F20110113_AABJNL grauel_w_Page_089.QC.jpg
1668560b64d4de0064abcfedfb3350dd
8e6fa4525998e17474ba4620b42fcc98276a683f
33069 F20110113_AABJMW grauel_w_Page_020.QC.jpg
1c34a333777c92d363c1c5f70e2d2077
da46e876596e558bd5f7511747830de8c50a003a
2008 F20110113_AABIKJ grauel_w_Page_073.txt
5aa7b7eeaaf4afbbcb57ab5ade299dae
322f40dc259f6dbcf10113ca1d45810ffb6410a7
50471 F20110113_AABIJU grauel_w_Page_136.pro
c75638767f5fba138b82fca2ecceda24
b1c830ca0e8c6a5f32ea6808ba32b2666d3ad8f1
31767 F20110113_AABJNM grauel_w_Page_151.QC.jpg
802c1d92b6f12a9e29a5e069b4421bbc
389f1cad6a569e4445f50b97b6a51bd0a0f1caa5
9293 F20110113_AABJMX grauel_w_Page_074thm.jpg
5cf41661cd273abb09590b2698d5bb59
339854ea783910393340a8dd5ece4b35a24b8faf
59594 F20110113_AABIKK grauel_w_Page_158.pro
aaf94cc505caa966dc04f39064ee4609
c8f3027c8dae9ee820ebaf34a5279d21f4c722cd
110169 F20110113_AABIJV grauel_w_Page_136.jp2
98daa31041f608d823213b87474d7b84
7006bbaaf6618d0ea1c99b844f0593d32846468a
21223 F20110113_AABJMY grauel_w_Page_010.QC.jpg
581a8c44093a5441ca7779c260699b4a
7c33ec0645efb9d1fb3e1d645ee19bb9e0096d3b
105776 F20110113_AABIJW grauel_w_Page_138.jp2
9220d8b0cada525313b3929b495d63f3
d092a5e756fcd6b5c55a1b842292d4147a439af2
3822 F20110113_AABJNN grauel_w_Page_178thm.jpg
a9deb4ae60fcf3add7197307f5d3eac1
ba5e5945dfea31242cc836249fb88a43f81ed940
31299 F20110113_AABJMZ grauel_w_Page_059.QC.jpg
6564c5b40c7aedb2c8e5da2e6727840e
235c3810f41a9cd03ebb700dc498040829d966f2
111127 F20110113_AABILA grauel_w_Page_134.jp2
7978bfa78c9861a3085de64bf9debc50
0189044465a2be2b01db2e89af37d8ed427c09e1
76049 F20110113_AABIKL grauel_w_Page_093.jp2
864fbfbc2cd2e4ad30f9795f1bbf66ef
397aa1212badc80c579adfc4e543346e94a64035
101535 F20110113_AABIJX grauel_w_Page_114.jp2
42936300bdb454556fd5379286104f03
9e08886753ddbb1203f5259a6ffd6d34da910f33
F20110113_AABILB grauel_w_Page_006.tif
4702eb60717da77350b750289ec681e7
ac428308509ad11d8247a70c6ef8e4d5adc51f37
108745 F20110113_AABIKM grauel_w_Page_078.jpg
0b30db61371186cb5e5f480380188947
d922f7cb19023c4fa75656d9f540db457deefc4f
35783 F20110113_AABIJY grauel_w_Page_161.QC.jpg
7001b2a13c7ab4ab9c3d6bbd782cb4c7
ddca5f548953de78a64fd08c5692c276beed2560
2038 F20110113_AABILC grauel_w_Page_110.txt
727dc0aee3f0f932d7a5c94660861477
794870762458d1e342810919e3fd96f0de9059d4
F20110113_AABIKN grauel_w_Page_004.tif
d5b6c52b19c38d0e68131c3cdeae0aa9
23e85deb4b9ccd34bbfa3b305be158b265dc3f41
42032 F20110113_AABIJZ grauel_w_Page_016.pro
e63c8b970fae4f0e5e2451a01f28678b
27104b1a65a8e16931328630a552f26cd7efcbc5
104322 F20110113_AABILD grauel_w_Page_058.jpg
23a7a197b71586a76f2728e79cef4106
aebfd229d8935d7fb9e5b0d8554a7082ac314523
47310 F20110113_AABIKO grauel_w_Page_039.pro
4e0aab980f48163dacfe7127f4a79e79
c754d224f5c7a7e9ed1c1de36ad740294a0912a7
51065 F20110113_AABILE grauel_w_Page_151.pro
12267cd2ae81822d5d2a6ff0f5561111
5710f3ef35269498297a8a6c34aef1832cea9824
F20110113_AABIKP grauel_w_Page_065.tif
bb167e371a996842c31bc6ae1fc175ad
4c70f0cfce62c076f2657d850432679a9c91c30a
103983 F20110113_AABILF grauel_w_Page_143.jp2
a3b0d7bac17367ec898d96a6d7fcbbe7
4df793cf97168748f92405156db6f1fc5c1ce05c
6067 F20110113_AABIKQ grauel_w_Page_051.pro
aadd823afb6cd39048778d7a40c5cf94
6238a365426765df365e58ce29c906bfb5bfeb71
1981 F20110113_AABILG grauel_w_Page_029.txt
9ad2db8e664a11e8bbc9974bc6d36bf6
957fdddd677bef78f7ac8d6c43b39b79dcdf2e49
8245 F20110113_AABIKR grauel_w_Page_064thm.jpg
fb101643c6a1f7c92897830a65c3116f
f51c4411f0d500a697685a2a36ce7558fd23c89f
5844 F20110113_AABILH grauel_w_Page_101thm.jpg
dc78a7dd1bfdcfc9583b483820046e80
d459ab4ca460701635a2e155045576fc1512b2aa
107448 F20110113_AABIKS grauel_w_Page_116.jp2
82f53f9a79d6f5a7a8e57ae4ca1949cd
8ffb9f1794dc82650ae3d1d3547121bc0832fb73
F20110113_AABILI grauel_w_Page_115thm.jpg
2a58a523ec9784b9b002b40f99184e4e
0d9f38ed6be6477457248c7214ab8c70775e0144
49315 F20110113_AABIKT grauel_w_Page_115.pro
d577a0cbe8079d6d0ff8bcd0c1887153
91e8be946cf52d4779f82bb39d747bbef01b5bd3
127416 F20110113_AABILJ grauel_w_Page_157.jp2
a11721ab2ef0f7e2c1c549516050126f
098f0df2f09564fbd690ff48a5856b448e98d9c3
41875 F20110113_AABIKU grauel_w_Page_154.pro
3addd709d0b101f67e737f431cbe440b
5c63cf78965ddab086859a366c402d66e2394767
F20110113_AABILK grauel_w_Page_109.txt
757e86ce9bf8f08b11d2956d147904bf
2a3a49828f456004a13387f4aa879db7cd941eeb
1868 F20110113_AABIKV grauel_w_Page_022.txt
fe351cbf21426714e15d79dc1c67ad75
874c8caffe5cfd6da8fb9325f8c5f40240d90dae
130864 F20110113_AABILL grauel_w_Page_168.jpg
2ea503ab03b656b185007f2d4c25c00a
0ce6c6d4e9b0ade2d3e9e12ab8029f4427af46ac
128423 F20110113_AABIKW grauel_w_Page_175.jpg
c53b970e6c3f9f3ead2710bcef52538e
b6616ab7491d791f2554c0ace4f7b3503c260d8b
56897 F20110113_AABIKX grauel_w_Page_099.jp2
b917899638e452c5d2e76c8e6cd93406
df197f3a91ccb166a2381f8bdedae54625b3329d
110080 F20110113_AABIMA grauel_w_Page_062.jp2
7cf2a3e137dbc9ae5686b4dd7081571b
f6cd77d168fdb9584b076629439ef69d102b64bc
104110 F20110113_AABILM grauel_w_Page_142.jpg
fcfbbc3f2d59d5a48026accc502df126
717da0be6b05246832da4b010bc5beb46ae001e7
51219 F20110113_AABIKY grauel_w_Page_018.pro
f315d9f98a710ad5afcdb67aa1162898
82a7e5b3777eda129d6a3605d02e33088d00b22a
F20110113_AABIMB grauel_w_Page_072.tif
2bcc275d84434e4c47b967655c52431d
2f4e1424bc2eae92435ef18f7666ca0c3e683ed2
59577 F20110113_AABILN grauel_w_Page_167.pro
5ebf3c88e84d83a7aabb53615b24ac4a
e5a3ee50a4df769b5dd297504f8bfb271d38573a
96604 F20110113_AABIKZ grauel_w_Page_044.jpg
fa227b0d1421fc9cf3376841f6bfae7d
985142d496b4f99d8790531efc9ac8900f6f4c78
76355 F20110113_AABIMC grauel_w_Page_111.jpg
60445b39a540b22f7e35189ab4e09322
548fdf8c1715c63ea5fe913e9e787220668a20ca
96271 F20110113_AABILO grauel_w_Page_153.jpg
9180e0ca70a58403aef6fefafcd55cd3
3fbd135597b4ba89f0018fda53144ccf4ebb2a79
F20110113_AABIMD grauel_w_Page_050.tif
2b65fb50306474679a084001cf1dee36
9726022d90121aa0bbc593684aedd3e18ba26440
49815 F20110113_AABILP grauel_w_Page_064.pro
1637d70446471ca97e278f8658029f78
ce207f70d7756a30c885e7b880c2b165acc44f80
33933 F20110113_AABIME grauel_w_Page_120.QC.jpg
66b6258b8e1401efeff75677ab09d372
81e78658fd98151e41fcea3613113879be0226e4
25087 F20110113_AABILQ grauel_w_Page_014.QC.jpg
7627af597ca505da9e6ea59b174fa154
94a8a9c6e1fc3772ef4a4fe4b6da68a1daf49c21
19825664 F20110113_AABIMF cativodata.xls
e6d18711619e2ccb18e79e6fc3f0d053
fa203d2b53898a4b4c637a9b944b07e0a0c84402
7941 F20110113_AABILR grauel_w_Page_133thm.jpg
f8c084dfe27916b11b4de715eb099acc
3e4b053746d58e1a737ec11190ecbb3216dcef49
2303 F20110113_AABIMG grauel_w_Page_123thm.jpg
1ca67aeec11010ef491199644c715b2f
f96c8259188ce7d585afaccafb6272e056a091f6
2044 F20110113_AABILS grauel_w_Page_037.txt
d120f988a2820d7cb0a471e9a95eba7b
af4d940882d01a4099ae22656f3d216c68f74f39
1703 F20110113_AABIMH grauel_w_Page_154.txt
03d8358562bc609c2a8b79f1f92b1652
5695812331c186499b35cf63cda5cde7757d57ec
12516 F20110113_AABILT grauel_w_Page_129.QC.jpg
4af7468923aa0f4c00414bd6a56c3ac0
6216c41b1c0cc0b2244cbf022302be55b0debb3a
128543 F20110113_AABIMI grauel_w_Page_166.jp2
6700052e7290371e514027f2bbfc5f1a
d94ac2b8b951eb9609504da25f284f0b68e404a4
2539 F20110113_AABILU grauel_w_Page_159.txt
2a5f1cb4e779f2286182e4051c9788ad
0139ddddb07d0b429fdc4eaeac23eb72b28abaf5
1992 F20110113_AABIMJ grauel_w_Page_085.txt
956b25311f089cd71c9aec8c01fb95c3
0805270ac2547a20a53c40f3f5e3c64e16f588b8
34177 F20110113_AABILV grauel_w_Page_058.QC.jpg
f8ee48b31527086c8ec1468c936f4f89
b633d6b3b3d8d88cd5256013dd1146bcfebd8a4a
49884 F20110113_AABIMK grauel_w_Page_119.pro
b009d16c6561fb46b0dc24d507f6a4f5
960dcc1eac7c389d8089d9115d57a52658fe6636
7536 F20110113_AABILW grauel_w_Page_055thm.jpg
f19a1f7e1c0a28119359ef55d28f5a3d
b48febb727f6dcc5b3331f9d50cc34fe432b7a7d
33349 F20110113_AABIML grauel_w_Page_176.QC.jpg
916714f6333d0e6d9fd1aa582a5e3759
429a7f05332a654c6e87780c464e496b66cf16ea
F20110113_AABILX grauel_w_Page_009.tif
91cc98a43f4b874d7f617c867bc60480
6f7ee21fb2925592d05d464e46cda5fba367ea12
36280 F20110113_AABINA grauel_w_Page_101.pro
10a25ec5975d988854a01d1e3360ca44
39f53ad3f52f04a629dbdea3576d7971ef2664ed
30851 F20110113_AABIMM grauel_w_Page_138.QC.jpg
6dfafecbb40e3228811a7df52fc8ce0a
23976d88f0a7538ed924a3ad0bc8647f89b748f4
87665 F20110113_AABILY grauel_w_Page_154.jpg
952c41ce0b0eaa10ab562de6444c153c
24f4bf7f338328a3b86adb7632166cc59448a0ec
57953 F20110113_AABINB grauel_w_Page_095.jp2
acad58c915abe17e537e7dee1b693aa0
21ceb3847429398e06757e8b88006999a4416dad
2585 F20110113_AABILZ grauel_w_Page_175.txt
a5308419dbe06375334c138d5c52fd9b
540155ee3890332787a6186cce11fe499d2fcdb9
6985 F20110113_AABINC grauel_w_Page_016thm.jpg
580ee1a3f658244613a4f8c1121fa2d4
c04264be0bc59c2f3d3ce62d64e44be5b5c5cf0e
129478 F20110113_AABIMN grauel_w_Page_160.jpg
1d44c1358d59671c7e82d97d424e9f1a
fcb39e423acabf3459c84eedb8e4dc2839ab5107
2023 F20110113_AABIND grauel_w_Page_070.txt
7ce6dee1a80204e774bfe1e31105cae1
ade210a8003cb1642aac742a4ae610d5e8c23eab
15510 F20110113_AABIMO grauel_w_Page_012.pro
7b9b12765e45b73d07f0856fb3ddfaf4
89e5914667d16481f6c9d61eade964cd223f8afb
F20110113_AABINE grauel_w_Page_107.txt
ce3631f17f198557c0fc8b2a81e79949
5169fa7e254f770d9e1180a0ea0b60a91d4681c2
6427 F20110113_AABIMP grauel_w_Page_126.pro
93dad371fd789be5bd964fc43c52bbb6
5fecee02d5566bc308332ba61c166e896a253b6c
32185 F20110113_AABINF grauel_w_Page_133.QC.jpg
baf27d51bf799b2f1638904d957d21ec
04a712289d051398abd759c31279f995e7272964
89330 F20110113_AABIMQ grauel_w_Page_004.jpg
ae8078f360f579e4f3a17a88a4940d83
d935ff3bf551b03aa604ea41434a4101942a2b9a
F20110113_AABING grauel_w_Page_119.tif
97d6ded74bec5e01a5389bec713c110f
05a750d6caed5954b51c11f6aacfffd74e49d44c
108461 F20110113_AABIMR grauel_w_Page_061.jp2
845b2ef435678ed41e3f91f46577d294
8ccbc3e1977a5292b36473294eebf8a5ac9289f1
2004 F20110113_AABINH grauel_w_Page_108.txt
18a7158ab828ac2cdf71c9b357aaad71
62d7027c3157a78c74c78d5dda7f7d6f837625cb
F20110113_AABIMS grauel_w_Page_146.tif
27eeebabc1150a4ad1e5413d8cbec5f1
592f209284bb02cadc971d8dc545eef5161b3625
4145 F20110113_AABINI grauel_w_Page_100thm.jpg
811cc600a8b45f25c23c01d47e2ccec3
d9fe8c30e1ac7618f07a63a52809e63241760051
F20110113_AABIMT grauel_w_Page_091.tif
e8df93733f452e8336c06a823f6879ae
9707f2be2b3e499745704a78fe72a641c0d26d60
91485 F20110113_AABINJ grauel_w_Page_150.jpg
8a3f7a644e0a44f71ba2b6379999390a
8f4fc271141d99e92258aaa7668af91ca67548d4
92315 F20110113_AABIMU grauel_w_Page_025.jpg
080683c652694d32ec71aa92c4b8584e
d605fdde584a2c5662dab9c844e1f771e41fc82f
F20110113_AABINK grauel_w_Page_143.tif
0aca6c3b61f8cd861919bf9409c1909d
84fbc951561b32ee8eb7750fd034b8ccd3d3033c
1682 F20110113_AABIMV grauel_w_Page_014.txt
cfacce8efd47345723e09149b0d9a3de
824dcfb27708a3a264bd32ddc415bc92792f346c
2557 F20110113_AABINL grauel_w_Page_170.txt
758cd271fcea4c0398bcd40bd173d3a2
00b7b6178555351cc6bf9ff4ce2e5a281c4f8808
8285 F20110113_AABIMW grauel_w_Page_165thm.jpg
8ba5c264d354204659d5f135bf15cbb2
6bc863c101dd95483dde2710ac19c1c5df35b970
8430 F20110113_AABIOA grauel_w_Page_059thm.jpg
1a57d36cda455b89f0975fbcc74345a6
f41879e48b08488250871e4c268fb2d014ddb0a9
96989 F20110113_AABINM grauel_w_Page_039.jpg
f94b95ba2abc3c9323a7df7d480fc39d
609cf50ec7bcf3ea08c2349e523150341ac6fc7e
196 F20110113_AABIMX grauel_w_Page_003.txt
55dee1e44dc948f561695e2755f62b33
89b907e8972947d01af10b19dc8a9b36880a5f30
7906 F20110113_AABIOB grauel_w_Page_039thm.jpg
4a406d65912482776df0bc3c1f9bd42a
62b412ca2ddab65d6d291d4eb30ef49c278ed76d
2030 F20110113_AABINN grauel_w_Page_035.txt
e7b4c6daafc668060606b83cc1d70725
e2e8ad819eac3b0ecccb8b48fc1875f2a0db77d2
128237 F20110113_AABIMY grauel_w_Page_161.jpg
e28af2a11e11f7cfdcdd352af9f6bd41
0f422fc20f712eab2c29c914582129d4c9b6eaae
F20110113_AABIOC grauel_w_Page_103.tif
ea461f511976757e04ba4f135f848dca
866dc3f9b1dc1385d770ff2332177785a99513e1
F20110113_AABIMZ grauel_w_Page_081.tif
5af28f293316fb2f5f65e8cd96127bc8
8b7283b5c72e1456dcdaf8ddc0989b8fa38468bf
F20110113_AABIOD grauel_w_Page_115.tif
713dc77eed3c39e30abcf9b38f1115cf
d0b763e5234ffcf1f9d16b6f54864db3e6f27bf4
47943 F20110113_AABINO grauel_w_Page_145.pro
95be632cecc1e9239dbc00a64a1fe019
9125a557e12c450f67acad70cd3afff9e7ec4ab8
11648 F20110113_AABIOE grauel_w_Page_177.pro
787c43bcb426f1043ee454b504389f5b
8f5b420f89640deb5f6e618bfcc759a91edebacb
101622 F20110113_AABINP grauel_w_Page_029.jpg
7c7343b304c38612074929884d3c280d
47522600379cb5d1416819e0f3cf012d27434f73
2724 F20110113_AABIOF grauel_w_Page_169.txt
28699cd4792838b9066f1503e83d79f1
b07ff1dfad76d638e39cdb2e4ae4551d736f782a
45620 F20110113_AABINQ grauel_w_Page_150.pro
976856baaa6b45bc1a122a110b2d373e
f6b4df808c7f49bc2f4ec9e440be9cb478ea5b86
1867 F20110113_AABIOG grauel_w_Page_131.txt
8d8da71e661ba348eb6d80d63e4ea86c
ecfe4405fb17d0911cb87501afa81b2291dea6dd
2479 F20110113_AABINR grauel_w_Page_003.QC.jpg
2599e498b7e0cf0683ba56cc4d51cf93
3a5ab2bf2a48a7fce9ec234e1b854eb46d0a1693
28089 F20110113_AABIOH grauel_w_Page_076.QC.jpg
66b3491ee447c4ea5493b61fc167bed9
75fa89eace542a6d109a13ce22833c1fb4c08975
52487 F20110113_AABINS grauel_w_Page_140.pro
e9a85e914ac358d7fef0b05e87c69e83
ff7d4fb4f4bc55d1bad5a8a0b2ce139b2196e915
98197 F20110113_AABIOI grauel_w_Page_059.jpg
b3ec57160abeff2c8b799b7f8520a387
c2bb6ef3ca204f4959f6ccd83a69690536d27343
111811 F20110113_AABINT grauel_w_Page_037.jp2
9c6135bb19c9b65b533828372a7977e8
275cec6da5cbd93b2d2a3cf79de7d11162912271
32996 F20110113_AABIOJ grauel_w_Page_088.QC.jpg
431513e59a0e609ab35e21fde3d598bf
1ec33ddfd6ac3bbdad19c0a49a916081e530dda8
F20110113_AABINU grauel_w_Page_120.tif
1f46967fe64def9fd2aed3658667ce1c
a84494fb93935e90d615c9c267dd94def7a394a9
1955 F20110113_AABIOK grauel_w_Page_036.txt
a297ff9aa0b812df5dc3bcac0e946d7d
730d001f740a7c8a96bfbfe2118fe688173f2af8
F20110113_AABINV grauel_w_Page_167.tif
87f29eaa271c2dc44388e2ae2494a986
f7a7099f05eca17140cf128647e9c8a3018bba49
F20110113_AABIOL grauel_w_Page_070.tif
c72a4e654cfa5106a6498a7ac3005adf
672061139a91676ad608ba744ec499ed78885150
F20110113_AABINW grauel_w_Page_079.tif
cba113dd3ce7d820db3f92250f779145
a777591641e4167c98c0cab914606b79e7817853
50881 F20110113_AABIOM grauel_w_Page_073.pro
b44399b77c0e19c99f95d5b3e72ad27d
3c8fb77d358e2dc63cd7b7aabd339fbc6eb8e7b6
51106 F20110113_AABINX grauel_w_Page_057.pro
ed0c82a5979fb91b8ba6d9d7c4d3e21b
d05659d4fe941de20366290a4d0319dfd2a494e4
F20110113_AABIPA grauel_w_Page_052.tif
0906e6039664a4dc062f1018e66715ac
b4a1e7b82352827a19453957d6348deabcd6400e
43642 F20110113_AABION grauel_w_Page_025.pro
99441eca6373d191f29abf5edf2b9d80
8b5a6d9be3d0d9db5049e374b3a030b80d702f0c
28914 F20110113_AABINY grauel_w_Page_084.QC.jpg
4b1feda3cf094a354401595efc518a63
dec6790292095d751bc625f36479eb369f851baa
17112 F20110113_AABIPB grauel_w_Page_005.pro
e92b006904db204cb7fc4fd706e0579e
baa879ad8f402214a89ce2a56bc9e0328f22ba73
F20110113_AABIOO grauel_w_Page_021thm.jpg
300780d2c1cd7d1807d8e804d6ac1857
10f97d04e2c48decfa57b415e324ef21f7cec995
94698 F20110113_AABINZ grauel_w_Page_023.jpg
28ac57d8c79818097f2eea0c23b61bfa
948066dcf3340ef731d718f09798caf5c8abae59
32399 F20110113_AABIPC grauel_w_Page_064.QC.jpg
bf0912f6d678714116dc033d1beb34de
fb6bcf44c27e6a2502837f36a4153e71e18382fa
33431 F20110113_AABIPD grauel_w_Page_139.QC.jpg
45afad48ab5942fee8222ac0b448cc93
d26b86e19031f9700d0ed14ef149c4d74c8d0f52
8058 F20110113_AABIOP grauel_w_Page_112thm.jpg
1049810fad9ad4a252ef5caf115bf9c2
f69e8233d70b7711d59a509525e4c02e87f65878
F20110113_AABIPE grauel_w_Page_057.tif
df2f1cf5c4ca994f4a9f8ade05efcf46
8059243e91a7e886427da5037f96226971a303ea
32873 F20110113_AABIOQ grauel_w_Page_115.QC.jpg
4b89a65af200f577b897caa94b33c247
e0f995e465f4c8e5c67a7e4b66e3c883bc6eb237
365 F20110113_AABIPF grauel_w_Page_126.txt
9d83953e90dc17b5b93196eeff7ea775
840df529284cb0cce04d9a17be5ab055cbe52f74
31073 F20110113_AABIPG grauel_w_Page_117.QC.jpg
162091af4213a3c26cf14a778e20a149
20845a9196d04f25ca4f5364c62e04fb578ee89c
2951 F20110113_AABIOR grauel_w_Page_003.pro
d4033debcb84acf0097d41f7c3af7070
cf51c1e6d759169b6d3405d959208bd92360404a
F20110113_AABIPH grauel_w_Page_073.tif
43c7eb907c76f9870a4101dacc8c4e18
4823ea1128788473ca1c6570a3af73dc86547af6
43357 F20110113_AABIOS grauel_w_Page_084.pro
d8e758a30ea420abe5c6538638bacaa2
7ea3a12034a6b8c0a245b5165ec01c952e2a22b6
34557 F20110113_AABIPI grauel_w_Page_037.QC.jpg
2481bb998ae4b1c8b73c74e4e1f6752b
7b5a1ca1ed39559142b1dbbfc8f5d9dfd171e235
F20110113_AABIOT grauel_w_Page_142.tif
2ace7443b351a831032d435a0815e76f
2a8ba86a8745c3b8a7921dd5f98962baff3eada6
56860 F20110113_AABIPJ grauel_w_Page_009.pro
78b45f14012498ca34e65fc1d5207f5f
26bc8c659cbba6677a5bf393ccc3d3fc98587e44
F20110113_AABIOU grauel_w_Page_054.tif
6c0a175ecde7e3b2de029da12729459f
d19d601a00f721abd1f5c55d7bb64b2e7a427bac
33509 F20110113_AABIPK grauel_w_Page_070.QC.jpg
c3cb44591fd3982b5380568d427b7925
7cdff46ee99a3350f3edfe5d3ce094aca09858ad
106146 F20110113_AABIOV grauel_w_Page_059.jp2
3d0e1ec6dbb8e67910944d218ad40903
6098f1dcc79d2244458505aca6ce85ec30f54e89
F20110113_AABIPL grauel_w_Page_178.tif
f3b47dfd496fe31eec804a32e26562a5
5e723a22f2715ee65438c866ebeaffc8de08c4e3
116558 F20110113_AABIOW grauel_w_Page_173.jpg
7d8ab10f24ba6b831d579e96543d5444
c53022439c9499e57f41c0bad21af57b5a8404d7
1958 F20110113_AABIQA grauel_w_Page_056.txt
646a4f106d1a0ce44063aa8023ea80cb
b45ad98503dd2ce49432eab292cf30cb86435a5d
49690 F20110113_AABIPM grauel_w_Page_061.pro
9cc61bcc07cb3a0689ec71814d180253
d36f69ab1b9561992d04043d044c6ab356deba81
94912 F20110113_AABIOX grauel_w_Page_032.jpg
5e28ae07d386520017011c2d83b78238
8f936b71c439c5e3e262b8c1526a4416d016d213
8675 F20110113_AABIQB grauel_w_Page_174thm.jpg
0873625f2e720abe3e6ee61a894d3f03
16ddf9419dced9a55d8186308ce5f040af42b0e5
F20110113_AABIPN grauel_w_Page_085.pro
e5b00b2b817fe9316b9e6e6206c32411
a38a7cb742d1410b79ad8bca3c87ff200282d24f
8708 F20110113_AABIOY grauel_w_Page_160thm.jpg
e43f3d8a5ab6f21969f4e8a5ff423bb5
8b854d4b20ee7e8083604aa9579212733dade0a2
32312 F20110113_AABIQC grauel_w_Page_091.QC.jpg
6726b4b2e9c5212c27e7ac7c6f02844f
9ab5cd6c5597c5708cb598b7a6ef3a203dfb1ca4
44816 F20110113_AABIPO grauel_w_Page_083.pro
fc239bed230c5ac1c68482149b66a4c5
f31dc78f97b6e4d331870207b9cb3365ec99fb50
8064 F20110113_AABIOZ grauel_w_Page_001.QC.jpg
3e2e46762641675b556a232a4a4511c9
2429b57dc8ffb3d063f773e986f4cd645b993190
100709 F20110113_AABIQD grauel_w_Page_082.jpg
f9d601953cc5fe394f21d295940e7e49
0f5b1bbee3bd387fbbf32cebe8335c77b062a5af
1071 F20110113_AABIPP grauel_w_Page_017.txt
b39eaeb4dada872503c14761ba262df3
5c8581e93ca285d1b888ac91476fc8249c525ceb
14372 F20110113_AABIQE grauel_w_Page_178.QC.jpg
811454dc5e137fd191d4c80619837a91
fe9ed7edf6f1e105aff958f662970bfa434b9275
1916 F20110113_AABIQF grauel_w_Page_030.txt
811e449ec3b0d4c7ff478854f38aef8c
d547d2524f0769180c94fb344e93c5e5e6af9b36
8005 F20110113_AABIPQ grauel_w_Page_060thm.jpg
28ff5ba8bab68eadd9bcc1b58a0ed17a
3cbfdf5ba405ec594a2d7487a0a7a957e146386a
129164 F20110113_AABIQG grauel_w_Page_170.jpg
efaf88268263226727b883365da37f80
d20e11dd513559364fb891d9be36cc1b43b06a94
9940 F20110113_AABIPR grauel_w_Page_105.QC.jpg
010d3de6d1b95ebadb9d4c6bd3ed9adf
d22c73fb0f1e9c6fbdcb2fae312e7029fb6a0156
109113 F20110113_AABIQH grauel_w_Page_141.jp2
85440baf7ea6a22f2b34adfaf9811d53
c93a9522abe6142bc4820dc2c756b73c26eca59b
31894 F20110113_AABIPS grauel_w_Page_121.QC.jpg
7118ecd709a96a806d5c6f19c2833185
27740042c9cd99552bf2f767fd4097a0af3b436d
2472 F20110113_AABIQI grauel_w_Page_051thm.jpg
1243b8dac5d5186759b31eced8fa431e
6c1550509910ed2b8d750941586d75da81c82919
5122 F20110113_AABIPT grauel_w_Page_002.jpg
c423820d202459ff4fc07ea4018c6e37
2bbed57c9634bffe1836c2457c4cbad11f7bf448
366 F20110113_AABIQJ grauel_w_Page_155.txt
d9d89755cc360b5cd29ac705b86bb935
8e1a5715607fa82245625050ff0fe45e3ce5b2ba
11443 F20110113_AABIPU grauel_w_Page_092.pro
14647a0fd5ca24f77a741c27352c6cd6
653d12d2dc75af995639f1a0357309d4b9645359
2128 F20110113_AABIQK grauel_w_Page_021.txt
0b0c9782a5bf3aa7940294d4bc1bfe0d
e34bfc797a88393533e52b3f6afce95feaeb0987
115945 F20110113_AABIPV grauel_w_Page_021.jp2
3491bf8e251dd9d2e03bb7c5b5954445
a80ff1d0ce4d300635137f2865199065623c9ee2
33678 F20110113_AABIQL grauel_w_Page_172.QC.jpg
2b15e2a03773b998ddbb56b18293e67c
1d98ca8a06898b968da42974dc59348fb0dd9a54
108845 F20110113_AABIPW grauel_w_Page_139.jp2
30152d84dddb7a009db96af329be20c4
4fbef12b53a9428659044f08a483356d45550232
29175 F20110113_AABIQM grauel_w_Page_130.QC.jpg
3672a3d16d8c9c87f493ec09cd59b220
738e20ceda389e227742dac3f237ed95246797c9
130781 F20110113_AABIPX grauel_w_Page_169.jpg
fc635299b7792f5c057e27de04643567
af0716ea97b137c25f5bde172298a3dcb6c18bc1
1582 F20110113_AABIRA grauel_w_Page_066.txt
c1ab14615d6a481153bbfb817d2e86b9
600c66a3f039bb7d9c983dc1c413d14a448c0a6a
48520 F20110113_AABIQN grauel_w_Page_067.jpg
9f361f37131861b957721951cfe865e0
225453f964529429ada674cde4f891c638f08b1b
8435 F20110113_AABIPY grauel_w_Page_043thm.jpg
35f1abafc2fdf4aa552b5dc51fcaf45c
8e09fac0330f9638f3e8a5a7c5401dbdaf7e2a8e
F20110113_AABIRB grauel_w_Page_024.tif
f8d3f4d628e153c6dd9b2d6e894dad86
f3f5baf5e7e6ac62a2cd6a6bf4d8b7a5446f1475
31175 F20110113_AABIQO grauel_w_Page_075.QC.jpg
205d006c55de205ddaa02ab93ecaf4a6
bbb44976898567f662bd5a2108f1e40378844542
1903 F20110113_AABIPZ grauel_w_Page_081.txt
b50fc21020dfa0e70db7c61b93df063a
31cdcad495381a49e5560bca0cbebdf572a21c3b
95451 F20110113_AABIRC grauel_w_Page_150.jp2
bf1fc9dbec6c4171ee0d34ae6a780a60
18893ffdfc8a56ba402c58b0b02628eb0585f555
34430 F20110113_AABIQP grauel_w_Page_043.QC.jpg
e859521bcfba9a2e4ffba549b0b6c9f3
784ebdd60453bbc53f771309e091146f7cadbfa9
82398 F20110113_AABIRD grauel_w_Page_111.jp2
9dafd348b86cfb86f5ae539b3315308a
b62bf1ac7316745fbd4ac0dd8ce2c62ac0a079be
2181 F20110113_AABIQQ grauel_w_Page_092thm.jpg
cc54f0eced81c689958129d01b65b0d0
d59acda6b01b95f0f8def597a1d68b586957ada1
100556 F20110113_AABIRE grauel_w_Page_056.jpg
d59517d078d68d473cd34d81917f8eb9
dcc242bffba728eb1deb32725a691c3a2d79fa00
34823 F20110113_AABIRF grauel_w_Page_165.QC.jpg
8997036b562a6be4ac142a287c2a9659
2cbbd5b637d95b6f734b87e8f9d623ba802f8147
2367 F20110113_AABIQR grauel_w_Page_006.txt
4693c75b94f08284a17ab55489f46c00
e7e10bcd044b79d8067f6d709e550482e4c88522
9260 F20110113_AABIRG grauel_w_Page_123.QC.jpg
6cf42bc25c71fb9d83e2f26920d71ed9
d98c626718b1de3d63a9734c73301bd37228ba49
101703 F20110113_AABIQS grauel_w_Page_071.jpg
9baee6cf804098e86a55236569b0b621
48fe01878656eee57348aaba0d2ad5e895558d54
7452 F20110113_AABIRH grauel_w_Page_151thm.jpg
d5a7ecdc4b4d8e828e692d26f987b29d
1a419827f2fabd28b46656c4862187836cddac84
50932 F20110113_AABIQT grauel_w_Page_062.pro
2b5f1f3ebc099003fdc02b75765bea62
ae381bafe6a31c6d812f243b8639e058c80d1205
F20110113_AABIRI grauel_w_Page_112.tif
bc61381e7bbf0b337eb2e0c7ef7a7085
ebe90957713d40b396dce005d73b6e71954ba30b
40749 F20110113_AABIQU grauel_w_Page_005.jp2
44cc4de212dc16b4dbe38c154869d5a2
233f4d6f2b01e3ca0c961d418265a50d3788891a
17834 F20110113_AABIRJ grauel_w_Page_033.pro
8cddef38701b00aea27b787be9d7ce69
95eea98dff024e09d62181e50e884fd6865b4e56
8201 F20110113_AABIQV grauel_w_Page_110thm.jpg
19b9e650ef69b26d906495c92ec69a5d
defe9fc076b86c5d786fa1f291971d3c8849c2d9
32009 F20110113_AABIRK grauel_w_Page_038.QC.jpg
e369b2dad9ec15a5542d416f86ba7540
7c48150735c9706d525a045b7cbbdbf4a66b5390
33904 F20110113_AABIQW grauel_w_Page_027.QC.jpg
2186a1f6ce86153bfbb3a85929b0d0c4
0a64726f9d2749ee0f541613a9126a7775c3395c
1980 F20110113_AABIRL grauel_w_Page_082.txt
293c92ab04370f37c02d7f52e38c5b38
e72906e3531e1f3a1d74182a964cc6e598c58905
64531 F20110113_AABIQX grauel_w_Page_161.pro
8902c400509a69a83e90662069687d4e
889c0fdeaadf8941a2673192eae5a1f0aeb0ccd1
1598 F20110113_AABISA grauel_w_Page_008thm.jpg
5b5404c1ebeacfa307ba85ca9ef46033
1d0fd80cdc6fc76916fe216678cb0e0829086240
F20110113_AABIRM grauel_w_Page_019.tif
02312623e18aa878ddb052d80fa6fb0c
7f3b517012e0950d3abe28ebdfecf0e5c8a2d0f3
F20110113_AABIQY grauel_w_Page_095.tif
3c5e593125df7b3531bfde15b11e719b
945b8994df7465c9b96bc39c44bc70d41f2bdd34
6751 F20110113_AABISB grauel_w_Page_149thm.jpg
c78b6c1fa943bafc1df46023fbb4796d
1145c93ed9a422fc213e8514926246d8d488ecff
1051986 F20110113_AABIRN grauel_w_Page_011.jp2
f8e31cfce265670b57c0e982a95e1a38
232530115e3dcf11a1c37833657898b6382171a5
17930 F20110113_AABIQZ grauel_w_Page_103.pro
c11acc2af423064125db2d7846d56161
ef93abbd352364a3d52d5d9f9260e8fac8870cdf
7827 F20110113_AABISC grauel_w_Page_145thm.jpg
677a19585b607b247293199df250a707
d7e64bf6e76084f1c00e6e287281b76267fa12ea
34277 F20110113_AABIRO grauel_w_Page_140.QC.jpg
cab669e464a4279e1a48a735ccb7ad24
e81d95a3cfc25b1840e2288f671c4ec245f8f2ea
75373 F20110113_AABISD grauel_w_Page_101.jp2
433ee5300afa34cac1c38faa51855320
d2226f553c77f7b00446a3b32144a1e0ca3b4cc1
8399 F20110113_AABIRP grauel_w_Page_120thm.jpg
9cccbde494e49e1e51eed3de91631aaf
57da2d71cf509301b621fc581b544c7c1ed61000
8211 F20110113_AABISE grauel_w_Page_073thm.jpg
79b6fa0c7df1692f75475bd50aa4d8d1
2c354cc6502fbce348f44c116a7901771829484a
F20110113_AABIRQ grauel_w_Page_002.tif
6ee5d97e5268a69a6eb9ac907ca80466
35359e5648c53fc5dc8f97d0a9700bb199324117
13145 F20110113_AABISF grauel_w_Page_147.jp2
3059fca2df33bbd0cf7d39b10a01aa82
a1efd66355c34a0c8fbaba4593b6e6783b870df2
F20110113_AABIRR grauel_w_Page_067.tif
24e562a39e1af8eaaa6a5d5857b0761e
6b5047b7341885b52a88a16219d85f010cb9ff77
96575 F20110113_AABISG grauel_w_Page_131.jpg
35a882ad37861f148b38772d7cc226b4
a191595c2f56f357ed0e69f5ac04b124144e07df
469 F20110113_AABISH grauel_w_Page_001.txt
3cb9b8c4a905f1467dec0149f0a5d68a
f6f0cf852b59459a7588bf301918cd620cbf5a9e
F20110113_AABIRS grauel_w_Page_031.tif
d86054def468a9f0807d864e6ade39cd
7da7d4c5509f58a0e189cd636127e12f30efb554
114294 F20110113_AABISI grauel_w_Page_009.jpg
9008f8a1d3096df3ef1d1571261410bb
61ba3906c1520b9d01c7635412754f783bc3679e
33946 F20110113_AABIRT grauel_w_Page_062.QC.jpg
5a85c494cd996af5437e5016684669fb
869a95bf3f9d11de585793cbe5fa5a3c161140b5
48788 F20110113_AABISJ grauel_w_Page_117.pro
c2754257bc14302e733f632f10e58bed
a5977304910f457eff0e1a4a2eaccc3d4a33de98
6735 F20110113_AABIRU grauel_w_Page_007thm.jpg
78894e824b2bb26a6ab2ccbd0cbf79cc
637d21c380391171f06d1acec438e5f240a2c8b1
1907 F20110113_AABISK grauel_w_Page_086.txt
212250ce6cb426b6c00fd2da24f9e041
708cc097654a417a9faf38be1e54f118c24688fa
1977 F20110113_AABIRV grauel_w_Page_027.txt
f038e056be290f3c493bb8e4cff088fa
970f6724bdf527986e65bda578574ea374f5ba9b
30895 F20110113_AABISL grauel_w_Page_011.QC.jpg
0057eef1a20e2394109307639660a174
005efcfc4691d815a8b03d1a9e3dd3780b22cdae
100515 F20110113_AABIRW grauel_w_Page_061.jpg
253bbb0006ef5aa51b6f3bb58eaa4c0c
74a2ff1e54acad7722594eecaa487e768d30181c
103350 F20110113_AABITA grauel_w_Page_075.jp2
6d8621aef5cfd701647b6df8173a8ef8
299c942a48655969968ead458e7b5ee46288586d
F20110113_AABISM grauel_w_Page_170.tif
3358e33874d6aa89059ce17e01dec230
492284ca02d4f9fa4dec51a276a4472cdeffff4c
36488 F20110113_AABIRX grauel_w_Page_174.QC.jpg
a234aebc9ccb8643c23592d4e94c965a
c0a05196a9cd8ca204a9877c7e5154aad4ba4c1b
54495 F20110113_AABITB grauel_w_Page_011.pro
b56a3a3d37f2cde17cb2e1b4168dc5af
420763e6bf1aec85773e145bf3dfc32c4328fe5b
34181 F20110113_AABISN grauel_w_Page_073.QC.jpg
8a59303d51f0b03c2b7aec4d1a0803a1
5e178dab45d9b0742bae5ae87d817c48e41b9a9f
4991 F20110113_AABIRY grauel_w_Page_010thm.jpg
df681efb797e13686bd21f95ebacfcb1
70480a093c2d65a3f0c827816c4766b57c3f4e3b
F20110113_AABITC grauel_w_Page_093.txt
bc4d7cf1b97076e747b683bd5e248167
dc50c6a4efa6e0c6c2a8849daa09ec2852f97e25
100072 F20110113_AABISO grauel_w_Page_116.jpg
50e4ea63af3a40db9978e222f2eb6504
a9e18af57859fd1f52ba0a7b2c1a58d5eec31d3d
F20110113_AABIRZ grauel_w_Page_016.tif
2da4c6715bc4ce77b201b8d78346fcf2
3b6620e92e3beb0d37cb72c13636e0b01544b463
2028 F20110113_AABITD grauel_w_Page_132.txt
2775ac0658d377c7b7a7ed88cbaa52a6
e352118d95ee4ce2d4ae4944707eb9286f4d5ca1
27537 F20110113_AABISP grauel_w_Page_123.jpg
353c5568fb86a182db7bd92d269a5d9e
b5dda78575d4e03197008b3d9525a8bbf0bea931
126886 F20110113_AABITE grauel_w_Page_171.jp2
aa7e1c836ed12335f242f0a62f565bd2
75e247a0d2eecdc6ee61ac9ef3364c2960b0f6cc
60739 F20110113_AABISQ grauel_w_Page_176.pro
2e5bba4b93adbee34d12d1ac13dc54be
b21aeca88dc94b018709bf6e8c2baa4b85ed23f4
49233 F20110113_AABITF grauel_w_Page_029.pro
d3919bdca6233b7b7a63f2e19a0b4348
e7a848ce48c1e607c5461e8f632ae73b059a1116
F20110113_AABISR grauel_w_Page_128.tif
060ac088bc0f60dce73ca34db0354855
3aa75f846a77c7944965ee9f980774a7b38d0949
392 F20110113_AABITG grauel_w_Page_128.txt
abea5b5aee6b12050318926fa8b66794
ef219873dd38ebf6be68c21fe5161463bd6143ca
F20110113_AABISS grauel_w_Page_019.txt
caea0f651d57e3744975d8741f338bac
772f767880169d878703aadf3f23e53e316e3d48
85141 F20110113_AABITH grauel_w_Page_014.jp2
d96d436ff535d5f2bed01ce0be02b5c3
67054210256dafd05a031331e86c16ecbd715b28
63929 F20110113_AABITI grauel_w_Page_175.pro
a7b75a6c137d13ef7bd5c984ecd235b6
c0a229300232847335ff1af074a3fe26db796bbe
89134 F20110113_AABIST grauel_w_Page_096.jp2
ea3a38d8433bf23025d85eec92466435
8f90a4050ba6e95098ba80dfcd5cd9dcf70f31c6
14245 F20110113_AABITJ grauel_w_Page_129.pro
ab73d6ef0e28237bfd362212a04f8ad5
ff8bdba680137acf7aae85718fa08ecaa791c6d5
107113 F20110113_AABISU grauel_w_Page_080.jp2
d996d48e4c1071c8e945374dfac69a2c
3b39e571080a539e97d531363d66b51c86edc4f9
58746 F20110113_AABITK grauel_w_Page_098.jpg
1377aa69a9894cafb5799c333758eb1e
1b40a77dc19589b887fc1d192f9dd8b5c0e72410
99124 F20110113_AABISV grauel_w_Page_025.jp2
2cebc27472c82f0b65c1e792da61f3d7
8ba7500f3279650e6f38dfd8ce1a8c2edac63dd3
F20110113_AABITL grauel_w_Page_046.tif
dc852a91cb73d4d5fbad3d44e73b91e5
050fa2266e43eafa5f00bf4fca7bac42b25969bf
2032 F20110113_AABISW grauel_w_Page_156.txt
fd5d24703b0430ce9e2bf1de4d328d08
e93d856f6e1c7c4101c5e6a3f369794bf844a8bf
8374 F20110113_AABITM grauel_w_Page_070thm.jpg
38f614932309fdb56ee2076eeee0f9c0
8386958ee1b708a4b9c28fa3c4e5e57931afc95e
8643 F20110113_AABISX grauel_w_Page_027thm.jpg
16485c658ebc0f651ef73d5b53aabc7e
fb94899c676f9e3b6160be836710e40ad44f4d95
3470582 F20110113_AABIUA grauel_w.pdf
8216f19490496baed2f10fc37f6f98fb
e48b0fbfbf403fe238ca7a2ef60fb7e367372e34
1792 F20110113_AABITN grauel_w_Page_024.txt
a18c8398a3b2db30a903a47ee986f0ef
39cc368b6f463e40ec9c67115b147976b3981da9
25604 F20110113_AABISY grauel_w_Page_048.QC.jpg
fa727c3afc6e08b75e4b758a93bea369
4f0f4ab0a142bb8f85fca5c31a04f14700ed32fd
3232544 F20110113_AABIUB cativodata.csv
fe2d4515f773115847a49060c5e81d91
d526bfa45d49d135d130e439a001bf851e5a1e9b
65693 F20110113_AABITO grauel_w_Page_163.pro
b4f4989e8a4e7cc1d42cb88fc93b37e6
6dc3277dab680b3bb9d8550e3270fb1e0c916047
22457 F20110113_AABISZ grauel_w_Page_149.QC.jpg
3ee25e5058047c7c954410da5a505e18
576827457d3df8239d02f4f2cbccb65fbc1fa758
2522 F20110113_AABIUC grauel_w_Page_162.txt
4fa0a01988fd10656f044c03335cf13e
6c21e750682762141f82ca90acc40bf353b5ece4
101483 F20110113_AABIUD grauel_w_Page_042.jp2
fb50d3126b87d4ac40f6ebbcb9085164
d9e34a2d5a9a4113eb6445a01901c381174cd336
8087 F20110113_AABITP grauel_w_Page_089thm.jpg
b5ecc33dbf6eeb8097dfb1e80afdf14f
bafdafe7606d2298cd67a4e573f8efd74a4dd088
33603 F20110113_AABIUE grauel_w_Page_110.QC.jpg
f46cbc371bc184d01f29b604418e472e
5fbca361bffa7fd51518feb909b6ce37c9370d3d
F20110113_AABITQ grauel_w_Page_177.tif
6ac75cf8f97c0cdea66b8ced588ca5e6
e13155ca11dc34ff9717d891e0fe816a138bb5dc
F20110113_AABIUF grauel_w_Page_077.tif
7cf336cbcb67a5c037ce2fd41a850077
9dcdf7d50832e98f11b02c1e53c3466098163571
101007 F20110113_AABITR grauel_w_Page_153.jp2
e14ab20cfb0b573bdc40727a9e3e46b2
0817d78bb8afb121b41d1af6e93916659e355665
F20110113_AABJAA grauel_w_Page_132.tif
65b025e0c6c8f0970ba745bd8fe45fc3
225333c2b2f7e045bf57509b211b70ff8f93dca0
35082 F20110113_AABIUG grauel_w_Page_074.QC.jpg
c3e5fa8ce698b2983f17a996f7700f99
5b9d7340010107f0595cb8916ae5e662db671f63
17187 F20110113_AABITS grauel_w_Page_095.QC.jpg
929ad690c771ab6503c6853c00c044ea
a42932754fcc463dbb98e131aa002cce03732d56
98028 F20110113_AABJAB grauel_w_Page_034.jp2
e1e3930e3423e5f8af0a0b190aecf7e7
b92a621c30ae31e049d8325ba8844346f0855579
96580 F20110113_AABIUH grauel_w_Page_075.jpg
20859974fdedb9f74678e1de691560c9
36513379675cb30ec2becd64a1fc4e1db8c3c87b
61360 F20110113_AABITT grauel_w_Page_160.pro
214db5fe002e6196cf3d746b3bed3a74
a2976f23a81a0cc87cfb8793f0b349e7c07633c1
105758 F20110113_AABJAC grauel_w_Page_091.jp2
84dcfeb2268b5976e97e06fd7c142e76
5f0c4267f57823fd2b5e7dc0a43f8e33662e3613
53126 F20110113_AABIUI grauel_w_Page_021.pro
99378628751d82dec111cd7614da6f6f
8354125a4f6d9e5f5774c377c2aec8c006532a06
52083 F20110113_AABJAD grauel_w_Page_067.jp2
5c6e60174dc3b3ff9b752c13c21e415b
15db7b18bb88f7a0c75df43a1a0e027c9bd9f0f4
F20110113_AABIUJ grauel_w_Page_039.tif
69012994a76bf72f04b627ec174de478
a06227795ae4be3427dfae77660cf5d71960e7c7
25571 F20110113_AABITU grauel_w_Page_017.pro
d4eb2c99cb3a1c1ff4a9a764b05ee326
1b90cdfb424343198529a2dfe147b3e041b48852
47928 F20110113_AABJAE grauel_w_Page_081.pro
1acdfe147fc04f0d14c360e4a455da8e
73c02dfcf791f102d5e0d4f34e0c01d49b5f5086
110544 F20110113_AABIUK grauel_w_Page_057.jp2
6907db09a5aaa0faa47c9c2829bd8ac9
809cb298ee30a0e8554140c41b0a0c2d06af714b
118319 F20110113_AABITV grauel_w_Page_157.jpg
24b425d9eb2f5cbcb56e5c53c7a1ffec
947a5e7189202f18d7ffcdbabc3db2be388a5ce7
41581 F20110113_AABJAF grauel_w_Page_069.pro
b5a6d38b5e84e7f468758408d43e188a
3b253687c82ed80086d2351854cbc3e5bd29cd7d
105380 F20110113_AABHRJ grauel_w_Page_146.jp2
03ce8d0a50d1669cabef09149c55fe65
04c89c2dfd5e3c295cc66884126ecdbb9d6cfdf1
F20110113_AABIUL grauel_w_Page_158.tif
56165d07d6acd09ef4d9e7f0b14cbe74
81a429a04b1a79977ad1cb5af4d1d05f3c3bc5d9
F20110113_AABITW grauel_w_Page_176.tif
d10fab3ccf7edac0b9a5bae2bbdf4098
5f4830c2f0fc24ae7b18bcc3bf603fd45b99055b
F20110113_AABJAG grauel_w_Page_104.tif
93a3a8fab2bdc5aca4cbd4c87f7f982b
57178be705681cd70fbd4d3fcaedd452211f7eb1
33326 F20110113_AABIVA grauel_w_Page_116.QC.jpg
f583e3bedf3321345d562914741b1fff
ca5688f663c33512f900ce114d6d4a7551b0927e
34791 F20110113_AABHRK grauel_w_Page_144.QC.jpg
02dcea32635c0ab464ee8e2a6c709aab
2f574d3519d0f3a627b32bb1f7c8ae4bff0777e2
2010 F20110113_AABIUM grauel_w_Page_040.txt
30dc8359857d48a5ea1a24fca625213f
ef4cfb9d7994dd69c59f0541251395924d63469e
4736 F20110113_AABITX grauel_w_Page_013.QC.jpg
60dd86ffe9d912b082e2591693c3806e
bc27f9f8b2213eb18e2360098a30e969ac82a03d
F20110113_AABJAH grauel_w_Page_116.tif
add7d4787631aa674bfb25cbf04cba65
9d089b552c00e40ad3e34d456bbf5778e221ffc2
7350 F20110113_AABIVB grauel_w_Page_083thm.jpg
04f334b57c05208a6e296ecc03b72898
1ac62df08201bace79604ba4581072be6cf0bd9a
49716 F20110113_AABHRL grauel_w_Page_116.pro
8dd3117b5d9da803ae0e430ef30e7f92
b6de0dddcb554573ce74857437183d7c3bb5ad29
1578 F20110113_AABIUN grauel_w_Page_013thm.jpg
0abb501cfe48dde82f309f97ce96e0c7
417408ac2524b738390e112e5500e865fde9b659
34437 F20110113_AABITY grauel_w_Page_112.QC.jpg
09d7c4eb321f5c9b27d7691e36792c51
b06030f4c66dacf486a3b4c2dc2371135cabe3c2
12169 F20110113_AABJAI grauel_w_Page_123.pro
d6e637b08898c4f15a3c2e693d33dd9b
61b20fd2969dba252d131ce304d57172abdcbc47
F20110113_AABHSA grauel_w_Page_017.tif
1519fa888f81aca2636011f6c6e81389
e4ab5fd07bd2c72b75f94524cde4eb8ef0afa9ff
F20110113_AABIVC grauel_w_Page_060.txt
d6012824c8637efd161ef7ed813e646c
3a40cee00fe3842a594e85daf6ac0baf004a2fc7
1926 F20110113_AABHRM grauel_w_Page_141.txt
8586507eec92498a9c881133eb889e26
7c9993a0709b87e17aa34652d002e6e6c3756af1
36197 F20110113_AABIUO grauel_w_Page_164.QC.jpg
16c391d504398c6abeab24571ffe2599
f0c9ae0d417d9430182deb5c47accd3fc507c83e
126768 F20110113_AABITZ grauel_w_Page_176.jp2
cd54527ada4dd83604618042f08cc329
87d70da1838cec1af521f59c70d009758faf7898
111955 F20110113_AABJAJ grauel_w_Page_018.jp2
a2eb66511d8856f78a4356709e37c96b
d56194cf851d5038f7c3aead4e344fc375a9b182
101353 F20110113_AABHSB grauel_w_Page_026.jpg
63f2d4cdea66996f42501df525b5296c
dd76838659d3774f8466ff871f221c31b165a24e
2499 F20110113_AABIVD grauel_w_Page_096.txt
e6dfab0e0abca0a4c717fcd74e96dce1
83f49f628bd91a62b7cc0c7344c8fdfb6932d896
47432 F20110113_AABHRN grauel_w_Page_042.pro
9198f001072f25d1cfe50f23c52e86c7
9ad355aeb0fa24e2397d0b6f9e11ba96cb8949ef
36392 F20110113_AABIUP grauel_w_Page_010.pro
5d2986cb2a176327ba590da917cf0438
68c2d69f12490e58364adecc1675a55b3839e86b
8060 F20110113_AABJAK grauel_w_Page_128.pro
8c97754961fe194678ffe8fe3f73fb54
9c1a456afb43983f238cd6ffe9f4923b675c7506
516 F20110113_AABHSC grauel_w_Page_177.txt
7505d6943965ca041efed76d1fe8679d
cadb7732c666bafb437be2095798af611a989867
49427 F20110113_AABIVE grauel_w_Page_036.pro
c34224c6cf0406f05431b07b1505d49b
35bae7e8161b87eba03f51620ea892207685b4b5
90078 F20110113_AABHRO grauel_w_Page_102.jpg
e020fcf21c9d911c138e6f49d23180a3
9bb0722b422e7a76beb3dc6408590d45d993a59e
F20110113_AABIUQ grauel_w_Page_060.tif
2c31fe224cc6521994c3cb4aa6ffb858
5619b567cc203542518a2e5be97eb4b501989d3f
33184 F20110113_AABJBA grauel_w_Page_135.QC.jpg
40666ebfe780454223374dbeae4bcaae
af52597a3c0601b72b314bdb3bb949733c13cd17
35585 F20110113_AABJAL grauel_w_Page_171.QC.jpg
21a28a94705fa98f1e85ea7e659c0860
f512f897605d91d715ffb7f3a0c87fab057082cb
1035 F20110113_AABHSD grauel_w_Page_149.txt
0976891486b42e2383ca00be31b3078d
e668fc2466f51bdb682d3698e792004f3fcffb10
1489 F20110113_AABIVF grauel_w_Page_010.txt
d8f815220ed94e90f073476e2328ceb8
70fc7e83afe7369e1aae3a2c634d347bf40a2ee8
32989 F20110113_AABHRP grauel_w_Page_136.QC.jpg
66fe08397271cf43b72f3ed1d676e18c
3e4185a14808f19f092494adc5ded90393705c51
32600 F20110113_AABIUR grauel_w_Page_122.QC.jpg
c8a483ee4638fcee13b65654ed4812ae
d2522a997b6ca5b8d07046e3a9d02418161d3a85
95483 F20110113_AABJAM grauel_w_Page_019.jpg
85de127bb82ec1e656cefbdd07695d69
b7ef06ef30d63d7808456012301939af9465ed8c
107319 F20110113_AABHSE grauel_w_Page_036.jp2
0f9a7e9a367e4744896431a5e6a73eaf
7f0d558da603790a24fa5cbe49c74ff22a7e181d
45806 F20110113_AABIVG grauel_w_Page_019.pro
dd5f8bdf2409d9d8abd8a6570db0fc8a
5cfdbd55f3b70ec4350ef6c1a32d47a2d8d43549
48674 F20110113_AABHRQ grauel_w_Page_087.pro
093db2cc586338a4d538ad633062660e
f67fd0569dcc1eee7bcc8be5aea6c301063413f7
F20110113_AABIUS grauel_w_Page_032.tif
71324d84d3006d41a5ac78085d9f49bd
09035fbb622eb1a8d7b7fd73c432c8db9d94eb9e
69387 F20110113_AABJBB grauel_w_Page_066.jp2
1b750cafc24ad1daba328666a20f22cc
a4768af8e01f57b99e253c773dd61414607913b4
108136 F20110113_AABJAN grauel_w_Page_079.jp2
39a1969564f9a0d104dbaf73e0b00515
8949461838817832df3701f152667c48f5fc0c08
F20110113_AABHSF grauel_w_Page_042.tif
51fbddca47a3e68e0d1d859afce2950e
687e2e9c73e2e3a9c90baf1ec960d96695e563c3
F20110113_AABIVH grauel_w_Page_124thm.jpg
a825b637c242b5892850420dcd24cc81
af11891799a545dd1a7a4da97a7b008cec9a2036
F20110113_AABHRR grauel_w_Page_134.tif
81d6632d100d3587178cf82b24eab2d0
2b4128e88f5beec6db7fe776dee3707be4dd6d95
1256 F20110113_AABIUT grauel_w_Page_103.txt
a5d0adcc52b7fe349d4670d8f9405653
6ae949b173246f67d7ba96d57c675f12d8eaf517
1985 F20110113_AABJBC grauel_w_Page_118.txt
7c4ca229a495d8a8a93ade26489221d5
cc7aa869f85b0b0fd6d810550c39960053a3b853
F20110113_AABJAO grauel_w_Page_088.tif
2ecfe7d9f24c93a41f90fed80cf32d7b
c3df244b26e91044cbf28ed6c1a00cfcb837b681
1927 F20110113_AABHSG grauel_w_Page_047.txt
55194b69bcd191791ef8c80dd8efd39c
72aa6327310ca49272ec3c96fd83af963f9b4ba6
47995 F20110113_AABIVI grauel_w_Page_038.pro
6cb418064a757826f186b59fae2d8874
a5602f046ddcca51ba473b9bc73d25929440cb26
111739 F20110113_AABHRS grauel_w_Page_058.jp2
3a1f359aea25abcf2b0e7aa5590c3c5e
abc417e345184e0138f9ea4a67c7eeb0b7932918
31769 F20110113_AABIUU grauel_w_Page_077.QC.jpg
f9c728ff1fdda61ead4c38c7fe46dbff
0a6a7b7c864ccee3460f51a6b52ef55a4dba51e0
F20110113_AABJBD grauel_w_Page_126.tif
5b1e08f1c06443a12b64ec3dd8add192
cccb752cb5dc2a3f85d644cce2c46477526b8298
5947 F20110113_AABJAP grauel_w_Page_096thm.jpg
da0f6ad02752980c4760a06d67f9771a
819b3616445563e14018e037afb6ac3dc2137473
1721 F20110113_AABHSH grauel_w_Page_002.QC.jpg
2258b6ab84ce86e14cbdede339b5e88a
73a0e032fa08d6242406386843117b1544ed009e
1991 F20110113_AABIVJ grauel_w_Page_079.txt
0db0a5e3f4d9622475b39d5d4a7e1dbc
33f22ecedfef470ef329a0c9312301b92211de77
2381 F20110113_AABJBE grauel_w_Page_009.txt
56a2a58ac9882b6d4b4d342a9f47c0e8
b72ff9fbb4e8197a0a4f8f0fad08ba9c74230204
45002 F20110113_AABJAQ grauel_w_Page_034.pro
5eb59f47c8c32320484a192689575f22
d052c0a156e22d47b16208ef4b13c0ceffc23080
8439 F20110113_AABHSI grauel_w_Page_132thm.jpg
069e3a8a5bc0f4f5a02f186b8f3bd637
535815f63536ad50d4171ac6059cf431968e7123
F20110113_AABIVK grauel_w_Page_008.tif
dd4ab96ff13289ed12c448a25ba9166a
231e7ab0b05575cadef32f92bd9540dc2c79d306
F20110113_AABHRT grauel_w_Page_075.txt
82c0c9caf7ef4c63e6a6d5c34a2397c4
b36180ebe3ab005546808718cc57c8fd859dd9c6
103650 F20110113_AABIUV grauel_w_Page_063.jpg
870fd2e7a1ee47acb2dfd17b7c045666
6a682af35a49e959633ea42a4649e7998c528a7a
32334 F20110113_AABJBF grauel_w_Page_079.QC.jpg
1af79f13476ebd4db03097fc4c6ea8f2
94f0b39bae7def3b247fb637c01de653568dd60a
109547 F20110113_AABJAR grauel_w_Page_071.jp2
fddc368a293822421d470318fdf0c332
6a5c873b3107f078135f218e071e60a6648f783c
F20110113_AABHSJ grauel_w_Page_064.tif
17aab0a886d9bff89ee364e1a2cf82ef
8c1222b1eb9efc6ae1c9a02b0f8fcc9a71c2ed17
7672 F20110113_AABIVL grauel_w_Page_131thm.jpg
3289567841fdc5f3afdd8386ae747eec
9804f8faa71f1248fc4f6aff7653a5ab8bdc158f
71981 F20110113_AABHRU grauel_w_Page_007.pro
f29af95e774640cf979762314b8214fa
e26f10115a441d398b1e183d93a8ae432e452120
109265 F20110113_AABIUW grauel_w_Page_027.jp2
9ad77e6f4ebeb4ed57fb92d440b1eb9c
4b3bf708276a0117e0596822034ddf33fef482a9
7410 F20110113_AABJBG grauel_w_Page_072thm.jpg
54b792598688e1e0b6d6b2101385affa
c14b222afc3979c199b128c6886c3b39a8dd047c
33094 F20110113_AABJAS grauel_w_Page_085.QC.jpg
1ef39882721b975c13da01ab5d7c8332
af4a6159015998e5ea4a78e1876ac2e8865723b8
8234 F20110113_AABIWA grauel_w_Page_058thm.jpg
0d43f1306bff9ba202c7e41ddf4c78b5
a8805c5cefc3c58181115832fddb346f600bcaed
30779 F20110113_AABHSK grauel_w_Page_109.QC.jpg
76c72e713cf9ee801f4786a5cdc7c845
7cc5845cf44bc956d093c0a90fba5902dd6d7c81
35406 F20110113_AABIVM grauel_w_Page_162.QC.jpg
fe7e9d82baa81cfc17b013dbe344caf4
d2c41f21b58ebee2583239e0fbe93dec55beec9f
F20110113_AABHRV grauel_w_Page_090.tif
a23338c929159897e4612eb68772dc1d
8fdb2c9b068c79750894ed7492aa75d4456f125d
28985 F20110113_AABIUX grauel_w_Page_156.QC.jpg
0686a9c504d6cced77f8852118d27cde
e7f8c797d2da806b73e4543477435cd6c1f3c4fb
62339 F20110113_AABJBH grauel_w_Page_103.jp2
6a24585347ff4ef3f8e2ffff73584d3e
17672a2838283b111a8930405b0d0dcd8aa20204
F20110113_AABJAT grauel_w_Page_131.tif
e88f0071b1235f87545a04ac03f20662
35d624436aae9fec2c255629adf6f7ee7b39e847
51216 F20110113_AABIWB grauel_w_Page_037.pro
8118cbc59e50447d3ca72869ffba8a00
5755f5d1f3406605068ed44ab3ccdc11bdf8cbde
54680 F20110113_AABHSL grauel_w_Page_096.pro
901ba46ac0c3b39a5dae069865ff2b63
2cd5d520faafcfe8e114c2de86023f1f58d7c919
8695 F20110113_AABIVN grauel_w_Page_171thm.jpg
6bcdf5baaea70a72a1f356240ab91f1c
a5af3d27b62dc66e9dc257439cb1c97d86f95f51
F20110113_AABHRW grauel_w_Page_162.tif
44361f4c9f72a7013807fcb9f928f10c
60524fe239b05a18953a1ae473b53a53c48442d6
526 F20110113_AABIUY grauel_w_Page_123.txt
85f8dfda025bdc46a87d2e9919e4c9b9
61dc4f0f6893d7c27f97f9822ad4d30562a0e9ea
49259 F20110113_AABJBI grauel_w_Page_031.pro
de1f08036fcc5497c7d379bb5c5c3c35
b2faf62d486530dc97b965b7280b7609c72f3c3a
37092 F20110113_AABJAU grauel_w_Page_126.jp2
cf1785fd49ce74b8de74fa6b1c4e0839
786ae53aa1fefcd87e8ac0849ab3bf412c5f2c2f
51976 F20110113_AABIWC grauel_w_Page_046.pro
ee211c657dfabeabc88c7ced858253ad
5a95bbfdf7ae9bab2eedea95167301ae34b208f8
128883 F20110113_AABHSM grauel_w_Page_162.jpg
ec13466cf8f293de645da47f71c0be45
a20001667760de122a4ca25d7a68a4db72579275
22564 F20110113_AABIVO grauel_w_Page_006.QC.jpg
ee497624d3616fa085f3b3d4726fbc7e
42ddd215081bc23e16d8a7e6baff3548dc2bdad4
95709 F20110113_AABHRX grauel_w_Page_084.jp2
b11c032c33e933f6af70608c40e1a7c1
443ee3518c50d742594cb7d4e27344bddeea4a8d
114147 F20110113_AABIUZ grauel_w_Page_011.jpg
20bc08d727f11baae36d8aeafe6e2f0c
5559265b5223de256f3492d45a46ba217b801004
1794 F20110113_AABHTA grauel_w_Page_130.txt
2eb991be8b21250118c518d37d595da6
86762872e5afa1b0ab4ed8c3ab41b0e35acaad6a
61206 F20110113_AABJBJ grauel_w_Page_171.pro
fc90c8fea7e78eef59028a129494972c
3055d4af60cbef55bed74fbfb4c2ce164f4adb28
F20110113_AABJAV grauel_w_Page_010.tif
a2370b964fd729aa8fd89090fe00b18a
10b4e17e8e198724da85f88cb3cd4e53efa6fa2c
F20110113_AABIWD grauel_w_Page_099.tif
d23e93375443ae4d8e4c2acfa3809b83
77118d1f03918327bd17ce595081b357dc216805
9746 F20110113_AABIVP grauel_w_Page_097.QC.jpg
12a4dc37beb2002c4995072454bc96d6
41f8b5ffd04ddd642a4d6b1932b35ab80b904533
109198 F20110113_AABHSN grauel_w_Page_031.jp2
3f24ef9d85c8e8882ff0986c04c30913
ea58a7c2347a7b140040861646fb99dd90d9f464
91210 F20110113_AABHRY grauel_w_Page_106.jpg
511a3c61faaaf8801b3449970e77ec49
30b0828547585718d842422cafaf767124a53247
F20110113_AABHTB grauel_w_Page_107.tif
ae3f419a9de352001d48d69a0d163d96
7e579652bd10acf3d3cef240306cc59ca9436ad8
F20110113_AABJBK grauel_w_Page_158thm.jpg
5c726b510af1fa815ea149a1e4b5014c
6ae5ece05cdc109772fbd74e35915d92df5e86c3
41273 F20110113_AABJAW grauel_w_Page_050.jpg
8198781aa511563a2b14af8418f1648c
7c0d1f7597a3fbaa41f6cd8e728c0ab0208b83b3
44274 F20110113_AABIWE grauel_w_Page_024.pro
8905e36d880b56678a0b3411c9c6bf05
f3b9896cacde63b353e4af5a43a904dc678863aa
109754 F20110113_AABIVQ grauel_w_Page_122.jp2
2839975efa9d3a2df84e94fbe5369e31
a54ad84632a4ddb55c739fe5b370c0d1edc670c2
F20110113_AABHSO grauel_w_Page_087.tif
20178c1b56ef37c7abf8739dc85cb1e6
f275f7c0e15ec5ac66fe41f47d0727c5b37db397
110338 F20110113_AABHRZ grauel_w_Page_026.jp2
7615b014db42cc79f55c430a1cf16316
2065b9df07c9a50873a32974a81665ded36dae0c
29316 F20110113_AABHTC grauel_w_Page_105.jpg
2e362ab1a522400a7f479e926d5b5ff1
8ddd97c3cdbaa7ab78f4bac9f7cf277230303d96
48940 F20110113_AABJCA grauel_w_Page_141.pro
e9f4c1dfe18d960148d360fae4c792c8
009d15b87cf82816c33d3b4b96042741fcddf52e
19394 F20110113_AABJBL grauel_w_Page_051.jpg
af8950aa5c0be3eec3e058a5f45a749f
87d1fe484de1a3862d7ee2c0812272ebbe9636a3
32803 F20110113_AABJAX grauel_w_Page_071.QC.jpg
514bafef38609835f9ef2a74c07b7d9b
bf195c27b666fb9fd3e799afcd04ff167c653d0d
772 F20110113_AABIWF grauel_w_Page_067.txt
eaed6b7ee213d82858b136fbff1fcbae
d7839c110178de3083bcd362755cc7a814db17c6
3751 F20110113_AABIVR grauel_w_Page_129thm.jpg
b8ca80ca0d312e25509609b4b2a3eedd
235716efb0a3c77eaa863ad124a038223cef6b52
8181 F20110113_AABHSP grauel_w_Page_117thm.jpg
2065d5855c4cb16ae3c35b6afcf3ce7e
693532212be00cb99fdb78cec7008a6d7beb1caf
7780 F20110113_AABHTD grauel_w_Page_143thm.jpg
34336d186eb84b926f3ecae58ae46d69
b3819a4ee60c47a0e223ac534a8eadc99837fce9
74920 F20110113_AABJCB grauel_w_Page_015.jpg
36047ab5f26a74c57fbcd3e575bc9dfc
f880a514d15c0c476ec45ba1c9ef28a3804c5346
1938 F20110113_AABJBM grauel_w_Page_045.txt
e012eebf0a5c28450c92b7fe165d94e9
055d8060b03ad7041a522bab670831f0a25fed5a
1084 F20110113_AABJAY grauel_w_Page_068.txt
68b552b2d29394b29ced54811d797de7
f09325522df74336fc334b916c87832e9a2fdf3b
47881 F20110113_AABIWG grauel_w_Page_143.pro
00dd666309cfaf8dafc6ce8f28ea3172
62d0d810b67734a5718e3c05c3ac59e90b5ece1a
10813 F20110113_AABIVS grauel_w_Page_099.QC.jpg
19a766885cbab7fdba06b3af4764a473
8323c4b47429b05c2ad3967851169f8a10018cf5
8739 F20110113_AABHSQ grauel_w_Page_078thm.jpg
3b1db9f4d28e1c7c82d6abf8a1838f63
9e93265ea5bc4d81b12e521318d89f2ee4e978d8
30812 F20110113_AABHTE grauel_w_Page_128.jpg
b39ca5a08475e8f9374f5f728046fb97
648eae3a13a069703df438f92112e4ca8546856a
5108 F20110113_AABJBN grauel_w_Page_067thm.jpg
87f5e8abfa3cabc51c293d784aa95f77
69b27f41a556d442ce3bcbbc905ee7a830bc1eb1
48181 F20110113_AABJAZ grauel_w_Page_075.pro
74b7d5ed34144b5b11e34a8297873ab2
f682cdc50f5f26b1e02ee4c570cbb74a3af0c6e3
77982 F20110113_AABIWH grauel_w_Page_010.jpg
2e360c63f52a83f283c7edf6afd46f7a
c40549349eed1693840fe94a8c16c3991aac8822
92486 F20110113_AABIVT grauel_w_Page_034.jpg
9bb86b1b3cd1a2271318ca524042369a
b7e3609c9263fa2a78ad0aab73551dafba8da254
29120 F20110113_AABHSR grauel_w_Page_083.QC.jpg
62e9186315361e68c6d46eb1db2d53db
51f23e038491abb98a964a16eacc3f274301a71c
109968 F20110113_AABHTF grauel_w_Page_088.jp2
719d3dd06defac96f7cdc9e1056b5b90
a069eec7f60eb97b0ef7d7c249debafadb828208
4802 F20110113_AABJCC grauel_w_Page_094thm.jpg
e1f563777e29c4c4ea099d0a99f3b769
fd6b29beb947b32bdf115def5f42a3d99f8e0e61
34082 F20110113_AABJBO grauel_w_Page_167.QC.jpg
3f54401a31843893a6d0592e445fa0f6
ca0e7868945695119db85b374da4180706209bcd
12615 F20110113_AABIWI grauel_w_Page_005.QC.jpg
a300da45ee70d514cffcf4c4fb73c924
ab5843f10efbea78f9abd5d806a57ccfd67cda39
109446 F20110113_AABIVU grauel_w_Page_064.jp2
f95e782663fac135547bd63edd194013
2b32ba6d6fe515e9c597fb1d5e21a9e0ea19f2bc
8088 F20110113_AABHSS grauel_w_Page_045thm.jpg
2acb36339dcb823dd72b7102d7aff8db
6f21d9ed93f249937cc49c597e407dc545296073
128496 F20110113_AABHTG grauel_w_Page_159.jpg
23cbf96e8655d4378ff700e31e333e6b
87337c2d68f7b79ac02cc57e52147f2a5cecf2ad
1811 F20110113_AABJCD grauel_w_Page_041.txt
7c9cc420a68341225a1f6f48d6c8a1eb
64f23db76806c960bf007a88f90e97ff4f9a9196
37231 F20110113_AABJBP grauel_w_Page_111.pro
d5cc42adb1e774c009bb396d2b54049b
9bd243fbe85031f2387717bbf948d1059e273f8f
133487 F20110113_AABIWJ grauel_w_Page_174.jp2
2ec7750269fc02e25ae9eb044898fbcf
9eca3ad9e45d834b6a10a259a120cabe112f4761
31985 F20110113_AABIVV grauel_w_Page_087.QC.jpg
7ceb21458e7a254a9525df3303d0c618
bcdca8fbcbf81619b5ca4c861e19ad511e6a1397
3043 F20110113_AABHST grauel_w_Page_097thm.jpg
ddff705f9196d8c40ca5674629866474
4f88ed58bae70c64fbdcb6e301a08c3ba21a0cc3
53628 F20110113_AABHTH grauel_w_Page_100.jpg
d9d494c978ed5421fb0d4e99455dbb2e
552292c7a06d0810d3ee4a90d26549f0d6031e8b
2658 F20110113_AABJCE grauel_w_Page_163.txt
979e5b74545cda84ba6064a1fe6d7568
f2568a8b6b0085f705de77af7c93a0f07b683eb1
8496 F20110113_AABJBQ grauel_w_Page_046thm.jpg
4603b660a3c688d5a7aa53371b6db01b
c086df67bcb946dd992788f222b2c1edab9babc1
42252 F20110113_AABIWK grauel_w_Page_076.pro
fc43f1834033ebe7d5556e8b6398e809
9f88064ed9a0e57378810b157ff33edb9013366c
9174 F20110113_AABHTI grauel_w_Page_053.pro
e9e6f317f2e606ad616835fb2cc733d6
3f1650a413b972232f5665bbcf40c83f5c4f786a
8879 F20110113_AABJCF grauel_w_Page_168thm.jpg
03f38a360e37be0b7123de23cad6ec87
d9558966a25d8579fdf0c017ee6abca30052afce
8099 F20110113_AABJBR grauel_w_Page_038thm.jpg
27e1e3938cc5bbfc511506708e44f588
46b571718c9b26df826ad4273655fcba074410ae
F20110113_AABIWL grauel_w_Page_035.tif
ed4c185c64bfb4fbb6965d188088dd50
f0b249236c610afcd10a0084636949bd87188221
49949 F20110113_AABIVW grauel_w_Page_071.pro
03b4cd69120e4c4cae5b0ade11279ff8
caddbffbbadd0d38ec62a24656b2b8a75c259f9d
105157 F20110113_AABHSU grauel_w_Page_018.jpg
e22937cb0408ad251efc13cfd035bf0b
9eedf9818656e198401a1dba2c0e12b3f3dc64be
96773 F20110113_AABHTJ grauel_w_Page_045.jpg
156c64b925107ce6036841a2b3cc9987
3f8646e68ba98067263048530fc8c5a36e4ecc04
F20110113_AABJCG grauel_w_Page_155.tif
20cef72ff94894ec7c6e9c8863fd0040
addd9fe555a33a7be6f91f6436e0e7c20611a925
32146 F20110113_AABJBS grauel_w_Page_099.jpg
ef4fb91f1599a238a0c21b611947c013
0855f5dae8ef41ed1524cfdc0cc0218142aed851
30889 F20110113_AABIXA grauel_w_Page_044.QC.jpg
c244ccd08dfffdc830591657a944e73b
8cfe912835bf2bf7ef531269e570cae49024ec67
18748 F20110113_AABIWM grauel_w_Page_094.QC.jpg
93ccadcaf9fea21a6f7ce62b1c3d1289
31ed33dadad71c0c4ac3eb27d8c2eb26229415ae
92238 F20110113_AABIVX grauel_w_Page_069.jp2
f955f39366c4d4f4c32547695ed82783
c2b6f5a5764a67bbd5435ded44c1ada8d478a42a
8490 F20110113_AABHSV grauel_w_Page_082thm.jpg
267e4688021b288b39a45ed891e1e9a6
fc4cc054de9307ea424191c9ae9392779cf60a1c
27910 F20110113_AABHTK grauel_w_Page_007.QC.jpg
5a9ad3afe09e7d453c152a1f0d42d2af
84472c58797665339092c15c6e839c5a225b4333
1840 F20110113_AABJCH grauel_w_Page_150.txt
278e362a7706fbf07792c0f0f745c367
dfad0652b1c8a1190f9385d3ab9e23b7798174af
18475 F20110113_AABJBT grauel_w_Page_068.QC.jpg
1fe25e972cfa8bbb6ace3875ef044d31
630bbbae9cbfe28dabe218fddb091e177d1e8ed8
F20110113_AABIXB grauel_w_Page_150.tif
9ba87fc5e3cb8f4bd8a878ff17de3b52
d82559e1c337aa8cfcc0d31af6537d4b96383f08
8151 F20110113_AABIWN grauel_w_Page_056thm.jpg
0c2eff68919f7ccddb32570b8e2d81c9
04f04ed3f98666b9d635e6ba8cae1209f7bec667
24951 F20110113_AABIVY grauel_w_Page_097.pro
7265f34147f92d3ab891b9199be7b85f
b9083c0b0e1cc5d6c69e38dc05b06e44d28cf37c
8152 F20110113_AABHSW grauel_w_Page_071thm.jpg
6469e1bebf3acc14adbf0cb5a8956713
471a509be0725dd3071e055fbf03f90b06b5b25f
56962 F20110113_AABHTL grauel_w_Page_095.jpg
3cf3a46eaacbeae80d669b2ec635423b
b24ed55eb8be50beb944bde4d69278bfda908dcb
28487 F20110113_AABJCI grauel_w_Page_017.QC.jpg
a4fb36f6e1dd4e49ee36b83ab7c123c2
4b13c6ad64190bb8a82b375354f1ac134295e5b5
50936 F20110113_AABJBU grauel_w_Page_088.pro
95fd1ebbcda3616a7d25a87e534ffe76
15405d724cc5100bdd7cfb968f3babee00b479fb
F20110113_AABIXC grauel_w_Page_149.tif
aab1d319bbc7538c83a31fdd99e4983d
cbfebcfc0445e5492d46dd2ba4f94c1350b212dc
103829 F20110113_AABIWO grauel_w_Page_070.jpg
420b25eae881ca05dcfade4b72d66617
589cbd3b5c35da280e7e070f486631801651b9b9
13032 F20110113_AABIVZ grauel_w_Page_033.QC.jpg
0e6001d7a7af9951a8b553ada5e66c94
cdf8fe25b802adda52362535d1b64152e8b074c4
124677 F20110113_AABHSX grauel_w_Page_167.jp2
e6d1dee9e30383b227e0da2f5cd71456
51cddbf7b6a14e1cb726d9b7c7aa6fc5ee77d6fb
66042 F20110113_AABHUA grauel_w_Page_168.pro
c5fa01206eceb8a87ab5a33cf3f53a2c
e959d70947b3c89d8f3e34ee8e6ef59533d59784
F20110113_AABHTM grauel_w_Page_127.tif
8b9c8061f2049ca95d48d2566a27702c
7431cce4b34bdc54485ecef0f5f96d1a0dbf1ed1
8373 F20110113_AABJCJ grauel_w_Page_176thm.jpg
7dd449f25b418525705fa22d86b6c421
099826aac8806e29ac28f88971a84267c67d5fb7
1932 F20110113_AABJBV grauel_w_Page_133.txt
637046ca2e7590129e0198482914bd35
a6699343b800f388c67700e6d66af3f017f7cae0
F20110113_AABIXD grauel_w_Page_048.tif
cd118c519ccc5b2c0bf35b9ae74a9802
5b3714ebc6af885ca99599ea1ad65f1d7af83d05
1305 F20110113_AABIWP grauel_w_Page_065.txt
10a4741f71fdda811f2f85594c341d27
07f2bea8a807c67f4e59d9692aeab913e74775cf
32941 F20110113_AABHSY grauel_w_Page_108.QC.jpg
1a0ffe75f2a22b602847ff8036b7c282
3148117d4ebf8ea85b2676403679c06f7f06989a
1952 F20110113_AABHUB grauel_w_Page_117.txt
6cc410e5074d9afea5e85053831976b5
357074a03774c437cbe57b6c0378122d8c3d79bf
103686 F20110113_AABHTN grauel_w_Page_134.jpg
081566c79d07fd558691f168ae874abe
3f78889e2b77576e501f9e09687a551f457a0b24
8769 F20110113_AABJCK grauel_w_Page_001.pro
bd09c6eba549e2b06dad92883459b3e5
6bf6e63ace9b7d035a6eab99ffb80683b1ada267
61997 F20110113_AABJBW grauel_w_Page_162.pro
06cdc2c1fef92dc3be47c0ea860ace23
740ae34bd8c89e661316209b2e2199cc97e21a9a
F20110113_AABIXE grauel_w_Page_015.tif
a01e98b4484dfa2385c6541df2f9a9e4
dc6c28be131fc659f8d1dc395e10309c12dfd16d
1982 F20110113_AABIWQ grauel_w_Page_115.txt
58925405c5a60a53628e76578bb015c6
013851483a673a188626b7ac72889bd650c611f3
47235 F20110113_AABHSZ grauel_w_Page_044.pro
2870ad35b4727fa9e21385a2803718e2
ea7cf4838278a824287dae7f271dcaadd5be4a4e
106302 F20110113_AABHUC grauel_w_Page_087.jp2
48ce425e13d19e72d2ebdbda6949e5df
2c81cd6378a1582038bd511fc643318779648a67
92515 F20110113_AABHTO grauel_w_Page_004.jp2
88f6e3ee5627b84d6d62cb15d3cb784e
b1d087fceecc7c0c997a9a2a50bd804ed0c26c2f
33329 F20110113_AABJDA grauel_w_Page_082.QC.jpg
9f24a68419ce4e8131246f38a3fd78cb
31e24cff6c9ee9c17265189c5bfc243af57ccb19
2842 F20110113_AABJCL grauel_w_Page_012thm.jpg
093f7f69eca8a5f8495e84501d9b516c
2532ca79e31943806a3cf8c910facb78b1f7e38a
2092 F20110113_AABJBX grauel_w_Page_043.txt
b7d7670e357d58da26244a891ac273f0
45788ce178b2106bfbd3c95e4e05c8fb37301ed9
F20110113_AABIXF grauel_w_Page_038.txt
bcf51704a8ffbdc392d5d5edf3bc89f9
b50641af1ffe6069f5152ade2a74ff1ae6031446
102497 F20110113_AABIWR grauel_w_Page_088.jpg
929872b3a0491c5a6103c71c85630572
338c00c63f216ab14dbd029303799985b3c2140c
F20110113_AABHUD grauel_w_Page_061.txt
ad6d5201e907c6f4e0561287e983a677
d76d630107415366d261db1fac4e4a1a07692b60
1556 F20110113_AABHTP grauel_w_Page_111.txt
13b97a4861ad2e52485e63d6a47bb952
39e7b26c89b72bb0cf967c2f5c95213c001aec65
F20110113_AABJDB grauel_w_Page_156.tif
324df546b82c1016cfefceab6432844d
cd2add9a4f0f05ff0e12d2237847ba321f3faafb
F20110113_AABJCM grauel_w_Page_030.tif
8bbd52ceaea35d9d0a61b9f17aa50d32
ea13aa519e718c926b1e0bf5b18db3bfd3a01c03
F20110113_AABJBY grauel_w_Page_032.QC.jpg
ce3cffb47927b3253e2a9061bfbe735b
57a2653fe963ad50dbd7cd4bf377894a8110f00e
16835 F20110113_AABIXG grauel_w_Page_103.QC.jpg
09d61ba2049f94ee5c815857da70ae89
25aa3279f36063d495fcb1a0db8543da6580f9d3
8626 F20110113_AABIWS grauel_w_Page_167thm.jpg
82888aa4b10e3fbaf421d932ca87b231
1f85ac54acd4369c15f43bd8bbe9647d38b0ede6
50060 F20110113_AABHUE grauel_w_Page_122.pro
7ffd7e5469212e3ce86ac2e061c76994
08fed2ff11abad28d94d973f2fefd487379700d1
50019 F20110113_AABHTQ grauel_w_Page_137.pro
bea704215eeb53ef1943fe2850da9f3b
4e1a47d3a62c43d11daad07bc56b639127d39838
100113 F20110113_AABJDC grauel_w_Page_089.jpg
8831f97093eb1a6ad7d9f8b0db590c09
4e33708c7adf6a903350dde632de425f3257ce2b
2715 F20110113_AABJCN grauel_w_Page_049thm.jpg
22ade5a3a7227cd62ee4099adeb32539
e37bec7e208372197cfe4c329f74158e0c03405e
8348 F20110113_AABJBZ grauel_w_Page_162thm.jpg
bf031de7cdeec2248b18731fd92e4dc6
4af50519990dfd4a8aba83a88a6c5317853e3788
106441 F20110113_AABIXH grauel_w_Page_115.jp2
d24873693d6b03163d44514487033e45
99e96360b1855ee13ce712627631a4ea42e45d17
8165 F20110113_AABIWT grauel_w_Page_144thm.jpg
c8eb7141180c50af9e08b32c87ff725f
ee5258ddf242a9943f92b0b05c5d3ecf85c2896c
101082 F20110113_AABHUF grauel_w_Page_064.jpg
c27b8959d81497aa2356719cb12070dc
a065d4c1aab727aaf8be796b2f8dadac37c135ad
F20110113_AABHTR grauel_w_Page_085.tif
1024aa22636caf690d9b04a18ad1396c
76f6dc7a4b650d77e8aa5058f9268c14dc361eab
F20110113_AABIAA grauel_w_Page_164.tif
33fc20f926712f6a4d126f6847e4ea4c
7d92ae066f60ce74d0772ad7946eda98b0b26af6
F20110113_AABJCO grauel_w_Page_071.tif
693029102560051bd5665edaf3f32d5d
dfa26118b78370fa68b348c5c49c3e4a37804f35
F20110113_AABIXI grauel_w_Page_023.jp2
9112c1bd75473843a68aae48fd1dacd2
7ad050db1596a40fcccb91c5c000ac321b92289b
26486 F20110113_AABIWU grauel_w_Page_052.jpg
f05302398ab7c04e32cce99af1e17ddd
adb33737c458d14d620e54fe848ac077ec8ebb62
7593 F20110113_AABHUG grauel_w_Page_017thm.jpg
104faf38c6fb59231f9eb32899503b67
ab812c66b7f7e0dd6a4688e696bb821bf6f1c6c8
8314 F20110113_AABHTS grauel_w_Page_134thm.jpg
3db516f60f702e8abcd2b1abb96aba72
575dc202e5530ac17b2ccdf8b6aa423de389a436
204698 F20110113_AABJDD UFE0004366_00001.mets
edfae58e95b2e18d91d7894e6e84953c
fb7235de3a5aaa0a761753903767592076615ea1
F20110113_AABJCP grauel_w_Page_014.tif
999ce88f0a36d8c72c178e46e7a4be18
ea7a1027bf8399fad739e5ab6a0479a297fdaae3
96558 F20110113_AABIXJ grauel_w_Page_143.jpg
ab6569ed750f177ae4d6c2d174433649
69c75705518d56b9c3ba34eb883a043956af538d
103275 F20110113_AABIWV grauel_w_Page_027.jpg
2e67a5e91c6965c275df9b991132f6d1
f21a026e41d7ba27fd79cff800248de7b17790be
8287 F20110113_AABHUH grauel_w_Page_054.pro
04147a541e371748f43f7d81b03551df
4840e8f99a00c77a6459095c78e3246546888bb8
2071 F20110113_AABHTT grauel_w_Page_112.txt
e4eb21575f2d68cecc15ace76a554ae1
a8e3984db186478958063b7caa4d152581177b10
137552 F20110113_AABIAB grauel_w_Page_163.jp2
bfef3590f92019195e267e7161357cf5
540258f7629a5559096f8f3148b70f6cc48258fe
130851 F20110113_AABJCQ grauel_w_Page_170.jp2
e6ae724624d0e36c55a8798bfe350c74
10e7703b05380471c3cdb7fe194a89124b186cb8
31847 F20110113_AABIXK grauel_w_Page_049.jp2
f4a74f567a1950a82b6a17dcb9aae11b
c9d422de323ff487da34ed8324b89de774a02dd7
21544 F20110113_AABIWW grauel_w_Page_101.QC.jpg
768637c9d0a6f2ca32b550d82d0c1618
562759043cd1d2ea920e36d43b4a61ab6bfee961
22764 F20110113_AABHUI grauel_w_Page_124.jpg
86803a2f84146f707b0a2e3536bdd902
3089e717bcab0dff2956fe5eaaf81907642383a4
8634 F20110113_AABHTU grauel_w_Page_169thm.jpg
7f33b415db229a412973db31010cf902
ab708b8b55ceff75fd4fac7d5d005d81fa97612b
7825 F20110113_AABIAC grauel_w_Page_031thm.jpg
b0cc303fda1e55da50a6e1ff9386e9d9
6c5cd13bd849fba020da64ecabc010bbc034a60b
1051893 F20110113_AABJCR grauel_w_Page_007.jp2
2777c262bddaf5a358de92ca5953a37d
a09577fad40ba30aa0177247e59059258666f652
F20110113_AABIXL grauel_w_Page_083.tif
8dbdfae20b577c4e07b38a5ef0600c16
5b7285ac110b753a059c90d1328dff917f5f6a97
48989 F20110113_AABHUJ grauel_w_Page_133.pro
0b252d0a6164e3d69f3600d61025254b
12c0e7da8d28d25ffe63e82b8b680c19fb22a4e7
103771 F20110113_AABIAD grauel_w_Page_077.jp2
a50ada0fa77a895f927959037d7840a9
9c413c7a4042375224d2007953092c75fbfa686d
38639 F20110113_AABJDG grauel_w_Page_005.jpg
bf1b9e6d1187392488139a49b35a5b3c
69984fb0b259202a1c683b0b83f14d14700cf293
105623 F20110113_AABJCS grauel_w_Page_022.jp2
99444bd49fe44238797e2287c101d417
e1f1aa2ddc9cdb604e0d7ae08a71f6c5e76d67fd
56558 F20110113_AABIYA grauel_w_Page_103.jpg
b9e6cf58add03c230721191d5b4696b3
d76f4b45fc662f429a0955fc326f9cd3a3cbbeb2
2686 F20110113_AABIXM grauel_w_Page_168.txt
64a4f9509f73dc69a3350ca35dd81bd3
57a48b635a269fb888ee48d0c0574b3908dfd964
7629 F20110113_AABIWX grauel_w_Page_025thm.jpg
43f2be2cb1cdb02aedf33095c487d3e6
edee458d066e2d1a121320e1837e0f4c03484880
15245 F20110113_AABHUK grauel_w_Page_050.QC.jpg
c9a3b1efc1d56ea6f3f2cc25b723b436
3fa3eb306dc64ec165c75c906e20bd13e214b751
123862 F20110113_AABHTV grauel_w_Page_158.jp2
3ec8913c374879a33ba0a7af32f66574
ced483bd5627be17f8517c4df03250116e4a6253
32990 F20110113_AABIAE grauel_w_Page_061.QC.jpg
825633dab518567f20d7f8d231352242
0cecc081424c47432e016e401226983d9fd2bdae
125483 F20110113_AABJDH grauel_w_Page_007.jpg
6331f9d6e172e5833ac0279dcb6f288d
7f80b86de4cce780b04010b71d28112e522bd809
123 F20110113_AABJCT grauel_w_Page_002.txt
eb382896c11251c6e107a3575b6ca1c8
bb6eeccfd5804a9b99b54e11a3a5e87a49dc3aa3
34242 F20110113_AABIYB grauel_w_Page_057.QC.jpg
187a666ea305814a1d58c0732c42e021
14d6838be3c8fbe913f5cd4c0f117a4d8b2dcadb
96479 F20110113_AABIXN grauel_w_Page_106.jp2
1bb0afea2c8ee81f5af3350d715df66f
1285845d4a812f75cc5e2b81528db0a90f47a1f8
103120 F20110113_AABIWY grauel_w_Page_039.jp2
d37d6720c2b6dcee8528ed0d4692d86d
63742b8fed41e5881800c1bf9c3060af7a7dee28
1125 F20110113_AABHUL grauel_w_Page_125.txt
6ba03a0ec8326623fc50c740fafaffa2
0e4a7d4a08655b0bb088ed196c74d72f96d5631e
1051970 F20110113_AABHTW grauel_w_Page_009.jp2
1c5199a8f85da03f4fa2e909db8dc479
0a92c192a46af1b628e9ce4576e7314e3d36cf84
8541 F20110113_AABIAF grauel_w_Page_161thm.jpg
130c61907d5a04dd8daea05a7ae91b2c
7bafa8dcc9516374e43742becac4b7cbc9ecf916
89111 F20110113_AABJDI grauel_w_Page_016.jpg
09746359b2ef43a287df522ccca8e675
18ef2ce9855a6f7b01574f05f885c7c847ddb431
5177 F20110113_AABJCU grauel_w_Page_006thm.jpg
5195f855692c28a7fb66930356099c7f
c5546116b37e932f296fc3a0ff9ce3a3d9e32506
8698 F20110113_AABIYC grauel_w_Page_175thm.jpg
0cbbb8f60d0a237811cb13707aa1d6d8
e8da2bb3cf7671464ab30c0c935b17cec3a010c9
32640 F20110113_AABIXO grauel_w_Page_021.QC.jpg
74e7616eb881b9a7f214b503585553c0
aaca1bf433dc455ec4bf816a08db0b1e6c2e8b6c
7989 F20110113_AABIWZ grauel_w_Page_091thm.jpg
77d5b8c9e99c08d5fcf155b560e27467
b4faa7e4437c919a5418268150a33df19cafc3f2
1890 F20110113_AABHVA grauel_w_Page_143.txt
4afb5cbad76654994adef24326ff8830
e88e83f394029e12f10deec9f17969166425cb2e
8268 F20110113_AABHUM grauel_w_Page_047thm.jpg
fa2436de68e7254537601207562b94d4
5cb1f1e5ad0d2cc80f613b97a0871dd5c0f8a8e8
F20110113_AABHTX grauel_w_Page_093.tif
caaf95122323e93e9eb2ee54a836278f
2e1648f3fd29d836b09b6159c096744ed85c6e41
F20110113_AABIAG grauel_w_Page_141.tif
2a05c2859784f4b1c72ba67d9a3c8883
94cdac7591847a2e28c0a1f2a35ebdf4d3676b57
103531 F20110113_AABJDJ grauel_w_Page_020.jpg
8469cd724139d2fa6a77565f01300851
aa27f6af8383d3b7b0b8adeb829355aaebdb087e
50925 F20110113_AABJCV grauel_w_Page_040.pro
1a4505ba8187deaaa7c80206f740d685
b2518fb56cdc88fa417d5c0d9d000b4e15b527e5
20980 F20110113_AABIYD grauel_w_Page_155.jp2
6d503710071dda0841d0ec9014de5157
bd751e5c27249da6d713728b8e46afe1a247864a
98989 F20110113_AABIXP grauel_w_Page_133.jpg
1bdc3de046b37c8ed9e0ff51c2e9e80d
b84e0c312ee2e10e83a309200f68ee1b834bff7d
F20110113_AABHUN grauel_w_Page_168.tif
ba96b970a6affaa13308d5428c11a8e1
4025d072429a1e51306002acdc79ac42b0d02720
942 F20110113_AABHTY grauel_w_Page_129.txt
15fe586c9a0960affe8809f6dc493cad
5636aa276982632285dc8d5994d63e0c231b4526
111842 F20110113_AABIAH grauel_w_Page_132.jp2
7a5d5ef27f4ff23f24b0669d98266c7f
e16fbf73dee0d3ef590689410a202a1a8b0a69f1
35747 F20110113_AABHVB grauel_w_Page_159.QC.jpg
ffa67eb9f5c2b970aaecd783490be9a1
a45a9c87c64123cf03eb19f860ba9c7d41b6b84d
99184 F20110113_AABJDK grauel_w_Page_030.jpg
338d2c02f378f493d2795c67b999c3c3
be4c4735cd4ad2a536a6b290ba3074336e959536
32419 F20110113_AABJCW grauel_w_Page_141.QC.jpg
f4ae9ba4ed2337aa21c8b09e3bd9bbca
7ef17c7b97b2864f920336f54b8dee51599181dd
F20110113_AABIYE grauel_w_Page_028.tif
eb236b623b445c71e75eb6a091932173
fe9094bccdbb217c0f34b13a8a43581e30ea5568
8096 F20110113_AABIXQ grauel_w_Page_122thm.jpg
268de6db263223b6965457fafed326e8
1ca768d8d09c40b9b08dcf2917b43a12f9048cb2
216735 F20110113_AABHUO grauel_w_Page_053.jp2
90747f842bf5bb0f725a42e283cbe2cc
48f92ed6167cc7b8ed1dec65da95cc655d945a37
103324 F20110113_AABHTZ grauel_w_Page_032.jp2
4efdbb4cb69087120bc4325dec5043a9
c592d97df70cf9d83716dcb71b4c8d17a5be1b99
4498 F20110113_AABIAI grauel_w_Page_095thm.jpg
5f350e0a1dc5cbfcb09f4ba49726d9d0
5d3af8c76a48f037d032380504b768e8b11d0a37
8054 F20110113_AABHVC grauel_w_Page_020thm.jpg
3a7f11f0c37f29ec1c301c4326aaac1f
4838e143a3b8fda359b89f97ffa77d9f73822115
25705 F20110113_AABJEA grauel_w_Page_092.jpg
6ad2971309964add3687952cc73c3c03
54124c1bfc2d309a631c5b1f981d7427dde46cf0
100926 F20110113_AABJDL grauel_w_Page_031.jpg
8f9ab934eb7b5f16bbd311eaa3fbc841
76e72e8f149d906df1a9083a7a20acc4bd2a7349
104037 F20110113_AABJCX grauel_w_Page_156.jpg
2d1b00b6f6ea6165a845335c0d5d2e71
76d39d76b8f87fb04c9eca88a7e6b69e65eafe07
19736 F20110113_AABIYF grauel_w_Page_178.pro
9407d8c199078fab16ee3d7775acaf3a
54f27e07b3a82ed89115ecd8f6fc25b51e18fa1d
50107 F20110113_AABIXR grauel_w_Page_156.pro
7e5500527ce046bc6ed773fa8830d980
0ee8a59396a0c16b68f7a6b945674984b1ff556e
7500 F20110113_AABHUP grauel_w_Page_054.QC.jpg
6cf90b0825993ab100b4c60e9cf37c54
1d449abf8ace41f1994277d830cbc33ad2e80e33
54937 F20110113_AABIAJ grauel_w_Page_006.pro
ccc32792a49f951ee517023c78c16d7c
eec029c79ff5c52fbdfd816fa15273672f5ce0a3
1466 F20110113_AABHVD grauel_w_Page_095.txt
054bc5a806233c1ebda9d39be746ad3f
19a42f72a1fb1121f74713c889ce234d5eaf9b7c
72653 F20110113_AABJEB grauel_w_Page_093.jpg
181edf7ef5685b324858b3697c3c1367
745c02beed9bcb78068b7eabd7ef2c45707d10f7
38967 F20110113_AABJDM grauel_w_Page_033.jpg
7e65c5c70cd222b7b518631f3a4ce3a1
4e56a82a71e96765187777d9507681ad5849f3f8
F20110113_AABJCY grauel_w_Page_101.tif
7fe66db52d061a179ff6a22599a0bd2b
7123a75543f18380e4d4e31df8b3a2a277fdee30
F20110113_AABIYG grauel_w_Page_102.tif
41ae8659f2882fc1b65d11f4129165af
e144fba0df31dc88120eeec16f43663cf10dae1c
F20110113_AABIXS grauel_w_Page_018.txt
1c3cb9750449a6f4cd348ed57c459274
ea469c7da8382eb627ef89e67460d6e8a1290e67
106181 F20110113_AABHUQ grauel_w_Page_028.jp2
098ddde219aa2a7011cb4099c26fefd6
2344ffaece7091edf41b4574191bd6bfaef3d06c
15089 F20110113_AABIAK grauel_w_Page_013.jpg
05f3c8cb0ecc707ddfffa1127bfa9983
348221290ee233b351d20e4305739abea516ac91
2053 F20110113_AABHVE grauel_w_Page_057.txt
4c1c0b5c495ff059e88ffe370265a15a
d491119348491542f6da8a31b72fdadcd0a8dd97
101973 F20110113_AABJEC grauel_w_Page_108.jpg
abc3b3cd1769332aca52129524339072
491b0b28854453ae01e5a648e1a54ff56fc95852
105792 F20110113_AABJDN grauel_w_Page_037.jpg
b10d488332c7d9562fc2f94a6e4f33e1
78cce9d4b8d723a1e98cc89980ce90c7c9a89ee5
1873 F20110113_AABJCZ grauel_w_Page_044.txt
18447a966e9d2e25376a633334b197b2
60b9bb034169716379e7668716eb66c6c3a25c83
F20110113_AABIYH grauel_w_Page_020.tif
c398e38aa514c849911861918608beb5
cfef25078f3fa8a35d57e697648d34460da4e6ea
2042 F20110113_AABIXT grauel_w_Page_062.txt
e468f5b58726e34b1af35f0cba52d700
8e227c70f6c0f380afecaa1024d896203f25114a
1937 F20110113_AABHUR grauel_w_Page_113.txt
c3d3e6abf99df35561cee01468d2cdfd
071e1f666c8ba87541e70a6ce6105a2ea87378a2
51418 F20110113_AABIBA grauel_w_Page_142.pro
51f09848709ee1f5e53cdd2ec87b545d
108cb676fe6b7b1ca1f99da9ad89ec3bfd6bc1f9
F20110113_AABIAL grauel_w_Page_159.tif
a7338a9b4fba407ae58ecd2c9b5a7825
7c6e0642127f5a2855c2e197fd25c862e4a34ef2
88986 F20110113_AABHVF grauel_w_Page_096.jpg
070797dc0a90796a971f27fe48af1d61
4245c56df915e5d7eb8351142e3244b9b63eff84
94429 F20110113_AABJED grauel_w_Page_109.jpg
651671900d06726a84cf405a9ef24bb8
03c1a512fc23d8f98f450484e5656c6c1ae7010f
103702 F20110113_AABJDO grauel_w_Page_043.jpg
d93059a8173b25c2364c798bcd762ebf
ab5ba202f4c2353ca2a33f4b14a919f73db47bb2
F20110113_AABIYI grauel_w_Page_047.tif
f741964f3720ce629b6e9241e0e0a374
8ed2c8d2a466d4d12d81ce8a52855f8cf79b4b63
6446 F20110113_AABIXU grauel_w_Page_111thm.jpg
8995274b50162e2cb6cf181bdf26b13e
efcc03be3eed4d1098a3a96d9d21694f1a35ca6d
F20110113_AABHUS grauel_w_Page_092.tif
519a4ce59c2d94cc6d6c77185a07aedf
edc8101245404d3f9267795358a3135a63be8b27
10254 F20110113_AABIBB grauel_w_Page_104.QC.jpg
2434b7882f8f8d7ea8875a3a93b78fd5
4e0cb2254834edd926b90cef2c3e728247fd33b9
46320 F20110113_AABIAM grauel_w_Page_065.jp2
8a223fe2ac386848af3ab760cf448d35
dbfe6740fac243d0aeb9734964cd0e202ce25cd7
1232 F20110113_AABHVG grauel_w_Page_147thm.jpg
c2a9dfc153b3d150c6e5a52e2c49822e
d6e0a3cdd5bb1487ac5dd9790b44bf67c10d3440
77816 F20110113_AABJDP grauel_w_Page_048.jpg
16e804b26da51211cbcf806ca4a82403
42dae56a8cadaa1129116ef038f376e33d4d9797
F20110113_AABIYJ grauel_w_Page_077.txt
2efc889baede12ee54a6780330fb40e2
343d856224882ef64b05c99057f503b1e2f1819e
99593 F20110113_AABIXV grauel_w_Page_115.jpg
2c4d2a4fcc97d3936e833248f86e5597
33a17e65521a82a1e7ef6a11b69db0b6ac003a2d
34155 F20110113_AABHUT grauel_w_Page_134.QC.jpg
a5fa5a00a91ac085fb101c07d71966ed
9f5be2c0ea944117a0a84cd595b12246c8f2dfa8
26287 F20110113_AABIAN grauel_w_Page_052.jp2
f436620fc4efa9b8c143d42ba671de1d
97af4df465e6d2d6cf290c5e9f068f28df760fef
61565 F20110113_AABHVH grauel_w_Page_068.jpg
15d7cfc505509b2a4a86f92cdfae2685
81b5249bd70345f7b5318b30f2ae80de127e6019
104332 F20110113_AABJEE grauel_w_Page_110.jpg
b9224dc8f1a2d64ae206f5c67f733ffe
189df8d98a35bb9d2652d1849cf5c69a472b77df
32180 F20110113_AABJDQ grauel_w_Page_049.jpg
9b859ba398407e637905495c1046636d
531bf0c5a11bede9f78e731f0f22bac69173d4b6
5109 F20110113_AABIYK grauel_w_Page_008.QC.jpg
8bbd48195e021df80643cf4d163687c2
3477e1bb24af81d4f6457b9952213fbe5f8e232e
130527 F20110113_AABIXW grauel_w_Page_152.jp2
acb62fca68a1e503914aab4c64f5e53a
a86e89de5cad82bdf9a33a4766a91b222a95af4e
1373 F20110113_AABHUU grauel_w_Page_100.txt
e256f12fed20c0c4c17591398da835e3
f3aceb5051e99ec2f57ab93eeb29432154185451
6652 F20110113_AABIBC grauel_w_Page_124.QC.jpg
50565df0767fd56eb13e1daa17f36333
d5d491d66d5057b8ebdaa1ee17ec9866b25a241c
37609 F20110113_AABIAO grauel_w_Page_129.jp2
a7e8f6c0807d81a4a7aa74d11abf5be1
ad3d1eb7140f3a46ed505748e1400724b2068bdc
32258 F20110113_AABHVI grauel_w_Page_056.QC.jpg
63d256ef08d3dae8786d768eb1d96d8e
4ebc7b108e3db3a026f02c0995eb593cfade9eb7
104469 F20110113_AABJEF grauel_w_Page_112.jpg
8c5a11b1bd8afc44b8537116d51f2842
6b92980a497d5e26e4f68706ba123dd2fb57b46b
98763 F20110113_AABJDR grauel_w_Page_060.jpg
724573f31e758ea499f27311ba7a89b5
67a1587c47653e141cc4f1f8875c82209fd470c7
37191 F20110113_AABIYL grauel_w_Page_093.pro
1177598c877e01e7143e1b989f98e49a
addb65987c3fcf33e4602555bd609c72b1204138
115138 F20110113_AABIXX grauel_w_Page_151.jpg
34ef25932f23e93e344f70490669c65a
14662d6319042accb9e899e2dad9092df2d0c7ea
6439 F20110113_AABHUV grauel_w_Page_155.QC.jpg
eba690677b291bccfc1b85af19b402c0
be3521697c55b296e11818bff0f11fe828597fe2
1646 F20110113_AABIBD grauel_w_Page_094.txt
e396860acbe189a33c809e9992ee61a3
241f0a408e8984e41b208e9a146c9f22ccdd1949
7897 F20110113_AABIAP grauel_w_Page_075thm.jpg
8136a55a013cf3c99bef0b7d4502ee3c
35da7b6bccd6d6ed41d35778e8e0b392868341e5
19931 F20110113_AABHVJ grauel_w_Page_066.QC.jpg
5e95b959c511de5515414ee7c4772b53
be50e787cd90e79d4361da2f289f4bf0309ed797
98774 F20110113_AABJEG grauel_w_Page_113.jpg
36eaab3d7d6c61967596ce59edd21485
20f5b5d891b54a05532d971f9a1d47f56f8d1a55
103222 F20110113_AABJDS grauel_w_Page_062.jpg
bef51a2b4ba86409ffb0639159b2e8d0
16d03e3d1503ba36cdb581a2cf69446f9f9fb51a
101908 F20110113_AABIZA grauel_w_Page_040.jpg
c2389e489ec28486db0571122fc2a6ec
6964e0eac70531c80f8cf751dfe54a3d7b62350d
108719 F20110113_AABIYM grauel_w_Page_056.jp2
e34b3c24f6643689d8dd64aaf48d0946
6ddc51e76f36ea22895197f42784d26d4f5d4b9d
1971 F20110113_AABIBE grauel_w_Page_119.txt
880c7c416524653320a4da1c8ee3fd17
66308df025d388a25e02457007432db5f31b18b8
46729 F20110113_AABIAQ grauel_w_Page_065.jpg
9c8b9b32fb7f513cadce417f22ccb4a8
6c8c2e032fa14eb31c76930e9b0b585e985d23e5
119153 F20110113_AABHVK grauel_w_Page_151.jp2
1697cca8959a4eb1942cca3698808d76
6edc85c60a890532a50cdd57fcf43856804d2521
98781 F20110113_AABJEH grauel_w_Page_117.jpg
f2ab010b94e470f129c2dde4abfe5358
dc3379e57ac65df15513d1b3a10443ed159e4e66
86470 F20110113_AABJDT grauel_w_Page_069.jpg
067957f49eb4826374326dacc5e9d068
ef62b0937b0fd084d8ed70c442d309ba961217cc
62039 F20110113_AABIZB grauel_w_Page_164.pro
44da9e6e9d971eb9ccef5398f4f49739
6fa7fe2e61681376fac94a4999305d86f4593666
105575 F20110113_AABIYN grauel_w_Page_030.jp2
486b698588b958150eb5c0a470c29290
92cbd936f7729bcf2cc16e0d811173df23d0bc89
131896 F20110113_AABIXY grauel_w_Page_162.jp2
a8cab93028f60c38b7e1f2e7622bab8d
de7e78c4a67e98f19486753e663d7fe7b8345f3e
2060 F20110113_AABHUW grauel_w_Page_140.txt
1e5a86bef2a5330261730d7677663b96
a2615a1957dd204615e00f90ea46242c9c499cbd
136597 F20110113_AABIBF grauel_w_Page_168.jp2
67100caaaa893eaf6ed567e9fbceb9ac
74b76328d187623e3e0f7cc91dc6687c35df9580
F20110113_AABIAR grauel_w_Page_039.txt
21d1ad38be226affff47027ad3924c4e
6bbe57801886e8d8977202d0a0f2e37993c6c3a1
35827 F20110113_AABHVL grauel_w_Page_163.QC.jpg
91b2599056eb8ffb512d6dba59e6c0ba
9ceb7d82ffa09c3575de76b3da36f5b3e0e280f2
40413 F20110113_AABJEI grauel_w_Page_126.jpg
8d5c68bd025e718a73eb09d56b83cdd0
dcfd21831d8077395d331f72fae37b5d06110962
103266 F20110113_AABJDU grauel_w_Page_073.jpg
d2d8a9d160aea3a279b42c6bcad19318
fc10c3312b2fa2685a4d5ed861e6b4236c1f222a
100612 F20110113_AABIZC grauel_w_Page_121.jpg
c4eb3dcec5d9e4a2f2b93ab749f3c4bc
a1ecc712799ff8278da94da9a1ddaad884415189
2505 F20110113_AABIYO grauel_w_Page_164.txt
af842202698bf7ac5878a85be6668621
bc8ac6a2abd0d59cef5c3709044955ee063e1d86
106444 F20110113_AABIXZ grauel_w_Page_156.jp2
432c55c849df2db8c2dd97d658845101
30bb7ad4a8594eb530f69c47ac8b90a3322429c6
3093 F20110113_AABHUX grauel_w_Page_128thm.jpg
b3e653487655eb5fa7b44b6935b69a35
c31489b20e996289e4dd7c2cdbe6e0378430a6aa
F20110113_AABIBG grauel_w_Page_025.tif
f342ac86ddb3d5f4976c7a92f4efc115
476bc59cb566631ef02c74c621d36c3c19dcd01e
98883 F20110113_AABHWA grauel_w_Page_083.jp2
770812da5a1d457215d19c9d0851c09d
f97849ab102679f4db1d643933bf37f71b2d99e6
110817 F20110113_AABIAS grauel_w_Page_137.jp2
2d022f55f4aaff24ed1033ae035822cd
7993772ffca9e83c7fad8c2752fcf1032d0928af
610529 F20110113_AABHVM grauel_w_Page_012.jp2
4ac5b63e67fa718ff30aacc994fa0721
e9b97c5966cd8d872c68f730d5956b36e1f19822
43096 F20110113_AABJEJ grauel_w_Page_127.jpg
0723f43c046fd3f3c7d38afde190385c
8f6e50b18ed1b5ab87cb8740b0c5405fdc845a4b
116664 F20110113_AABJDV grauel_w_Page_074.jpg
42043d46c99b2ff7e9fed0d35c71a99d
18a960799b247f25256a8c1048d158244dbf457d
26828 F20110113_AABIZD grauel_w_Page_104.jp2
0120946aa1d69f01b126afc664367257
3c6259408afa9e63bd74c44f32da09ae8fd6ca3d
F20110113_AABIYP grauel_w_Page_003.jpg
840ff428e67d65d081c921f554b0a90d
b9127dadf7db9bbe066ec71aa83104868fb00596
133898 F20110113_AABHUY grauel_w_Page_175.jp2
f3a629648d2aa8d9b7fe23414cb8d3a9
d1cf041f0bd5b97d1c5966ff75e3828b702607fc
99392 F20110113_AABIBH grauel_w_Page_146.jpg
fbcdb28ed566d4f85249f433cfa9f3d9
7c3c056c9db5289ebea2b23765520ec2cc7a6991
32251 F20110113_AABHWB grauel_w_Page_086.QC.jpg
b7afbf875400e9fa91cb9b5c182abc16
2421ee4a3b1bf0513bab22dcdd4c20eb60104bd3
F20110113_AABIAT grauel_w_Page_166.tif
a2025c720c96a5f62a8331bf1771a289
38f150b51ffe673c7ea89c4c541b048b64fc4c01
1509 F20110113_AABHVN grauel_w_Page_048.txt
5db062334d24ba42b4699438e77c4b24
64a13a7bff8357026f13bcda6ed3b2f21d9ff62e
91413 F20110113_AABJEK grauel_w_Page_130.jpg
7fd63f4466802d6cd481e17c34be71e3
645da7a82237d3261d0153f5230084438fd6e7fa
86104 F20110113_AABJDW grauel_w_Page_076.jpg
420bbe187a9c6b6a5ff079af6aecc633
e7fac0b62d84f5199809b46ad1b921e458443c3f
44431 F20110113_AABIZE grauel_w_Page_178.jpg
159d35a489eaffa988c906e5cdd5a123
2965288a88acef16466435c7111bb714a5acd332
F20110113_AABIYQ grauel_w_Page_018.tif
46b6a758501e323fc6049b889b8ba9e1
97cbcbd0f1d4edd7d00a72f5250fd766145a0e5c
51187 F20110113_AABHUZ grauel_w_Page_070.pro
d144fa4cc72fbee1a5f984acec7a9fec
1697a16f1626131073e9360001afb459b63859d8
709 F20110113_AABIBI grauel_w_Page_033.txt
74b9fcb3c6c730be6874406e3d03dd20
55965311fefaeae104dea444c8177a6f98c584c0
8405 F20110113_AABHWC grauel_w_Page_142thm.jpg
7077d1900575bd7b9b23d49a72eeab02
7d9c0dbae7d3209e0719a5fb14e46470e1a017b9
32798 F20110113_AABIAU grauel_w_Page_030.QC.jpg
d31542b0349f3d081c8e2ebcd6dac70f
246ca8f5d41c8d7d548811512507f30d7e3304e4
36391 F20110113_AABHVO grauel_w_Page_015.pro
6f29db6712841d2d7a22091a1fd25515
f2eeef885a448c0a4b3205cee4d6f1a0b42f27ea
211844 F20110113_AABJFA grauel_w_Page_013.jp2
d4bafe16f77b85619e91d0d7b7919f2b
c3feca9e28680898a1a87c11412c922283dd5e5b
102428 F20110113_AABJEL grauel_w_Page_135.jpg
c730cc285e84f3f95176e10c9a1878fb
213e0d589bc35630e7f03cca7fdf0f4aeefc5224
100076 F20110113_AABJDX grauel_w_Page_080.jpg
fc841cab27857ee95e4647cbe65b3e0f
3bcbf8b59f21b6b706e6d538c8c78d9549e4659d
28108 F20110113_AABIZF grauel_w_Page_105.jp2
44f76b3e24d74599714cd0fdfab126de
648a063c35bb2fb09299c2736638bfd98490564b
7575 F20110113_AABIYR grauel_w_Page_052.pro
25f0cc69798319ddc1ba27809a2aa948
2a8998dc0437fb11565b1b0fd384fcd05d67eb20
4433 F20110113_AABIBJ grauel_w_Page_103thm.jpg
e5398eff1c2d19a385ad0bf1037ee929
e53bb5864210e1cc577f42025064c519bd859d85
70777 F20110113_AABHWD grauel_w_Page_101.jpg
f10dd19fcc8900b36af2691b9a1b6ef6
b0f61f5d358b714846dac45eabe7c69f5517f1e1
8155 F20110113_AABIAV grauel_w_Page_079thm.jpg
a51428e93752a880bf3381b07073def4
c1d049ef7a17bc9dc01dbc11454bc52314da25cc
F20110113_AABHVP grauel_w_Page_118.tif
4a25dd51163042fd464f649f896c4601
246fc3a3d71701296385fec83d9f54981f425c44
101403 F20110113_AABJFB grauel_w_Page_019.jp2
678d62619a10cc3d36312a0611d26b13
5402945c16e47949db3574a8e291fa2c3d1d92ec
102619 F20110113_AABJEM grauel_w_Page_139.jpg
db302aaaf3f5e856e50396468e6435df
f9e43d60ee38da3d691820b0a3a52c7e72c3d91d
91780 F20110113_AABJDY grauel_w_Page_083.jpg
63dc9ab70d35c1accd410fcac9363f01
bf5ea07ce085efbe1be7b83e8252cddd550a84e9
1829 F20110113_AABIZG grauel_w_Page_055.txt
2f5c73705746cec0edcc59e82ccf9105
862baaec4a1f1270d942900b6fa1242e9832e5bf
700 F20110113_AABIYS grauel_w_Page_005.txt
8f04ac28e692cf5d5e73d769026736e3
90f1df103eb145bab85369b5890de734c76e823b
834 F20110113_AABIBK grauel_w_Page_178.txt
1deffd558f48e8daf8e824047bf313e3
9382b8be5bb686a26f57834b9b053a152866ad26
102315 F20110113_AABHWE grauel_w_Page_109.jp2
93c34f75e741ba12d3ccee14d1eaf931
9c5774448ee6e1b7154cdb8d294f8c5cf8dc16b5
73586 F20110113_AABIAW grauel_w_Page_148.jp2
c75678ecf928796a51caaa2de8ac364f
d22c50fafaad5b0bb4636b7802ee82b519ea603c
48990 F20110113_AABHVQ grauel_w_Page_045.pro
bbdbbc98697d02ba2f2ab3fedff854d0
ad7ebc788df6264fd8d13b1804ac299859d4a014
98621 F20110113_AABJFC grauel_w_Page_024.jp2
4c894a501482adaecfae90bd52c0fec2
1651b0d15718d9d6451aa20f9359edc411b75ca9
105492 F20110113_AABJEN grauel_w_Page_140.jpg
9032e3061f7d42a7d670fe73b8b66f0c
5170762d470e2b846c83c79238de566aaed0ed00
98036 F20110113_AABJDZ grauel_w_Page_091.jpg
9ad526c5bf83ebcdb67879dc86abaf24
434179a33a0d259cb631ba89b3b3a058c908fd3d
47008 F20110113_AABIZH grauel_w_Page_077.pro
292aae955ba5b24067109670a92ed428
77100e6e2ec305579d686d12bead221f90966f46
31441 F20110113_AABIYT grauel_w_Page_022.QC.jpg
648511a4fb7a5acc153eacb718c8493a
5b4268ba244a99ed4c6e2c8d922f681fd0e51340
7160 F20110113_AABIBL grauel_w_Page_154thm.jpg
10e9660d1142afb690430a677c492b7d
eb1ac7b3282f2799ff43e9e74cece814b5d7b8d6
6009 F20110113_AABHWF grauel_w_Page_002.jp2
900836910369299ae33a3a889ec469fb
4f17e51d05f1222d58bdcb0fea413233fecdd030
6035 F20110113_AABIAX grauel_w_Page_093thm.jpg
96ad254fa0c00f96b444a48f3629431d
805e6d126698b1062a3a502bb20b155ed6ae0cd6
50385 F20110113_AABHVR grauel_w_Page_139.pro
afa6c9671c1ecf24a6ef467a4c726f8d
57cb106490f5b7b3ecd2313480b4d695254d86c6
2074 F20110113_AABICA grauel_w_Page_058.txt
bf39d11ad3a246894870d1a9f16316ee
61ed4553d819d72a75ebaf41bb931bd50a2fb414
42202 F20110113_AABJFD grauel_w_Page_033.jp2
6be387b4d690312277935b80617425f5
9cd5e725be11744a176f0d6e828f00a41ee56e2e
101736 F20110113_AABJEO grauel_w_Page_141.jpg
8bff5b0902683adaad51e1b36c22ef53
1d5f5bbc38cd9b086aae39dabf16bb68d4abdb61
2446 F20110113_AABIZI grauel_w_Page_173.txt
ffb2310705b61c21e303c433bec8a6ec
0c3b174ec069e184cde570b9bfcca3b991f5cd0c
126530 F20110113_AABIYU grauel_w_Page_165.jp2
1a7b73876e9378caf6bb0bc106228809
7d678e504dca9053a60f99fa54805d6b01bc846c
30499 F20110113_AABIBM grauel_w_Page_114.QC.jpg
881166c7d4c83fc428331053bde16100
dd92907597bdab0723ef963b298c97d2c64f8de1
1921 F20110113_AABHWG grauel_w_Page_032.txt
46252646af4a32e4a74eb22d68d02750
72af98f2e8bbbccb6ceb2e10b7eeca136ed59944
6860 F20110113_AABIAY grauel_w_Page_102thm.jpg
f7b4331804ac6700c24d7211bf05aee6
c9dda7e8e99471f3dfb83e493dd4c0851679f272
F20110113_AABHVS grauel_w_Page_059.tif
2bdd309642b0920189cc11a8d0e3b77f
2266657bb7fc855408263f5134a5430cab515a75
31953 F20110113_AABICB grauel_w_Page_143.QC.jpg
ab63045208c9365dc652537f52a74add
36264d80ea515d83dab7373a1cb90c6fcad937cb
113441 F20110113_AABJFE grauel_w_Page_035.jp2
97e83b0dd33e924a35bfed17a0251d07
eab8ba810117053cab5ec7abb8aaf5090f380ba5
67138 F20110113_AABJEP grauel_w_Page_149.jpg
aa1aaa80bd7420dd12673c6a691ebd6b
ce7a61783bcf1bb4751281d31b53ed2555ccc100
1988 F20110113_AABIZJ grauel_w_Page_139.txt
ff75950ac6143ebcafc5590a865f4db9
9bf78a24b39c0999e05cce5d6f308563e2cf407c
F20110113_AABIYV grauel_w_Page_078.txt
7c333ab929cb8cf79967b5a04515aed0
4218c5e9ef43df559425f9bde5d4fe9046670c78
106363 F20110113_AABIBN grauel_w_Page_145.jp2
79331012ef4cb38042b66ba931abfae1
5f1f8f01c289dd2bc5e688ca321081b7a5c93ff7
1679 F20110113_AABHWH grauel_w_Page_153.txt
1033ae9a500d556199e6fd13213bec74
eaa3ffcb0738c03c2d9f3b9d6d9bbab121cf2098
112537 F20110113_AABIAZ grauel_w_Page_020.jp2
e5a5477059934753ec230b4b20daa643
402b1986dd5c079d627658ba2d36f50a62cc7863
79967 F20110113_AABHVT grauel_w_Page_014.jpg
04402e29490e3fe2341a8ca97a6ece02
2d95bc7124ba863d8a3deba81d633be4354b8979
42267 F20110113_AABICC grauel_w_Page_004.pro
2bb942076bafc3e922e632eba72d267e
182953a332ddc2ba7265b5f2e1a1666ae5bdec67
122989 F20110113_AABJEQ grauel_w_Page_152.jpg
a78a8dc5c36a3cbe29dbe9abff88a26b
9060f4d0cc4f22bed8023bc5f911f361e07a18df
85562 F20110113_AABIZK grauel_w_Page_017.jpg
fcd20c72e87f992c29bfd243dbb16b68
469dc70fc979c4e470ebc7a8fb568d4da5adbf5e
1722 F20110113_AABIYW grauel_w_Page_076.txt
600bb6acad3de574eec40770fd6b721a
31f133e0bfdc0e42bbe0110b0e646ab4113e40a8
F20110113_AABIBO grauel_w_Page_136.txt
43de21bd975db7e47408b6ed10bde718
04f8ddb105a5c1a3d6d4296db9ef678062578c76
F20110113_AABHWI grauel_w_Page_174.tif
8c0d612c98a7ddddbcc73dac5cab7718
b0be3b51f4a34b0b373c9ad43a5df02ac3f3c73a
129341 F20110113_AABHVU grauel_w_Page_160.jp2
49e94a9b02d626a77336c212144446d6
1593eafb4ddf9a3dbf891ef00c5f16ef9aa1dcab
106877 F20110113_AABJFF grauel_w_Page_047.jp2
230faf5553ba8e844ac9d317516a5cd1
6871ed70019e36450b59508ff2235d227ce8e7af
112757 F20110113_AABJER grauel_w_Page_158.jpg
49078aa9c1018e6226a8f9d16b1edb39
1d754f16e9253d3488b7b85891c1401df2a7234c
92610 F20110113_AABIZL grauel_w_Page_016.jp2
531b60a7acac9e60346bee23ea332c4a
a0454294255f7d3b015a0305a73325b32e94433d
2426 F20110113_AABIYX grauel_w_Page_167.txt
1b34975e3a5fc94819c8f3d8ee94b893
88b6517742b3817d76acf059ff921827c67b35b4
F20110113_AABIBP grauel_w_Page_033.tif
96fe34373540d72917a11a20ebd0d57b
cc89a10d625785e8bd039f9a2b8064b7b2cfd363
53001 F20110113_AABHWJ grauel_w_Page_152.pro
0ec30363a7e3d69e998bdf4556cb2fb5
bc0b5e357e9c6874b88f0cf7a5bac22fdc5dc2b5
31695 F20110113_AABHVV grauel_w_Page_090.QC.jpg
a54436b3c39be78c049c32b5c44edcfb
e981e9f7851e79f2ddc884b0712267fa0f3599bc
F20110113_AABICD grauel_w_Page_175.tif
17067f2b472656330715bfbaee7ce3a5
218906412efe97bde0d63c43b4f4f0003ced4086
83723 F20110113_AABJFG grauel_w_Page_048.jp2
c40a6f0aebb5f8be80c7e57c5b596f15
e0dc25cd988f38d70f2662ba4bc597c73b3d742b
128925 F20110113_AABJES grauel_w_Page_163.jpg
53ed8aafc07667d701ac056460597ccf
42826f5d3a6215d894d52871ae4223902ddf2b6b
60694 F20110113_AABIZM grauel_w_Page_173.pro
5b51c1f063c7c7c8789d2efeeba77b47
ad10cdae5ddd27f0e9060e23ce4e6bb117723835
7460 F20110113_AABIYY grauel_w_Page_011thm.jpg
74e325c95509e14a7248823cc00530ee
2bcd23143e36b6867782666f6d2edd9ea4d82ac6
F20110113_AABIBQ grauel_w_Page_146.txt
42da6dd7d41a2849b54c133d89512737
2667b2d00eb459a175748f282f511cf90d074bb2
102351 F20110113_AABHWK grauel_w_Page_107.jpg
b9417f74a77827a0ddc0356669a7fcee
03541ef0073a42bc8b3dbfdf83bf629813fae1f5
110148 F20110113_AABHVW grauel_w_Page_063.jp2
bb4ef75f1b87d34aa865d733e231239f
741b20ea63f18b0a659583ed3acef43f9d733056
831 F20110113_AABICE grauel_w_Page_124.txt
26e536e24bf9567c4dfc1f37ab0b53cb
ffee8b97baa2a2cfebcf9a838746bdad4645c610
19805 F20110113_AABJFH grauel_w_Page_051.jp2
57a3147506e0988ad27a5637a304f773
bd4b96d68e64ae8065593f2063aed5a9072ae409
129557 F20110113_AABJET grauel_w_Page_164.jpg
a5de860355e1ffccef109b66b9416c6e
b813188c5f10981d167f792c9ada93f66eb5a013
33145 F20110113_AABIZN grauel_w_Page_060.QC.jpg
8acefd5fe0322b3fd0292a7d0f2c75fb
64473f85ea0ce6d852f697b46f808bd84693b3ee
7800 F20110113_AABIBR grauel_w_Page_041thm.jpg
30a09854257fa3340a7dce30ca4bb5a9
e3c1ea9b7221c29ccdbf4c4ea223bf8450bca46b
51165 F20110113_AABHWL grauel_w_Page_107.pro
72222c406dba57616d0708d0d421006c
6e0ee69484940dfa72889f2b741789bb7f9483c9
1793 F20110113_AABICF grauel_w_Page_016.txt
61ed89f336a540c57b5e28cd92b67aa5
3f02162145784deaf200eae8caeb239be885c7af
97301 F20110113_AABJFI grauel_w_Page_055.jp2
8e10568a96763afd36139f3fe4b04047
f9bc1ab71351a6fb3fd951220a8ab1f320c63d94
129224 F20110113_AABJEU grauel_w_Page_171.jpg
e7e9b48d0ab925a9d4369c421c54094c
a6417a4f1842acdb81fdd4359f6e90efc9439929
33235 F20110113_AABIZO grauel_w_Page_040.QC.jpg
5857dfc7751478ec1eac84d6a6567991
0ebb1d79986ebd163a81aa180c86b460fa31fbfa
28730 F20110113_AABIYZ grauel_w_Page_097.jpg
d072c3e4893ce75223f32e20379ae417
8be796c4c017456f86abe3dba15b68c986fd7372
21521 F20110113_AABHXA grauel_w_Page_065.pro
4fdbdc2022bb814f09cb83c91c67b226
7de50a39e9d83c316c15336263bc143267bde13a
12421 F20110113_AABIBS grauel_w_Page_147.jpg
41637330fe12cc9b3ae2a3a69a664257
1506b98bc430b82464b366e5e6c840bd2bb5e69f
F20110113_AABHWM grauel_w_Page_152.tif
897772d12a47aeb297713f7dd13da800
bc2a886638ef45b326caa5fa55cca729acec4f1a
F20110113_AABHVX grauel_w_Page_089.txt
a60a99250119f927ab0c43e8da9d4a7c
872f45ac0c7c12add168b62395b2ef44c1917ee2
8069 F20110113_AABICG grauel_w_Page_141thm.jpg
dfa2ec1f02eca004441bc2d53293c82f
1273d2041760c4a1ef3c689a3355fef97b13ba55
59966 F20110113_AABJFJ grauel_w_Page_068.jp2
95688be84692dcde06556139fdefb2f5
6e4596091ff2869bc6c6d33d0f6b42555bb9a9f6
114885 F20110113_AABJEV grauel_w_Page_172.jpg
100ed28e4978f04d9523a51227b5cd68
92a0dc815728c8e2594454104d47a3efac9223b7
F20110113_AABIZP grauel_w_Page_131.jp2
f46c814c1b3cf0fff38bbac0dcdd6fa5
57588da1f236632cf4e8ad86e7b0f3dd8a89ba44
F20110113_AABHXB grauel_w_Page_022.tif
303c6e50000c7f1e74be03b77baae63b
8f53cedc1149664b9690c03791b386e5190aa50a
31436 F20110113_AABIBT grauel_w_Page_145.QC.jpg
0d7ac476f5a453746c5d71a852a85241
ba80dbbdb931e846275582d42d3875470e873e77
F20110113_AABHWN grauel_w_Page_068.tif
7b3834a5e664e674772b1d767dee625c
c5029f7f83774cd66934d30f39652fb4600dae81
5295 F20110113_AABHVY grauel_w_Page_068thm.jpg
39fd742015e9ae5606c4f3aed639b1a8
3892b370f4a6e558749a8ba6e722a847e6e959dc
108809 F20110113_AABICH grauel_w_Page_089.jp2
590dd47c1d2139d4fba4e158eda13fff
502509022e5f955abedc1166b9100de5884c399f
92055 F20110113_AABJFK grauel_w_Page_076.jp2
221b10c1040f995289c751bb991a470f
ee7f75e791dfe3e3d6215bfef05d5e5807ba423b
130894 F20110113_AABJEW grauel_w_Page_174.jpg
f0a320ac54679afdeac1d5df772aa04b
52024d174030723f816e355be38bd727d7a1584f
F20110113_AABIZQ grauel_w_Page_110.tif
beb797e77f22a27c507dcbe98b62ca42
74054577b0645456bbcfdb45e170591eae3b25c0
33451 F20110113_AABHXC grauel_w_Page_142.QC.jpg
79d9a91a2b5d0e98868a22b6d37f09c5
572126458887607c08d2ca3a55404a478414a64a
7466 F20110113_AABIBU grauel_w_Page_106thm.jpg
48457d6109e65eb32bb60884d452b01f
c4c75bff8dea8322fd1bd15a8141d0b257d73fca
3060 F20110113_AABHWO grauel_w_Page_007.txt
85ee2ddd7911aea7f3fe33cc5016243b
cf7e2385eeaa1adb419b47a7218955a4c38397fc
133269 F20110113_AABHVZ grauel_w_Page_161.jp2
d78c42189f5fbab0961c18e22103ec97
2911a52fab290c2189edda021fa1f87146823c16
1051984 F20110113_AABICI grauel_w_Page_078.jp2
15323ed4bb9a00a3df116b7d93020234
0ffa1f1c12bb4597639f504827b8a5d121213f21



PAGE 1

ECOLOGY AND MANAGEMENT OF WETLAND FORESTS DOMINATED BY Prioria copaifera IN DARIEN, PANAMA By WILLIAM THOMAS GRAUEL A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2004

PAGE 2

Copyright 2004 by William Thomas Grauel

PAGE 3

Dedicated to the memory of Jack Westoby, who knew that forestry is more about people than about trees.

PAGE 4

ACKNOWLEDGMENTS I express sincere gratitude to my committee chair, Jack Putz, for sharing many insights on ecology, forestry, and writing. I thank my committee members, Eric Jokela and Daniel Zarin of the School of Forest Resources and Conservation, Benjamin Bolker of the Department of Zoology, and Thomas Kursar from the Department of Biology at the University of Utah, for their time and guidance, and I am very grateful to Dr. Stephen Humphrey of the School of Natural Resources and Environment for providing funding at the University of Florida. I particularly thank Claudia Romero for providing me with much of the Colombian literature on these forests. Along with Claudia I am very thankful for friendship and moral support from Geoff Blate, Clea Paz, and Kevin Gould, who, as fellow graduate students, understand. In Panama I received support and guidance from scientists and administrators at the Smithsonian Tropical Research Institute, including Ira Rubinoff, Egbert Leigh, Leopoldo Len, Stanley Heckadon, Rick Condit, Jim Dalling, Elena Lombardo, and Raineldo Urriola. Manuel Rodes and Jos Sols administered the project and allowed me and my co-workers to concentrate on the enormous amount of fieldwork. Among the many people who assisted me in the swamps of Darien, I deeply thank Ricardo Pineda, Ivn Cabrera of La Chunga, and especially Delfn Jaramillo of Tucuti for their staunch and reliable fieldwork. Thanks to the personnel of the Panamanian Environmental Authority in La Palma and Panama City for all their work in keeping the project moving. John iv

PAGE 5

Leigh of the International Tropical Timber Organization agreed to allow me to use project data for this dissertation and provided timely direction during the four-year project. Special thanks to Sarah Dalle for helping me keep faith in the Macondo world that is Darin, and a thank you to Julie Velasquez-Runk for sharing the burden and for finding stuff. Data for this study were collected as part of a project funded by the International Tropical Timber Organization (ITTO). The Panamanian Environmental Authority (Autoridad Nacional del Ambiente, ANAM) and the Smithsonian Tropical Research Institute (STRI) provided logistical support. v

PAGE 6

TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES.............................................................................................................ix LIST OF FIGURES...........................................................................................................xi LIST OF OBJECTS.........................................................................................................xiii ABSTRACT.....................................................................................................................xiv CHAPTER 1 LITERATURE REVIEW OF Prioria copaifera..........................................................1 Introduction...................................................................................................................1 Distribution...................................................................................................................1 Species Description......................................................................................................3 Phenology.....................................................................................................................4 Ecology.........................................................................................................................4 Wood Uses and Properties............................................................................................9 Diseases and Insects...................................................................................................10 Yield...........................................................................................................................11 Growth and Mortality.................................................................................................14 Natural Regeneration..................................................................................................16 Artificial Regeneration...............................................................................................17 Conclusion..................................................................................................................17 2 EFFECTS OF LIANAS ON GROWTH AND REGENERATION OF Prioria copaifera IN DARIEN, PANAMA............................................................................19 Introduction.................................................................................................................19 Study Site....................................................................................................................22 Methods......................................................................................................................23 Results.........................................................................................................................26 Discussion...................................................................................................................28 3 GROWTH AND SURVIVAL OF Prioria copaifera SEEDLINGS PLANTED ALONG A HABITAT GRADIENT IN A PANAMANIAN SWAMP.....................40 vi

PAGE 7

Introduction.................................................................................................................40 Study Site....................................................................................................................42 Methods......................................................................................................................43 Results.........................................................................................................................45 Discussion...................................................................................................................47 4 STRUCTURE, COMPOSITION, AND DYNAMICS OF Prioria copaiferaDOMINATED SWAMP FORESTS IN DARIEN, PANAMA..................................54 Introduction.................................................................................................................54 Study Sites..................................................................................................................57 Principal Sites......................................................................................................58 Secondary Sites...................................................................................................60 Methods......................................................................................................................61 Plot Descriptions.................................................................................................61 Sampling and Analyses.......................................................................................61 Results.........................................................................................................................65 Tree Species Diversity and Stand Structure........................................................65 Cativo Growth, Mortality, and Recruitment........................................................67 Growth of other Tree Species..............................................................................69 Growthdependent Mortality of Cativo Trees....................................................69 Cativo Regeneration............................................................................................70 Discussion...................................................................................................................70 5 GROWTH AND YIELD PROJECTIONS OF Prioria copaifera FROM FOUR SWAMP FORESTS IN DARIEN, PANAMA...........................................................91 Introduction.................................................................................................................91 Study Sites..................................................................................................................94 Methods......................................................................................................................95 Scenarios.....................................................................................................................97 Results.......................................................................................................................100 Discussion.................................................................................................................102 6 GEOGRAPHICAL, ECOLOGICAL, SOCIAL, AND SILVICULTURAL CONTEXTS FOR CATIVO (Prioria copaifera) SWAMP CONSERVATION IN THE DARIEN OF PANAMA.............................................................................115 Introduction...............................................................................................................115 Timber Harvesting in Darien....................................................................................119 Forest Conservation Perspectives.............................................................................123 Conclusions...............................................................................................................129 APPENDIX A MODELING METHODOLOGY USED IN CHAPTER 5......................................133 vii

PAGE 8

B SOURCE DATA FOR CHAPTERS 4 AND 5.........................................................139 LIST OF REFERENCES.................................................................................................141 BIOGRAPHICAL SKETCH...........................................................................................163 viii

PAGE 9

LIST OF TABLES Table page 1. Cativo wood properties............................................................................................10 2. Mean ( SE) annual diameter growth (mm) of Prioria copaifera one, two, and two to four years after liana cutting.........................................................................34 3. Mean canopy openness at 1.3 m above the ground as estimated with a spherical densiometer, arranged by seedling age and habitat..................................................50 3. Initial mean seedling height and diameter (at 20 cm above the ground; standard errors noted parenthetically).....................................................................................50 3. Mean annual growth rates........................................................................................51 4. Total plot area measured for different minimum tree diameters and number of tree species found................................................................................................78 4. Species diversity indices and relative dominance of cativo (Prioria copaifera).....78 4. Stem density and basal area of all species (above) and cativo only (below)...........78 4. Incidence (%) of prostrate, inclined, broken stems and sprouts from prostrate trunks........................................................................................................................79 4. Forestwide annual treefall and tree incline rates (i.e., partial uprooting) for small (above) and large (below) trees for four sites...........................................................79 4. Mean annual diameter growth (mm/year) of cativo trees of three stem types, based on 1997, 1998, or 1997 census periods...........................80 4. Mean annual diameter growth (mm/year) of cativo trees........................................81 4. Ingrowth by stem type. Percentage of recruited individuals from broken stems, undamaged stems, or sprouts from prostrate and inclined trees..............................82 4. Cativo annual recruitment and mortality rates (%) by stem diameter class for four census periods..........................................................................................................83 4. Annual mortality rates (%) of cativo trees by stem type and stature for four census periods............................................................................................84 ix

PAGE 10

4. Mean annual growth (mm/year) of cativo trees that were alive at the end of the study and those that died during the study for which there was one or more years of growth data.................................................................................................85 4. Abundance of cativo trees < 1 cm dbh by height class..........................................86 4. Annual mortality rates (%) of cativo trees by height class for the period November 1998 November 1999..........................................................................86 4. Mean annual height (cm/year) and diameter (mm/yr) growth rates for cativo trees < 1 cm dbh, (sample sizes noted parenthetically).................................86 5. Characteristics of 4 cativo-dominated forests in Darien, Panama.........................109 5. Cativo volume (m 3 ha -1 ) and volume increment (m 3 ha -1 yr -1 ) of four cativo-dominated forests in Darien, Panama.....................................................................110 5. Total volume yield after 65 years of growth and harvest simulations at three different cutting cycles for three riverine swamp forests in Darien, Panama........110 5 4. Percentage total volume reduction due to logging-induced damage for three riverine forests at three cutting cycles....................................................................110 A. Parameter estimates for the four sites....................................................................134 x

PAGE 11

LIST OF FIGURES Figure page 1. Distribution of Prioria copaifera................................................................................2 2. Diameter distributions of ascending lianas in six 25 x 25 m plots in heavily infested riverine Prioria copaifera forest degraded by repeated entry logging........35 2. Mean ( 1 SE) density of Prioria copaifera regeneration (< 1 cm dbh) in areas of high (N = 10) and low (N = 6) liana densities.....................................................36 2. Mean ( 1 SE) Prioria copaifera seedling recruitment censused two years after liana cutting in three control and three treatment plots............................................37 2. Mean ( 1 SE) annual Prioria copaifera diameter growth based on five annual censuses of all trees 4 cm dbh in three control plots and three plots in which all lianas were cut at the beginning of the study......................................................38 2. Mean ( 1 SE) annual Prioria copaifera diameter growth of cativo based on five annual censuses according to liana infestation level in control and treatment plots..........................................................................................................................39 3. Mean diameters and heights ( 1 SE) of planted Prioria copaifera (cativo) seedlings...................................................................................................................52 3. Percent seedling survival, beginning with the first census (November 1997) after planting (September 1997).......................................................................................53 4. Principal study sites. a) Casarete, b) Sambu, c) Juanacati, and d) Naranzati...........59 4. Stem types: a) prostrate and inclined and b) vertical sprouts from a fallen stem.....63 4. Darien Province, Panama showing principal sites (S=Sambu, N=Naranzati, C=Casarete, J=Juanacati) and seven secondary sites...............................................87 4. Annual diameter growth (mm/year)..................... .....................................................88 5. Growth trajectories of cativo at four sites in Darien, Panama starting at 10 cm dbh...........................................................................................................111 xi

PAGE 12

5. Cativo volume projections for three previously logged riverine forests in Darien, Panama...................................................................................................................112 5. Year 2000 dbh for trees as they attain 60 cm dbh during harvest simulation of a 20-year cutting cycle.......................................................................................113 5. Cativo volume projections for an unlogged inland swamp forest in Darien, Panama. ...............................................................................................114 A-1. Quadratic regression curves fit to relative growth data for the four sites...............134 xii

PAGE 13

LIST OF OBJECTS Object page 1 Comma delimited variable Cativo data (as a text file)...........................................139 2 Cativo data (as an Excel spreadsheet)....................................................................139 xiii

PAGE 14

Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy ECOLOGY AND MANAGEMENT OF WETLAND FORESTS DOMINATED BY Prioria copaifera IN DARIEN, PANAMA By William Thomas Grauel August 2004 Chair: Francis E. Putz Major Department: Natural Resources and Environment The state of knowledge of Neotropical swamp forests dominated by Prioria copaifera (cativo) is reviewed based on the available literature. Results of two silvicultural experiments (liana-cutting and reforestation) are reported, forest-stand dynamics are described, and growth and yield projections are presented. Permanent sample plots at four sites in three watersheds in Darien, Panama were installed in 1997 and 1998. Woody plants 1 cm diameter at breast height (dbh, 1.3 m above the ground) were tagged, mapped and measured, and censused annually to gather demographic information on growth, recruitment, and mortality. Cativo regeneration (trees < 1 cm dbh) was also censused at short intervals to monitor seedling dynamics. Cativo growth was studied at an additional six sites along an inundation gradient that varied in water salinity and hydroperiod. Cativo growth was slow to moderate at sites flooded by tidal waters, moderate at inland swamps flooded for long periods, and moderate to fast at riverine sites flooded by xiv

PAGE 15

fresh water; mean annual diameter growth ranged from 0.05 > 0.8 cm/year. Mortality and recruitment rates of cativo varied widely among sites and years as well, with mortality exceeding recruitment at two sites for two of the five years of monitoring. At one slow growing site, trees that died during the study grew significantly slower prior to death than trees that lived, demonstrating the occurrence of species-level growth-dependent mortality for the first time in tropical forest. Unlogged inland swamps contained very large standing volumes in trees 60 cm dbh (up to 180 m 3 /hectare). Riverine forests that were repeatedly logged during 19502000 contained few large trees, but showed potential for future timber harvests in the form of abundant regeneration and large standing volumes in trees 40 cm dbh. Sixty year volume projections suggest that higher dbh cutting limits and longer cutting cycles reduce residual damage, and can produce high timber yields for inland swamps and riverine forests, respectively. Volatile world timber markets and log shortages may be reducing incentives for cativo logging (and thus swamp forest conservation) in the face of large development projects and increased colonization in Darien, Panama. Cativo swamp forests have important hydrological and carbon-sequestering values that should be incorporated in land-use decisions. xv

PAGE 16

CHAPTER 1 LITERATURE REVIEW OF Prioria copaifera Introduction Unlike the vast majority of tropical tree species, much is known about the ecology of the swamp species Prioria copaifera Griseb. (hereafter cativo). Its potential commercial value was recognized in the 1920s, but thousands of hectares of cativo-dominated forests had already been cleared to make way for the banana boom of the early twentieth century. By the end of the century much had been converted to agriculture and of the remainder, most had been cutover and degraded. Distribution Cativo is found from Nicaragua to Colombia, and is also present in Jamaica (Figure 1, Holdridge 1970). Although cativo was included in lists of the tree species of the coastal region of Ecuador by Rimbach (1932) and Acosta Sols (1947) its presence in that country is not confirmed. Barbour (1952) reported that the commercial range includes the Atlantic coast of Central America from Nicaragua to Panama, the watershed of the Bayano River, the rivers flowing into the Gulf of Darien (Golfo de San Miguel), and the area around the mouth of the Atrato River in northwest Colombia. Cooper (1928) referred to enormous stands of cativo in the Valle Estrella of Costa Rica and the Laguna de Chiriqui in Panama. In 1987, it was estimated that cativo forests covered 49,000 ha in Panama, with the major concentration in the easternmost Province of Darien but including eastern Panama Province ( INRENARE, Instituto Nacional de Recursos Naturales Renovables 1987). The same report noted 4000 ha of cativo on Coiba Island 1

PAGE 17

2 and 17,000 ha in the Chucunaque River watershed; 9000 ha in the combined watersheds of the Tuira, Balsas, and Marea Rivers; and an additional 4000 ha in adjacent swampy areas. By 1999, the National Ministry of the Environment in Panama estimated an area of only 15,000 ha of cativo in the country (ANAM 1999a). Figure 1. Distribution of Prioria copaifera. In Colombia, Linares Prieto (1988) described cativos range as including the watersheds of the Atrato and Leon Rivers in the Uruba region in the northwest of the country. According to Escobar and Vasquez (1987), cativo is also found in the Departments of Antioquia and Choco. Tosi (1976) stated that cativo was not found on the Pacific side of Colombia or south of Buchado in the Department of Choco, but Escobar and Vasquez (1987) reported cativo in the low-lying areas around the village of Santa Marta along the Nechi River and referred to a sample of cativo in the Gabriel Gutierrez Medel Herbarium of the Agronomy Department at the National University in Medellin collected in the Department of Magdalena in 1949, all of which are on the

PAGE 18

3 Pacific. Colombia contained 363,000 ha of cativo forest (Linares Prieto 1987b, c, Gonzlez Prez. et al. 1991). Subsequently, the area was estimated at 173,000 ha in 1978 (Linares Prieto 1987b); and in the late 1980s estimates ranged from 60,000 ha (Linares Prieto 1987c), to 90,000 ha (Linares Prieto 1987b), to163,000 ha (Linares Prieto 1988). In Costa Rica, the areas of cativo-dominated forests on the Pacific (Allen 1956) as well as on the Atlantic side of the country (Bethel 1976) have been severely reduced (Veiman 1982) and today the species is listed as threatened in the country (Jimnez Madrigal 1995). Species Description Prioria copaifera is the only species in the genus, and is in the subfamily Caesalipinioideae of the Fabaceae. Average adult height is 25 m, with a dbh (diameter at breast height, 1.3 m) from 45 to100 cm (INRENARE 1987a), although the trees reach 180 dbh (Grauel, unpublished data) and 40 m in height (Del Valle 1972). The branches of mature trees are somewhat arched and the foliage is distributed uniformly in a round, thick crown (Escobar and Vasquez 1987). The bark is smooth, brownish-gray to gray in color, and with abundant lenticels found in continuous horizontal bands (Echavarria A. and Varon P. 1988). Cativo has no buttresses, and its compound leaves generally have four opposite leaflets, elliptic-lanceolate in shape, with a swollen petiole. The leaflets are 16 cm long and 8 cm wide, asymmetric, rounded at the base and with an acuminate apex. The tiny, white flowers develop in panicles (Muoz Valencia 1966). The flowers have 10 stamens and no petals, but the sepals resemble petals (Gentry 1996). The fruit is a flat indehiscent pod, with one side slightly convex and the opposite side concave, 10 cm long, 7 cm wide, 3 cm thick, woody, with a single seed (Mahecha Vega et al. 1984). Seeds are large (mean fresh weight = 48 g, Lopez 2001, to 96 g, Dalling et

PAGE 19

4 al. 1997) and dispersed by water. In addition to the common name cativo, P. copaifera has been known as cautivo, kartiva, trementino, floresa, tabasara, amanza mujer, camibar, murano, and Spanish walnut (Schmieg 1927, Anonymous 1933, Harrar 1941, Hess et al. 1950, Escobar and Vasquez 1987). Phenology Del Valle [cited in Escobar and Vasquez (1987)] noted that cativo leaf-drop is uniform throughout the year and the species is evergreen. Cooper (1928) found that the species flowers generally in March and April and fruits in October and November, but also found a tree with flowers and immature fruits in February. Linares Prieto (1988) reported that flowering typically begins in June and peaks in August and September. Fruiting then begins in September and October and peaks in April and May. In Colombia, seedling recruitment peaks at the beginning of the rainy season from April to June (Linares and Martinez Higuera 1991). Although cativo produces seeds twice a year, large seed crops seem to be produced once every 2 years (Pizano SA 1995, Grauel in review 2004a). Ecology Holdridge (1964) reported that throughout its range from eastern Nicaragua into northwestern South America, cativo is found in the Tropical Moist, Wet, and Rain Forest lifezones. According to Tosi (1976), the cativo forests of Colombia are found in the Tropical Moist Forest and Tropical Wet Forest lifezones of the Holdridge classification system; annual precipitation ranges from 2000 to 8000 mm with average annual temperature of 24 to 28 C (Gonzlez Prez et al. 1991), while in Darien, Panama, cativo is found in the Tropical Moist Forest lifezone (Holdridge and Budowski 1956). Cativo is found in four distinct habitats:

PAGE 20

5 On the Atlantic coast of Costa Rica, Panama, and Colombia, it is found just inland from mangrove forests where salt water does not intrude. Along the many rivers of southern Central America and northwest Colombia, cativo is found in alluvial valleys that are flooded periodically, generally in the rainy season. Away from major rivers cativo is found in low-lying areas that are inundated for extended periods, up to the entire 9-month rainy season in Darien, Panama. Cativo is also found in upland forests, but never as abundant and dominant as in flooded habitats. Although Consultores Ambientales LTDA (1995) declared that cativo cannot germinate or develop in well-drained soil, it is a common component of the upland, mixed-species forest on Barro Colorado Island in Panama (Condit et al. 1993b, 1995a, Sheil and Burslem 2003). Colombian researchers have classified several types of cativales based on landscape position and duration of inundation. Linares Prieto (1988) used landscape position to classify cativo forests as low, medium, and high alluvial plains. The National Corporation for Forest Research and Promotion (CONIF, Corporacin Nacional de Investigacin y Fomento Forestal, cited in Echavarria A. and Varon P. 1988) classified different types of cativo forest according to length of inundation as greater than 6 months, 3 to 6 months, and less than 3 months. Gonzlez Prez et al. (1991), referring to the Instituto Geogrfico Agustin Codazzi cited in Gmez (1990), defined the inundation periods as permanent, 6 to 8 months, 3 to 6 months, and less than three months. In a study of early natural cativo regeneration, Martnez Higuera (1989) defined their two study sites as low alluvial plain frequently flooded and low alluvial plain infrequently flooded. Linares Prieto (1987b) offered various criteria for the classification of cativo-dominated forests: three types of forest distinguished by landscape position (plain, terrace, and alluvial fans.

PAGE 21

6 six types of cativales defined by the combination of life zone and landscape position. number of cativo stems per hectare; median cativo basal area, average distance between trees, number of trees of other species, the number of other species apart from cativo, median basal area of other species than cativo, and median total basal area per hectare. phytosociology of Prioria in terms of abundance, frequency, dominance, importance, and productivity. common associates being Cynometra spp, Pterocarpus officinalis, Gustavia spp, Carapa guianensis, Anacardium excelsum, Eschweilera spp, nunamo (Myristicaceae), Castilla elastica, and Lecythis spp., the number of species increasing with elevation and with better drained soils. In a study that used life zone and landscape position as classification criteria, Escobar and Vasquez (1987) proposed nine types of cativo forest but concluded that their mathematical analysis failed to support the categories. A defining ecological characteristic of cativo-dominated forests is their monodominance or low species diversity within stands. Maximum homogeneity is found in the riverine forests inundated by high Pacific Ocean tides in Darien, Panama. In these sites, cativo can comprise more than 95% of the stems 1 cm dbh and the forest contains less than ten woody species per hectare (Grauel and Kursar 1999, Grauel and Putz 2004). Along the Marea River in Darien, cativo had a relative abundance of 91% in a 10 x 1000 m transect that included five other tree species and one palm ( 10 cm dbh, Mayo Melendez 1965). Away from the influence of tides, floristic diversity increases. Linares Prieto (1988), citing the thesis of Escobar and Vasquez (1987), reported the relative basal area dominance of cativo as 50 to 92%. In Colombia, Gonzlez Prez et al. (1991) found that cativo-dominated forests contain approximately 60 tree species, 15 of which comprise 95% of the individuals and are in the families Fabaceae, Bombacaceae, and

PAGE 22

7 Sterculiaceae, although the minimum diameter of the study was not stated. In cativo-dominated forests in the area of Domingodo-Truando in the Colombian Department of Choco, Consultores Ambientales LTDA (1995) found 86 tree species per hectare (10 cm dbh). In Darien, Panama, Golley et al. (1975) found 44 species of trees per hectare in a cativo-dominated forest along the Chucunaque River. Elsewhere in Darien, Holdridge (1964) found Carapa guianensis to be the only other large tree associated with cativo, although he noted that Pterocarpus officinalis was restricted to the edges of small streams in the same forest. The dominance of cativo in seasonally flooded habitats was suggested as being attributed to the better competitive ability conferred by ectomycorrhizae (EM) compared to vesicular-arbuscular mycorrhizae (VAM, Connell and Lowman 1989); but Torti et al. (1997) showed the existence of VAM in cativo. Lopez and Kursar (1999) demonstrated that 3 tierra firme species survived inundation as well as flood-tolerant cativo, and suggested that a cycle of inundation and drought-induced water stress may better explain patterns of tree diversity than inundation alone. An inventory of animal diversity in a cativo forest in Colombia reported 24 species of mammals, 16 species of birds (principally in the families Sittacidae, Cracidae, and Ramphastidae), 6 species of fish, and 7 species of reptiles (Martnez Higuera 1989). Several of these mammal, bird, and reptile species are becoming increasingly rare due to hunting pressure, including Tapirus bairdii, Mazama americana reperticia, Penelope purpurascens wagler, Crax rubra, Lutra longicandis, and Caiman sclerops (Linares Prieto 1988). Some other studies on wildlife in cativo forests include those of

PAGE 23

8 Mondragn et al. (1994), Ospina Torres (1994, 1995a, 1995b, 1996, 1997), and Orozco Rey (1995). Holdridge (1964) described how cativo forests alternate with monospecific forests dominated by Mora oleifera along the Tuira and Tucuti (Balsas) Rivers. Mora-dominated forests are typically found at slightly lower elevations where the effects of brackish water from high tides are greater, but cativo trees can be found in Mora-dominated forests (Porter 1973). Cativo forests are characterized by a distinct microtopography where adult trees are surrounded by mounds 20 to 30 cm in height and 5 m in diameter (Duke 1964). Duke (1964) also mentioned that fallen cativo trees were very common. Although adult cativo trees are shallow-rooted, in a comparison of cativo-dominated gallery forests with tropical moist, premontane, and mangrove forests. Golley et al. (1969) found belowground biomass to be greatest in mangrove forests, and similar among the other forest types. Overstory biomass, however, was notably higher in gallery (cativo) forests, with over 100 Mg/ha dry weight in stems alone. Holdridge (1964) estimated a leaf area index of 6.1 in a cativo forest along the Tuira River in Darien, Panama. In Colombia, soil fertility of cativo forests varies from very low to moderate (Martnez Higuera 1989) with fine to medium texture, pH from 5.1 to 6.0, and poor drainage in general (Linares Prieto 1987c). Soils of riverine cativo forests in Panama are composed of clay to loamy clay with pH 5.2 to 6.8 (Mariscal et al. 1999, Tapia 1999) while soils of inland swamps are more clayey and acidic (pH 4.4.0, Tapia 1999). The Colombian Institute of Hydrology and Soil Use, cited in Linares Prieto (1988) and

PAGE 24

9 Martnez Higuera (1989), classified soils of cativo forests as Inceptisols (63%) and Entisols (37%). Wood Uses and Properties The value of cativo wood lies in its historic abundance and accessibility, not necessarily in its inherent properties. Early descriptions noted cativo for its cylindrical form and general abundance (Kluge 1926, Cooper 1928) as well as its potential for supplying raw material for architects and interior decorators (Schmieg 1927), although Pittier and Mell (1931) considered the wood to be of little or no use. Later, cativo was included in a series of studies before (Kynoch and Norton 1938) and during WWII (Harrar 1941, 1942a, b) that sought to provide technical information on the physical and mechanical properties of foreign and domestic woods. Further research beginning in 1947, funded by the U.S. Office of Naval Research, resulted in recommendations of cativo for plywood, cabinetry, and furniture (Hess et al. 1950). Similar studies of potential applications of cativo occurred later in Colombia (Hoheisel and Lpez G. 1972, Universidad Nacional de Colombia 1984). Several authors have mentioned the abundant resin that bleeds from freshly cut logs and can make sawing difficult (Cooper 1928, Hess et al. 1950, Barbour 1952, Del Valle 1972). The copious resin of cativo was used by indigenous groups for such diverse uses as repairing boats and for medicine (Cooper 1928, Duke 1986). By using high temperatures during kiln-drying, appreciable amounts of resin can be removed from the lumber with the additional benefit of relieving some of the internal stresses that are caused by the presence of tension wood (Kukachka 1965).

PAGE 25

10 Table 1. Cativo wood properties Author Specific gravity (g cm -1 ) Shrinkage % Harrar (1941) 0.48 9.87 Hess et al. (1950) 0.40 8.9 0.41 sapwood 9.2 sapwood Barbour (1952) 0.50 heartwood 22.9 heartwood Kukachka (1965) 0.40 8.8 Cativo wood properties have been investigated extensively (Kynoch and Norton 1938, Hernandez Hurtado 1984, Jaramillo Gallego and Velasquez Salazar 1992, Escobar C. and Rodriguez 1993). It is moderately light in weight (Table 1) and although it is relatively nondurable with respect to both fungal decay and insect attack, cativo has good dimensional stability and was used as a base for piano keyboards for that reason (Kukachka 1965). Cativo veneer from Panama was marketed in Canada and the United States in 1933 (Anonymous 1933). Large-scale imports of cativo to the United States occurred in the mid to late 1940s from Costa Rica (Hess and Record 1950). By 1952, almost 9500 m 3 /yr were exported from Colombia and Costa Rica to the US, the figure increasing to over 47,000 m 3 /yr by 1958 (Kukachka 1965). While some cativo from the Caribbean side of western Panama was exported, it eventually supplied 90% of the raw material for the domestic plywood industry and 50% of sawn-wood production in the country (FAO, Food and Agriculture Organization 1982). The Panamanian Institute of Renewable Natural Resources recommended using cativo for furniture, packing crates, and cabinetry (INRENARE 1987a). Diseases and Insects Cativo was classified as moderately to non-durable in its resistance to the white-rot fungus Polyporus versicolor (Trametes versicolor) and durable to non-durable for the

PAGE 26

11 brown rot fungus Poria monticola (Hess et al. 1950). Ferrer (1999 ) collected 615 fungi associated with living and dead cativo trees in five different forests, and found 58% Ascomycetes and 42% Basidiomycetes. Apparently, it is not known how many of these are pathogens, saprohytes, or mutualists. In a cativo forest along the Sambu River in Darien, Panama, Ferrer (1999) found that 27% of the Basidiomycetes belong to the genus Phellinus, one of which is among the most important tree pathogens of temperate forests (Phellinus weirri). Hess et al. (1950), Barbour (1952), and Kukachka (1965) mention the susceptibility of the boles of recently felled trees to attack by ambrosia beetles. Insects that perforate recently cut logs belong principally to the families Scolytidae and Platypodidae, and occasionally Brentidae and Tenebrionidae (Romero 1982). The species most commonly found on recently cut logs, but not specific to Prioria, are Platypus parallelus Fabricius and Xyleborus affinis Eichhoff (Estrada Lpez and Gmez Quiceno 1988). Some protection from attack is rendered by direct sunlight, immersion of the logs in water, and the presence of bark; one application of insecticides may prevent attack for 6 to 15 days (Romero 1982, Estrada Lpez and Gmez Quiceno 1988). Yield Early research on cativo as a timber source stressed the large sizes and clear boles of the trees. Cooper (1928) noted an average size of 60 to 90 cm dbh. Barbour (1952) found the commercial size range of the species to be 60 to 120 cm dbh, with maximum sizes of 150 to 180 cm. Barbour (1952) emphasized the straight form of the trunks, without branches for 12 m (and many times branchless up to 30 m in height). From 1951 to 1953 Bruce Lamb studied the forests of Darien for the Panama Forest Products Company to develop log-supply sources and determine available timber

PAGE 27

12 volumes for both upland and lowland forests. Lamb estimated cativo wood volume along the Chucunaque, Tuira, Balsas, Sambu, Congo, and Cucunat Rivers and around the Laguna de la Pita (today called Matusagarat). Along the Balsas River, Lamb found pure stands of cativo for a distance of 20 km and up to a km in width on each side of the river. He estimated an average volume of 71 m 3 /ha and a total of 141,600m 3 for the watershed (Lamb 1953). On the Tuira River, Lamb encountered cativo forests 25 km upriver from the Tuiras confluence with the Balsas River up to the mouth of the Chucunaque River. In this area of approximately 4000 ha, volumes averaged 24 m 3 /ha. Although Lamb did not examine the forests upriver from the mouth of the Chucunaque, there were reports of cativo forests up to Boca Cupe, and he estimated a total of 23,600 m 3 for the Tuira watershed. According to Lamb, the highest-quality cativo wood came from the Chucunaque River watershed, and he estimated a total volume of 47,200 m 3 along the 80-km course of the river. Along the Sambu River, Lamb found cativo 10 km from the rivers mouth at the confluence of the Jess River up to the Sambus confluence with a small stream called Morobichi (8 km further upriver). This cativo forest extended up to 1500 m inland from the river, and Lamb estimated an average volume of 35 m 3 /ha and a total of 23,600 m 3 for the Sambu watershed. Lamb estimated a total of 1,180,000 m 3 for the entire province (Lamb 1953). Cativo forests are known to contain large wood volumes per hectare, but comparison of different estimates is difficult where the minimum diameter is not specified; in addition, some estimates are for commercial volume and others for total volume. In a previously unlogged inland swamp in the Balsas River watershed (near a small stream called Naranzati), 96 m 3 /ha of commercial ( 60 cm dbh) cativo wood was

PAGE 28

13 measured in 1999 (Grauel, unpublished data). Also in 1999, a 100% inventory was carried out of a 50-ha plot along the Sambu River in Darien where cativo comprises 95% of the species diversity. Using a form factor specifically developed for Prioria, a mean volume of 65 m 3 /ha was calculated for trees 60 cm dbh; while across the river in a series of smaller permanent plots, mean volume totaled only 40 m 3 /ha for the same forest type. The difference can probably be attributed to different management histories under different ownership regimes: the latter being found on open-access public land that is subject to frequent, low-intensity timber harvesting by local loggers; while the 50-ha plot is located on land belonging to Embera-Wounaan indigenous communities who harvest cativo much less frequently. When considering a minimum diameter of 40 cm dbh and the commercial height to the lowest branch, this 50-ha plot contains 190 m 3 /ha (Grauel, unpublished data). Recent volume measurements in cativo forests along the Balsas River ranged from 20 m 3 /ha ( 60 cm dbh, Grauel, unpublished data) to 25 m 3 /ha (total volume, Mariscal et al. 1999). In Colombia, Linares Prieto (1987b) stated that a cativo forest contained more than 150 m 3 /ha in commercial wood and a mean of 80 to 100 m 3 /ha for trees 52 cm dbh. In a cativo forest with 60 tree species where cativo comprises 60% of the basal area, Linares Prieto (1988) measured a total volume of 123 m 3 /ha and 46 m 3 /ha in trees 40 cm dbh. In a cativo forest with 5 other commercial tree species in Podega, Colombia Escobar Munera (1981) calculated a mean of 36 m 3 /ha for all species. In another forest inventory of trees 49 cm dbh of 15 tree species, a mean of 7.3 individuals and 27 m 3 /ha, cativo comprised 36% of the total volume (Consultores Ambientales LTDA 1995).

PAGE 29

14 Growth and Mortality As with volume estimates, growth estimates vary and depend on the methodology, age and size of the trees, management history of the forest, and site-specific biotic, abiotic, and climatic factors. Comparisons of growth estimates are difficult where different field methodologies and modeling approaches are used. Like many tropical trees, cativo produces growth rings, but no dendrochronology based on crossdating has been performed to show that the rings are produced annually. Using the pinning technique (Kuroda and Shimaji 1984) where wood is wounded and subsequent growth is measured with destructive harvesting, however, McKenzie (1972) concluded that cativo produces annual rings. I strongly suspect that cativo may produce one or more rings per year. Cativo diameter growth is probably influenced by many variables. Londoo Londoo and Gonzalez Prez (1993) reported that crown area, crown position, and Hegyis diameter-distance competition index had significant effects on growth of cativo in less diverse forests but not in the more diverse sites. In an unlogged cativo forest with 50 to 60 tree species, Del Valle (1979) found that maximum diameter increment was attained by trees approximately 70 cm dbh. Two studies in Panama found maximum diameter increment in medium-sized trees, from 20 to 50 cm dbh depending on the site. In an upland forest in Panama, Condit et al. (1993a) measured maximum annual diameter increments of 2 to 4 cm, while cativo from flooded forests showed maximum annual growth rates of 1.5 to 2.0 cm, with means of 0.6 to 1.0 cm (Grauel 1999). Several modeling approaches have been used to estimate lifetime growth trajectories based on short-term growth rates. Del Valle (1979) used a matrix modeling approach to produce an estimate of 98 years for a 10 cm dbh tree to reach 60 cm.

PAGE 30

15 Gonzlez Prez (1995) developed a von Bertalanffy growth model for cativo, and produced an estimate of 90 years for a 14.5 cm dbh tree to reach 60 cm. In a comparative study of two sites where diameter structure, floristic diversity, spatial distribution, and growth were contrasted, Gonzlez Prez et al. (1991) found growth to be three times greater in the more diverse forests. Although the authors admit to small sample sizes, they estimated 168 years in less diverse forests and 77 years in more diverse forests for a 10 cm dbh tree to reach 60 cm (Gonzlez Prez et al. 1991). There was no difference in growth between diverse and cativo-dominated forests in another study in Colombia, where Linares Prieto (1987b) estimated that a tree would reach optimum harvest size (60 cm dbh) in 55 years in less diverse forests and in 60 years in the more diverse forests. In a different study, the same author estimated that a tree could reach 60 cm dbh in only 38 years, and suggested that this time period could be reduced with adequate silvicultural techniques (Linares Prieto 1988). In a study of growth and yield potential of cativo on an upland site in Panama, Condit et al. (1995b) estimated a period of 130 years for a stem to grow from 1 cm dbh to 60 cm, based on mean growth using data from 1982-1985, and 180 years based on data from 1985-1990. Using the same modeling approach and based on data from 1997-2001, I found two sites with similar inundation regimes to vary considerably; one requiring 315 years, and the other 179 years for a 1 cm dbh stem to grow to 60 cm. At another forest farther upriver, only 80 years would be required for a 4 cm dbh tree to reach 60 cm. For an inland swamp, based on growth data from 1997-2000, 157 years would be required for a 1 cm dbh tree to reach 60 cm, and 186 years to reach 80 cm.

PAGE 31

16 Natural Regeneration In a demographic study of cativo in Colombia, Montero (1996) estimated 30,490 seeds/ha were produced during the 6-month period from December to May. Montero (1996) noted that trees with relatively low seed production tended to produce greater numbers of established seedlings than trees that had greater seed production and hypothesized that there was some optimal period for seedfall that resulted in higher probability of survival and maximum recruitment rates. Linares and Martinez Higuera (1991) found lower densities of cativo natural regeneration in frequently inundated forests than in less frequently flooded forests, although the tendency toward monodominance was greater where flooding was more frequent. Linares and Martinez Higuera (1991) also found a strong correlation between mean monthly precipitation and seedling density, and determined that 2% of the large initial seedfall became established. Lopez (2002) followed a 1997 cohort of cativo seedlings and found 2.5% survivorship after 3 years. Dalling et al. (1997) found that less than 10% of cativo seeds were viable 2 months after seedfall, and that 30% of the viable seeds had suffered damage by insects or pathogens. Even with up to 60% of the seedmass damaged, however, there was no reduction in the probability of germination; and seeds with up to eight insect larvae germinated as often as seeds with no infestation (Dalling et al. 1997). Furthermore, cativo seeds have the ability to produce an average of 2.1 additional, sequential resprouts after the initial sprout is damaged or lost (Dalling et al. 1997). Tamayo Velez (1991) determined that a germinating seed required 48 days to develop into a 29-cm tall seedling, and declared that height growth of cativo plants between 30 and 150 cm in height was 50-60 cm/month. Martnez Higuera (1989)

PAGE 32

17 suggested an annual growth rate of 2.4 m for trees in the same height range. In two strongly monodominant cativo forests in Darien, Panama, density of cativo natural regeneration (trees < 1 cm dbh) varied widely, from less than 5,000/ha at one site to over 17,000/ha at the other. Mean annual height growth for trees between 30 and 150 cm tall varied little, from 2-5 cm, with maximum annual growth rates of 15-25 cm (Grauel, in review 2004a). Artificial Regeneration In a study of artificial regeneration of cativo carried out in Urub, Colombia, Linares Prieto (1987a) tested five planting methods and two planting seasons, and determined that average survival and height were greatest for bare root seedlings planted during the dry season. After 4 years, these seedlings had reached 2.7 m in height (Linares Prieto 1987a). Caycedo (1988), cited by Martnez Higuera (1989), measured annual height growth in seedlings of 70 cm. Cativo seedlings of two ages were planted in three habitats in Darien, Panama. Seedling mortality after 4 years was highest in the natural habitat of the forest understory (89%) and lowest in partial sun on the edge of the forest bordering a treeless marsh (54%, Grauel in review 2004b). Maximum height growth was observed in the full sun, where seedlings of both ages grew approximately 50 cm/yr. Conclusion Relatively few tropical tree species have been studied as extensively as cativo. Because of its accessibility, abundance, form, and wood properties, cativo logging has provided livelihoods for thousands of rural Latin Americans as well as for forest industries in North, Central, and South America. Little research has been carried out on the importance of the ecosystem services that cativo forests provide, however.

PAGE 33

18 Unfortunately, the existence of technical, silvicultural, and ecological knowledge of cativo and the forests where it is abundant has not resulted in sustainable management. As is frequently the case in natural resource management, technical knowledge is insufficient when social and economic forces can influence the strength of forest policies that determine the quality of management carried out on the ground. Nevertheless, technical knowledge can form the foundation for research that links ecological dynamics with the social and economic policies that affect forest management, with the aim of promoting socioeconomic development that conserves natural ecosystems.

PAGE 34

CHAPTER 2 EFFECTS OF LIANAS ON GROWTH AND REGENERATION OF Prioria copaifera IN DARIEN, PANAMA Introduction The abundance and ecological roles of lianas in tropical forests have long attracted the attention of tropical silviculturists (Fox 1968, Appanah and Putz 1984, Chaplin 1985, Putz 1991, Vidal et al. 1997, Carse et al. 2000). Because lianas are a major component of woody plant diversity and provide important food sources for wildlife, they play critical roles in maintaining biological diversity (Nabe-Nielsen 2001, Burnham 2002, Schnitzer and Bongers 2002). Unfortunately, where sustainable forest management is the primary tool for forest conservation and the primary objective is timber production, lianas can be a major impediment. Given that the likelihood of forest conversion to more profitable land uses than forestry is enhanced if prospects for subsequent timber harvests are not economically competitive, liana proliferation can contribute indirectly to forest loss. The large trees that provide the timber value of a forest are more likely than smaller trees to be infested with lianas (Putz 1984, Putz and Chai 1987, Nabe-Nielsen 2001, Perez-Salicrup et al. 2001), and lianas can have various silvicultural implications for forest management. During harvesting operations for example, felling of liana-laden trees can induce excessive stand damage, because their crowns are likely to be connected to their neighbors (Putz 1984). Avoidance of this accessory damage has frequently, but not always (Parren and Bongers 2001), been accomplished through pre-felling liana cutting (Fox 1968, Appanah and Putz 1984, Johns et al. 1996). An additional benefit of pre19

PAGE 35

20 felling liana cutting is the post-harvest reduction in liana proliferation in logging gaps (Alvira et al. 2004, Gerwing and Vidal 2002). Reducing post-logging liana infestations is desirable, because lianas can seriously impede succession in gaps (Schnitzer et al. 2000) and diminish opportunities for rapid recruitment and growth of desirable timber species. In addition to physically impeding establishment of seedlings and saplings of tree species in logging gaps, lianas can reduce host tree fecundity (Stevens 1987), lowering the reproductive output of valuable timber species in forests where natural regeneration is the only cost-effective silvicultural option for stand perpetuation. Heavy liana infestations can also substantially reduce diameter growth of adult trees (Whigham 1984, Gerwing 2001, Clark and Clark 1990), which lowers the net present value of future timber yields by prolonging cutting cycles. Prioria copaifera (hereafter cativo, Fabaceae), a canopy tree found in freshwater wetland forests from Nicaragua to Colombia, has been exploited for timber for decades (Barbour 1952), with little apparent concern for long-term management. Today, commercial stands are found principally in eastern Panama and northwest Colombia. Repeated logging of monodominant cativo stands during 40 years of exploitation testifies to the regenerative capacity of the species. Nevertheless, large areas of cativo forest have been converted to agricultural production or to mixed-species secondary forest and liana tangles as a result of overharvesting. Of the original 363,000 ha of cativo in Colombia for example, less than 90,000 ha remain (Linares Prieto 1987b). Similarly, extensive stands of cativo were once found in western as well as eastern Panama, but today commercial stands are found only in Darien Province. Of 30,000 ha of cativo-dominated forest in Darien in 1987 (INRENARE 1987a), an estimated 15,000 ha remained in 1999

PAGE 36

21 (ANAM 1999a). Increasingly, Panamanian foresters as well as local Darien community members desire to promote sustainable logging of the remaining cativo forests. The stands of almost pure cativo that are found as bands along the principal rivers of Darien vary between 100 m and 1 km in width. Behind the forest, treeless wetlands composed of the palms Elaeis oleifera and Oenocarpus mapora and various lianas including Dalbergia brownei, Combretum sambuensis, Elachyptera floribunda, Tetrapteris macrocarpa, Allamanda cathartica, Phryganocydia corymbosa, Cydista diversifolia, Smilax spinosa, Banisteriopsis spp., and Heteropteris spp. often dominate the landscape. In the absence of silvicultural interventions other than logging, high-statured riverine forests are likely to be converted into palmand liana-dominated vegetation. Present day stand structure of many riverine cativo forests in Darien is a result of traditional logging methods that do not employ heavy machinery (Grauel and Pineda M. 2001). Instead, logs are levered or rolled by hand towards the river on roads constructed from 15 to 30 cm dbh (diameter at breast height, 1.3 m) cativo trunks cut and laid end to end to form two parallel rails. In many riverine cativo forests, the combination of removing all harvestable-size trees as well as many subcanopy individuals for rail building has left a very discontinuous canopy and large multiple-tree gaps, which are habitats favorable for liana proliferation. The leguminous liana Dalbergia brownei proliferates abundantly in disturbed cativo forests. A principal component of the treeless wetlands found behind the natural river levees where cativo dominates, this liana uses cativo forest edges to climb into the forest canopy. Although this species does not establish in the deep shade of the cativo

PAGE 37

22 forest understory, large stems (up to 20 cm diameter) are commonly found hanging from the 30-40 m high canopy in many of the cativo forests of the lower Balsas, Sambu, and Tuira Rivers (Grauel and Pineda M. 2001). Areas with high liana densities seem to have developed in large logging gaps created 20-30 years ago. Many mature cativo trees in heavily infested areas are visibly deformed, apparently from having developed while carrying large liana loads or from having been damaged during logging. In other areas that have been continually and recently subjected to small scale harvesting, D. brownei is proliferating on the ground in large canopy gaps and appears to delay cativo regeneration. In the present study I measured, by observation and experimental liana removal, the effect of lianas on cativo adult stem growth as well as on seedling height growth, recruitment, and mortality. Study Site The study was conducted in a riverine cativo forest along the Balsas River in eastern Panama (8 o 07' N, 77 o 52' W). Mean annual precipitation at Camogant, the nearest town (approximately 8 km from the study site), is 2457 mm (based on Government of Panama published reports for 1978-1982, 1984, 1986, and 1988-1994) while rainfall measured at the study site in 1998 and 1999 totaled 2970 mm and 2758 mm, respectively. The forest is inundated periodically with rainwater during the 9-month wet season from April to December. In addition, it is flooded twice per day for about five days during the monthly spring tides known locally as the aguaje. The freshwater backup caused by the Pacific spring tides affects the riverine forests as far as Camogant, 73 km from the mouth of the Tuira River at the Gulf of San Miguel. Although at the study site the tidal flooding is mostly the freshwater backup, soil samples show a slight brackishness (electrical conductivity 5.0 mmhos/cm) and mangrove forests are found

PAGE 38

23 only 7 km downriver from the study site. Soils at the study site are heavy clays classified in the suborders fluvent and aquept, are acidic to slightly acidic, and poorly drained (Tapia 1999). The study site is on private land owned by a logger and is next to an operating sawmill. The owner, who has been logging cativo in Darien since 1960, is currently logging further upriver and has protected the forest where the study took place because he values it for hunting and aesthetics, although he told us that he had harvested a few scattered trees about ten years prior to the study. This cativo forest is composed of about 95% Prioria copaifera of all size classes (Grauel and Kursar 1999). Other tree species include Pterocarpus officinalis, Mora oleifera, and Carapa guianensis. Results from a 1 ha permanent plot show 10 cativo trees per hectare 60 cm dbh, the legal cutting limit, but the majority of these were left due to bad form or hollowness. Regeneration of cativo of all sizes is abundant. Methods In September 1997 six 25 x 25 m plots were installed in a line at 50-75 m intervals in areas with intact canopies but with relatively high densities of lianas compared to the forest overall. Each plot was subdivided into twenty-five 25-m 2 subplots to facilitate stem mapping. Inside the plots I measured all trees 4 cm dbh as well as the diameters of all ascending liana stems 1 cm at breast height. I did not attempt to differentiate genetically distinct lianas; every stem encountered at 1.3 m above the ground was measured. To increase the sample size for the growth analyses of cativo, additional trees were measured up to 5 m outside of each plot, but no lianas were measured outside the plots. Every plant was tagged and mapped and subsequent censuses were carried out in

PAGE 39

24 1998, 1999, and 2001. For the growth analysis, diameter classes for trees were selected based on relative canopy position; 15 cm dbh was used as the cutoff between canopy and understory individuals. Due to the low canopy of the forest where lianas are abundant, even trees 15-30 cm dbh may receive substantial direct illumination, while trees < 15 cm dbh are generally in the understory. During the initial measurements, each tree was classified as severely or lightly infested by lianas. Severely infested trees had at least five individual liana stems hanging from the crown and some stems or branches apparently deformed by lianas. Lightly infested trees had fewer than five liana stems hanging from the crown and no visible deformations. For the growth analyses, growth rates of liana-free trees were included in the lightly infested category. All lianas were cut with a machete inside and up to 10 m outside of three randomly chosen plots. In 10 randomly chosen 25m 2 subplots in each plot, all natural regeneration of cativo from seedlings to small trees 1 cm in diameter were counted, tagged, and measured (height) before the vine cutting treatment and two years later. Where necessary to reduce heteroskedasticity, seedling frequency data were natural log-transformed. Mean relative height growth for seedlings in treated and control plots was compared with a two-sample t-test and the difference of mean absolute height growth was tested by ANOVA using initial height as a covariate. Mortality of these seedlings and small trees in treatment and control plots was also compared. Two years after the initial census, the same subplots were surveyed for new cativo regeneration. Treatment differences in mean density of seedlings recruited per plot was compared with t tests.

PAGE 40

25 For several reasons, including the observation that increases in cross sectional area of lianas are associated with much larger increases in leaf area than in trees (Putz 1983), it is desirable to estimate diameter growth rates of lianas. In 2001, four years after the initial measurements, 56 Dalbergia brownei lianas in the control plots were again measured to estimate stem diameter growth. Individuals < 6 cm dbh were measured with dial calipers; a mean diameter was calculated from measurements of the long and short axes. Lianas 6 cm dbh were measured with a diameter tape. Wood density was estimated using ten bark-free stem samples, to allow comparisons with other studies. While growth rates of trees are often negatively correlated with wood density, this pattern may not hold for lianas that do not produce structural wood for support. Canopy openness above 2 m was measured immediately before and two months after liana cutting in all plots with a vertical densitometer (Stumpf 1993). Both measurements were made during the rainy season. This instrument projects a point vertically upward that encounters either canopy or open sky at each evenly spaced sample point along a linear transect. Canopy openness is estimated as the proportion of points of open sky along three transects in each plot. To compare rates of cativo growth and regeneration in heavily vine-infested areas with forest with low liana infestation, data from the six plots of the present study were compared with data from plots selected at random for a demographic study of cativo in the same forest. The demographic study was based on five 20 x 20 m and five 40 x 40 m plots established in March 1997. All trees 10 cm dbh were tagged, mapped, and measured (dbh), while trees 1 cm dbh were measured in all five 20 x 20 m plots and in five randomly chosen 20 x 20 m subplots in each of the 40 x 40 m plots. All trees were

PAGE 41

26 measured annually from 1997 to 2001. In addition, all trees < 1 cm dbh in eight randomly chosen 5 x 5 m subplots of each plot were tagged and mapped and were measured (height only) in November 1997. This population of seedlings and saplings was censused approximately every two months for two years whereas height was measured annually. Results Two months after cutting lianas, significant but modest increases in canopy openness were observed in the treated plots. There was no difference in the before and after canopy coverage in the three control plots, while the three treated plots showed a mean increase of 7% (p < 0.01) in canopy openness. Mean annual diameter growth of Dalbergia brownei was 1.3 mm yr -1 (n = 56, sd = 1.4, range = -0.8 to 5.5 mm). Mean wood density of D. brownei (dry weight/fresh volume) based on ten samples was 0.38 g cm -3 (sd = 0.047). Based on the mean number of stems 4 cm from the six 25 x 25 m plots, cativo dominated the forest with 1320 stems ha -1 (sd = 212), virtually identical to nearby areas of riverine forest with lower liana densities (1338 stems ha -1 Grauel and Kursar 1999). Pterocarpus officinalis the only other abundant tree species, was represented by 51 stems ha -1 4 cm dbh. The 35.1 m 2 ha -1 of cativo basal area represents 96.6% of the total basal area of trees 4 cm dbh. For all cativo trees 4 cm dbh, 71% had lianas hanging from the crowns, while 93% of midand upperstory trees ( 15 cm dbh) had lianas. There were 1757 ascending liana stems ha -1 1 cm dbh (sd = 270), with a mean liana basal area of 3.40 m 2 ha -1 (sd = 0.8). Of the two liana species found, Dalbergia brownei comprised 96.9% of the basal

PAGE 42

27 area. The only other liana encountered, Elachyptera floribunda (Hippocrateaceae), was mainly represented by small stems, with 75% of the ascending stems < 3 cm in diameter, while over 80% of the D. brownei stems were 3 cm in diameter (Figure 2). Prior to liana cutting, in the six heavily liana-infested plots the mean density of cativo seedlings and saplings (< 1 cm dbh) was 707 ha -1 (sd = 1154). In contrast, in the ten randomly located plots for the demographic study of cativo at the same site, the mean density of cativo seedlings and saplings was 6350 ha -1 (sd = 12882, Figure 2). Although this was almost an order of magnitude difference and is plainly discernible in the forest, variability was large due to the clumped distributions of seedlings, but the difference was significant (t = 3.12, df = 14, p = 0.008). For cativo regeneration present at the beginning of the study, relative and absolute height growth over two years did not differ between liana-cut and control plots. Although mean initial height for seedlings and saplings happened to be significantly greater in the three control than in the three treatment plots, there was no difference in initial height of only those trees that survived to produce growth records. Cativo seedling and sapling mortality was nearly double in treated than in control plots (63% vs. 36%, Pearson 2 = 6.2, p = 0.01). Cativo seedling recruitment during two years after liana cutting was more than three times greater in the treated than in the control subplots but, due to large variability, was not statistically significant (t = 1.30, df = 4, p = 0.26, Figure 2). On a per ha basis, over 7700 cativo seedlings recruited during two years after lianas were cut compared to just over 2200 seedlings for the control plots.

PAGE 43

28 Mean annual diameter growth of cativo trees during 1997-2001 was about twice as rapid in the liana-cut compared to the control plots (Figure 2). For trees 15 cm dbh the difference was significant (t = 3.41, df = 4, p = 0.03), while for trees between 4 and 15 cm the difference was not statistically significant (t = 2.61, df = 4, p = 0.06). Surprisingly, severity of liana infestation had little apparent effect (no significant differences found) on cativo diameter growth for either control or treated plots (Figure 25). The largest difference was for canopy trees in the control plots, where severely infested trees grew slightly slower than lightly infested trees. Discussion Despite their abundance, liana cutting had only a slight (7%) but statistically significant (p < 0.01) effect on canopy openness of the cativo forest, because most of the liana foliage is displayed on the tops of tree crowns. Liana-infested cativo forests look feathery at the top of the canopy, due to the abundant emergent branches of small-leaved D. brownei searching for higher trellises. Two years after liana cutting, canopy openness was similar for all plots, perhaps because the cativo canopies increased leaf production after liana cutting. In contrast, Gerwing (2001) found that increases in canopy light transmittance persisted for two years following vine cutting, and Perez-Salicrup (2001) measured no change in canopy openness four months after vine cutting in a Bolivian lowland forest but an increase in openness two years later. Both the Bolivian and Brazilian studies took place in much drier forests and probably on less fertile soils than the present study; perhaps the cativo trees were better able to take advantage of the removal of lianas and produce foliage rapidly. The low diameter growth rate of Dalbergia brownei is similar to the growth rate found by Putz (1990, 1.4 mm yr -1 ) for fifteen liana species from a tropical moist forest in

PAGE 44

29 Panama. In a lowland wet forest in Ecuador the most abundant liana, Machaerium cuspidatum had an average annual growth rate of 1.4 mm yr -1 for stems between 30 and 50 mm (Nabe-Nielsen 2002). In contrast to these low diameter growth rates, a single shoot of D. brownei was observed to grow 1.24 m in length in 71 days. Increased mortality of cativo seedlings in liana-cut plots compared to control plots appeared to be due to numerous large liana stems falling from the canopy, most within the first year following cutting. In addition, floodwaters commonly move coarse woody material around on the forest floor, which frequently results in the bending and breakage of seedlings. This additional impact may explain why tree seedling mortality increased significantly following liana-cutting here but not in a tropical tierra firme forest in Bolivia with higher liana densities (Perez-Salicrup 2001). Enhanced seedling recruitment in liana-cut plots (Figure 2) more than compensated for increased mortality of the initially scarce regeneration in these heavily liana-infested areas. Large cativo seed crops were produced in 1997 and 1999, and the germinated seedlings from the May-June 1999 seedfall were captured in the plot census in November 1999 before dry season (January-April) mortality occurred. Three possible mechanisms for this increased seedling recruitment following liana removal include: 1) reduced seed movement out of the liana-cut plots; 2) enhanced seedling survival; and, 3) increased seed production. Reduced seed movement could result because cativo seeds are large (mean fresh wt = 48 g, Lopez 2001) and mainly dispersed by water. Seeds that fall and that are not partially buried in the heavy wet soil may be transported by the monthly spring tides or

PAGE 45

30 the periodic flooding caused by wet season rains. Fallen liana stems could have acted as small dams inhibiting cativo seed movement during inundation. If seed production and retention as well as capture of water dispersed seeds were equal in all plots, high early mortality rates in the control plots would explain the difference in seedling densities between control and treatment plots. But most cativo seedling mortality occurs during the short dry season from January to April (Lopez 2002), and the plots were censused after seedfall but before the onset of the dry season of the second year after liana cutting. Cativo trees newly liberated from lianas may have produced more seeds. A possible mechanism that could help explain an increased production of seeds by cativo is that lianas interfere with cativo flower or seed production, either physically or through competition for light. D. brownei produces long recurved spines that wrap around small diameter objects that they encounter, such as flower-bearing branches. A drain on host resources caused by the constriction of vascular elements in branches or twigs was proposed by Stevens (1987) to explain the negative effect of lianas on the fecundity of Bursera simaruba trees in Costa Rica, and a similar mechanism may be at work in the cativo/Dalbergia canopy. Another possible mechanism for the proposed increased seed production in the treated plots is the removal of belowground competition after the death of the lianas, resulting in increased nutrient availability. Increased water availability may also have been a factor, since seasonally flooded cativo forests can experience severe short-term annual droughts. Putz (1991) suggests that because lianas do not need to produce large diameter structural roots, root systems of lianas may be more efficient in water and

PAGE 46

31 nutrient uptake. Lianas were also experimentally shown to be effective belowground competitors in a study with north temperate vine species (Dillenburg et al. 1993). Although liana cutting resulted in a notable positive increase in Prioria copaifera stem diameter growth, lianas may not be solely responsible for low forest-wide growth rates, as proposed by Gerwing (2001) for a seasonal Amazonian forest in Brasil. In the riverine cativo swamps in Panama discussed in this study, mean annual growth of trees in the control plots was not significantly different than similar sized trees in the nearby permanent plots where lianas were generally less abundant (data not shown). Mean annual growth of cativo trees of all sizes in both treatment and control plots was greatest during the second year following liana cutting and then declined (Table 21). Although the liana leaves fell during the first two months after cutting and the hanging stems fell within the first year following treatment, this pattern of increased growth followed by decline is probably not due to a fertilizer effect from the fallen liana material; forest-wide growth rates for cativo, based on permanent plots at this site and three others in different watersheds in Darien Province, showed the same pattern, suggesting a correlation with climate. The observation that the growth rates of liana-infested control plot trees 15 cm dbh did not differ from growth rates of trees of the same dbh in adjacent permanent plots with few lianas (data not shown) could be attributed to the lower overall canopy height in the area of the liana-cutting plots. With fewer large trees in the liana-abundant areas of the forest, trees that otherwise would be subcanopy individuals may receive more light than a similar sized tree in adjacent non-liana forest. This interpretation suggests that these liana-abundant areas are old canopy gaps in the process of recovery, similar to the

PAGE 47

32 vine-dominated disclimax of Whigham (1984) or the stalled gap of Schnitzer et al. (2000). The profound negative effect of lianas on cativo growth and reproduction probably results from a combination of aboveground and belowground influences. Lianas were estimated to occupy 31% of the forest canopy surface area during the wet season in a seasonally dry tierra firme forest in central Panama (Avalos and Mulkey 1999) and liana leaves might significantly reduce light availability for cativo leaves. Belowground competition for nutrients and water in the dry season could also be a factor. Given that rooting depth in cativo forests is limited by the high water table (Lopez 2002), belowground competition for water may be severe during the dry season because root systems die back during wet season flooding (Lopez 2002). In a semi-deciduous lowland forest in Bolivia, Perez-Salicrup and Barker (2000) found significantly less negative water potentials in Senna multijuga trees where lianas were cut as well as increased tree diameter growth. In contrast, in the same forest after liana cutting, Barker and Perez-Salicrup (2000) found no difference in water status of mahogany trees with and without lianas and concluded that lianas and trees had access to different sources of water due to different rooting depths. Two issues pertinent to considerations of liana cutting as a silvicultural tool are cost of implementation and biodiversity impacts. Based on the experimental plots, it would require 16 person-hours to treat one hectare of forest. Although the experiment took place on private land, the majority of these degraded forests are on state land managed by the Panamanian Environmental Ministry (ANAM). Currently, ANAM is working with several communities in Darien to develop a partnership whereby forest areas are

PAGE 48

33 identified for potential timber production and legal tenure is transferred to local community groups. The government intends to train community members in mapping, inventory, and other management activities; liana cutting could be one of the recommended silvicultural treatments. Conservation of biodiversity is a critical issue in our study area, especially because of the presence of Darien National Park and the current improvement of the Pan-American highway that will doubtlessly increase colonization rates and forest clearing. Liana-cutting in seasonally flooded cativo forests is not expected to have severe effects on species diversity for several reasons. Tree species diversity is extremely low in tidally-flooded cativo forests (Grauel and Kursar 1999) and only one species of liana seems to have proliferated excessively as a result of several decades of uncontrolled timber exploitation. Because this species, Dalbergia brownei, is also a common component of nearby treeless wetlands, attempts at controlling its proliferation in cativo forests suited for timber production probably will never eliminate it. Promoting sustainable management of cativo forests for timber production could actually serve as a biodiversity conservation tool. By providing local communities with viable economic activities in the areas surrounding Darien National Park, pressure to exploit the natural resources of the park could be reduced.

PAGE 49

34 Table 2. Mean ( SE) annual diameter growth (mm) of Prioria copaifera one, two, and two to four years after liana cutting. 1997-1998 1998-1999 1999-2001 Treatment Liana Infestation N Mean Annual Increment (mm) N Mean Annual Increment (mm) N Mean Annual Increment (mm) light 171 1.5 (0.2) 168 2.2 (0.3) 152 1.6 (0.2) Control severe 204 1.6 (0.2) 196 3.1 (0.3) 181 1.8 (0.2) light 166 2.2 (0.2) 164 4.1 (0.4) 144 2.7 (0.3) Lianas Cut severe 180 3.3 (0.3) 172 6.6 (0.4) 154 4.5 (0.3)

PAGE 50

35 0100200300400500600Number of Stems ha-1 Elachyptera floribunda Dalbergia brownei 00.10.20.30.40.50.61-22-33-44-55-66-77-88-99-1010-1111-12>12Diameter Class (cm)Basal Area (m2 ha-1) Elachyptera floribunda Dalbergia brownei b a Figure 2. Diameter distributions of ascending lianas in six 25 x 25 m plots in heavily infested riverine Prioria copaifera forest degraded by repeated entry logging. a) Number of stems per hectare. b) Basal area in square meters per hectare.

PAGE 51

36 High Low Liana Density 0 2000 4000 6000 8000 10000Seedling Density / ha Figure 2. Mean ( 1 SE) density of Prioria copaifera regeneration (< 1 cm dbh) in areas of high (N = 10) and low (N = 6) liana densities.

PAGE 52

37 Control Vines Cut Treatment 0 5 10 15 20 25 30 Seedling Density (# / 25 m2) 10000 8000 6000 4000 2000 12000 0 Seedling Density (# / ha) Figure 2. Mean ( 1 SE) Prioria copaifera seedling recruitment censused two years after liana cutting in three control and three treatment plots.

PAGE 53

38 4-15 > 15 DBH (cm) 0 1 2 3 4 5 6 7 8Annual Diameter Growth (mm) Lianas Cut ControlTreatment Figure 2. Mean ( 1 SE) annual Prioria copaifera diameter growth based on five annual censuses of all trees 4 cm dbh in three control plots and three plots in which all lianas were cut at the beginning of the study.

PAGE 54

39 4 -15 > 15 DBH ( cm) Severe LightInfestation Level 4 -15 > 15 DBH (cm) 0 2 4 6 8 Annual Diameter Growth (mm) Control Lianas cut Figure 2. Mean ( 1 SE) annual Prioria copaifera diameter growth of cativo based on five annual censuses according to liana infestation level in control and treatment plots.

PAGE 55

CHAPTER 3 GROWTH AND SURVIVAL OF Prioria copaifera SEEDLINGS PLANTED ALONG A HABITAT GRADIENT IN A PANAMANIAN SWAMP Introduction Until relatively recently, native tropical species have been little utilized for reforestation programs (Evans 1992). For plantation forestry in the past, a general lack of information led to a reliance on few familiar exotic species with high growth rates. Concern over the loss of biodiversity as well as recognition of other production systems besides monospecific plantations has increased attention on native species for reforestation (e.g., Butterfield 1995, Haggar et al. 1998). Furthermore, much effort has now been expended in acquiring knowledge about growth and mortality rates, as well as propagation techniques, of native tree species in the tropics, often with the specific aim of identifying promising candidates for reforestation (Condit 1995, Foroughbakhch et al. 2001, Wightman et al. 2001, Moulaert et al. 2002). Reforestation with native tropical species may be carried out for timber production (Keenan et al. 1999), site restoration (Parrotta and Knowles 1999, Engel and Parrotta 2001, Montagnini 2001), fuelwood production (Kataki and Konwer 2002), carbon sequestration (Silver et al. 2000), biodiversity conservation (Blakesley et al. 2002a, Blakesley et al. 2002b), or other reasons. For a given end use, species choice should not be based soley on site characteristics or growth rates because the purposes for which trees are planted may vary not only with the type of tree but also with the type of user (Raintree 1991). Certain landowners may be more amenable to reforestation with native 40

PAGE 56

41 species where opportunity costs for labor are low or motivation consists of a wider range of benefits than only financial profitability (Putz 2000). In this study, a valuable native timber tree, Prioria copaifera (hereafter cativo), was planted to evaluate growth and mortality rates in different habitats in swamp forests in eastern Panama. I hypothesized that both mortality and growth would be greatest in the high-light environment and lowest in the forest understory where cativo natural regeneration was abundant. Cativo is a large Caesalpinoid timber tree that has been harvested commercially for decades (Anonymous 1933, Hess and Record 1950, Kukachka 1965). Originally distributed from Nicaragua to Colombia, logging and conversion to banana plantations have severely reduced the area of cativo forests in Costa Rica and western Panama (Veiman 1982, Jimnez Madrigal 1995). Today, commercial stands are restricted to eastern Panama and northwest Colombia. Cativo is known principally from single species or monodominant stands in swamp and riparian habitats, but is also found, less frequently, in upland soils where it sometimes dominates (Condit et al. 1993b). In Darien, Panama, the area of cativo-dominated forests has been reduced to less than 15,000 ha from an original coverage of 60,000 ha (Grauel and Pineda M. 2001). Many riverine cativo forests are adjacent to extensive marshes composed of palms, dense liana tangles, and occasional vine towers where trees still stand from the remnant forest. In the early 1950s Lamb (1953) described a belt of commercial cativo forests along the Balsas River that averaged 1km in depth on each bank of the river, but in 2000 the belt was only 100m deep in some areas. Decades of logging has left many of these cativo forests badly liana-infested and otherwise severely degraded (Grauel and Putz

PAGE 57

42 2004). Hence, an important objective of the study was to determine the feasibility of restoring treeless marshes to cativo forests. Study Site The study was conducted in a forest on private land along the banks of a tributary of the Tuira River, the Balsas River, in Darien Province in eastern Panama (8 o 07' N, 77 o 52' W), 48 km from the mouth of the Tuira River at the Gulf of San Miguel. The landowner has been logging cativo in Darien for forty years, and although he has cut only a few scattered trees at the study site within the last ten years, it is likely that this forest was subjected to an initial selective cutting around thirty years ago. In an early survey of cativo-dominated forests Barbour (1952) noted that commercial-sized trees ranged from 60 to 120 cm in dbh (diameter at breast height, 1.3 m), with occasional specimens of 150 to 180 cm dbh. Today, few large trees are found in these easily accessible riverine forests (Grauel and Kursar 1999). The arboreal component of the study site is composed of 95% cativo (basal area and stems ha -1 ) of all sizes (Grauel and Kursar 1999). Other tree species that occur in the stands include Pterocarpus officinalis, Mora oleifera, and Carapa guianensis. Marshes are found adjacent to the cativo forest and are composed of scattered palms (Elaeis oleifera ) liana tangles, and occasional vine towers, suggesting that the tall forest has been displaced. The lianas of the marsh, principally Dalbergia brownei, climb the tree canopy at the well-defined edge of the forest where there is abundant light, but seldom colonize the understory of the high-statured forest where little light penetrates. Mean annual precipitation at Camogant, the nearest town (approximately 8 km from the study site), is 2457 mm (based on Government of Panama published reports for 1978, 1984, 1986, and 1988) while rainfall measured at the study site in 1998

PAGE 58

43 and 1999 totaled 2970 mm and 2758 mm, respectively. The study site is subjected to periodic flooding from rain events during the 9-month wet season (April-December) and is also flooded when river water is backed up by high monthly tides. The slightly brackish soils at the study site (electrical conductivity 5.0 mmhos/cm) are heavy clays classified in the suborders Fluvents and Aquepts, are acidic to slightly acidic, and are poorly drained (Tapia 1999). Methods In August and September 1997, cativo wildings of two ages were dug up in the forest and planted in three different habitats with distinct light environments. First year seedlings, seeds that matured in May-June of 1997, were identified by having an attached seed and little woody tissue. Individuals referred to as older seedlings had no attached seed and had woody stems. These latter seedlings may have been 3 years or older given that cativo seedlings can survive for a number of years with little or no growth (Lopez 2002). These latter seedlings were likely 2 years old. I tested the effects of 3 levels of canopy cover on two ages of seedlings in four sites (= blocks) separated by 50-200 m along forest-marsh interfaces. In each of four blocks, side-by-side pairs of 7 x 7 m plots were installed in the shade of the forest, on the edge between the forest and treeless marsh, and in the marsh in full sunlight. Site preparation in the marsh involved extensive liana cutting to establish plots and facilitate access but only moderate liana cutting in the edge habitat where liana density was not as abundant due to the partial shade of the adjacent forest canopy. Seedlings were carefully dug up and bare root planted at 1 x 1 m spacing. For each plot pair in a given light environment, seedling age was randomly assigned and 49 seedlings of a given age were planted in each plot. Therefore, for each habitat seedling age combination, 196 seedlings were planted

PAGE 59

44 for a total of 1176 seedlings. In the first plot that was planted, the seedlings (older, shade) were transplanted with entire blocks of soil to protect the roots. This practice proved to be excessively laborious and was abandoned. All subsequent results exclude these 49 seedlings that had high survivorship but little height growth at the end of the study. Canopy openness is defined as the proportion of the sky hemisphere that is not obscured by vegetation when viewed from a single point (Jennings et al. 1999). At the center of each plot, canopy openness at 1.3 m above the ground was measured with a spherical densiometer (Lemmon 1957) by averaging four readings taken in the cardinal directions. After planting in August and September 1997, seedlings were tagged, each stem was marked at 20 cm above the ground, and two perpendicular diameter measurements were taken with Vernier calipers and averaged. Total height of each seedling was also measured. Both height and diameter were recorded at approximately six month intervals for two years; final measurements were made after an additional 20 months had elapsed (July 2001). Repeated measures analysis of covariance was used with either initial height or diameter as the covariate to test the significance of seedling age (between-subjects), habitat (within-subjects), and their interaction. In addition, final mean and maximum height and diameter were each compared between seedling ages within habitats. Height and diameter data were natural log-transformed to comply with the ANOVA assumption of normality. For tests on seedling sizes after the initial measurements, Geisser-Greenhouse adjusted p-values were used because the assumption of homogeneity of variances of treatment-differences was not met (Maxwell and Delaney 1999).

PAGE 60

45 Seedling survival was censused five times in the first two years (1997-1999) with a final tally in July 2001. Survival among the three habitats was compared for each seedling age with one-way ANOVA, then Bonferroni post-hoc tests were used to determine differences among means. All data were arcsine square root-transformed to reduce the unequal variances found in a few plots. Annual rates of absolute and relative growth in height and diameter were calculated for one year, two years, and four years after planting. Growth was first examined for each seedling age by comparing performance among the three habitats with ANOVA and Bonferroni comparisons. Younger and older seedling growth was then compared with t-tests for each habitat. For growth over the entire four years of the study, as well as final seedling size, comparisons could only be made between the edge and sun habitats because of high seedling mortality in the shade. Results Canopy openness among the three habitats ranged from 10% (Table 3). The high variability for the readings in the edge environment resulted from one reading capturing the forest canopy, one capturing the open sky of the marsh, and the other two including both canopy and sky. Younger and older seedlings differed significantly in both height and diameter at the time of planting (Table 3). Older seedlings were twice as large in diameter and about 25% taller than first year seedlings. The repeated measures ANCOVA revealed that only habitat, not seedling age, had significant effects on height and diameter (both G-G adjusted p < 0.001) during the six measurements, and there was no age-habitat interaction (Figure 3). Maximum heights for both seedling ages were found in the full sun, where mean maximum height among

PAGE 61

46 the four blocks for younger seedlings was 382 cm and for older seedlings 350 cm. Within the edge and sun habitats, however, there was no significant difference in maximum or mean height attained between seedling ages at the end of the four year study. Because of high mortality of first-year seedlings in the shade, no comparisons could be made regarding final size attained in that habitat. The planted seedlings were also largest in diameter in the full sun habitat. The only significant difference in all comparisons of diameter and DBH, however, was found in mean stem diameter, where older seedlings were slightly larger than younger seedlings in the sun (38.6 mm vs 32.2 mm, t = 2.5, df = 6, p = 0.047). For the edge and sun habitats, 88% 99% of total four year mortality occurred in the first seven months after planting (Figure 3). Seedlings initially survived better in the shade but at the end of the study mortality was highest for both seedling ages in the shaded understory. Results from the ANOVAs showed that survival was consistently highest for both seedling ages in the edge habitat, but the Bonferroni tests revealed no significant differences in survival at the end of the study between the edge and full sun habitats for either seedling age. Furthermore, for the older seedlings there was no significant difference between survival in the shade and full sun habitats. Variation among plots was fairly modest (Figure 3). Maximum mean annual height and diameter growth for both seedling ages occurred in the sun habitat during 1999, when seedlings grew about 1 cm in diameter (at 20 cm stem height) and from 80 cm in height (Table 3). For each seedling age, absolute and relative height and diameter growth were significantly different in most habitat comparisons. The few exceptions occurred when growth in the edge habitat was

PAGE 62

47 similar to growth in either the full sun or the shade habitat for a few growth intervals. In comparisons of seedling ages within each habitat, annual relative height growth was never significantly different for any growth period. During the 12 months after planting, younger seedlings grew relatively faster in diameter than older seedlings in the shade and edge habitats. In the second year after planting, younger seedlings relative diameter growth was faster only in the shade. Annual relative diameter growth rates for the period 1997 were significantly greater for younger seedlings than older seedlings in both the edge and sun habitats. Discussion This experiment was motivated by the continuing decrease in the area of cativo-dominated swamp forests due to degradation from repeated entry logging. Planting cativo wildings in the understory of the forest provided a baseline for comparison of growth and survival with wildings planted in two habitats that might be considered when the restoration of degraded swamps is a land management objective. There is a tradeoff in terms of growth and survival and the labor requirements of site preparation and maintenance. Initial clearing of dense, extensive liana tangles by hand proved to be much more difficult than clearing areas in the edge habitat, and periodic cleaning was also much easier in the edge habitat because lianas did not resprout as vigorously there. Cativo has often been described as shade tolerant (Linares Prieto 1987a, 1988, Tamayo Velez 1991) or as shade tolerant early in life but requiring high light levels for further development (Linares Prieto et al. 1997). Younger seedlings had much higher mortality in the shade of the forest understory than in the high light habitat, while older seedlings had similar mortality rates in these two habitats, suggesting that seedling establishment requires more than a single year.

PAGE 63

48 Mortality rates observed in this study were generally higher than in a cativo reforestation experiment in Colombia that examined the effects of different types of planting techniques and timing of planting (Linares Prieto 1987a). That study occurred on abandoned agricultural soils while the study site in Darien is less suited for agriculture due to a slight brackishness in the soil and long hydroperiods (Tapia 1999). Although mortality was highest in the shaded understory of the forest, seedling deaths were not soley attributable to shade or root competition. During periodic inundations, fallen branches float around and damage small cativo seedlings. Several planted seedlings in the understory were found bent over completely by coarse woody debris. Liana tangles in open habitats, in contrast, prevent this woody debris from moving. Another agent of mortality present in the forest but not likely in liana-infested areas was trampling by people, particularly hunters. These findings add to the already highly variable published growth rates of cativo seedlings and saplings. Maximum mean annual height growth after four years in the present study was observed in the high light environment, where younger and older seedlings grew 48 and 54 cm yr -1 respectively. These growth rates are similar but predictably lower than those from a reforestation experiment in Colombia on less acidic soils and in high light, where Linares Prieto (1987) reported annual rates of height growth ranging from 67 cm yr -1 In that study, slow plantation height growth was attributed to a lack of suitable mycorrhizae, infertile soils, or poor plantation management. Still, plantation growth greatly exceeds seedling growth in natural forest. In natural cativo-dominated forest in Darien, Panama, a large population (~ 1300 trees) of natural regeneration, from newly germinated seedlings to small saplings 150 cm tall, revealed

PAGE 64

49 essentially zero mean height growth for trees between 60 cm and 150 cm tall. Mean annual height growth of seedlings less than 30 cm tall was 15 cm yr -1 and for seedlings between 30 and 60 cm tall mean annual height growth was 5 cm yr -1 (Grauel 1999). Cativos fastest height growth seems to occur after germination until seed reserves are exhausted (Tamayo Velez 1991). In certain cases cativo may have the desirable socioeconomic and biophysical attributes that would make it the opportune choice for reforestation. Small landholders or cooperatives may have more reasons than just timber production for reforestation. During the course of this study the author was asked by a member of a small-scale loggers cooperative about the feasibility of restoring degraded swamps by planting cativo. A comment was also made regarding the lack of wildlife in the increasingly extensive teak plantations in Panama, implying that native species attract wildlife. Cativo seeds are consumed by agoutis (Dasyprocta aguti) and its leaves by land crabs (possibly Gecarcinus spp), a species that also is harvested by local people. Cativo is the principal raw material for the domestic plywood industry in Panama but is also used locally for furniture and construction; its value lies in its relative abundance and accessibility. Native timber species such as Tabebuia rosea, Dalbergia retusa, Astronium graveolens, and Pachira quinata are currently being planted on a commercial scale in Panama (Mariscal et al. 2002, Wishnie et al. 2002a, Wishnie et al. 2002b), but it remains to be seen if cativo can compete with other, more valuable hardwoods, both native and exotic, for planting on upland soils. This study demonstrates that cativo is particularly suited to reforesting severely degraded sites that are unsuited for agriculture due to flooding and that previously were in cativo.

PAGE 65

50 Table 3. Mean canopy openness at 1.3 m above the ground as estimated with a spherical densiometer, arranged by seedling age and habitat. Seedling Age Habitat % Openness Standard Deviation shade 10.7 0.7 Younger edge 38.7 12.3 sun 83.9 4.1 shade 11.1 1.6 Older edge 27.8 13.4 sun 85.2 9.2 Table 3. Initial mean seedling height and diameter (at 20 cm above the ground; standard errors noted parenthetically). All differences between seedling ages are significant (p < 0.01) based on t-tests. N = 4 plots in blocks 2-4 and N = 3 plots in block 1. Age Younger Older Younger Older Block H e i g h t (cm) D i a m e t e r (mm) 1 41.4 (0.75) 56.9 (1.56) 2.7 (0.05) 5.6 (0.25) 2 40.0 (0.82) 55.1 (1.94) 2.6 (0.08) 5.8 (0.41) 3 43.3 (0.39) 60.5 (2.60) 2.7 (0.02) 5.9 (0.21) 4 50.4 (2.73) 65.8 (1.33) 3.0 (0.19) 6.5 (0.45)

PAGE 66

51 Table 3. Mean annual growth rates. Units of absolute growth are millimeters for diameter and centimeters for height. Comparisons are among habitats within each seedling age for different time periods throughout the study. All comparisons are significantly different (P < 0.05) except where noted, a indicates no difference between edge and sun, and b indicates no difference between edge and shade. Diameter Growth Y e a r 1997-1998 1998-1999 1999-2001 1997-2001 Seedling Age Habitat Relative Absolute Relative Absolute Relative Absolute Relative Absolute Sun 1.27 4.0 1.11 8.0 0.62 9.6 2.44 7.5 Edge 0.83 a 2.6 a,b 0.54 3.0 b 0.28 b 3.1 1.03 3.3 b Younger Shade 0.27 0.7 0.14 0.6 0.04 0.1 0.05 0.2 Sun 0.54 3.5 0.98 9.7 0.51 10.4 1.30 8.3 Edge 0.33 1.9 0.46 3.7 b 0.24 3.2 0.53 3.1 Older Shade 0.02 0.1 0.04 0.2 0.05 0.3 0.05 0.3 Height Growth Y e a r 1997-1998 1998-1999 1999-2001 1997-2001 Seedling Age Habitat Relative Absolute Relative Absolute Relative Absolute Relative Absolute Sun 0.36 18.2 0.45 31.4 0.80 79.7 0.99 48.4 Edge 0.15 b 7.3 a,b 0.18 11.6 0.27 b 22.0 b 0.32 b 16.0 b Younger Shade -0.01 -0.6 -0.10 -5.5 0.04 1.8 0.02 0.8 Sun 0.25 15.8 0.48 39.4 0.74 87.2 0.87 54.0 Edge 0.09 b 4.9 0.25 17.4 0.23 22.5 b 0.26 16.1 Older Shade -0.03 -2.2 0.01 0.4 -0.02 -1.4 -0.01 -0.07

PAGE 67

52 Seedling Diameter SunDate Aug 97Apr 98Nov 98May 99Nov 99Jul 01 Diameter (mm) 01020304050 Younger Seedlings Older Seedlings Seedling Height Sun Seedling Height EdgeDate Aug 97Apr 98Nov 98May 99Nov 99Jul 01 Seedling Height ShadeDate Aug 97Apr 98Nov 98May 99Nov 99Jul 01 Seedling Diameter EdgeDate Aug 97Apr 98Nov 98May 99Nov 99Jul 01 Diameter (mm) 0510152025 Seedling Diameter ShadeDate Aug 97Apr 98Nov 98May 99Nov 99Jul 01 Diameter (mm) 246810 Figure 3. Mean diameters and heights ( 1 SE) of planted Prioria copaifera (cativo) seedlings. Repeated measures analysis showed significant differences in size among habitats but not between seedling ages when initial size (at time of planting) was included as a covariate.

PAGE 68

53 Younger Seedlings Edge Survival % 020406080100 C1 D2 C3 C4 Older Seedlings Edge D1 C2 D3 D4 PlotPlot Younger Seedlings Sun Survival % 020406080100 E1 E2 E3 F4 PlotPlot Older Seedlings Sun F1 F2 F3 E4 PlotPlot Younger Seedlings ShadeDate Nov 97Apr 98Nov 98May 99Nov 99Jul 01 Survival % 020406080100 B1 B2 A3 A4 Plot Older Seedlings ShadeDate Nov 97Apr 98Nov 98May 99Nov 99Jul 01 A2 B3 B4 Plot Figure 3. Percent seedling survival, beginning with the first census (November 1997) after planting (September 1997). Each point pertains to percent survival of the original 49 seedlings planted. Plot letters refer to pairs within habitats (A and B shade, C and D edge, E and F sun), plot numbers refer to blocks.

PAGE 69

CHAPTER 4 STRUCTURE, COMPOSITION, AND DYNAMICS OF Prioria copaiferaDOMINATED SWAMP FORESTS IN DARIEN, PANAMA Introduction Unplanned selective timber harvesting over time results in a pattern of chronic disturbance that strongly shapes forest structure and composition (Kittredge et al. 2003). Timber harvesting is a discrete event that, alone, does not necessarily lead to forest degradation. But where logging is poorly done or is too frequent, forests may become susceptible to fires and liana infestations (Nepstad et al. 1999, Pinard et al. 1999, Gerwing 2002). Given the unfortunate commonness of this disturbance regime, an understanding of forest stand development in response to chronic degradation is critical to the pursuit of sustainable forest management because the diverse values of forests depend largely on forest structure and species composition (Oliver and Larson 1996). The disturbance history of a site substantially influences present day forest stand structure and productivity. Unfortunately, detailed site histories are usually unavailable and evidence of past disturbances may not be obvious. Given that stand structure alone is insufficient to indicate population trends in natural forests (Condit et al. 1998), understanding patterns of recovery in degraded forests requires demographic information as well. Without knowledge of how species, cohorts, and even individual trees respond to disturbances such as logging, predictions cannot be made regarding the likely responses of forest stands to further management interventions. 54

PAGE 70

55 Neotropical swamp forests dominated by Prioria copaifera, a Caesalpinoid legume, were long ago noted for their theoretical ease of management due to their low diversity and the ability of this species to regenerate naturally (Barbour 1952, Holdridge 1964). These traits have apparently allowed Prioriadominated forests to be especially resilient to repeatedentry logging. While potentially easy to manage, managers of Prioriadominated forests suffer from a lack of demographic information. In more diverse forests where trees of most species are scarce, predictions are often based on small sample sizes, although recent innovations such as large plots (Condit et al. 1999) and landscapescale sampling (Clark and Clark 1994, 1999) are addressing this limitation. But with few exceptions (but see Favrichon 1998, Finegan and Camacho 1999, Fredericksen and Mostacedo 2000), most information has been derived from unlogged forest preserves that bear little resemblance to the actual working forests where harvesting occurs. And rarely have studies of a particular forest type been replicated at different sites. Although forest stand structure can reveal clues about disturbance regimes and past uses, simple size distributions of stems often obscure underlying dynamic processes. In particular, resprouting from snapped or partially uprooted trees can greatly influence demographic parameters such as growth, recruitment, and mortality. Although resprouting is increasingly recognized as an important component of forest dynamics, many forest dynamic models omit this mechanism of regeneration and consequently overestimate forest recovery rates (Paciorek et al. 2000). Foresters typically focus on growth rates and on the adequacy of regeneration, overlooking the importance of mortality rates of target species in the development of management plans, often because they lack data. Although correlations have been made

PAGE 71

56 between mortality and stand density (Lugo and Scatena 1996), light environment (Davies 2001), forest fragmentation (Mesquita et al. 1999), and climate (Condit et al. 1995b, Aiba and Kitayama 2002), the estimation of a given species mortality rate is usually difficult due to small sample sizes and limited study periods. Although my study is based on only five annual censuses, I had the advantage of fairly large sample sizes that permitted examination of mortality by stem type (fallen, inclined, resprout from erect stem, stem sprout from prostrate or inclined stem) as well as evidence of growthdependent mortality. Although growthdependent mortality has been reported for temperate forests using growth estimates derived from tree rings (Kobe et al. 1995, Pacala et al. 1996, Kobe and Coates 1997, Wyckoff and Clark 2000, Caspersen and Kobe 2001, Lin et al. 2001, Bigler and Bugmann 2003, van Mantgem et al. 2003), a frequent lack of tree rings, inadequate sample sizes, and insufficient numbers of censuses have precluded investigations of growthdependent mortality in tropical forests (but see Finegan and Camacho 1999 for a standlevel analysis). Studying forests dominated by Prioria copaifera allowed collection of a large dataset for a single, commercially important species and its main associates. In this paper I describe the stand structure of five P. copaifera dominated forests in eastern Panama and report on stand dynamics of four of those five sites based on five years of monitoring data; I also supply tree growth data from an additional six sites. All trees were described on the basis of evident lean, breakage, and resprouting. Because P. copaifera sprouts from erect broken stems as well as from inclined and partially uprooted trees, I differentiate between these sprout types in presentations of data on growth, recruitment, and mortality rates.

PAGE 72

57 Distributed from Nicaragua to Colombia and also found in Jamaica, Prioria copaifera (hereafter cativo) was recognized as a potential source of commercial timber in the first half of the twentieth century (Kluge 1926, Schmieg 1927, Anonymous 1933). During and after WWII, interest in cativo wood increased (Harrar 1941, 1942a, b, Hess and Record 1950, Hess et al. 1950, Barbour 1952). Much of the cativo forest in Costa Rica was exploited to the point of current scarcity (Veiman 1982, Jimnez Madrigal 1995), while in the more remote parts of Panama and Colombia cativo forests were and continue to be subjected to silvicultural experimentation and intensive timber harvesting (Lamb 1953, Mayo Melendez 1965, Christiansen 1980, FAO 1982, INRENARE 1987, Linares Prieto 1988, FAO 1990, Alvarado Q. et al. 1996, CONIF 1997, Mariscal et al. 1999, Grauel and Pineda M. 2001). Although limited in extent, wetland forests dominated by cativo are valued as sources of timber because of their large commercial volumes (Golley et al. 1969, Grauel and Pineda M. 2001) and their ready accessibility by river. Study Sites Permanent plots were installed in 1997 at four sites in three watersheds in Darien, Panama (Figure 4). Small populations of cativo trees at an additional six sites provided additional information about growth of this species under a range of inundation regimes. Eight of the ten sites were previously logged at various intensities and frequencies, most recently three years before this study began. Plots were also installed at a remote site near the Colombian border in Darien National Park where there was no evidence of cativo logging.

PAGE 73

58 Principal Sites Casarete is located along the banks of the Balsas River (8 o 07' 114 N, 77 o 52' 1947W), 19 km upriver from its confluence with the Tuira River and 48 km from the Tuiras mouth at the Gulf of San Miguel (Figure 4). Soils are heavy clays classified in the suborders Fluvent and Aquept, are acidic to slightly acidic (pH 5.2.5), poorly drained, and slightly brackish (electrical conductivity 5.0 mmhos/cm, Tapia 1999); mangrove forests are found only 7 km downriver. Rainfall measured at the study site in 1998 and 1999 was 2970 mm and 2758 mm, respectively. Mean annual rainfall at Camogant, the nearest town, (8.5 km from the study site), is 2457 mm (Repblica de Panama 1995). Because the forest owner values the area for hunting and aesthetics he protected it from logging for approximately 25 years (but did harvest a few large trees approximately ten years before the study). The Sambu River site (8 o 03' 49 06 N, 78 o 13' 16 W) at the mouth of the small Chunga River, is 17 km upstream from the Sambus mouth at the Gulf of San Miguel. This site is approximately 4.5 km from Boca de Sabalo, also on the Sambu River, and 9.5 km from the Wounaan village of Taimati, on the coast of the Gulf of San Miguel, where mean annual precipitation is 1342 mm and 1592 mm, respectively (Repblica de Panama 1995). Soils are similar in texture but slightly less acidic than Casarete (pH 5.7.8), and although it is nearer to the ocean, there was no evidence of salinity (Tapia 1999). The forest is located on openaccess, public land and was repeatedly logged, the most recent entry three years before the study began. Juanacati is 65 km from the mouth of the Tuira River near the town of El Real (8 o 04' 38 N, 77 o 46' 378 W). Mean annual rainfall at El Real (5 km from the site) is 2096 mm (Repblica de Panama 1995). Although Juanacati, like the previous riverine

PAGE 74

59 sites, is flooded by monthly spring tides, no soil salinity is evident and soil pH is higher than the other sites (pH 6.2.3). There is no evidence of recent logging, but anecdotal evidence and the forests proximity to El Real suggest that the site was repeatedly logged in the past. Indeed, L.R. Holdridge noted the presence of stumps only a few kilometers downriver from this site in 1962 (Holdridge 1964). The fourth site is an inland location near a small intermittent stream called Naranzati (8 o 2' 58" 26 N, 77 o 5' 40" 02 W), approximately 7 km west of the town of Camogant. Unlike the three riverine study sites, this forest is flooded for the entire nine month wet season (April to December). Due to its inaccessibility, this site and other inland swamps were subject to logging only recently as riverine cativo forests became increasingly depleted of large trees. a b c d Figure 4. Principal study sites. a) Casarete, b) Sambu, c) Juanacati, and d) Naranzati.

PAGE 75

60 Secondary Sites To compare cativo growth in different landscape positions and flooding regimes, I chose three sites adjacent to mangrove forests and subject to brackish water inundation. Bajo Grande (8 22' 23" N, 78 09' 24" W) is an area near La Palma on the coast of the Gulf of San Miguel; the study site is a few hundred meters inland from the coast, behind the coastal Rhizophora mangle forests. The other two tidal sites are slightly upriver from the transition zone of red mangrove to cativo forest, one along the Tuira River (8 o 10' 1333" N, 77 o 50' 1823" W) and the other along the Balsas River (8 o 09' 037" N, 77 o 53' 0410" W). All three tidal sites are monodominant, unevenaged cativo stands. Additionally, I chose three sites that are unaffected by brackish water for monitoring cativo growth. Two of these freshwater sites are along the Balsas River, between the principal Balsas River site (Casarete) and the community of Camogant (Bongales: 8 o 05' 30" N, 77o 51' 35" W and Limn: 8 o 04' 13" N, 77o53' 16" W). Although these sites are occasionally inundated by high tides, the floodwaters are comprised of freshwater tidal backup and are not brackish. The final freshwater site is an inland swamp far up the Amarraderro River, a small tributary of the Balsas River (8 o 00' 485" N, 77 o 49' 38" W). All six secondary sites, except possibly the latter inland swamp, were subjected to intermittent logging during the last decades of the 20 th Century. A seventh site is found near the headwaters of the Balsas River inside Darien National Park near the Colombian border (7 o 34' 23" 32 N, 77 o 47' 021" W), 130 km upriver from the mouth of the Tuira River. This site was probably never logged for cativo due to its remoteness, a condition that limited my access to a single visit.

PAGE 76

61 Methods Plot Descriptions Because of the low tree species diversity of cativodominated forests, I was able to use small plots and still gather sufficient demographic information for cativo and several common associates. To capture landscape heterogeneity, and because some cativo forests are limited to narrow bands along rivers, many small plots were installed at each site instead of single large plots. Casarete has five 40 x 40 m and five 20 x 20 m plots, Sambu has five 40 x 40 m plots, and at Naranzati there are five 40 x 40 m and three 40 x 20 m plots. At Juanacati, all trees 4 cm dbh were measured in six 50 x 50 m plots. At the other three sites, all trees 10 cm dbh were measured in all plots and trees 1 cm dbh were measured in a randomly chosen 20 x 20 m quadrant of each plot, or in the entire plot if it was 20 x 20 m (Table 4). Most plots were installed in early to mid, but the work was interrupted at Naranzati and Juanacati and was completed in early 1998. All trees were tagged and mapped at the time of plot installation and all plots were subdivided into 5 x 5 m subplots to facilitate mapping. Trees 7 cm dbh were measured with a diameter tape to the nearest millimeter while smaller trees were measured with calipers, with two perpendicular measurements being averaged. Sampling and Analyses To increase the sample size for large cativo trees, additional trees outside the permanent plots were tagged and measured at Casarete and Naranzati. This approach was not feasible at Juanacati and Sambu due to the overall scarcity of large cativo trees.

PAGE 77

62 Three of the eight plots at Naranzati were mistakenly logged two months before the 1999 census. Analyses after that time were done using data from only the unlogged plots. In late 1997 and early 1998 all cativo stems < 1 cm dbh in 80 randomly chosen 5 x 5 m subplots within the larger plots at Casarete and 28 subplots at Sambu were tagged, mapped, and measured (height); dbh was also measured for those saplings > 1.5 m tall. These two populations of seedlings as well as new cativo recruits at these sites were tagged, measured, and mapped approximately every two months for two years and marked seedlings were measured annually. The permanent plots were censused annually until 2001, except the Naranzati site which was last censused in 2000. At the secondary sites trees 20 cm dbh were measured in 1997, 1998, and 1999, except the Amarradero site, which was logged after the 1998 census. Intercensus intervals were always nearly annual to minimize seasonal effects on growth. Methods of censusing and measuring trees as well as the approach to data checking generally followed Condit (1998). During each census, in addition to recording each trees status (alive, dead, recruit), diameter (trees 1 cm dbh), and height (trees < 1 cm dbh), codes were assigned to denote if a stem was prostrate, inclined > 45 from vertical but not lying on the ground, broken above or below the point of measurement (POM), resprouting from a broken stem, or sprouting from a prostrate or inclined stem. I report the incidences of these stem types, but for growth analyses I exclude fallen stems and combine erect and inclined stems. I refer to stems that show evidence of previous breakage and subsequent resprouting as broken stems. Although I separately coded trees

PAGE 78

63 that had broken depending on if the break was above or below the POM, in this paper I combine the two because there were only a few cases where a substantial recorded loss in diameter was the result of stem breakage and subsequent resprouting. Living fallen stems were measured if possible but were always tagged and mapped because they served as hosts for vertical sprouts. References to prostrate stems refer to uprooted, not snapped, stems. I measured all vertical sprouts from prostrate stems and from inclined stems if they emerged from < 1.3 m from the ground. a b Figure 4. Stem types: a) prostrate and inclined and b) vertical sprouts from a fallen stem. Two professional foresters performed all the ~ 22,000 dbh measurements in the permanent plots. One forester measured trees only in 1997 and 1998, while the other (the author) measured half the trees in 1997 and 1998 and all trees from 1999-2001, overall measuring 80% of the trees in the study. I was also responsible for all error checking and database management. All data were independently entered into a computerized database by two different people immediately upon return from the field. Any discrepancies between the dbh measurements that could not be corrected in the office were noted, and follow-up field trips a few weeks after the principal census were carried out to remeasure trees or confirm codes.

PAGE 79

64 Within sites, I first examined variability among plots of each years annual cativo growth of erect, broken, and vertical sprouts with ANOVA and Tukey posthoc tests. Data were natural logtransformed if variances were substantially unequal. For these comparisons, the longest annual growth record available was used. Of 3065 cativo trees in the four principal sites, 80% of the annual growth records were for the period 19972001, 14% were for 1998, and 6% were for 1997. For cativo, growth was then compared among normal stems, broken stems, and sprouts from inclined/prostrate stems within five diameter classes using ANOVA and Bonferroni posthoc tests. To explore interannual growth of cativo I used five diameter classes and I combined all stem types but excluded those that were prostrate on the ground. I also report annual diameter growth of cativos three principal arboreal associates. I report annual recruitment and mortality rates for cativo at the four principal sites using four diameter classes for trees 1 cm dbh and five height classes for smaller trees at the two sites where cativo regeneration was studied. In each census I used the totals of stems from the previous census, that is, I do not calculate demographic parameters using only the originallytagged 1997 population (see Sheil and May 1996). The monitored large trees outside the plots at Casarete and Naranzati were included in calculations of mortality rates. Furthermore, in the few cases where trees were not found in a particular years census, those trees were subtracted from the previous years total number of trees to exclude them from calculations of recruitment rates, instead of assuming the trees had died. I evaluated the extent to which cativo mortality varied with recent growth rates. With five censuses at most sites, I was able to compare annual growth for a maximum of

PAGE 80

65 three years between trees that were alive at the end of the study and those that died during the study. I used two size classes (< 10 and 10 cm dbh) and performed onetailed ttests on populations with at least five dead stems to test the hypothesis that slower growing trees suffered a higher probability of mortality. I report species diversity using Fishers alpha and the ShannonWeiner (SW) index in two ways, first with only all stems 4 cm dbh so that all four sites can be compared. Then I also calculated the SW index for the three sites with minimum dbh of 1 cm. Voucher specimens were collected from the four principal sites in 2000; unidentified species were not separated into morphospecies; I noted the number of unidentified species at each site and considered all unknown trees as a single species in the diversity calculations. Results Tree Species Diversity and Stand Structure Although cativo dominated all four principal sites, species diversity varied substantially. The riverine sites on the Sambu and Balsas (Casarete site) Rivers showed the greatest cativo dominance, with cativo comprising 95 and 96% of the stems of all size classes, respectively (Table 4). Diversity indices grouped the four sites into pairs, with Casarete and Sambu being strongly monodominant and Juanacati and Naranzati being relatively more diverse. Only seven tree species were tallied at each of the former two sites, while 48 and 54 species were identified at Juanacati and Naranzati, respectively. For trees 10 cm dbh at the five sites with plots, cativo comprised from 46% of the stems and from 33% of the basal area (Table 4). Pterocarpus officinalis was cativos principal associate common to all four sites, making up 2% of the stems. Other common overstory trees at Juanacati were Pentaclethra macroloba (10.2%),

PAGE 81

66 Carapa guianensis (7.2%), Licania platypus (3.3%), and Mora oleifera (1.3%). At the Naranzati inland swamp, cativo tended to dominate the overstory with P. officinalis and P. macroloba, but understory associates included Andira inermis (1.3%), Eschweilera integrifolia (4.7%), Gustavia nana (2.0%), and Brownea rosademonte (2.5%). The palms Oenocarpus mapora and Astrocaryum standlyanum were common understory species at the two more diverse sites as well, comprising 2% of the stems. Stand density and basal area varied considerably among sites (Table 4). Stand density of trees 10 cm dbh ranged from 328 trees/ha (Darien NP) to 757 trees/ha (Casarete). Stand basal area varied less markedly, but most cativo basal area was represented by trees 60 cm dbh at the inland swamp (Naranzati) and in Darien NP, and by stems 10 cm dbh in the riverine forests. Cativo made up ~ 95% of stand basal area at the riverine Sambu and Casarete sites and ~ 83% at the inland Naranzati swamp (Table 4). The Juanacati site was visually dominated by 14 huge, emergent Mora oleifera trees per hectare that made up 21% of stand basal area, the same percentage as Pterocarpus officinalis, while cativo comprised 33% of basal area and Pentaclethra macroloba 14%. For all species, substantially more large prostrate and severely inclined trees were found at the three riverine sites than at the inland swamp or the remote Darien NP site. No prostrate, living trees were noted at either Naranzati or Darien NP. In contrast, large prostrate stems were fairly common at the three riverine sites, where 1% of live stems 10 cm dbh were on the ground (Table 4). Large inclined stems were also more prevalent in the riverine forests, where 4% of all stems were partially uprooted and leaning > 45.

PAGE 82

67 The rates at which trees partially uprooted and fell to the ground or leaned > 45 were generally higher at all sites for trees 10 cm dbh than for smaller trees (Table 4). Consistent with the proportion of live fallen and inclined stems recorded at the beginning of the study, rates of falling and inclination were higher at the riverine sites than the inland swamp, but rates varied greatly among years within sites. At the time of plot installation a modest proportion of stems showed signs of previous stem breakage at most sites. Where trees < 10 cm dbh were measured, between 6% were broken, and approximately 3% of large stems had suffered but recovered from stem breakage. No broken and resprouted stems were noted in the plots at Darien NP. Sprouts from prostrate and inclined stems were much more prevalent in the riverine forests than inland swamps. Less than 2% of small stems (< 10 cm dbh) at the inland Naranzati swamp consisted of these sprouts, and no larger sprouts were found. In contrast, 6% of smaller stems were sprouts at the riverine sites. Among the riverine sites, Casarete stood out by having more large than small stems classified as sprouts, where 12% of all stems 10 cm dbh were sprouts from prostrate or inclined parent trees. The 23.5 m long stem of a 33 cm dbh cativo that partially uprooted and fell to the ground at Casarete between the 1999 and 2000 censuses produced 175 vertical sprouts 1.5 m tall by the 2001 census. Fifteen of these shoots were inside the plot (within 5.7 m from the root system of the parent tree) and were tallied as recruits in 2001, with mean and maximum diameters of 1.7 and 3.2 cm dbh, respectively. Cativo Growth, Mortality, and Recruitment Relative rates of cativo growth of three stem types (undamaged, broken/resprouting, or sprouting from inclined and prostrate trunks) are unique to each

PAGE 83

68 site. Sprouts grew faster than either undamaged or broken stems at Casarete for stems <40 cm dbh, while at Sambu the smallest sprouts grew slower than undamaged stems and only sprouts 4 cm dbh from inclined or prostrate stems grew faster than undamaged stems of the same size. Undamaged stems grew faster than broken stems at Juanacati but growth rates of normal stems and sprouts from prostrate/inclined trunks were similar (Table 4). Cativo annual diameter growth varied considerably among both sites and years, but a few patterns were apparent. Growth was slowest, with some exceptions, during the El Nio year of 1997 and fastest for the following census period (1998). For trees < 10 cm dbh, the Casarete and Naranzati sites were both characterized by very slow growth rates, while stems of this size class grew significantly faster in almost all years at Sambu and Juanacati (Table 4). Mean diameter growth rates within the larger size classes ( 10 cm dbh) varied notably among the 9 sites (10 sites for 1997), ranging from only 0.5 mm to > 8 mm per year. Sprouts from prostrate or inclined stems made up a significant portion of forestwide recruitment in many cases (Table 4). About 50% of saplings entering the 1 cm dbh size class at Casarete were sprouts of this type. Also at Casarete, these sprouts consistently comprised substantial portions of the recruitment into the 10 cm dbh size class. Recruitment and mortality rates for cativo show similar intersite patterns to those observed for growth. Mortality equaled or exceeded recruitment at Casarete and Naranzati for trees <10 cm dbh in the first two years of the study (Table 4). In general,

PAGE 84

69 recruitment of cativo at Sambu and Juanacati greatly surpassed mortality for all size classes and years. Mortality rates of different cativo stem types varied among sites, but in general, sprouts from prostrate/inclined trunks and undamaged stems showed the highest mortality rates at Casarete and Sambu (Table 4). No sprouts from prostrate/inclined trunks died at either Juanacati or Naranzati; at these two sites either broken or prostrate stems had mortality rates that approached or occasionally exceeded those observed for undamaged stems. Growth of other Tree Species Pterocarpus officinalis was the fastest growing tree species at all four sites, with mean annual diameter growth of 9 mm for trees 10 cm dbh in three of the four forests (Figure 4b). The other two most common associates of cativo, Carapa guianensis and Pentaclethra macroloba had mean annual growth of large trees 10 cm dbh that approached 5 mm at the two more diverse sites where they were found. Growthdependent Mortality of Cativo Trees Surviving trees grew significantly faster (p 0.05) than trees that died for nine of ten ttest comparisons (Table 4). The nonsignificant case revealed significance when the size class was further divided. For 1997 growth, survivors at Casarete <10 cm dbh grew equally slowly as those trees that subsequently died, but when these smaller trees were analyzed as two size classes, surviving trees between 1 cm dbh grew significantly faster than those trees that died (mean 0.1 vs. .1 mm, df = 90, t = 3.1, p = 0.002), while trees 4 cm dbh in both groups had essentially zero growth.

PAGE 85

70 Cativo Regeneration Regeneration was much more abundant at the more recently logged Sambu River site than at Casarete (Table 4). Small seedlings (< 30 cm tall) were not abundant at either site, suggesting that they grow rapidly after germination. Seedlings 30 cm tall were most abundant at Casarete, while at Sambu seedlings 60 cm tall were the most common. Due to a large seedfall in April and May 1999, annual cativo seedling recruitment rates based on the period November 1998 November 1999 for Casarete and Sambu were 136.6% and 29.1%, respectively. Annual mortality for the same period was similarly high for seedlings < 30 cm tall but differed substantially for taller trees between the two sites (Table 4). Mean annual height growth was generally < 5 cm at each site (Table 4). Exceptions were the relatively small number of seedlings < 30 cm tall at both sites and saplings > 90 cm tall at Sambu, both of which grew rapidly. Discussion Forests dominated by cativo in Darien, Panama vary substantially in structure, species composition, and stand dynamics. Flooding regimes and management histories appear to be the main determinants of presentday structure and dynamics. Slight soil salinity, in particular, seems to favor cativo dominance (Mayo Melendez 1965). Cativo forests near mangrove forests exhibit almost total dominance by cativo, whereas inland swamps and riverine forests that escape tidal flooding with brackish water contain relatively high tree species diversity. Cativo dominance can probably be attributed to a variety of mechanisms. Flood tolerance alone is an insufficient explanation because other flood tolerant species are generally rare in cativo forests (Lopez and Kursar 1999). Cativos root system dies back to a much lesser extent than other flood tolerant species and gives the species competitive

PAGE 86

71 advantage in seasonally flooded forests that are subject to short but severe annual droughts (Lopez 2002). High leaf area index, as shown by Holdridge (1964), may modify the understory environment to be more favorable for cativo seedling survival than for other species. Such a modification was attributed to Gilbertiodendron dewevrei, a tropical tree species that forms monodominant stands in West Africa. Although the crab species common to riverine cativo forests have not been studied in Darien, the land crab Gecarcinus quadratus was shown to affect species diversity in a coastal Costa Rican forest by selective seedling consumption (Sherman 2002). The degree of cativo dominance varies widely in Panama. Cativo is a locally common species in the 50 ha forest dynamics plot on Barro Colorado Island, with a mean of 27 trees ha 1 cm dbh and a maximum of 223 (Condit et al. 1993b). The three sites in Darien for which I have comparable data have densities of cativo 5 times greater than on BCI. Similar to cativo swamps in Darien, the relative basal area dominance of cativo in Colombia is 50% (Escobar and Vasquez 1987). Anecdotal evidence suggests that Carapa guianensis, a species that produces wood of a similar quality as mahogany, was probably much more common in the past in some cativo forests and L.R. Holdridge (1964) identified C. guianensis as cativos sole associate in a 1962 transect very near the Juanacati site on the Tuira River. The histories of timber harvesting of these forests undoubtedly influence present day stand structure and dynamics, but details on logging frequencies and intensities are largely unknown. It is likely that all the riverine forests were repeatedly logged since the 1950s (Repblica de Panam 1978). Although I characterized the inland swamps and the

PAGE 87

72 Darien NP site as intact forests, it is possible that even these remote forests were logged for mahogany several decades ago. Stand structure analyses of cativo forests reveal high timber volumes or at least the potential for high volume production. The forest at Darien NP stands apart from all other sites with the lowest density of stems but the largest stand basal area. The other sites (except Juanacati) contain higher basal areas than most other tropical forests (Leigh 1999). This is notable because the riverine cativo forests of Darien were identified for their timber potential in the 1950s (Lamb 1953) and L.R. Holdridge noted stumps from harvesting activities in the early 1960s near the Juanacati site (Holdridge 1964). The history of logging is undoubtedly responsible for the paucity of trees 60 cm dbh (the legal cutting limit), but these riverine forests still contain a similar number of stems 10 cm dbh as most other lowland tropical forests (Leigh 1999). High stem density at Casarete may be a result of the sites management history. Having been protected by its owner from the repeated logging that typically occurs on open access state land, this forest may have passed through a period of enhanced recruitment after the first wave(s) of logging in the 1950s and 1960s. Increased recruitment and growth of undamaged residual trees after logging is a welldocumented phenomenon (e.g. Magnusson et al. 1999, Parrotta et al. 2002). The Casarete forest 40 years ago may have been similar to the presentday Sambu forest that was recently logged and exhibits high seedling and sapling densities, fast sapling growth, and low sapling mortality. Sprouting is a wellrecognized regeneration strategy in tropical forest (Putz and Brokaw 1989, Rijks et al. 1998, Gavin and Peart 1999, Kammesheidt 1999, Negreros

PAGE 88

73 Castillo and Hall 2000, Yamada et al. 2001) but sprout density varies from being common (Paciorek et al. 2000) to absent in mature forests (Kammesheidt 1998). Vegetative sprouts from various sources may be the principal colonizers of gaps (Putz and Brokaw 1989, Negrelle 1995), but mortality rates of resprouted broken stems are generally higher than nonsprouts (Guariguata 1998, Paciorek et al. 2000, Ickes et al. 2003). In cativo dominated forests it is important to distinguish between resprouts originating from erect, broken trees and those that emerge from fallen and inclined trees. Sprouts from fallen or inclined trunks are frequently much more common, often grow more rapidly than trees originating from seed but have the highest mortality rates of any stem type, including trees that have uprooted and are lying on the ground. The tree recruitment assemblage in newly formed treefall gaps in cativo forests may not be dominated by true seedlings. In some cativo forests the principal gap colonists were not newly germinated seedlings or established shade tolerant saplings; instead, regeneration was dominated by sprouts from inclined or prostrate stems. For example, at least half the recruitment of 1 cm dbh stems every year at Casarete was comprised of sprouts from fallen stems, which also grew faster than their conventionally rooted counterparts. Sprouts from prostrate stems develop their own root systems composed of roots that emerge from the bottom of the parent stem. At Casarete, these sprouts continued their superior growth rates at least into the subcanopy after which it was difficult to determine their mode of regeneration. The three riverine sites share a somewhat similar history of logging as well as a higher proportion of live prostrate and inclined trees than the inland swamps that have not been logged. Logging and inclined or prostrate trees may not necessarily be related,

PAGE 89

74 however. Riverine cativo forests were characterized by a large number of fallen trees at about the time that widespread commercial logging was beginning in Darien (Duke 1964, Holdridge 1964), but recent logging has been sporadic due to scarcity of commercialsized trees. Cativo sawnwood and plywood production during the late 1990s was only a quarter of its peak in the late 1960s (Romero M. et al. 1999). I conclude that fallen and inclined cativo trees may be a common feature of riverine forests due to their shallow root systems and saturated soils, with logging being a lesser factor. Growth rates of the various stem types at different sites may vary with forest structure but may also be limited by different factors depending on stem type. Although sprouts from inclined/prostrate trunks at Sambu and Casarete grew at similar rates, undamaged stems and broken stems at Sambu grew four to five times faster than their counterparts at Casarete, presumably because the recent logging at Sambu left a more open canopy. Growth of all cativo stem types increased with increasing distance from tidal influences but decreased with increasing hydroperiods. In general, cativo trees in the inland swamps that are flooded continuously during the rainy season grew more slowly than in the riverine forests, and the riverine sites further upriver (Juanacati, Bongales, Limn) showed faster mean cativo growth than downriver sites. Diameter increment of canopy trees varied by site and year, but the patterns of variability differed among sites. Canopy tree growth at Casarete varied up to 2-fold, with 1997 as the slowest growing year. At the other three principal sites, canopy tree growth varied less, and the census period that spanned the 1997 El Nio was not always the slowest growing year. These findings stand in contrast to the strong reductions in tree

PAGE 90

75 growth in Costa Rica during the 1997 El Nio, which were negatively correlated with daily minimum temperatures (Clark et al. 2003). The fastest growing tree species in my sample plots was Pterocarpus officinalis, which is not considered a timber tree. Cativos two other most common associates, Carapa guianensis and Pentaclethra macroloba are harvested in both Panama and Costa Rica (Webb 1997, Sitoe et al. 1999). C. guianensis in particular is valued by Darien loggers, while P. macroloba produces less valuable wood. C. guianensis is more abundant in riverine forests flooded only with freshwater, and has moderate growth rates and relatively high densities in some Darien cativo forests. This species may have been locally extirpated in some riverine forests but could be reintroduced by seed scattering as was recommended by Webb (1997) for logging gaps in swamp forest in Costa Rica. In the absence of spatiallyexplicit data that allow for the development of competition indices and the construction of distancedependent forest dynamic models, higher survival of faster growing trees (growthdependent mortality) should be taken into account when projecting growth trajectories. When mean growth of a large population of small trees is low, modeling lifetime growth based on mean growth may result in unrealistically long trajectories (Grauel and Kursar 1999), especially if a large proportion of slow growing trees die before reaching commercial size. I noted heavy seedfall during plot installation in April 1997, but the regeneration study began in September 1997 at one site and May 1998 at the other, so I measured recruitment in 1998 at one site and 1999 at the other. Although cativo produces some seeds twice a year, large seedfalls seem to occur once every two years

PAGE 91

76 (Pizano SA 1995), and I also measured a large pulse of seedling recruitment from July to November 1999, two years after the large observed seedfall in AprilMay 1997. When stand structures and dynamics vary so markedly among forests, it seems inadvisable to extrapolate results widely. Witness the difference in cativo dynamics between these Darien cativo forests and the population of cativo on Barro Colorado Island. With much lower mortality rates and generally higher mean and maximum growth rates for cativo on BCI, use of their data for management tasks such as timber harvest scheduling or yield projections would justify overharvesting of most Darien cativo forests. Mortality, more than growth, was shown to be a pivotal factor in the simulated sustainability of cativo harvest potential based on BCI data (Condit et al. 1995a), but annual mortality rates of Darien cativo forests varied greatly and were sometimes much higher than on BCI. A critical aspect for understanding present day forest structure and dynamics is the history of use that has resulted in what are now degraded forests. Increasingly, these degraded forests will be a source for wood and nonwood products as the area of intact, mature tropical forest declines. For example, the forest most recently logged in this study (Sambu) is one of the most dynamic and resilient, with relatively high growth and recruitment rates, low mortality, and sufficient densities of advance regeneration to theoretically provide additional timber harvests. Forest history, although it may only be inferred, can yield insights into todays forest and perhaps help to guide management direction. This study highlights the importance of examining stand development patterns at a variety of sites, even when a single forest type is identified based on species

PAGE 92

77 composition and landscape position. Only where variability is recognized can it be considered when making forest management decisions. Anthropogenic disturbance may have been the primary factor in shaping the structure and function of many cativodominated forests in Darien, but the persistence of these forests after decades of harvesting attests to their resilience and should serve as inducement for better management.

PAGE 93

78 Table 4. Total plot area measured for different minimum tree diameters and number of tree species found. Plot Area (ha) Number of Tree Species Site 1 cm 4 cm 10 cm 1 cm 4 cm 10 cm % unidentified Casarete 0.4 0.4 1.0 8 7 6 < 0.1 Sambu 0.2 0.2 0.8 7 5 5 0.0 Juanacati 1.5 1.5 48 24 0.4 Naranzati 0.32 0.32 0.96 54 42 28 1.5 Table 4. Species diversity indices and relative dominance of cativo (Prioria copaifera). DBH 1cm Site Fisher's SW Evenness Simpson Cativo dominance Casarete 1.10 0.20 0.09 0.93 0.96 Sambu 1.00 0.22 0.11 0.91 0.95 Naranzati 12.37 2.10 0.53 0.32 0.55 DBH 4cm Site Fisher's SW Evenness Simpson Cativo dominance Casarete 1.01 0.22 0.10 0.92 0.96 Sambu 0.76 0.28 0.16 0.87 0.93 Juanacati 9.52 2.09 0.54 0.25 0.47 Naranzati 7.19 1.81 0.52 0.36 0.59 Table 4. Stem density and basal area of all species (above) and cativo only (below). Site Stems/ha 14 cm Stems/ha 4 cm Stems/ha 10 cm BA/ha (m 2 ) 10 cm Casarete 1460 705 757 43.3 Sambu 2095 785 484 48.9 Juanacati 500 426 31.4 Naranzati 1675 1057 339 47.1 Darien NP 328 71.1 Cativo Stems/ha 14 cm Cativo Stems/ha 4 cm Cativo Stems/ha 10 cm Cativo BA/ha (m 2 ) 10 cm Cativo Stems/ha 60 cm Cativo BA/ha (m 2 ) 60 cm Casarete 1415 668 727 41.5 8 2.5 Sambu 2030 690 463 39.9 12 4.4 Juanacati 229 195 10.3 3 0.9 Naranzati 732 457 240 39.3 51 29.2 Darien NP 160 48.8 58 45.4

PAGE 94

79 Table 4. Incidence (%) of prostrate, inclined, broken stems and sprouts from prostrate trunks. Other stems showed no signs of earlier breakage and presumably regenerated from seed. Prostrate Stems Inclined Stems Broken Stems Sprouts from Inclined or Prostrate Stems Site <10 cm 10 cm <10 cm 10 cm <10 cm 10 cm <10 cm 10 cm Casarete 0.0 5.0 3.0 7.7 8.2 4.5 8.3 12.0 Sambu 0.0 3.1 1.9 7.2 6.1 4.0 16.7 1.6 Juanacati 2.1 1.1 4.2 4.1 10.9 5.6 5.9 0.7 Naranzati 0.0 0.0 3.5 0.9 9.7 3.1 1.8 0.0 Darien NP 0.0 0.0 0.0 0.0 Table 4. Forestwide annual treefall and tree incline rates (i.e., partial uprooting) for small (above) and large (below) trees for four sites. < 10 cm dbh 1997 1998 199900 2000 Incline Fall Incline Fall Incline Fall Incline Fall Casarete 0.40 0 0 0 3.22 0.57 0.23 0.23 Sambu 0.17 0 0 0 0.13 0.13 0.15 0 Juanacati 1.48 0.37 0.97 0 1.18 0.34 0.49 0 Naranzati 0.87 0 0.55 0 1.16 0.93 10 cm dbh 199798 199899 19992000 200001 Incline Fall Incline Fall Incline Fall Incline Fall Casarete 0.56 0 0.26 0 2.38 1.68 0.46 0.58 Sambu 1.43 0 1.08 0.81 1.87 0.42 0.46 0 Juanacati 0.88 1.32 0.73 0.36 0.53 1.24 0.26 0.38 Naranzati 0 0 0.29 0.58 0.33 0.33

PAGE 95

80 Table 4. Mean annual diameter growth (mm/year) of cativo trees of three stem types, based on 1997, 1998, or 1997 census periods. Within diameter classes, different letters denote significant differences (p0.01) among stem types, (sample sizes noted parenthetically). C a s a r e t e S a m b u DBH cm Undamaged Broken Sprouts from prostrate/inclined trunks DBH cm Undamaged Broken Sprouts from prostrate/ inclined trunks 14 0.35 a (427) 0.3 a (33) 0.9 b (25) 14 1.5 a (301) 1.5 a,b (24) 1.0 b (66) 4 0.7 a (205) 0.9 a (20) 1.2 a (14) 4 2.0 a (117) 4.1 b (20) 1020 1.8 a (288) 1.6 a (22) 4.0 b (63) 1020 4.6 a (140) 6.5 a (6) 6.6 a (6) 2040 3.5 a (204) 1.2 a (4) 5.4 b (19) 2040 4.2 a (101) 2.2 a (4) 40 2.6 (120) 0.6 (3) 40 2.8 (103) 0.8 (2) N a r a n z a t i J u a n a c a t i DBH cm Undamaged Broken DBH cm Undamaged Broken Sprouts from prostrate/ inclined trunks 14 0.9 a (92) 2.7 a (8) 4 1.2 a (79) 2.6 a (5) 4 3.0 a (288) 2.0 b (34) 2.7 a,b (15) 1020 2.7 a (61) 1.7 a (5) 1020 5.3 a (164) 2.8 a (9) 5.7 a (3) 2040 4.7 (43) 2040 8.5 a (70) 2.5 b (7) 40 4.5 (140) 40 7.1 (32)

PAGE 96

81 Table 4. Mean annual diameter growth (mm/year) of cativo trees. Statistical comparisons are vertical, within diameter classes and among sites; different letters denote significant differences (p0.01; sample sizes noted parenthetically). All stem types except prostrate are grouped. 1997 14 cm 4 cm 100 cm 200 cm 40 cm Casarete 0.1 b (485) 0.0 b (242) 1.2 b (374) 2.3 c (230) 1.7 b (123) Sambu 1.8 a (396) 2.2 a (138) 4.4 a (153) 4.3 b (108) 3.3 a,b (107) Naranzati 0.3 b (64) 0.2 b (52) 1.0 b (37) 3.0 b,c,d (20) 3.5 a (23) Juanacati 3.1 a (118) 4.7 a (53) 5.4 a,b (26) 7.5 a (9) Bajo Grande 2.0 a,b (9) 1.7 c,d (70) 1.9 b (48) Rio Amarradero 0.7 a,b (11) 0.5 c,d (51) 2.0 b (92) Rio Balsas1 0.5 d (96) .5 b (21) Bongales 3.1 a,b (11) 5.8 a,b (62) 5.9 a (34) Limon 5.0 a,b (4) 8.4 a (36) 8.1 a (8) Rio Tuira 3.1 a,b (11) 2.0 c,d (151) 1.2 b (38) 1998 14 cm 4 cm 100 cm 200 cm 40 cm Casarete 0.5 b (491) 0.9 b (237) 2.5 c (376) 4.3 c (227) 3.5 c (126) Sambu 1.6 a (410) 3.7 a (148) 6.4 b (156) 5.6 b,c (105) 3.9 c (104) Naranzati 0.6 b (185) 1.2 b (117) 2.4 c (97) 4.2 c (67) 3.5 c (176) Juanacati 4.5 a (332) 8.2 a (179) 10.5 a,b (78) 9.4 a,b (31) Bajo Grande 3.7 a,b,c (8) 8.0 b (68) 11.0 a (49) Rio Balsas1 3.2 c (98) 4.1 b (20) Bongales 4.5 a,b,c (9) 8.7 a,b (57) 7.2 b (36) Limon 6.8 a,b,c (3) 12.0 a (34) 12.7 a (10) Rio Tuira 4.6 a,b,c (11) 7.2 b (144) 5.7 b,c (39) 1999 14 cm 4 cm 100 cm 200 cm 40 cm Casarete 0.5 b (483) 1.1 b (238) 1.7 b (372) 5.0 b (226) 3.2 c (128) Sambu 1.3 a (427) 2.6 a,b (162) 4.2 a,b (157) 3.8 b (110) 2.3 c (104) Naranzati 1.6 a (100) 1.7 b (83) 2.6 b (66) 5.4 b (43) 5.1 b (139) Juanacati 3.0 a (338) 5.1 a (190) 8.4 a (81) 7.8 a (39) 2000 14 cm 4 cm 100 cm 200 cm 40 cm Casarete 0.8 a (479) 1.0 b (243) 2.1 b (355) 3.6 b (224) 2.0 b (126) Sambu 1.1 a (459) 1.9 a (170) 4.0 a (169) 3.3 b (116) 1.6 b (107) Juanacati 1.3 b (332) 2.6 b (197) 5.1 a (83) 5.2 a (40)

PAGE 97

82 Table 4. Ingrowth by stem type. Percentage of recruited individuals from broken stems, undamaged stems, or sprouts from prostrate and inclined trees. 1997 1998 1999 Site DBH cm Sprouts from inclined/prostrate stems Broken Undamaged Sprouts from inclined/prostrate stems Broken Undamaged Sprouts from inclined/prostrate stems Broken Undamaged Casarete 50 0 50 63 6 31 53 33 14 Sambu 14 5.5 5.5 89 11 4 85 0 0 100 Naranzati 0 0 100 50 0 50 0 20 80 Casarete 0 0 100 11 0 89 0 0 100 Sambu 29 0 71 7 0 93 21 0 79 Juanacati 4 0 50 50 6 0 94 25 6 69 Naranzati 0 0 100 0 0 100 0 0 100 Casarete 50 0 50 38 0 62 20 0 80 Sambu 0 0 100 30 0 70 0 0 100 Juanacati 1040 0 0 100 0 0 100 0 0 100 Naranzati 0 0 100 0 0 0 0 0 100 Casarete 0 0 100 0 0 100 0 0 100 Sambu 0 0 100 0 0 100 0 0 100 Juanacati 40 0 0 100 0 0 100 0 0 100 Naranzati 0 0 0 0 100 0 0 100

PAGE 98

83 Table 4. Continued 2000 Site DBH cm Sprouts from inclined/prostrate stems Broken Undamaged Casarete 89 2 9 Sambu 61 6 33 Naranzati 14 Casarete 25 0 75 Sambu 11 0 89 Juanacati 50 5 45 Naranzati 4 Casarete 57 0 43 Sambu 14 0 86 Juanacati 20 0 80 Naranzati 1040 Casarete 50 0 50 Sambu 0 0 100 Juanacati 0 0 100 Naranzati 40 Table 4. Cativo annual recruitment and mortality rates (%) by stem diameter class for four census periods. R e c r u i t m e n t R a t e s M o r t a l i t y R a t e s Site DBH cm 199798 199899 199900 200001 199798 199899 199900 200001 Casarete 1.2 3.0 3.2 10.7 8.3 4.6 2.8 2.1 Sambu 8.4 11.6 3.0 3.7 1.9 1.6 1.7 0.6 Naranzati 14 3.9 1.7 4.8 3.9 4.3 3.0 Casarete 1.0 3.6 3.1 2.8 7.2 2.8 0.4 0.7 Sambu 11.2 10.2 6.9 4.9 0 0 0.5 0 Juanacati 1.6 5.5 5.2 4.6 1.6 0.4 0 0.7 Naranzati 4 5.0 2.1 8.1 2.5 2.1 1.2 Casarete 0.3 1.3 0.9 1.0 1.4 1.0 1.1 1.1 Sambu 2.1 3.9 2.1 2.2 0.4 0.8 0 0.6 Juanacati 4.8 7.7 5.9 1.4 0 0 1.6 0.6 Naranzati 1040 2.4 0 4.4 0 0.7 0.8 Casarete 0.8 2.7 1.0 1.6 0 1.5 0.8 0 Sambu 1.7 1.0 0.8 0.9 0 1.0 0 0 Juanacati 7.3 8.0 4.9 8.4 0 0 0 0 Naranzati 40 0 1.9 8.4 0 0 0.7

PAGE 99

Table 4. Annual mortality rates (%) of cativo trees by stem type and stature for four census periods. 84 199798 1998 DBH cm Broken Sprouts from inclined/prostrate stems Prostrate Inclined Undamaged Broken Sprouts from inclined/prostrate stems Prostrate Inclined Undamaged Casarete 0.83 2.51 0.21 0.83 3.56 0.13 1.16 0 0.39 2.33 Sambu 0.18 1.06 0 0.18 0.19 0.57 0 0 0.38 Juanacati <10 0 0 0.81 0 1.62 0 0 0.35 0 0.70 Naranzati 3.32 0 0 0 0 1.42 0 0 0 1.89 Casarete 0 0.23 0.23 0.46 0.23 0.13 0.13 0.26 0.13 0.40 Sambu 0 0 0 0.25 0 0 0 0.56 0 0.28 Juanacati 10 0 0 0 0 0 0 0 0 0 0 Naranzati 0 0 0 0 0 0.33 0 0 0 0 19992000 2000 Broken Sprouts from inclined/prostrate stems Prostrate Inclined Undamaged Broken Sprouts from inclined/prostrate stems Prostrate Inclined Undamaged Casarete 0.29 0.58 0 0 1.16 0 0.70 0 0.35 0.59 Sambu 0 0.98 0 0 0.42 0 0.45 0 0 0 Juanacati <10 0 0 0 0 0 0.48 0 0 0 0.24 Naranzati 0.54 0 0 0 1.63 Casarete 0.15 0.15 0 0.29 0.44 0 0 0.12 0.35 0.47 Sambu 0 0 0 0 0 0 0 0 0 0.47 Juanacati 10 0 0 0 0 1.39 0 0 0 0.49 0 Naranzati 0 0 0 0 0.75 0

PAGE 100

85 Table 4. Mean annual growth (mm/year) of cativo trees that were alive at the end of the study and those that died during the study for which there was one or more years of growth data. 1997 growth Site DBH cm Alive 2001 Dead 2001 df t p 110 0.04 (683) .09 (44) 65 1.6 0.05 Casarete 10 1.7 (712) 0.3 (17) 25 5.1 < 0.001 110 1.9 (519) 0.5 (14) 16 3.4 0.002 Sambu 10 110 0.2 (107) 0.6 (9) 9 0.5 0.32 Naranzati 10 410 Juanacati 10 1998 growth Site DBH cm Alive 2001 Dead 2001 df t p 110 0.6 (707) 0.2 (21) 33 4.4 < 0.001 Casarete 10 3.3 (718) 1.6 (13) 13 2.5 0.01 110 2.2 (545) .03 (12) 14 6.3 < 0.001 Sambu 10 110 Naranzati 10 410 Juanacati 10 8.7 (294) 3.6 (6) 5 2.5 0.025 1999 growth Site DBH cm Alive 2001 Dead 2001 df t p 110 0.7 (711) .1 (10) 11 4.9 < 0.001 Casarete 10 3.6 (722) .8 (7) 15 17.2 < 0.001 110 Sambu 10 110 Naranzati 10 410 Juanacati 10

PAGE 101

86 Table 4. Abundance of cativo trees < 1 cm dbh by height class. Trees were tallied but not measured at Casarete in December 1999. C a s a r e t e S a m b u Height (cm) Nov 1997 Nov 1998 Dec 1999 June 1998 July 1999 June 2000 < 30 485 110 222 341 526 300 3995 2275 5956 6281 3937 600 1210 1195 6889 6548 6063 90150 745 860 2667 3363 3558 >150 <1 cm dbh 390 215 1378 993 1326 total 6825 4655 12045 17111 17526 15411 Table 4. Annual mortality rates (%) of cativo trees by height class for the period November 1998 November 1999. Height (cm) Casarete Sambu < 30 51.4 63.3 3060 30.0 9.9 6090 13.2 1.2 90150 3.2 0.0 >150 <1 cm dbh 6.6 2.0 Table 4. Mean annual height (cm/year) and diameter (mm/yr) growth rates for cativo trees < 1 cm dbh, (sample sizes noted parenthetically). Casarete Nov 97Dec 98 Dec 98Dec 99 Height (cm) Height Growth (cm) Dbh Growth (mm) Height Growth (cm) Dbh Growth (mm) < 30 18.5 (25) 8.3 (4) 3060 4.9 (432) 1.9 (117) 6090 2.1 (155) 1.4 (94) 90150 2.1 (106) 1.4 (101) 0.3 (24) >150 <1 cm dbh 0.3 (71) 3.0 (26) 0.4 (24) Sambu June 98July 99 July 99June 00 Height (cm) Height Growth (cm) Dbh Growth (mm) Height Growth (cm) Dbh Growth (mm) < 30 1.9 (8) 13.3 (8) 3060 2.5 (338) 2.8 (157) 6090 2.4 (416) 2.2 (242) 90150 4.4 (168) 5.3 (140) 0.8 (17) >150 <1 cm dbh 0.9 (70) 10.1 (43) 0.6 (38)

PAGE 102

87 Figure 4. Darien Province, Panama showing principal sites (S=Sambu, N=Naranzati, C=Casarete, J=Juanacati) and seven secondary sites.

PAGE 103

88 CasareteDBH (cm) 020406080 Annual Growth (mm) 0102030 SambuDBH (cm) 0204060 80 Annual Growth (mm) 0102030 JuanacatiDBH (cm) 020406080 Annual Growth (mm) 0102030 NaranzatiDBH (cm) 020406080100120 Annual Growth (mm) 0102030 a Figure 4. Annual diameter growth (mm/year). a) Prioria copaifera (cativo) b) Pterocarpus officinalis, c) Carapa guianensis, d) Pentaclethra macroloba.

PAGE 104

89 CasareteDBH (cm) 020406080 Annual Growth (mm) 0102030 SambuDBH (cm) 02040608 0 Annual Growth (mm) 0102030 JuanacatiDBH (cm) 020406080 Annual Growth (mm) 0102030 NaranzatiDBH (cm) 02040608 0 Annual Growth (mm) 0102030 b Figure 4 Continued.

PAGE 105

90 JuanacatiDBH (cm) 0102030405060 Annual Growth (mm) 0102030 NaranzatiDBH (cm) 0102030405060 Annual Growth (mm) 0102030 c JuanacatiDBH (cm) 020406080 Annual Growth (mm) 0102030 NaranzatiDBH (cm) 02040608 0 Annual Growth (mm) 0102030 d Figure 4 Continued.

PAGE 106

CHAPTER 5 GROWTH AND YIELD PROJECTIONS OF Prioria copaifera FROM FOUR SWAMP FORESTS IN DARIEN, PANAMA Introduction Tropical forests historically were viewed by foresters in temperate countries primarily as sources of valuable woods (Whitford 1921, Record 1925). Only relatively recently have concerns about biodiversity and other less financially-tangible values become prominent (Putz et al. 2000b). Research has shown that tropical forests have many diverse values that differ among stakeholders (Godoy et al. 2002, Kainer et al. 2003, van Beukering et al. 2003). Despite growing awareness of the values of tropical forests, forest degradation as well as deforestation continue due to a wide variety of factors such as rising agricultural and timber prices, low rural wages, and few employment opportunities other than forestry or agriculture, among others (Kaimowitz and Angelsen 1998, Barbier and Burgess 2001). In many countries, when forests have been depleted of marketable wood, their perceived value is lost and the land is converted for agricultural uses that may be no more sustainable than the timber mining that preceded them. Increasingly recognized is the need for permanent forest estates and for policies that promote not just a forest products industry, but forest management that is more than just one step in the process of deforestation (Verissimo et al. 2002). Although forests have many values, timber is still the forest output that is in greatest demand worldwide. Unfortunately, continuing demand for timber has not been sufficient to lead to forest conservation for a variety of reasons. Often, the benefits of 91

PAGE 107

92 intact forests have been undervalued and the benefits of forest conversion overestimated (Repetto 1988). Logging, at least as it is conventionally carried out, not only depletes the forest of the available valuable timber but causes residual damage to soils and vegetation that reduces subsequent harvests. Improved forest management techniques are infrequently employed because they are perceived as being more expensive than conventional logging (Putz et al. 2000a), despite accumulating evidence to the contrary (Holmes et al. 2002, Boltz et al. 2003). Other variables that limit commercial interest in sustaining timber yields from natural forests in the tropics are high discount rates, unknown or slow volume increments, tenurial insecurity, and fluctuating timber prices (Pearce et al. 2003). Scarcity of trees of valuable species and low rates of increment in commercial volumes are the principal biological constraints on timber production from tropical forest and necessitate long cutting cycles. Plantations often produce more timber more often than natural forests not only because of intensive silvicultural practices that increase growth, but also because every stem on a given hectare is available for harvest. Natural forest management for timber production would be more competitive if commercial species were more abundant or if the timber was more highly valued. Monodominant forests composed of a commercially valuable species meet the former requirement. This study examines the wood producing potential of a unique Neotropical forest type for which species composition and regeneration capacity are such that, despite only moderate stem diameter growth rates, many forests of this type have sustained several decades of timber harvesting and still contain an abundance of potentially valuable trees. Using data on forest structure and stand dynamics from permanent plots at four sites, I estimate

PAGE 108

93 future commercial timber volumes under a variety of cutting rotations (years between harvests) and diameter limits. Neotropical swamp forests dominated by Prioria copaifera (cativo) have long been valued for their timber (Barbour 1952, Lamb 1953). Indeed, the common pattern of a timber species being depleted and its use replaced by other species occurred with cativo in at least part of its range (Veiman 1982). In Darien, Panama, in contrast, sites that started being logged for cativo in the 1950s and 1960s are still cativo-dominated forests, albeit degraded to different degrees (Holdridge 1964, Grauel and Putz 2004). Timber harvesting in Darien can be characterized as either laboror capital-intensive. Labor-intensive logging occurs in riverine forests by local community members as part of a diverse livelihood strategy that may also include farming, hunting, non-timber forest product collecting, and fishing. Permits are obtained for harvesting individual trees, and heavy machinery is not used for road building or log extraction. After target trees are felled, the loggers fell additional trees 25 cm dbh and position them end-to-end to form two parallel rails. This practice facilitates the manual rolling or levering of the harvested logs to the river. Easily-accessed forests are repeatedly logged, and well-formed trees larger than the legal cutting limit of 60 cm dbh are consequently scarce. Other evidence of repeated logging including the frequency of damaged crowns among the residual trees, liana tangles, and common, large canopy gaps. Capital-intensive timber harvesting in Darien occurs mostly in a mosaic of inland cativo swamps and tierra firme (unflooded) forests. Logging concessionaires float heavy machinery as far upriver as is feasible and then build roads and logging camps; logging is restricted to the short annual dry season ( 3 months). Although forest

PAGE 109

94 management plans are required by law, most consist of volume and species estimations based on a few sample plots. Because logging concessions are limited to five years and renewal is uncertain, there is little incentive for the long-term planning of subsequent timber harvests and roads are poorly constructed and maintained. After cativo, Anacardium excelsum is the most common commercial species harvested in this landscape. Study Sites Cativo-dominated swamp forests are found both along rivers and in interior lowlands of Darien Province of Eastern Panama. Inland swamps are flooded continuously over the 9 month wet season whereas forests adjacent to major rivers are flooded periodically by wet season rains and by monthly spring tides that can reach 70 km or more upriver. Forests closest to the influence of tides are strongly monodominant (> 95% cativo in all size classes) and may contain < 10 species ha -1 whereas riverine and inland forests outside of tidal influence are relatively more species rich. These forests may contain 40 species ha -1 although cativo can still comprise over 80% of the overstory (Linares Prieto 1988). The four sites in this study vary by landscape position, flooding regime, species diversity, and management history (Table 5). Soils at all sites are fine textured and poorly drained. Slight brackishness was detected in the soil only at the Casarete site (Tapia 1999). The three riverine sites have all been logged in the past and although cativo regeneration of all sizes is abundant, large trees are relatively scarce. Although there was a logging concession nearby at the time of this study (1997), the inland swamp in this study had not been logged at the time of plot installation (1997).

PAGE 110

95 Methods The stand structure, growth, and mortality data for this study were collected between 1997 and 2001. All plots were installed in 1997 or 1998; trees were marked, mapped, and measured annually up to 2001. The minimum diameter measured was 1 cm dbh at three of the sites (Casarete, Sambu, Naranzati) and 4 cm at Juanacati, but in this paper only data for trees 10 cm was used. I excluded trees < 10 cm dbh because in preliminary analyses no trees smaller than 10 cm dbh entered the harvest projections during the 60 year simulation, and because variability in growth rates is much higher for small trees than large ones. Data on large trees was generally scarce at the riverine sites, and few trees attain large sizes due to intense logging pressure. For the riverine sites I used only data for trees 60 cm dbh, the legal cutting limit, because data for larger trees were scarce. At the inland swamp, trees 60 cm are abundant, but because the harvest simulation uses a maximum diameter cutting limit of 100 cm dbh, and because growth data for trees > 100 cm dbh were scarce, I used data for trees between 10 and 100 cm dbh. Data on diameter growth rates were derived from the longest possible measurement interval for each tree. Of 3065 cativo trees in the four sites, 80% of the annual growth records were for the period 1997, 14% were for 1998, and 6% were for 1997. The wood volume of each tree 40 cm dbh was estimated by multiplying its basal area by a species-specific form factor (Philip 1994) and then by commercial height, defined as the height to the lowest branch. Commercial height was estimated for trees 40 cm dbh, 50 cm dbh, and 60 cm dbh after measuring about two dozen trees 40 cm dbh to the lowest branch with a telescoping measuring pole at both Casarete and Sambu. Commercial height estimates from the Casarete site were used for the Naranzati

PAGE 111

96 and Juanacati sites. Volume estimates were based on a form factor specifically developed for cativo using 23 trees felled along the Balsas River near the Casarete site by Mariscal et al. (1999, Grauel and Pineda M. 2001). Volume of those trees was determined by the method of height accumulation by which successive diameters are measured at known heights along the stem (Philip 1994). Volume (V) was calculated using Smalians formula: V = (h/2)*(A b + A u ) where h = height A b = cross sectional area at base A u = cross sectional area at top. Form factor (F) is defined as F = V/((A bh )*h) where V = volume A bh = cross sectional area at breast height, 1.3m h = height. The mean form factor for the 23 trees 49 cm dbh was 0.641513 (standard deviation = 0.0916; (Grauel and Pineda M. 2001). During each simulation, as individual trees grew into the 40, 50, and 60 cm dbh size classes, different appropriate commercial heights were applied to their volume calculation. Growth projections were made using the method described in Condit et al. (1993) in which growth is expressed as a function of dbh with a polynomial equation and the resulting parameters are used to develop a diameter trajectory through time (see Appendix A for details). The resulting growth curve expresses dbh as a function of time with the initial condition (time = 0) as the minimum diameter (10 cm dbh).

PAGE 112

97 Volume was projected by calculating the age (the time since the minimum diameter, 10 cm dbh) of each individual stem in the plots (year 2000 dbh), projecting its growth, and calculating future diameter and associated volume (Condit et al. 1995a). I estimated volume in this way for all trees as they attained 40 cm dbh in five year increments beginning with the year 2000. This approach allowed me to simulate a timber harvest at 2005 and then at any interval I chose by selecting trees and their associated volumes that surpassed a given diameter limit in any future year. For all projections I applied the site-appropriate annual mortality rate (m) to the volume (v) at a given year (t) with v e -mt where t is the number of years beyond the year 2000. Based on five census years at the permanent plots, annual mortality rates of cativo trees 10 cm dbh at Casarete, Sambu, Juanacati, and Naranzati were 1.15%, 0.45%, 0.55%, and 0.50%, respectively. Scenarios For all sites I projected timber volume out to the year 2065, but I modeled timber harvests differently for capital-intensive logging (inland swamp, Naranzati) and labor-intensive logging (riverine forestsCasarete, Sambu, and Juanacati) for several reasons. Stand structures were quite different between the previously logged riverine sites focused on by community loggers and the unlogged inland swamp typical of concession logging. Furthermore, the labor-intensive logging that is carried out in riverine forests occurs at higher frequencies but lower intensities than the capital-intensive logging of inland swamps. At inland swamps with extremely large standing volumes in commercial-sized trees, the logging concessionaires fellers seek out the largest stems since they are paid by the amount of volume felled. For purposes of comparison with the riverine sites, I simulated a 60 cm dbh cutting limit at the inland swamp, but higher diameter cutting

PAGE 113

98 limits probably better represent reality and would be silviculturally more appropriate if the aim is to perpetuate the stand. In addition to the legal cutting limit (60 cm dbh), I simulated cutting limits of 80 cm, 90 cm, and 100 cm dbh every 20 years beginning in 2005 because many Panamanian foresters feel that the 5 year concession period will soon be legally extended to 20 years. In addition to the four fixed dbh cutting limits, a fifth scenario adjusted the cutting limit according to stand structure after an initial 100 cm dbh cutting limit in 2005, the cutting limit was reduced to 90 cm for the final three harvests. The labor-intensive timber harvesting common in the riverine forests is quite haphazard because individual local community members are free to seek out harvestable trees without the spatial constraints of a logging concession, and harvesting can potentially occur year round. Due to easy accessibility of riverine forests and low capital requirements for this sort of logging, the return time for harvesting to occur on a given hectare is probably less than for inland swamps where roads and skid trails have to be constructed for log extraction. The legal diameter cutting limit (60 cm dbh) is generally adhered to because log rafts are checked at government stations downriver. For the three riverine sites I simulated only the legal cutting limit at 5, 10, and 20 year cutting cycles. Residual stand damage resulting from logging can vary greatly depending on harvest intensity, stand structure, and the machinery employed, as well as worker training and supervision. Because I had no data on logging damage in Darien, for all scenarios and sites I modeled logging damage as a linear relationship with harvest intensity that was developed by Webb (1997) from a comparison of tropical logging operations. In this

PAGE 114

99 way, future yields were reduced according to the cumulative damage in all previous harvests. Both natural and logging-induced mortality were applied after the total harvest volume was calculated. This approach eliminated the need to simulate the death of individual stems because mortality was simulated as a reduction in potential harvest volume (Condit et al. 1995a) For the 2005 harvest simulation in the unlogged inland swamp, I included trees that were greater than or equal to the simulated diameter cutting limit in 2000. In contrast, for all riverine sites I excluded the few trees that were 60 cm in the year 2000 from subsequent harvests because observations indicate that they were not harvested previously due to poor form or hollowness. Hence, for all riverine sites, only trees that were < 60 cm dbh in 2000 but 60 cm in 2005 are included in the first simulated harvest of 2005. In the harvest projections for the riverine, labor intensive logging, I monitored the year 2000-diameter of all trees when they reached harvestable size (60 cm dbh) in future years. During each harvest, community loggers cut some 25 cm dbh trees to position as parallel rails for log extraction, so it was necessary to reduce the number of trees in that size class in any future harvest that included them (i.e. when they had attained 60 cm dbh). For instance, in the 2005 simulated harvest, loggers would cut some trees 25 cm dbh to use as rails, so when that size class reached harvestable size (60 cm dbh) in say, 2045, there would be fewer trees available for harvest. Modeling the reduction of future yields due to the felling of trees for rails was problematical because the number of trees felled depends on distance from the nearest river rather than on the number of

PAGE 115

100 commercial-sized trees harvested or total volume available. Based on observations of previously used roads (parallel, end-to-end rails) and numbers of smaller stumps at the most recently logged site, I reduced future volumes by 10% for every harvest that includes trees that were 25 cm dbh in the year 2000. Results In 2000, standing volume in commercial-sized trees 60 cm was 10 m 3 ha -1 at the riverine sites and approached 200 m 3 ha -1 at the inland swamp (Table 5). Two riverine forests had substantial volumes in trees 40 cm dbh. Mean annual volume increment of large trees ( 60 cm dbh) was greatest at the inland swamp and up to two orders of magnitude greater than the slowest volume-accreting site, but increment for trees 40 cm dbh was lowest at the inland swamp. Commercial volume increments of large trees at all three riverine sites were rather low and never exceeded 0.6 m 3 ha -1 yr -1 For trees 40 cm dbh, however, the three riverine sites had annual volume increments 1.1.2 m 3 ha -1 yr -1 Lifetime growth trajectories also varied considerably among sites, with the time to grow from 10 cm to 60 cm dbh ranging from 72 to 203 years (Figure 51). All three riverine forests yielded the most total volume per hectare after 65 years in the 20-year cutting cycle scenario (Table 5), and maximum yield in this scenario was at year 65 at all sites (Figure 5). All sites yielded little wood at the first harvest in 2005, but production increased thereafter. Volume yield after year 20 fluctuated at Casarete, was relatively constant at Sambu, and increased at Juanacati. For Casarete and Sambu, there was about a 25% higher total yield per ha in the 20-year compared to the 5-year cutting cycle scenario, while at Juanacati the 20-year cutting cycle yielded > 40% more wood volume than the 5-year scenario (Table 5).

PAGE 116

101 Future yields were reduced at the Juanacati site because of the practice of cutting trees for road building. For those harvests that included trees that were about 35 cm dbh in the year 2000, yields were reduced 21% in the five year cutting cycle scenario and 12% and 11% in the 10 and 20 year cutting cycles, respectively. Trees of the size used for rails for log extraction began to enter the legal diameter cutting limit of 60 cm around 2045 and 2050 at Juanacati. (Figure 5). At the slower growing Casarete and Sambu sites, no trees of suitable size for rails attained legal harvest size during the simulation to the year 2065. Total volume yields per hectare for the 20 year cutting cycle (4 harvests) from the inland swamp under the 60, 80, 90, and 100 cm dbh cutting limits were 227, 191, 150, and 72 m 3 respectively, while the variable cutting limit scenario yielded 141 m 3 Using the legal 60 cm dbh cutting limit, 98.6% of the total 60 year yield was contained in the first harvest. Yields using the variable dbh cutting limit were the most consistent throughout the 65 year simulation compared with the fixed dbh limits, with no single harvest extracting more than 44% of the 60 year total yield. With the exception of the initial harvest in 2005, the variable dbh limit yielded the most volume at each future harvest (Figure 5). Simulated logging damage for a given harvest at the riverine forests increased with cutting cycle length because longer cutting cycles allowed for greater accumulation of volume, and damage was modeled as a linear function of volume. The largest reductions of yield due to residual logging damage over the 65 year simulation were for the 5-year cutting cycle for all three riverine sites. Volume reductions due to logging damage ranged from 40% for the riverine sites under the 5-year cutting cycle and 15% for

PAGE 117

102 the 20-year cutting cycle (Table 5). For the inland swamp, yield reduction due to logging-induced damage was similar for the 60, 80, 90, and variable dbh cutting limit scenarios and ranged from 26% for the 90 cm limit to 31% for the 80 cm limit. Overall yield reduction due to residual damage under the 100 cm dbh cutting limit was only about 7%. Discussion All four sites in this study show reasonable potential for wood production, but the sources of that potential vary. The inland swamp contains large standing stocks of commercial sized trees while one riverine forest has very high cativo stem diameter growth rates and the other two have the advantage of very low species diversity. Standing commercial volumes of cativo wood in trees 60 cm dbh in two of the three riverine forests (20 m 3 ha -1 ) seem to indicate that little degradation has occurred. Indeed, the forest most recently logged, Sambu, contained more volume in trees 60 cm dbh in the year 2000 than a 1953 estimate for the cativo forests along the Sambu River by F. Bruce Lamb, a professional forester who surveyed the major watersheds of Darien for potential log sources (Lamb 1953). His surveys spanned all three logged sites included here. For the cativo forests that included the Casarete, Sambu, and Juanacati sites, he estimated mean volumes of 71, 35, and 24 m 3 ha -1 respectively (Lamb 1953). Although these forests are often described today as degraded, the Sambu and Casarete forests show a tremendous resilience in the form of large volumes contained in trees 40 cm dbh (Table 5). A major caveat, however, is that tree form is not taken into account in this study, and the abundance of trees with small, broken crowns and deformed stems testify to the repeated entries that are typical to the labor-intensive logging common in the

PAGE 118

103 riverine forests. In addition, many riverine forests are susceptible to infestations by lianas that can negatively affect stem form, regenerative capacity, and diameter growth (Grauel and Putz 2004). Besides assuming that all trees have merchantable stems, the model assumes constant mean growth throughout the 65 year time period, regardless of harvest frequency or intensity. Conventional wisdom holds that growth of residual trees increases after logging because of increased light availability, but this may not always hold. At the Sambu site for example, growth of trees 40-60 cm dbh seems to be better correlated with crown form than with crown illumination (Grauel, unpublished data). Severe crown damage from repeated logging entries slow the growth of trees, even those with full crown illumination. Interannual climate variability was large during the five years when data were collected for this study, and mean diameter growth rates are reasonable to use in projecting future growth where climate-induced effects on forest productivity are unknown. In addition, variability in relative diameter growth rates were greatest in trees < 10 cm dbh, and by using data 10 cm dbh, this source of uncertainty is reduced somewhat. Estimates of growth, volume, and both natural and logging-induced mortality are all sources of uncertainty, but the model nevertheless serves as a useful tool for comparing future wood yields both among sites and among logging scenarios. The inland swamps that were the object of most capital-intensive, mechanized logging of the 1990s and early 2000s are quite different in terms of stand structure compared to the riverine forests. Although the inland swamp forests are rarely found in patches > 10 ha, commercial volumes are typically very large due to the dominance of Prioria and historical inaccessibility. The 182 m 3 ha -1 of commercial wood volume

PAGE 119

104 found in this study is similar to the standing volume of some southeast Asian tropical forests (Sist et al. 1998) and is probably among the largest commercial volumes per area found in the Neotropics. Per-area volume increments of large cativo trees ( 60 cm dbh) are more a reflection of stem density than of growth rates of individual trees. All three logged sites have annual increments substantially less than the volume increment of the unlogged inland swamp, even though cativo in two of the three logged sites has shorter lifetime growth trajectories than cativo in the unlogged swamp. Volume increments of trees 4060 cm dbh reflect stand density, species composition, and also growth rates. The annual increments ranging from 1.6.2 m 3 ha -1 at Casarete and Sambu are in large part due to the fact that these forests are composed of 95% cativo of all size classes. With such large standing volumes of the commercial species, even moderate growth rates can result in large volume increments. Juanacati, with much less standing volume in cativo but the fastest diameter growth rates of all sites, still had annual volume increment of 1.1 m 3 ha -1 In the riverine forests, longer cutting cycles yielded more wood over the 65 years than shorter cycles because there was less frequent damage to the residual stand from logging. Even though the average amount of residual damage per entry was higher for the longer cutting cycles, higher yields resulted because total damage, as estimated here with reduction of yield, was lower. Another timber harvest simulation for Panamanian tierra firme forest also found that longer cutting cycles yielded more cativo timber than shorter cycles. In a comparison of timber yields of seven commercial species, Condit et al. (1995) concluded that low natural mortality rates and abundant advance regeneration of cativo in unlogged

PAGE 120

105 forest imparted an advantage in sustaining timber yields, but in that study residual damage from logging was held at 15%. By modeling damage as a function of available volume for harvest, I conclude that mortality of residual trees due to logging has a greater influence on future yields than natural mortality. My estimates of logging-induced mortality are probably conservative because, although not explicitly included in this study, tree mortality resulting from injury during logging may also increase in the months and years immediately following logging (Pinard and Putz 1996, Sist and Nguyen-The 2002). There should generally be less damage from timber harvesting in the riverine forests subject to labor-intensive harvesting than where heavy machinery is used during logging. Depending on harvest intensity, road building and the maneuvering of heavy machinery are usually the principal causes of canopy cover reduction and ground area disturbance (Jackson et al. 2002). Log extraction with skidders may even be the principal cause of tree death (Bertault and Sist 1997), although Johns et al. (1996) found that tree felling caused more damage than any other single aspect of logging in the eastern Amazon. A source of uncertainty regarding the amount of damage caused by labor-intensive logging in riverine cativo forests is the question of the number of trees used for rail-laying. In forests that are composed of 95% of a single commercial species, it might seem that repeatedly thinning a subcanopy size class would severely restrict opportunities for future harvests. Although trees of the size used for rail building did not enter the simulated harvests at Casarete and Sambu, the present day stand structures have probably been influenced by this practice. At the site with the fastest mean cativo diameter growth (Juanacati), the potential for yield reductions due to the depletion of smaller trees for rails

PAGE 121

106 would be even greater, but higher species diversity suggests that non-commercial species could be used for rails there. At the more species-diverse inland swamps (Naranzati) where capital-intensive, mechanized logging is employed, the amount of residual damage may determine whether sustainable timber yield is possible. Many studies have found a positive correlation between logging damage and the volume extracted (e.g., Bertault and Sist 1997, Webb 1997, Panfil and Gullison 1998, Sist et al. 1998, Pereira et al. 2002), but differences in logging techniques may reduce such a correlation (Pinard and Putz 1996). At low harvest intensities, road building with heavy machinery may be the principle cause of residual damage but as extracted volumes increase, secondary damage from tree felling and skidder operation cause a higher proportion of damage (Gullison and Hardner 1993). With locally large volumes, the potential for severe damage and greatly reduced subsequent yields is great at inland cativo swamps (Figure 5). Without improved forest management in the form of better harvest planning, worker training, and careful supervision, diameter cutting limits alone can allow massive damage to the residual stand (Sist et al. 2003). Maintenance of species composition is desirable in cativo forests because their homogeneity imparts a substantial advantage in terms of wood production. Although species composition of cativo-dominated swamp forests is largely determined by environmental factors such as the interaction of flooding and drought (Lopez and Kursar 1999, Lopez 2002), logging might lead to shifts in species composition. Simulation models of other tropical forests have shown shifts in species composition after logging, particularly with shorter cutting cycles (Kurpick et al. 1997, Favrichon 1998, Huth and

PAGE 122

107 Ditzer 2000, Phillips et al. 2003). The potential for changes in species composition after logging inland swamp forests that are relatively more diverse than riverine forests is unknown. But in a descriptive study of relatively species-rich cativo forests in Colombia 21 years after logging, Linares Prieto et al. (1997) suggest that management should increasingly focus on second growth cativo forests. The variable dbh cutting limit scenario in this study is probably most likely to perpetuate a cativo-dominated forest because over-harvested or cleared cativo forests have been observed to shift species composition away from cativo (Barbour 1952, Holdridge 1964). Apparently, species composition has not changed substantially in at least two of the riverine forests as a result of their logging history; they remain 95% cativo of all size classes although they are increasingly susceptible to degradation by lianas (Grauel and Putz 2004). The third riverine forest (Juanacati) is composed of 50% cativo stems but is dominated by an overstory of a few, scattered, emergent Mora oleifera trees. Cativo growth and recruitment rates at this site were among the fastest of ten sites and M. oleifera regeneration was scarce, suggesting a species shift towards cativo. A key disincentive to improved concession-based forest management in Darien is the current forest law that limits concession periods to 5 years, with uncertain possibilities of renewal. Longer concession periods by themselves may not prevent timber mining, but given the adequate stocking and moderately fast growth rates of most cativo forests, they could help to give concessionaires financial interests for maintaining forest productivity (Gillis 1992). This study shows the potential for continuous flows of timber from cativo-dominated swamp forests in Darien, Panama, even after 50 years of repeated-entry

PAGE 123

108 logging in easily-accessed riverine forests. Inland swamps only relatively recently subjected to logging contain wood volumes rivaling those once harvested from Asian dipterocarp forests (Manokaran 1998). Despite the inherent advantages for forest management of these low diversity forests, the key to sustainability may lie in keeping societys demands for timber within the range of the forests capacity to produce it (Johnson and Cabarle 1993).

PAGE 124

Table 5. Characteristics of 4 cativo-dominated forests in Darien, Panama. Flooding Regime refers to the 9 month rainy season. Precipitation data are from the nearest measuring station, < 10 km away in all cases. 109 Site Coordinates Landscape Position Flooding Regime Relative Species Diversity Management History Annual Precipitation (mm) Ownership Casarete 8 o 07' N, 77 o 52' W riverine periodic low logged 2457 private Sambu 8 o 04' N, 78 o 13' W riverine periodic low logged 1342 public Juanacati 8 o 05' N, 77 o 47' W riverine periodic higher logged 2096 private Naranzati 8 o 03' N, 77 o 57' W inland continuous higher intact 2457 public

PAGE 125

110 Table 5. Cativo volume (m 3 ha -1 ) and volume increment (m 3 ha -1 yr -1 ) of four cativo-dominated forests in Darien, Panama. 400 cm 60 cm Site Standing Volume Volume Increment Standing Volume Volume Increment Casarete 131.9 1.56 19.8 0.09 Sambu 163.5 2.23 40.0 0.58 Juanacati 30.5 1.12 9.8 0.36 Naranzati 33.0 1.00 182.2 1.52 Table 5. Total volume yield after 65 years of growth and harvest simulations at three different cutting cycles for three riverine swamp forests in Darien, Panama. Volume ha -1 (m 3 ) Cutting Cycle (years) Casarete Sambu Juanacati 5 36.7 74.8 92.4 10 44.3 91.3 135.5 20 48.5 103.2 165.0 Table 5 4. Percentage total volume reduction due to logging-induced damage for three riverine forests at three cutting cycles. Cutting Cycle (years) Casarete Sambu Juanacati 5 39.9 47.4 57.1 10 25.8 36.1 46.4 20 14.6 27.9 38.0

PAGE 126

111 Years Since 10 cm DBH 050100150200 DBH (cm) 0102030405060 Casarete Sambu Juanacati Years Since 10 cm DBH 050100150200250300 DBH (cm) 020406080100 Naranzati Figure 5. Growth trajectories of cativo at four sites in Darien, Panama starting at 10 cm dbh.

PAGE 127

112 5 yr cutting cycle10 yr cutting cycle20 yr cutting cycle Volume ha-1 (m3) 05101520 Casarete Sambu Juanacati Volume ha-1 (m3) 010203040 Year 20002010202020302040205020602070 Volume ha-1 (m3) 020406080 Figure 5. Cativo volume projections for three previously logged riverine forests in Darien, Panama. A fixed 60 cm dbh cutting limit simulated for 5-, 10-, and 20-year cutting cycles.

PAGE 128

113 Year 20002010202020302040205020602070 Year 2000 DBH (cm) 10203040506070 Casarete Sambu Juanacati Naranzati Figure 5. Year 2000 dbh for trees as they attain 60 cm dbh during harvest simulation of a 20-year cutting cycle. Error bars are standard deviations.

PAGE 129

114 Year 20002010202020302040205020602070 Volume ha-1 (m3) 050100150200250 60 cm 80 cm 90 cm 100 cm 100-90 cm Diameter Cutting Limit Figure 5. Cativo volume projections for an unlogged inland swamp forest in Darien, Panama. Four fixed dbh cutting limits simulated for a 20-year cutting cycle beginning in 2005. The fifth projection is based on a 100 cm dbh cutting limit in 2005 followed by a 90 cm dbh cutting limit at subsequent harvests.

PAGE 130

CHAPTER 6 GEOGRAPHICAL, ECOLOGICAL, SOCIAL, AND SILVICULTURAL CONTEXTS FOR CATIVO (Prioria copaifera) SWAMP CONSERVATION IN THE DARIEN OF PANAMA Introduction The cativo (Prioria copaifera)-dominated swamp forests of Darien, Panama that have been repeatedly subjected to uncontrolled logging since the 1950s occur in rich ecological, social, and cultural landscapes. These landscapes offer opportunities but also present constraints for cativo swamp forest conservation through active management. Although many of the ecological characteristics of cativo swamp forests could foster natural forest management for timber, the forests are subject to social, economic, and political forces that have created disincentives for sustainable forest management and led to forest degradation and conversion. This paper explores the settings in which cativo forests are embedded in Darien, and tracks the historical trends of cativo timber production and swamp forest conversion. The abundance of large, cylindrical-trunked cativo trees in the forests of Darien was noted in the 1920s (Kluge 1926, Cooper 1928), but large-scale commercial exploitation of cativo wood began in the early 1950s (Lamb 1953, OEA, Organizacin de Estados Americanos 1978). From the 1950s through the 1990s, cativo supplied about 50% of all sawn wood in Panama and represented 50% of the production from concessions in Darien (Castillo M. 1999). The Panamanian government has long recognized that cativo logging generates many livelihoods for people in Panamas largest and poorest province, Darien (Castillo M. 1999). 115

PAGE 131

116 The original distribution of cativo forests has been greatly reduced by logging, development, and conversion to agriculture. In the 1950s-1970s, much cativo wood was harvested from Caribbean coastal cativo forests located behind mangrove forests in Costa Rica, Panama, and Colombia. Now, in the early 2000s, commercial stands of cativo are found only in eastern Panama and northwestern Colombia. Although cativo is also abundant in some upland forests, it never dominates the forest to the degree found in inundated forests (Condit 1993b). Flooded forests dominated by cativo, called cativales, are found either as narrow bands along rivers or as inland swamps. In Darien, riverine forests are flooded periodically during the wet season (9 month) by rain and by mostky freshwater backed up by the large Pacific tides that can reach 70 km upriver. Where tidal flooding is common, many forests are strongly monodominant, with cativo comprising 95% of the woody species of all size classes. Tree growth under these conditions is slow to moderate, possibly due to the mild brackishness of floodwaters. Riverine forests far enough upriver to escape the salinity of tidal waters are more species diverse, especially in the understory, and tree growth can be quite rapid. The average size of adult trees was estimated to be 90 cm dbh (Lamb 1953), and because of their accessibility, most riverine cativo forests have been repeatedly logged. As a result, very few well-formed trees 60 cm in diameter at breast height (1.3 m, dbh) can be found in riverine cativales. Generally, cativo regeneration is very abundant even in forests that have been repeatedly logged, although overharvesting can result in vine infestations that can impede cativo regeneration (Grauel and Putz 2004).

PAGE 132

117 Inland cativales are flooded for longer durations, up to the entire rainy season. While the forest overstory is composed of around 80% cativo, the understory is more diverse and overall tree growth is moderate. Unlogged inland swamps contain large wood volumes in well-formed, large trees. Because of their isolation and inaccessibility, many of these swamps have only started being logged in the early 1980s. Cativo forests are found in one of the most biologically diverse regions in Central America (Herrera-MacBryde and ANCON 2001), and any approach to cativo forest conservation and management should consider the potential impacts on this biodiversity. The Gulf of San Miguel, for example, holds 46% of the mangroves of the entire country and the nearly pure stands of Rhizophora brevistyla rank among the worlds tallest mangroves at 41m. Evergreen riparian forests are found along the major rivers Tuira, Balsas, Chucunaque, as well as many smaller streams. Cativo can be found in nearly pure stands, although the riverine forests may contain up to 66 tree species (Duke 1975). Cativo forests are also found in and around Darien National Park, one of the largest protected areas in Central America (579,000 ha) that was established in 1980 and designated as a World Heritage Site and Biosphere Reserve shortly thereafter. Its unique flora and fauna have varying affinities due to the locations mixed biogeographical and geological history of isolation and connection. Other protected areas include various indigenous and forest reserves that, together with the national park, comprise the more than half of the province that is under some sort of management category involving environmental protection (Herrera-MacBryde and ANCON 1992). The three dominant ethnic groups that influence the fate of cativales in Darien can be distinguished on the basis of race, culture, and historical origins (Heckadon M. et al.

PAGE 133

118 1982), and have divergent views of the utility of cativo forests. Mestizo descendents of the Spaniards historically dominated Panamanian history and culture and are the most recent arrivals in Darien. These colonists from Panamas western, and largely deforested provinces, generally view forests as obstacles to agricultural and livestock production. Widening deforestation in the northern part of Darien testifies to their increasing numbers. Clearing forest and producing crops or cattle helps colonists establish land tenure rights, but cativo forests that are cleared are eventually abandoned and become mixed-species secondary growth. African-American descendents of escaped slaves (Darienitas) dominate the politics and culture of many of the principal towns in Darien, many of which are along rivers where cativo forests are common. Slaves in increasing numbers were brought by the Spaniards to work in gold mines as indigenous populations diminished; substantial populations of Darienitas occupied Darien as early as 1770 (Cansari et al. 1993). Other African-Americans who are more recent immigrants from Colombia are referred to as Chocoanos. Cativo logging has been a principal livelihood for the black (Darienita and Chocoano) inhabitants of riverine communities such as Yaviza, El Real, Camogant, and Sambu. They also practice subsistence agriculture and have converted many riverine cativo forests for production of plantains, rice, and root crops, especially yams and cassava. The three indigenous groups who inhabit the province are forest-dependent groups who have generally conserved cativo forests where they have legal land tenure. Many Kuna, Embera, and Wounaan live in established, politically-recognized areas (comarcas) but others are scattered throughout the province where they have less certain land tenure.

PAGE 134

119 Indigenous groups seldom engaged in cativo logging in the past, at least partly due to lack of capital. In the 1990s and early 2000s, however, they increasingly started to sell harvesting rights to cativales on the comarcas to logging companies. Even indigenous communities outside the politically-recognized lands negotiate for payment from loggers for access rights to nearby forests, despite the fact that their land claims are not yet legally recognized. Timber Harvesting in Darien The southern portion of Darien Province is not connected to the national road system and rivers serve as the primary transportation routes. The lack of roads and the high cost of river transport have almost certainly slowed rates of colonization and consequently forest conversion (Kursar and Grauel 2002). Population density in this area in 1990 was only around 2 inhabitants per square kilometer (Cansari et al. 1993). On the other hand, the proximity of many cativo forests to rivers has undoubtedly facilitated their logging by forest product companies because transporting logs by water eliminates the need to construct roads. Cativo log rafts are either floated to La Palma for loading on barges or hauled as rafts all the way to sawmills in Panama City. Although limited in extent, cativo forests have historically been highly valued for their timber. Cativo wood is not particularly hard, durable, or attractive; its value lies in its historical abundance and accessibility. The coastal cativo forests on the Caribbean side of Costa Rica, Panama, and Colombia supplied large quantities of raw material to the United States plywood market in the years after World War II. The cativo forests of Darien, in contrast, supplied raw material primarily for the Panamanian domestic plywood market, and it had other general uses in construction and even furniture-making. Cativo supplied 90% of the raw material for the domestic plywood industry in Panama

PAGE 135

120 and 50% of sawn wood production in the country (FAO 1982). Wood production from cativo declined after reaching a peak in the late 1960s, and by the late 1990s was only a quarter of its peak (Romero M. et al. 1999). The estimated original 60,000 ha of cativo forests has been reduced to 15,000 at the end of the 20 th century (ANAM 1999) and much of what remains is degraded from repeated-entry logging. Large scale cativo harvesting in the Darien started in 1950, long before any emphasis was put on sustainable forest management (Bachmura 1972, Christiansen 1980, 1984, Martn Nuez 1984). At the time, laws and regulations relating to forestry focused on timber extraction and reforestation rather than forest management. The policies that promoted timber harvesting were tailored to different stakeholders, but the end result of non-sustainable extraction was the same. There are three types of tree harvesting permits that can be obtained through the Panamanian environmental ministry, ANAM (Autoridad Nacional del Ambiente). Subsistence permits are available to individuals that allow them to harvest individual trees. Community permits are limited to maximum areas of 100 ha, while concession permits allow companies to harvest timber within 2000 ha areas for five years with possibilities of extension. Many of Dariens inhabitants can fit the definition of persons of scarce resources in Panamas Forestry Law No. 1 of February 3, 1994 and can obtain subsistence permits to harvest timber. Subsistence permits allow the holder to harvest a few individual trees each month. If the wood is destined for the plywood market, the 60 cm dbh minimum legal cutting limit is generally adhered to because the plywood factories in Panama City cannot utilize smaller logs (Castillo M. 1999). Logging with subsistence permits is carried out in the more accessible riverine forests. The local

PAGE 136

121 community members cut harvestable trees as well as many smaller trees to lay end to end as a series of parallel rails. The rails facilitate the rolling of the harvested logs over the wet, soft ground to the river. This labor-intensive type of timber harvesting is potentially less damaging to forests because no heavy machinery is employed for road building or log extraction. Often however, middlemen visit the riverine communities, buy many subsistence permits from their holders, and supply chainsaws and gasoline so that the riverine cativo forests have been logged repeatedly. This practice increases the frequency with which the riverine forests are logged because community members who would not otherwise engage in logging are encouraged and paid to do so by the middlemen. Private Panamanian companies, usually from outside Darien Province, solicit permits for forest concessions of 2000 ha. Such concessions are generally located away from the major rivers and include upland forests as well as inland cativales. Currently, the maximum duration of these concessions is five years, but they are typically renewed multiple times with two-year extensions. This capital intensive logging is generally beyond the capacity of Darienitas or indigenous communities, but some Darien inhabitants obtain employment with concession operators. Management plans are required of the concessionaires by law, but they generally consist of volume and species estimates based on a few small plots and are essentially short-term harvesting plans. Concession owners operate with little incentive to plan for multiple rotation-forestry. In addition, by using heavy equipment operators who often have more experience in urban construction than in forest operations, as well as employing unsupervised fellers with informal training at best, many concessions have caused unnecessary forest damage and waste.

PAGE 137

122 Community permits are typically obtained by indigenous groups for timber harvesting on the politically recognized comarcas. Although indigenous communities can obtain logging permits, they usually contract out the actual harvesting to commercial-scale concessionaires because the indigenous communities lack the capital for mechanized logging. They are free to harvest timber from riverine cativo forests using labor-intensive methods. Indigenous communities that are outside of the comarcas are unable to obtain community logging permits because their land tenure is uncertain. Land claims by indigenous communities outside of the comarcas are not yet politically recognized and obtaining legal land tenure is one of their principal desires (TechnoServe-Panam 1996). A common practice is for these tierras colectivas (collective lands) communities to make agreements with commercial-scale logging concessionaires for access to logging concessions where the access routes pass through land claimed by the indigenous community. In Darien, the amount of all timber harvested, including cativo, under the three different types of permits, and consequently by the different stakeholders, has varied considerably in recent years. Between 1982 and 1990, 79% of the volume of all species reportedly extracted from Dariens forests was through concession-based logging, but in the early 1990s more wood was extracted with subsistence permits than through concessions (Torrealba 1996). Wood harvested under community permits fluctuated in the 1990s but remained a minor contribution in comparison to the other two types of permits, but this may be changing. No logging concessions have started or been renewed in the early 2000s, and Panamas principal environmental non-government organization, Association for the Conservation of Nature (ANCON) is increasingly concerned about

PAGE 138

123 the impact of community-based logging by indigenous groups (Bethancourt 2004). Because indigenous communities lack the capital necessary for mechanized logging, they typically contract out the work to concessionaires (Gonzlez Apolayo 2003). Private companies who may be facing log shortages from their own poorly managed logging concessions are often willing to enter somewhat informal agreements with indigenous communities to extract logs. Such agreements usually involve cash payments as well as promises to provide infrastructure improvements such as wells or aqueducts. Problems arise particularly when agreements are made with tierras colectivas communities. Loggers reportedly often harvest outside of agreed upon boundaries and damage agricultural fields, actions against which indigenous communities typically find that they have little legal recourse (Vargas 2003). Cativo forests and other forest types have been degraded through all three types of permits by different stakeholders, but in general few benefits of timber harvesting have remained with Darien inhabitants (Vargas 2003, Bethancourt 2004). Forest Conservation Perspectives Cativo forests as well as other forest types in Darien Province have been under substantial pressure from both logging and agriculture since the 1950s. The province, Panamas largest, provides the vast majority of the timber from natural forests in the country but has also experienced rapid deforestation (Torrealba 1996). Annual rates of population increase in Darien between 1980 and 1990 were greater than 5% (ANAM 1999), and from 1992-2000 the Province suffered the highest rates of deforestation in the country as 172 km 2 of forest were converted annually, mainly to pasture (Repblica de Panam 2003).

PAGE 139

124 The challenges for conservation of cativo forests are similar to those facing many tropical forests. Forest degradation and deforestation increase with rising income (up to a point), rising agricultural prices, and greater accessibility (Kaimowitz and Angelsen 1998, Contreras-Hermosilla 2000). Darien Province has been a focus for colonization and development in Panama since 1950 (Hernndez 1970). Now, in the early 2000s, the colonization of Darien continues, with major infrastructure developments and other projects that could result in substantial reductions in the area of cativo forests and in further degradation of the forests that remain. Surfacing the Pan American Highway to its end in the town of Yaviza, at the confluence of the Chucunaque and Tuira Rivers, is predicted to more than halve the area of cativo forest by lowering transportation costs for timber whose principal transportation route is the Pan American Highway. The primary effect on logging of high-value species probably resulted when the road to Yaviza was opened in the 1980s. It is likely that most of the Province has been selectively cut for high value timber already. Nelson et al. (1999) concluded that the cutting of high value timber such as Swietenia macrophylla and Pachira quinata would probably not increase because their profitability of extraction would be unchanged. In other words, any reduction of transportation costs from surfacing the road would not induce increased logging of those high value species that are now only found in remote locations. Where high value species are still found, the major cost of transportation is in moving the logs from the forest to the highway. In the case of cativo forests, however, the model shows that the profitability of other land uses, especially agriculture or pasture, would increase and create financial incentives to convert some cativo forests.

PAGE 140

125 Much discussion in the environmental literature has focused on the costs, benefits, and tradeoffs between natural area preservation and sustainable forest management as viable conservation tools in the tropics (e.g., Dickinson et al. 1996, Rice et al. 1997, Bowles et al. 1998, Lugo 1999, Rice et al. 2001, Wilshusen et al. 2002, Romero and Andrade 2004) but strict preservation of the remaining cativo forests in the buffer zones of Darien National Park would be an inappropriate conservation strategy. Where the goal is limited to biodiversity conservation, protected areas such as national parks and biosphere reserves can be effective in stopping land clearing and in mitigating the occurrence of other uses (Bruner et al. 2001). Even where biodiversity has been affected by logging, some argue for a cessation of attempts at forest management and the designation of such sites as protected areas (Reid and Rice 1997, Rice et al. 1997). Some cativo forests were never logged due to their remoteness and are now protected in Darien National Park, but the majority of wetland forests dominated by cativo are found in the buffer zones surrounding the park. Such areas were so designated specifically to provide livelihoods to local communities, and low diversity cativo swamps are less valued for biodiversity conservation by most people in any case. The Panamanian Environmental Ministry has designated cativo forests of all types in the buffer zone of Darien National Park as production forests, and their conservation should be based on their capacity to provide benefits to the local communities that live near them. Furthermore, sustainable forest management and strict preservation need not be viewed as exclusive strategies; they can be viewed as complementary ones when the perspective is on the scale of landscapes (Cabarle 1998, Lugo 1999, Whitmore 1999). A regional management approach addresses the conservation value of even human-affected areas and embeds

PAGE 141

126 sustainable forest management in a diverse cultural and ecological landscape (Cannon et al. 1998, Chazdon 1998). Cativo forest management is promising because it would be applied in conjunction with protected areas. Cativo forests are particularly suited to being actively managed within their landscape due to their resilience and potential for wood production, even after repeated-entry logging. Disincentives for cativo forest conservation may result if prices for plywood made from cativo remain low in Panama. Reduced wood production from cativo forests is probably due in part to decreasing log supplies, but dynamic international tropical hardwood markets probably also played a role. Little plywood made from Panamanian cativo is exported, but the prices for plywood imported into Panama have dropped and created disincentives for Panamanian producers to continue to harvest cativo since plywood consumers can buy cheaper imported plywood. At the end of May 2003, prices for tropical hardwood plywood were only 50-60% of prices in 1997 (Adams 2003, ITTO Secretariat, International Tropical Timber Organization 2003). For various reasons, tropical plywood prices are among the most volatile of any commodity (ITTO Secretariat 2003), and if international prices rise, incentives for cativo harvesting will also increase. But as long as imported plywood is cheaper than domestically produced plywood in Panama, decreased timber harvesting from cativo forests may present an obstacle for their conservation. Logging concessions might be neglected or abandoned, for example, and left open to clearing and conversion to agriculture by colonists. Degraded riverine forests might be seen by local community members not as future sources of timber but as more valuable for subsistence crop production and consequently cleared. Alternatively,

PAGE 142

127 higher prices alone may not lead to conservation of cativo forests but instead simply result in continued, unsustainable logging. Strict financial arguments almost always favor rapid exploitation of marketable forest resources and the conversion from natural forest management to more intensive land uses (Pearce et al. 2003). Because time, more so than most other production processes, is a major input for wood production from natural forests, short-term land uses are usually more financially remunerative (Leslie 1977). Increasingly, economic arguments now include the concepts of economic equity and economic sustainability as well as simple efficiency in terms of minimizing costs and maximizing profits (Ruitenbeek and Cartier 1998). It is now generally agreed that the economic arguments against tropical forest management and conservation are incomplete unless they include non-revenue producing benefits and external effects (Leslie 1987, Barbier 1995, Pearce et al. 2003). Direct financial benefits for Darien communities from timber harvesting have apparently been limited, but cativo forests also provide benefits such as non-timber forest products and watershed services that are seldom explicitly valued by policy makers. Historically, the economic importance of cativo swamp forests has been tied to their value for wood production, but little is known about the role they play in the hydrological cycle. Most Darien riverside communities depend to various extents on fishing for subsistence or income, but apparently no research has been done on the use of flooded cativo forests by fish. Changes in water quality or timing of flows may strongly affect fish populations and consequently those whose livelihoods or diets depend on fish (Aylward 2000). Logging has the potential to have profound effects on the physical and biological structure of streams (Campbell and Doeg 1989), especially logging of riparian

PAGE 143

128 forests. Floodplain forests in Brasil were shown to be critical feeding habitats as well as refuges for many species of fish in both whitewater and blackwater rivers (e.g., Saint-Paul et al. 2000). Furthermore, increased sedimentation and turbidity resulting from deforestation can dramatically decrease fish biomass productivity (e.g., Johnson and Kolavalli 1984). Valuation of fisheries is problematical, particularly when subsistence is the end use, but undervaluing fisheries could result in an underestimation of the value of intact flooded forests and the overestimation of the value of competing land uses (Ronnback and Primavera 2000). Despite their limited extent, cativo swamps may have an inordinate value for carbon sequestration because of their large biomass per unit area, but have not been considered separately from other forests in the same ecological lifezone (Corrales 1998). Forest conservation and land use change have major implications when formulating strategies for mitigating future carbon emissions, but patterns of deforestation and forest regrowth can vary dramatically and make estimating carbon fluxes at the regional level difficult (Saleska et al. 2003). Although it is not yet agreed upon that tropical forests are net carbon sources or carbon sinks (Clark 2002), estimates of the amount of biomass in standing forests are necessary in order to estimate the amount of carbon that would be released if forests are converted to other land uses. In tropical moist forest in Panama, Chave et al. (2003) estimated 281 Mg ha -1 of aboveground biomass. In a comparison of tropical moist forest in Darien with riparian forests dominated by cativo, Golley et al. (1969) estimated over 1000 Mg ha -1 in stems alone in a 0.25 ha plot in riparian forest along the Chucunaque River..

PAGE 144

129 Using a regression equation published by Chave et al. (2001) from a pantropical dataset, I estimated 486 Mg ha -1 of aboveground biomass for an unlogged inland swamp and 360 Mg ha -1 in a riverine logged forest along the Sambu River in Darien. The inland swamp is composed of 80% cativo while the riverine forest in 95% cativo, and both estimates are for only cativo trees 10 cm dbh. Natural forest management on communal lands is certainly a more cost efficient manner of sequestering carbon than forest conversion for short-term agriculture, pasture, or fallows (De Jong et al. 2000). Conclusions In addition to explicitly assigning values to environmental services of forests, a key to slowing conversion of forests designated for wood production to other land uses is to maintain their value for timber production. This idea presumes that production forests are found on land suitable for dryland agriculture or pasture, and at least some cativo forests are not so suited. Riverine cativo forests that are near mangrove forests may be inundated by slightly brackish water, while some inland swamps are probably unsuitable for the sorts of agriculture practiced in Darien due to prolonged flooding. Unfortunately, the former are degraded from repeated-entry logging under the subsistence permit system and are increasingly infested by lianas (Grauel and Putz 2004). Inland swamps and most other forest types in Darien that are logged by private companies with either concession or community permits, are subjected to timber harvesting but not forest management. Current practices, under all permit types, degrade production forests by damaging advanced regeneration. Logging of riverine forests by labor-intensive methods and capital-intensive, mechanized logging of inland swamps may result in similar trajectories of land use change if the species of timber sought are the same. Similar to the landscape in the

PAGE 145

130 Colombian province of Choco where these two types of timber harvesting occur, the southern half of Darien is characterized as a region of small agricultural holdings embedded in a relatively stable landscape of heavily degraded forests (Sierra et al. 2003). Although it appears that the boom period of cativo logging has passed, sustainable forest management by smallholders is still possible in such a landscape (Pinedo-Vasquez et al. 2001). The Panamanian Environmental Ministry (ANAM) has taken measures to improve forest management in Darien, but a few additional steps could substantially increase progress towards the goal of sustainable forest management of cativo forests. With various international funding sources, particularly the International Tropical Timber Organization (ITTO), ANAM has installed permanent plots in different forest types to gather basic demographic and forest dynamic information, a critical prerequisite for realistic setting of harvest cycles and intensities (INRENARE 1982, Mariscal et al. 1999, Grauel and Pineda M. 2001). ITTO funded projects in the late 1990s and early 2000s to promote sustainable management of cativo and mangrove forests as well as the tagua palm (Phytelephas seemanii), a key non-timber forest species. Now that important baseline information is available, ITTO could fund efforts at strengthening the capacities of the various stakeholders involved in timber harvesting in Darien to employ reduced-impact harvest techniques such as management planning and improved tree felling and log extraction techniques. Even indigenous communities who dont carry out logging themselves would benefit. Although many such communities have legal land tenure and contract out the actual logging, they often lack the information necessary to be able to

PAGE 146

131 distinguish good logging practices from bad. Such training efforts could steer what is now essentially timber mining towards true forest management. Foresters called for a change in policy (INRENARE 1987) and the Panamanian government designated forests not suitable for agricultural use as production forests in the Forestry Law of 1994. Many Panamanian foresters recognize the potential benefits of extending the current concession period from five to twenty years. Combining longer concession periods with regulations stipulating improved forest management methods could help assure that production forests are maintained under forest cover and reduce the possibility of forest conversion after timber mining. The goal of forest conservation is to optimize in a sustainable and equitable manner the contribution of forests to the prosperity and well-being of a wide variety of stakeholders (Poore et al. 1998). Forest degradation is not caused by local populations disinterest in conservation but in part by historic centralization of control over forest resources, and the resulting problems of enforcing property rights while promoting sustainable livelihoods (Forest Trends 2003). The Panamanian Environmental Ministry is actively pursuing innovative partnerships with riverine communities in Darien to promote cativo forest management by transferring certain tenure rights to the communities over what is now public land. Although cativo forests seem to be in a transition phase during which their value as sources of timber is declining, this could change suddenly given that their main wood output (plywood) is produced for a highly volatile market. The watershed and carbon storage benefits that result from conserving cativo forests accrue to society as a whole, but the costs are borne by local communities

PAGE 147

132 in the form of foregone agricultural production. The challenge is to create incentives for swamp forest conservation that benefit the people who determine their fates.

PAGE 148

APPENDIX A MODELING METHODOLOGY USED IN CHAPTER 5 Regression analyses are used to fit growth estimates as a continuous function of diameter, and then instantaneous changes in dbh are calculated by treating the growth curve as a differential equation (Condit et al. 1993a). The method begins by fitting a quadratic regression to data on relative diameter growth. For each site, I expressed growth as a function of diameter with: g = aL 2 + bL + c Eq. 1 where g is instantaneous growth, L is natural log-transformed dbh, and a, b, and c are parameters. Instantaneous growth as defined by Condit et al. (1993) is: [ln(dbh t ln(dbh 0 )]/t Eq. 2 where the subscripts t and zero denote measurements taken at time t and time 0. At low growth rates, instantaneous growth differs little from relative growth (Condit et al. 1993a). Note that all diameters in my database are in mm, although I present results in cm. Variable names are lower case and Systat commands are upper case. The command in Systat 9.01 to express growth as a function of diameter is: NONLIN MODEL g = a*(LOG(dbhmm)^2)+b*LOG(dbhmm)+c ESTIMATE / GN and results in parameter estimates (Table A) and scatter plots with fitted lines (Figure A). 133

PAGE 149

134 Table A. Parameter estimates for the four sites Parameter Estimates Site a b c Casarete -0.009143 0.094461 -0.228965 Sambu 0.006201 -0.090492 0.327912 Juanacati -0.008008 0.073781 -0.133809 Naranzati -0.005378 0.054901 -0.122269 Casarete Sambu Juanacati Naranzati Figure A-1. Quadratic regression curves fit to relative growth data for the four sites. An explicit solution to Eq. 1, found by direct integration, takes on different forms for different parameter values (Condit et al. 1993a). If b 2 /4a 2 < c/a and a 0, then, letting k 2 = c/a-b 2 /4a 2 the solution is: t = (1/ak)*arctan[(L+b/2a)/k] + m Eq. 3 If b 2 /4a 2 > c/a and a > 0, letting k 2 = b 2 /4a 2 -c/a, the solution becomes:

PAGE 150

135 t = (1/2ak)*ln[(L+b/2a-k) / (L+b/2a+k)] + m Eq. 4 The third solution applies if b 2 /4a 2 > c/a but a < 0, and, letting k 2 = b 2 /4a 2 -c/a, is: t = (1/2ak)*ln[(L+b/2a+k) / (-L-b/2a+k)] + m Eq. 5 where t is time and L is natural log-transformed dbh. The constant m is the constant of integration and is found at the initial conditions of t = 0 at L = ln(10 cm). For Casarete, Juanacati, and Naranzati, the parameter estimates resulted in the solution of Eq. 5, while the parameter values for the Sambu data led to Eq. 4. Solving the respective equations for m at time t = 0 is the next step, and the resulting constants of integration were: Casarete 38.612; Sambu 60.940; Juanacati 0.042; and Naranzati 28.700. The growth trajectories in Figure 5 are the result of solving the site-appropriate equation (in this case either Eq. 4 or Eq. 5) for the entire range of diameters in 1 cm steps. In order to project future volumes with this method, the age, that is, the time-since-minimum-diameter (in this case 10 cm) must be estimated for each stem. Using the parameters for Casarete as an example, the Systat code to calculate this starting age (variable initime) is LET a = -.009143 LET b = .094461 LET k = 1.302807 LET m = 38.612355 LET L00 = LOG(dbhmm00) LET eq5= -(1/(2*a*k))*(LOG((L00+(b/(2*a))+k)/(-L00-(b/(2*a))+k))) LET initime = eq5+m The variable which I called initime is necessary in order to project future diameters using either equation 3, 4 or 5 above. Next, the appropriate equation is rearranged to solve for the future diameter. In the case of Casarete, rearranging Eq.5 to estimate the diameter 5 years in the future is accomplished with the following line of Basic code:

PAGE 151

136 LET dbh5yr = exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+5)-m)))+(k*exp((-2*a*k)*((initime+5)-m))))/(1+exp((-2*a*k)*((initime+5)-m)))) I used this code in a single short program that uses height estimates and basal area to estimate the commercial volume of each tree. The program applies different height estimates depending on the diameter of the tree at any given 5 year point in time. For example, to estimate volume of trees 40 cm in the year 2000: IF dbhmm00>=600 THEN LET ht00=12 IF (dbhmm00=>400) AND (dbhmm00<600) THEN LET ht00=11.02 IF dbhmm00=>400 THEN LET vol00=.64151329*ba00m2*ht00 For Casarete, the remainder of the program estimates volume every 5 years up to 65 years in the future: LET dbh5yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+5)-m)))+(k*exp((-2*a*k)*((initime+5)-m))))/(1+exp((-2*a*k)*((initime+5)-m)))) LET ba5=dbh5yr^2*.0000007854 IF dbh5yr>=600 THEN LET ht5=12 IF (dbh5yr=>400) AND (dbh5yr<600) THEN LET ht5=11.02 IF dbh5yr=>400 THEN LET vol5=.64151329*ba5*ht5 LET dbh10yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+10)-m)))+(k*exp((-2*a*k)*((initime+10)-m))))/(1+exp((-2*a*k)*((initime+10)-m)))) LET ba10=dbh10yr^2*.0000007854 IF dbh10yr>=600 THEN LET ht10=12 IF (dbh10yr=>400) AND (dbh10yr<600) THEN LET ht10=11.02 IF dbh10yr=>400 THEN LET vol10=.64151329*ba10*ht10 LET dbh15yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+15)-m)))+(k*exp((-2*a*k)*((initime+15)-m))))/(1+exp((-2*a*k)*((initime+15)-m)))) LET ba15=dbh15yr^2*.0000007854 IF dbh15yr>=600 THEN LET ht15=12 IF (dbh15yr=>400) AND (dbh15yr<600) THEN LET ht15=11.02 IF dbh15yr=>400 THEN LET vol15=.64151329*ba15*ht15 LET dbh20yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+20)-m)))+(k*exp((-2*a*k)*((initime+20)-m))))/(1+exp((-2*a*k)*((initime+20)-m)))) LET ba20=dbh20yr^2*.0000007854 IF dbh20yr>=600 THEN LET ht20=12 IF (dbh20yr=>400) AND (dbh20yr<600) THEN LET ht20=11.02 IF dbh20yr=>400 THEN LET vol20=.64151329*ba20*ht20

PAGE 152

137 let dbh25yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+25)-m)))+(k*exp((-2*a*k)*((initime+25)-m))))/(1+exp((-2*a*k)*((initime+25)-m)))) LET ba25=dbh25yr^2*.0000007854 IF dbh25yr>=600 THEN LET ht25=12 IF (dbh25yr=>400) AND (dbh25yr<600) THEN LET ht25=11.02 IF dbh25yr=>400 THEN LET vol25=.64151329*ba25*ht25 LET dbh30yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+30)-m)))+(k*exp((-2*a*k)*((initime+30)-m))))/(1+exp((-2*a*k)*((initime+30)-m)))) LET ba30=dbh30yr^2*.0000007854 IF dbh30yr>=600 THEN LET ht30=12 IF (dbh30yr=>400) AND (dbh30yr<600) THEN LET ht30=11.02 IF dbh30yr=>400 THEN LET vol30=.64151329*ba30*ht30 LET dbh35yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+35)-m)))+(k*exp((-2*a*k)*((initime+35)-m))))/(1+exp((-2*a*k)*((initime+35)-m)))) LET ba35=dbh35yr^2*.0000007854 IF dbh35yr>=600 THEN LET ht35=12 IF (dbh35yr=>400) AND (dbh35yr<600) THEN LET ht35=11.02 IF dbh35yr=>400 THEN LET vol35=.64151329*ba35*ht35 LET dbh40yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+40)-m)))+(k*exp((-2*a*k)*((initime+40)-m))))/(1+exp((-2*a*k)*((initime+40)-m)))) LET ba40=dbh40yr^2*.0000007854 IF dbh40yr>=600 THEN LET ht40=12 IF (dbh40yr=>400) AND (dbh40yr<600) THEN LET ht40=11.02 IF dbh40yr=>400 THEN LET vol40=.64151329*ba40*ht40 LET dbh45yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+45)-m)))+(k*exp((-2*a*k)*((initime+45)-m))))/(1+exp((-2*a*k)*((initime+45)-m)))) LET ba45=dbh45yr^2*.0000007854 IF dbh45yr>=600 THEN LET ht45=12 IF (dbh45yr=>400) AND (dbh45yr<600) THEN LET ht45=11.02 IF dbh45yr=>400 THEN LET vol45=.64151329*ba45*ht45 LET dbh50yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+50)-m)))+(k*exp((-2*a*k)*((initime+50)-m))))/(1+exp((-2*a*k)*((initime+50)-m)))) LET ba50=dbh50yr^2*.0000007854 IF dbh50yr>=600 THEN LET ht50=12 IF (dbh50yr=>400) AND (dbh50yr<600) THEN LET ht50=11.02 IF dbh50yr=>400 THEN LET vol50=.64151329*ba50*ht50 LET dbh55yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+55)-m)))+(k*exp((-2*a*k)*((initime+55)-m))))/(1+exp((-2*a*k)*((initime+55)-m)))) LET ba55=dbh55yr^2*.0000007854

PAGE 153

138 IF dbh55yr>=600 THEN LET ht55=12 IF (dbh55yr=>400) AND (dbh55yr<600) THEN LET ht55=11.02 IF dbh55yr=>400 THEN LET vol55=.64151329*ba55*ht55 LET dbh60yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+60)-m)))+(k*exp((-2*a*k)*((initime+60)-m))))/(1+exp((-2*a*k)*((initime+60)-m)))) LET ba60=dbh60yr^2*.0000007854 IF dbh60yr>=600 THEN LET ht60=12 IF (dbh60yr=>400) AND (dbh60yr<600) THEN LET ht60=11.02 IF dbh60yr=>400 THEN LET vol60=.64151329*ba60*ht60 LET dbh65yr=exp((-(b/(2*a))-k-((b/(2*a))*exp((-2*a*k)*((initime+65)-m)))+(k*exp((-2*a*k)*((initime+65)-m))))/(1+exp((-2*a*k)*((initime+65)-m)))) LET ba65=dbh65yr^2*.0000007854 IF dbh65yr>=600 THEN LET ht65=12 IF (dbh65yr=>400) AND (dbh65yr<600) THEN LET ht65=11.02 IF dbh65yr=>400 THEN LET vol65=.64151329*ba65*ht65 Once the variables volxx were created, for each 5, 10, or 20 year harvest, I selected only trees 60 cm dbh, but < 60 cm dbh 5, 10, or 20 years previously (depending on the cutting cycle being simulated), calculated the total volume, and estimated their average diameter as it was in the year 2000. For example, the following two blocks of code simulate a harvest at years 15 and 20, in which only trees that attained 60 cm dbh in a given 5 year period can be included in a harvest: SELECT (dbh15yr=> 600) AND (dbh10yr< 600) STATS vol15 / SUM N STATS dbhmm00 / MEAN SD N SELECT (dbh20yr=> 600) AND (dbh15yr< 600) STATS vol20 / SUM N STATS dbhmm00 / MEAN SD N In this way I simulated a timber harvest every 5 years (as well as 10 and 20 years for the riverine sites) with a 60 cm dbh cutting limit (as well as 80, 90, and 100 cm dbh limits for the inland swamp site).

PAGE 154

APPENDIX B SOURCE DATA FOR CHAPTERS 4 AND 5 The data in identical files cativodata.xls and cativodata.csv are the source data for Chapters 4 and 5. These files contain all diameter measurements, associated codes and notes, and annual growth and basal area calculations for each individual stem. The species names recorded in the year 2000 are the most reliable, as earlier years contain some common names or misidentifications. Codes for individual stems, initially designated in Spanish, are: S = suelo; a prostrate stem. I = inclinado; an inclined stem. M = mltple; erect stem sprouts from an inclined or prostrate stem. R = rebrote; resprouted erect stem. B = bifurcado; a forked stem. Q = quebrado; a broken stem. E = herido; a wounded trunk; H = hinchado, swollen trunk; A = arqueado, a stem arched over, by a fallen tree or branch. Most column titles are self-explanatory. The column titles agroXX-XX and rgroXX-XX refer to annual absolute growth and annual relative growth, respectively, for given census intervals. The column title POMm contains the Point Of Measurement above the ground of individual stems in meters; M distance from root m (resprouts) pertains to stems coded as M and is the distance in meters that these stems are found from the roots of the parent, inclined or protrate, stem. The crown illumination index in this data is from Clark and Clark (1999). The columns longagro and longrgro contain annual growth increments (absolute and relative growth, repectively) based on the longest census interval available 139

PAGE 155

140 for each tree. Because plot installation work was interrupted in 1997 at Juanacati (paramilitary incursion) and Naranzati (I contracted dengue), and some plots were installed in 1998, the last column origdbhmm contains the first diameter measurement, whether from 1997 or 1998, in a single column.

PAGE 156

LIST OF REFERENCES Acosta Sols, M. 1947. Commercial possibilities of the forests of Ecuadormainly Esmereldas Province. Tropical Woods 89:1. Adams, M. 2003. Is this the end for thin-panel tropical plywood? Tropical Forest Update 13:18. Aiba, S. I., and K. Kitayama. 2002. Effects of the 1997 El Nino drought on rain forests of Mount Kinabalu, Borneo. Journal of Tropical Ecology 18:215. Allen, P. H. 1956. The Rain Forests of Golfo Dulce. University of Florida Press, Gainesville. 415 p. Alvarado Q., R., E. Rodrguez R., R. Samaniego, and C. Guerrero. 1996. Situacin actual de los bosques de cativo (Prioria copaifera) en la provincia de Darin, Panam. Instituto Nacional de Recursos Naturales Renovables/Fundacin PANAMA/Unin Internacional para la Conservacin de la Naturaleza, Panam. Alvira, D., F. E. Putz, and T. S. Fredericksen. 2004. Liana loads and post-logging liana densities after liana cutting in a lowland forest in Bolivia. Forest Ecology and Management 190:73. ANAM. 1999. Panam Informe Ambiental. Autoridad Nacional del Ambiente, Panam. Anonymous. 1933. New trade names for cativo. Tropical Woods 35:48. Appanah, S., and F. E. Putz. 1984. Climber abundance in virgin dipterocarp forest and the effect of pre-felling climber cutting on logging damage. The Malaysian Forester 47:335. Avalos, G., and S. S. Mulkey. 1999. Seasonal changes in liana cover in the upper canopy of a neotropical dry forest. Biotropica 31:186. Aylward, B. 2000. Land-use, hydrological function, and economic valuation. Pages 47 in M. Bonell and L. A. Bruijnzeel, editors. Forest-Water-People in the Humid Tropics. Cambridge University Press, Kuala Lumpur, Malaysia. Bachmura, F. T. 1972. La Economa Forestal de la Repblica de Panam. Informe Tcnico 16, La Organizacin de las Naciones Unidas para la Agricultura y la Alimentacin, Panama. 141

PAGE 157

142 Barbier, E. B. 1995. The economics of forestry and conservation: Economic values and policies. Commonwealth Forestry Review 74:26. Barbier, E. B., and J. C. Burgess. 2001. The economics of tropical deforestation. Journal of Economic Surveys 15:413. Barbour, W. R. 1952. Cativo. Journal of Forestry 50:96. Barker, M. G., and D. Perez-Salicrup. 2000. Comparative water relations of mature mahogany (Swietenia macrophylla) trees with and without lianas in a subhumid, seasonally dry forest in Bolivia. Tree Physiology 20:1167. Bertault, J. G., and P. Sist. 1997. An experimental comparison of different harvesting intensities with reduced-impact and conventional logging in East Kalimantan, Indonesia. Forest Ecology and Management 94:209. Bethel, J. S. 1976. Forests in Central America and Panama: Which kind, how large and where? Revista De Biologa Tropical 24:143. Bethancourt, G. 2004. Disminuacin de bosques preocupa a ANCON. in El Panam Amrica, Panam, March 3. Bigler, C., and H. Bugmann. 2003. Growth-dependent tree mortality models based on tree rings. Canadian Journal of Forest Research 33:210. Blakesley, D., S. Elliott, C. Kuarak, P. Navakitbumrung, S. Zangkum, and V. Anusarnsunthorn. 2002a. Propagating framework tree species to restore seasonally dry tropical forest: implications of seasonal seed dispersal and dormancy. Forest Ecology and Management 164:31. Blakesley, D., K. Hardwick, and S. Elliott. 2002b. Research needs for restoring tropical forests in Southeast Asia for wildlife conservation: Framework species selection and seed propagation. New Forests 24:165. Boltz, F., T. P. Holmes, and D. R. Carter. 2003. Economic and environmental impacts of conventional and reduced-impact logging in tropical South America: A comparative review. Forest Policy and Economics 5:69. Bowles, I. A., R. E. Rice, R. A. Mittermeier, and G. A. B. da Fonseca. 1998. Logging on in the rain forests Response. Science 281:1455. Bruner, A. G., R. E. Gullison, R. E. Rice, and G. da Fonseca. 2001. Effectiveness of parks in protecting tropical biodiversity. Science 291:125. Burnham, R. J. 2002. Dominance, diversity, and distribution of lianas in Yasun, Ecuador: Who is on top? Journal of Tropical Ecology 18:845.

PAGE 158

143 Butterfield, R. P. 1995. Promoting biodiversity Advances in evaluating native species for reforestation. Forest Ecology and Management 75:111. Cabarle, B. J. 1998. Logging on in the rain forests. Science 281:1453. Campbell, I. C., and T. J. Doeg. 1989. Impact of timber harvesting and production on streams a review. Australian Journal of Marine and Freshwater Research 40:519539. Cannon, C. H., D. R. Peart, and M. Leighton. 1998. Tree species diversity in commercially logged Bornean rainforest. Science 281:1366. Cansari, R., D. Castaeda, and W. Harp. 1993. Estudio Socio-Cultural en Tres Regiones del Darin: Ro Balsas, Samb, y Garachine. Instituto Nacional de Recursos Naturales Renovables, Organizacin de las Naciones Unidas para la Educacin, la Ciencia, y la Cultura., Panam, Repblica de Panam. Carse, L. E., T. S. Fredericksen, and J. C. Licona. 2000. Liana-tree species associations in a Bolivian dry forest. Tropical Ecology 41:1. Caspersen, J. P., and R. K. Kobe. 2001. Interspecific variation in sapling mortality in relation to growth and soil moisture. Oikos 92:160. Castillo M., S. 1999. Estudio Socio-Econmico del Cativo. Documento para Discusin Autoridad Nacional del Ambiente. Direccin Nacional Forestal., Panam. Chaplin, G. E. 1985. An integrated silvicultural solution to weedy climber problems in the Solomon Islands. Commonwealth Forestry Review 64:133. Chave, J., B. Riera, and M. Dubois. 2001. Estimation of biomass in a neotropical forest of French Guiana: Spatial and temporal variability. Journal of Tropical Ecology 17:79. Chave, J., R. Condit, S. Lao, J. P. Caspersen, R. B. Foster, and S. P. Hubbell. 2003. Spatial and temporal variation of biomass in a tropical forest: Results from a large census plot in Panama. Journal of Ecology 91:240. Chazdon, R. L. 1998. EcologyTropical forestsLog 'em or leave 'em? Science 281:1295. Choi, S. D., and Y. S. Chang. 2004. Factors affecting the distribution of the rate of carbon uptake by forests in South Korea. Environmental Science & Technology 38:484488. Christiansen, P. 1980. Anlisis de los Recursos Forestales del Darin y su Aprovechamiento Actual y Futuro. Documento de Trabajo 1, FAO, Panama.

PAGE 159

144 Christiansen, P. 1984. Anlisis de Costos de Apeo y Transporte para Abastecer el Aserradero y la Planta de Chapas Planificados en Darin. Documento de Trabajo 10, La Organizacin de las Naciones Unidas para la Agricultura y la Alimentacin, Panama. Clark, D. A. 2002. Are tropical forests an important carbon sink? Re-analysis of the long-term plot data. Ecological Applications 12:3. Clark, D. A., and D. B. Clark. 1994. Climate-induced annual variation in canopy tree growth in a Costa-Rican tropical rain-forest. Journal of Ecology 82:865. Clark, D. A., and D. B. Clark. 1999. Assessing the growth of tropical rain forest trees: Issues for forest modeling and management. Ecological Applications 9:981. Clark, D. B., and D. A. Clark. 1990. Distribution and effects on tree growth of lianas and woody hemiepiphytes in a Costa Rican tropical wet forest. Journal of Tropical Ecology 6:321. Clark, D. A., S. C. Piper, C. D. Keeling, and D. B. Clark. 2003. Tropical rain forest tree growth and atmospheric carbon dynamics linked to interannual temperature variation during 1984-2000. Proceedings of the National Academy of Sciences of the United States of America 100:5852. Condit, R. 1995. Research in large, long-term tropical forest plots. Trends in Ecology & Evolution 10:18. Condit, R. 1998. Tropical Forest Census Plots: Methods and Results from Barro Colorado Island, Panama and a Comparison with Other Plots. Springer-Verlag, Berlin. Condit, R., P. S. Ashton, N. Manokaran, J. V. LaFrankie, S. P. Hubbell, and R. B. Foster. 1999. Dynamics of the forest communities at Pasoh and Barro Colorado: comparing two 50-ha plots. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences 354:1739. Condit, R., S. P. Hubbell, and R. B. Foster. 1993a. Identifying fast-growing native trees from the Neotropics using data from a large, permanent census plot. Forest Ecology and Management 62:123. Condit, R., S. P. Hubbell, and R. B. Foster. 1993b. Mortality and growth of a commercial hardwood "el cativo", Prioria copaifera, in Panama. Forest Ecology and Management 62:107. Condit, R., S. P. Hubbell, and R. B. Foster. 1995a. Demography and harvest potential of Latin American timber species: Data from a large, permanent plot in Panama. Journal of Tropical Forest Science 7:599.

PAGE 160

145 Condit, R., S. P. Hubbell, and R. B. Foster. 1995b. Mortality rates of 205 neotropical tree and shrub species and the impact of a severe drought. Ecological Monographs 65:419. Condit, R., R. Sukumar, S. P. Hubbell, and R. B. Foster. 1998. Predicting population trends from size distributions: A direct test in a tropical tree community. American Naturalist 152:495. CONIF. 1997. Manejo y Conservacin del Ecosistema Catival. Corporacin Nacional de Investigacin y Fomento Forestal, Pizano, S.A. Connell, J. H., and M. D. Lowman. 1989. Low-diversity tropical rain forests: some possible mechanisms for their existence. American Naturalist 134:88. Consultores Ambientales LTDA. 1995. Proyecto de aprovechamiento integral sostenible con silvicultura comunitaria para los bosques del area Domingodo-Truando en el Choc. Pizano S.A., Santa Fe de Bogat. Contreras-Hermosilla, A. 2000. The underlying causes of forest decline. Occasional Paper 30, CIFOR, Jakarta. Cooper, G. P. 1928. Some interesting trees of western Panama. Tropical Woods 14:1. Corrales, L. 1998. Estimacin de la Cantidad de Carbono Almacenado y Capturado (Masa Area) en los Bosques de la Repblica de Panam. Comisin Centroamericana de Ambiente y Desarrollo, Panam. Dalling, J. W., K. E. Harms, and R. Aizpra. 1997. Seed damage tolerance and seedling resprouting ability of Prioria copaifera (el cativo). Journal of Tropical Forest Ecology 13:481. Davies, S. J. 2001. Tree mortality and growth in 11 sympatric Macaranga species in Borneo. Ecology 82:920. De Jong, B. H. J., R. Tipper, and M. Gmez. 2000. An economic analysis of the potential for carbon sequestration by forests: Evidence from southern Mexico. Ecological Economics 33:313. Del Valle, J. I. 1972. Introduccin a la dendrologa de Colombia. Facultad de Ciencias Agrcolas, Universidad Nacional de Colombia, Medelln. Del Valle, J. I. 1979. Curva preliminar de crecimiento del cativo (Prioria copaifera) en bosque virgen empleando el mtodo de los tiempos de paso. Revista Facultad Nacional de Agronomia 32:19. Dickinson, M. B., J. C. Dickenson, and F. E. Putz. 1996. Natural forest management as a conservation tool in the tropics: Divergent views on possibilities and alternatives. Commonwealth Forestry Review 75:309.

PAGE 161

146 Dillenburg, L. R., D. F. Whigham, A. H. Teramura, and I. N. Forseth. 1993. Effects of belowground and aboveground competition from the vines Lonicera japonica and Parthenocissus quinquefolia on the growth of the tree host Liquidambar styraciflua. Oecologia 93:48. Duke, J. A. 1964. Botany. Page 335 in N. Wilson, Raimond Engineers Inc., editor. El Real Environmental Survey, Darien Province, Republic of Panama, 1962. U.S. Army Transportation Research Command, Fort Eustis, Virginia. Duke, J. A. 1975. Plant species in the forest of Darien, Panama. Pages 189 in F. B. Golley, J. T. McGinnis, R. G. C. Clements, and M. J. Duever, editors. Mineral Cycling in a Tropical Moist Forest. University of Georgia Press, Athens, Georgia. Duke, J. A. 1986. Isthmian Ethnobotanical Dictionary. Scientific Publishers, Jodhpur. Echavarria A., J. A., and T. Varon P. 1988. Estudio dendrolgico de la asociacin catival en La Balsa, Choc. Ingeniero Forestal. Universidad Nacional de Colombia, Medellin. Engel, V. L., and J. A. Parrotta. 2001. An evaluation of direct seeding for reforestation of degraded lands in central Sao Paulo State, Brazil. Forest Ecology and Management 152:169. Escobar, O., and J. R. Rodriguez. 1993. Las Maderas en Colombia. Medelln. Escobar, J. A., and I. F. Vasquez. 1987. Caracterizacin de tipos de cativales. Ingeniero Forestal. Universidad Nacional de Colombia, Medellin. Escobar Munera, L. 1981. Anlisis estructural, estudio de la regeneracin y tratamientos silviculturales en un bosque de catival. 7, Empresa Maderera del Atrato S. A., Medellin. Estrada Lpez, J. R., and J. Gmez Quiceno. 1988. Reconocimiento, identificacin, y control de perforadores de cativo (Prioria copaifera Griseb.) de la zona de Urab. Ingeniero Forestal. Universidad Nacional de Colombia, Medelln. Evans, J. 1992. Plantation Forestry in the Tropics: Tree Planting for Industrial, Social, Environmental, and Agroforestry Purposes, 2nd edition. Clarendon Press, Oxford. FAO. 1982. Caractersticas y Usos de 19 Especies con Valor Comercial en Panam. Centro de Tecnologa de la Madera de la Direccin Nacional de Recursos Naturales Renovables, Panama City. FAO. 1990. Plan de Manejo para la Produccin Sostenida de los Cativales del Darin. Pages 263 in Instituto Nacional. de Recursos Naturales Renovables, editor. Plan de Accin Forestal de Panam. Food and Agriculture Organization, Panam.

PAGE 162

147 Favrichon, V. 1998. Modeling the dynamics and species composition of a tropical mixed-species uneven-aged natural forest: Effects of alternative cutting regimes. Forest Science 44:113. Ferrer, A. (1999). Comparacin de las comunidades de hongos asociadas con Prioria copaifera (Cativo) en Panam. In Anexo 2: Actividades del Componente de Cativo, Reunin del Comit Directivo, Proyecto Manejo de Cativales y Productos No Maderables con Comunidades Campesinas e Indgenas en Darin, Panam, Proyecto PD 37'95 Rev. 2(F), Panama City. Finegan, B., and M. Camacho. 1999. Stand dynamics in a logged and silviculturally treated Costa Rican rain forest, 1988. Forest Ecology and Management 121:177. Forest Trends. 2003. Strategies for Strengthening Community Property Rights over Forests: Lessons and Opportunities for Practitioners. Forest Trends, Washington, D.C. Foroughbakhch, F., L. A. Hauad, A. E. Cespedes, E. E. Ponce, and N. Gonzalez. 2001. Evaluation of 15 indigenous and introduced species for reforestation and agroforestry in northeastern Mexico. Agroforestry Systems 51:213. Fox, J. E. D. 1968. Logging damage and the influence of climber cutting prior to logging in the lowland dipterocarp forest of Sabah. Malaysian Forester 31:326. Fredericksen, T. S., and B. Mostacedo. 2000. Regeneration of timber species following selection logging in a Bolivian tropical dry forest. Forest Ecology and Management 131:47. Gavin, D. G., and D. P. Peart. 1999. Vegetative life history of a dominant rain forest canopy tree. Biotropica 312:288. Gentry, A. H. 1996. A field guide to the families and genera of woody plants of northwest South America (Colombia, Ecuador, Peru), with supplementary notes on herbaceous taxa. University of Chicago Press, Conservation International, Chicago. Gerwing, J. J. 2001. Testing liana cutting and controlled burning as silvicultural treatments for a logged forest in the eastern Amazon. Journal of Applied Ecology 38:1264. Gerwing, J. J. 2002. Degradation of forests through logging and fire in the eastern Brazilian Amazon. Forest Ecology and Management 157:131. Gerwing, J. J., and E. Vidal. 2002. Changes in liana abundance and species diversity eight years after liana cutting and logging in an eastern Amazonian forest. Conservation Biology 16:544.

PAGE 163

148 Gillis, M. 1992. Forest Concession Management and Revenue Policies. Pages 139 in N. P. Sharma, editor. Managing the World's Forests: Looking for Balance Between Conservation and Development. Kendal/Hunt Publishing Company, Dubuque, Iowa. Godoy, R., H. Overman, J. Demmer, L. Apaza, E. Byron, T. Huanca, W. Leonard, E. Perez, V. Reyes-Garcia, V. Vadez, D. Wilkie, A. Cubas, K. McSweeney, and N. Brokaw. 2002. Local financial benefits of rain forests: comparative evidence from Amerindian societies in Bolivia and Honduras. Ecological Economics 40:397. Golley, F. B., J. T. McGinnis, R. G. Clements, G. I. Child, and M. J. Duever. 1969. The structure of tropical forest in Panama and Colombia. Bioscience 19:693. Golley, F. B., J. T. McGinnis, R. G. C. Clements, and M. J. Duever. 1975. Mineral Cycling in a Tropical Moist Forest Ecosystem. University of Georgia Press, Athens. Gmez, H. D. 1990. Algunos aspectos estructurales de los cativales de la regin de Urub. Ingeniero Forestal. Universidad Nacional de Colombia, Medellin. Gonzlez Apolayo, G. 2003. Extraccin ilegal e informal de madera en Darin. in El Panam Amrica, Panam. May 12. Gonzlez Prez, H., H. D. Gmez T., and F. J. Arteaga C. 1991. Aspectos estructurales de un bosque de cativo en la regin del Bajo Atrato, Colombia. Revista Facultad Nacional de Agronoma, Medelln 44:3. Gonzlez Prez, H. 1995. Anlisis del crecimiento diamtrico de Prioria copaifera en condiciones naturales por medio de un modelo matemtico determinstico. Crnica forestal y del medio ambiente 10:101. Grauel, W. T. 1999. Avances del Componente de Cativo del Perodo junio 1998 a marzo 1999 en, Anexo 2: Actividades del Componente de Cativo, Reunin del Comit Directivo. Proyecto Manejo de Cativales y Productos No Maderables con Comunidades Campesinas e Indgenas en Darin, Panam, Proyecto PD 37 Rev. 2(F), Panam. Grauel, W. T. 2004a. Structure, composition, and dynamics of Prioria copaifera-dominated swamp forests in Darien, Panama. In review. Grauel, W. T. 2004b. Growth and survival of Prioria copaifera (cativo) seedlings Planted along a habitat gradient in a Panamanian swamp. In review. Grauel, W. T., and T. A. Kursar. 1999. Species diversity and stand dynamics of cativo (Prioria copaifera Griseb.) forests in Darien Province, Panama. Pages 69 in C. Kleinn and M. Khl, editors. IUFRO S4.11 Long-Term Observations and Research in Forestry. CATIE, Turrialba, Costa Rica.

PAGE 164

149 Grauel, W. T., and R. Pineda M. 2001. Manual Tcnico para el Manejo Sostenible de los Cativales en Darin, Panama. Autoridad Nacional del Ambiente, Panama City. Grauel, W. T., and F. E. Putz. 2004. Effects of lianas on growth and regeneration of Prioria copaifera in Darien, Panama. Forest Ecology and Management 190: 99108. Guariguata, M. R. 1998. Response of forest tree saplings to experimental mechanical damage in lowland Panama. Forest Ecology and Management 102:103. Gullison, R. E., and J. J. Hardner. 1993. The effects of road design and harvest intensity on forest damage caused by selective logging Empirical results and a simulation model from the Bosque Chimanes, Bolivia. Forest Ecology and Management 59:114. Haggar, J. P., C. B. Briscoe, and R. P. Butterfield. 1998. Native species: A resource for the diversification of forestry production in the lowland humid tropics. Forest Ecology and Management 106:195. Harrar, E. S. 1941. Some physical properties of modern cabinet woods I. Hardness. Tropical Woods 68:1. Harrar, E. S. 1942a. Some physical properties of modern cabinet woods II. Screw-holding power. Tropical Woods 70:1. Harrar, E. S. 1942b. Some physical properties of modern cabinet woods III. Directional and volume shrinkage. Tropical Woods 71:26. Heckadon M., S., F. Herrera, and A. Pastor. 1982. Breve Estudio de los Grupos Humanos del Darin. Pages 81 in S. H. Moreno and A. McKay, editors. Colonizacin y Destruccin de Bosques en Panama. Asociacin Panamea de Antropologa, Panam. Hernndez, A. 1970. Migracin de colonos en Darin. Hombre y Cultura 1:81. Hernandez Hurtado, V. 1984. Propiedades fisico-mechanicas y trabajabilidad de la madera de cativo (Prioria copaifera Griseb) en condicin verde. Ingeniero Forestal. Universidad Nacional de Colombia, Medelln. Herrera-MacBryde, O., and ANCON. 2001. Darien Province and Darien National Park. in. Smithsonian Institution, The World Conservation Union, World Wildlife Fund. Centers of Plant Diversity Project. Accessed April, 2004. http://www.nmnh.si.edu/botany/projects/cpd/ma/ma20.htm Hess, R. W., and M. E. Record. 1950. Foreign wood imports. Tropical Woods 96:21. Hess, R. W., F. F. Wangaard, and F. E. Dickinson. 1950. Properties and uses of tropical woods, II. Tropical Woods 97:1.

PAGE 165

150 Hoheisel, H., and O. Lpez G. 1972. Ensayos preliminares sobre la aptitud del cativo (Prioria copaifera) para la elaboracin de tableros de pajilla-cemento. Universidad Nacional de Colombia, Laboratorio de Productos Forestales, Medelln. Holdridge, L. R. 1964. Ecology. Pages 335 in N. Wilson, Raimond Engineers Inc., editor. El Real Environmental Survey, Darien Province, Republic of Panama, 1962. U.S. Army Transportation Research Command, Fort Eustis, Virginia. Holdridge, L. R. 1970. Manual Dendrolgico para 1000 Especies Arboreas en la Repblica de Panam. Programa de las Naciones Unidas para el Desarrollo, Panam. Holdridge, L. R., and G. Budowski. 1956. Report of an ecological survey of the Republic of Panama. Caribbean Forester 17:92. Holmes, T. P., G. M. Blate, J. C. Zweede, R. Pereira, P. Barreto, F. Boltz, and R. Bauch. 2002. Financial and ecological indicators of reduced impact logging performance in the eastern Amazon. Forest Ecology and Management 163:93. Huth, A., and T. Ditzer. 2000. Simulation of the growth of a lowland Dipterocarp rain forest with FORMIX3. Ecological Modelling 134:1. Ickes, K., S. J. Dewalt, and S. C. Thomas. 2003. Resprouting of woody saplings following stem snap by wild pigs in a Malaysian rain forest. Journal of Ecology 91:222. INRENARE. 1982. Parcelas Permanentes para el Estudio de Crecimiento en Bosque Natural, Darin. Instituto Nacional de Recursos Naturales Renovables, Panama. INRENARE. 1987. Situacin General de los Bosques de Cativo y su Utilizacin. Instituto Nacional de Recursos Naturales Renovables, Panama. ITTO Secretariat. 2003. Reviving plywood. Tropical Forest Update 13:16. Jackson, S. M., T. S. Fredericksen, and J. R. Malcolm. 2002. Area disturbed and residual stand damage following logging in a Bolivian tropical forest. Forest Ecology and Management 166:271. Jaramillo Gallego, E. A., and L. G. Velasquez Salazar. 1992. Factores fisico-anatmicos que inducen el hundimiento de trozas de cativo (Prioria copaifera G). Ingeniero Forestal. Universidad Nacional de Colombia, Medelln. Jennings, S. B., N. D. Brown, and D. Sheil. 1999. Assessing forest canopies and understorey illumination: canopy closure, canopy cover, and other measures. Forestry 72:59.

PAGE 166

151 Jimnez Madrigal, Q. 1995. Familia Caesalpiniaceae. Pages 43 in T. N. C. Instituto Nacional de Biodiversidad, National Heritage Foundation, editor. Arboles maderables en peligro de extincin en Costa Rica. INCAFO S.A., San Jose, Costa Rica. Johns, J. S., P. Barreto, and C. Uhl. 1996. Logging damage during planned and unplanned logging operations in the eastern Amazon. Forest Ecology and Management 89:59. Johnson, N., and B. Cabarle. 1993. Surviving the Cut: Natural Forest Management in the Humid Tropics. World Resources Institute. Washington, D.C. Johnson, S. H., and S. Kolavalli. 1984. Physical and economic impacts of sedimentation on fishing activities; Nam Pong Basin, Northeast Thailand. Water International 9:185. Kaimowitz, D., and A. Angelsen. 1998. Economic Models of Tropical Deforestation: A Review. CIFOR, Bogor, Indonesia. Kainer, K. A., M. Schmink, A. C. P. Leite, and M. J. D. Fadell. 2003. Experiments in forest-based development inWestern Amazonia. Society & Natural Resources 16:869. Kammesheidt, L. 1998. The role of tree sprouts in the restoration of stand structure and species diversity in tropical moist forest after slash-andburn agriculture in Eastern Paraguay. Plant Ecology 139:155. Kammesheidt, L. 1999. Forest recovery by root suckers and above-ground sprouts after slash-and-burn agriculture, fire and logging in Paraguay and Venezuela. Journal of Tropical Ecology 15:143. Kataki, R., and D. Konwer. 2002. Fuelwood characteristics of indigenous tree species of north-east India. Biomass & Bioenergy 22:433. Keenan, R. J., D. Lamb, J. Parrotta, J. Kikkawa, P. Corona, and B. Zeide. 1999. Ecosystem management in tropical timber plantations: Satisfying economic, conservation, and social objectives. Journal of Sustainable Forestry 9:117. Kittredge, D. B., A. O. Finley, and D. R. Foster. 2003. Timber harvesting as ongoing disturbance in a landscape of diverse ownership. Forest Ecology and Management 180:425. Kluge, H. C. 1926. Prioria copaifera. Tropical Woods 5:8. Kobe, R. K., and K. D. Coates. 1997. Models of sapling mortality as a function of growth to characterize interspecific variation in shade tolerance of eight tree species of northwestern British Columbia. Canadian Journal of Forest Research 27:227.

PAGE 167

152 Kobe, R. K., S. W. Pacala, J. A. Silander Jr., and C. D. Canham. 1995. Juvenile tree survivorship as a component of shade tolerance. Ecological Applications 5:517532. Kukachka, B. F. 1965. Cativo, Prioria copaifera Gris. Research Note FPL-095, USDA Forest Service, Forest Products Laboratory, Madison, Wisconsin. Kuroda, K., and K. Shimaji. 1984. The Pinning Method For Marking Xylem Growth in Hardwood Species. Forest Science 30:548. Kurpick, P., U. Kurpick, and A. Huth. 1997. The influence of logging on a Malaysian dipterocarp rain forest: A study using a forest gap model. Journal of Theoretical Biology 185:47. Kursar, T. A., and W. T. Grauel. 2002. Logging from rivers may control human invasions. Conservation Biology 16:285. Kynoch, W., and N. A. Norton. 1938. Mechanical properties of certain tropical woods, chiefly from South America. Bulletin 7, University of Michigan School of Forestry and Conservation, Ann Arbor. Lamb, F. B. 1953. The forests of Darien, Panama. Caribbean Forester 14:128. Leigh, E. 1999. Biomass and productivity of tropical forest. Pages 120 in Tropical Forest Ecology: A View From Barro Colorado Island. Oxford University Press, New York. Lemmon, P. E. 1957. A new instrument for measuring forest overstory density. Journal of Forestry 55:667. Leslie, A. J. 1977. Where contradictory theory and practice co-exist. Unasylva 29:2. Leslie, A. J. 1987. A second look at the economics of natural management systems in tropical mixed forests. Unasylva 39:46. Lin, J., P. A. Harcombe, and M. R. Fulton. 2001. Characterizing shade tolerance by the relationship between mortality and growth in tree saplings in a southeastern Texas forest. Canadian Journal of Forest Research 31:345. Linares Prieto, R. 1987a. Determinacin del tipo de plantn y la poca del ao adecuados para la plantacin de cativo. Serie Tcnica 23, Corporacin Nacional de Investigacin y Fomento Forestal, Bogot. Linares Prieto, R. 1987b. Estudio del catival en Colombia. Pages 54 in Reunin Nacional de Silvicultura. Memoria: Impacto de la Investigacin Silvicultural Tropical en el Desarrollo Econmico Forestal Colombiano. Corporacin Nacional de Investigacin y Fomento Forestal., Bogot.

PAGE 168

153 Linares Prieto, R. 1987c. Uso actual y potencial de los suelos de la asociacin catival en Colombia. in IV Congreso Colombiano de la Ciencia del Suelo. Linares Prieto, R. 1988. Estudio preliminar de la asociacin catival en Colombia. Serie documentacin 17, Corporacin Nacional de Investigacin y Fomento Forestal, Bogot. Linares Prieto, R., and H. Martinez Higuera. 1991. La regeneracin natural temprana del bosque de cativo en Choco-Colombia. Serie Tcnica 30, Corporacin Nacional de Investigacin y Fomento Forestal, Bogot. Linares Prieto, R., J. R. Romero, G. P. Palacio, and R. S. d. Acosta. 1997. Bases Ecolgicas para la Silvicultura del Catival (Priorietum copaiferae). Informe Preliminar del Proyecto sobre Publicacin Tcnica del Catival Corporacin Nacional de Investigacin y Fomento Forestal, Santaf de Bogot. Londoo Londoo, F. L., and H. Gonzlez Prez. 1993. Identificacin de variables relacionadas con el crecimiento diametrico del cativo (Prioria copaifera). Crnica Forestal y del Medio Ambiente 8:25. Lopez, O. R. 2001. Seed flotation and postflooding germination in tropical terra firme and seasonally flooded forest species. Functional Ecology 15:763. Lopez, O. R. 2002. Mechanisms Explaining Low Diversity and Monodominance in Seasonally Flooded Forests. PhD Dissertation. University of Utah, Salt Lake City. Lopez, O. R., and T. A. Kursar. 1999. Flood tolerance of four tropical tree species. Tree Physiology 19:925. Lugo, A. E. 1999. Will concern for biodiversity spell doom to tropical forest management? Science of the Total Environment 240:123. Lugo, A. E., and F. N. Scatena. 1996. Background and catastrophic tree mortality in tropical moist, wet, and rain forests. Biotropica 28:585. Magnusson, W. E., O. P. de Lima, F. Q. Reis, N. Higuchi, and J. F.Ramos. 1999. Logging activity and tree regeneration in an Amazonian forest. Forest Ecology and Mangement 113: 6774. Mahecha Vega, G., R. Rodrguez Soto, and L. E. Acero Duarte. 1984. Estudio dendrolgico de Colombia. IGAC, Universidad Distrital Francisco Jos de Caldas, Bogot. Manokaran, N. 1998. Effect, 34 years later of selective logging in the lowland dipterocarp forest at Posah, Peninsular Malaysia, and implications on present-day logging in the hill forests. Pages 41 in S. S. Lee, Y. M. Dan, I. D. Gauld, and J. Bishop, editors. Conservation, Management and Development of Forest Resources. Forest Research Institute Malaysia, Kuala Lumpur.

PAGE 169

154 Mariscal, E., R. Martnez, and T. Hagiwara. 1999. Distribucin diametrica y estimativa del volmen de la especie Prioria copaifera (cativo) a partir de modelo de regresin. Pages 78 in Proyecto de Desarrollo Tcnico de la Conservacin de los Bosques (CEMARE), editor. Proyeccin de la Investigacin Forestal Hacia el Siglo XXI. Centro para el Desarrollo Sostenible, Ro Hato, Panam. Mariscal, E., M. H. Wishnie, and J. Deago. 2002. Crecimiento comparativo de especies nativas y Teca plantadas en parcelas puras y mixtas en reas invadidas por la graminea Sacharum spontaneum (L.) Gramineae, utilizando mtado de quema y sin quema: Informe del establecimiento del ensayo. ECO-07-Esp, Proyecto de Reforestacin con Especies Nativas (PRORENA), Yale University, Smithsonian Tropical Research Institute, Panama City. Martn Nuez, I. 1984. Estudio de prefactibilidad de desarrollo industrial forestal del rea comprendida entre los ros Chico, Tupisa, y Tuquesa (Antecedentes y situacin forestal). Documento de Trabajo 8, INRENARE, FAO, Panam. Martnez Higuera, H. 1989. Algunas experiencias de investigacin en la asociacin catival en Colombia. Pages 25 in CATIE, editor. Curso Intensivo Internacional en Silvicultura y Manejo de Bosques Naturales Tropicales. CATIE, Turrialba, Costa Rica. Martin-Smith, K. M. 1998. Effects of disturbance caused by selective timber extraction on fish communities in Sabah, Malaysia. Environmental Biology of Fishes 53:155167. Maxwell, S. E., and H. D. Delaney. 1999. Designing Experiments and Analyzing Data: A Model Comparison Perspective. Lawrence Erlbaum Associates, Inc., Mahwah, N.J. Mayo Melendez, E. 1965. Algunas caractersticas ecolgicas de los bosques inundables de Darin, Panam, con miras a su posible utilizacin. Turrialba 15:336. McKenzie, T. A. 1972. Observations on growth and a technique for estimating annual growth in Prioria copaifera. Turrialba 22:352. Mesquita, R. C. G., P. Delamonica, and W. F. Laurance. 1999. Effect of surrounding vegetation on edge-related tree mortality in Amazonian forest fragments. Biological Conservation 91:129. Mondragn T., F., N. Prez, and F. E. Pez P. 1994. Estudio del habitat de la Tortuga Dulceacuicola "Jicotea" Trachemys scripta callirostris. Gray (1855) en la regin del Medio Atrato, Choc, Colombia. Convenio Pizano S.A. CONIF, Sante Fe de Bogot. Montagnini, F. 2001. Strategies for the recovery of degraded ecosystems: Experiences from Latin America. Interciencia 26:498.

PAGE 170

155 Montero G., M. I. 1996. Tabla de Vida y Aplicacin de un Modelo Matricial al Estudio de la Demografa de Prioria copaifera. Universidad Nacional de Colombia, Medellin. Moulaert, A., J. P. Mueller, M. Villarreal, R. Piedra, and L. Villalobos. 2002. Establishment of two indigenous timber species in dairy pastures in Costa Rica. Agroforestry Systems 54:31. Muoz Valencia, A. 1966. Algunos aspectos de los bosques de cativo en la regin de Urub. Ingeniero Forestal. Universidad Distrital Francisco Jos de Caldas, Bogot. Nabe-Nielsen, J. 2001. Diversity and distribution of lianas in a neotropical rain forest, Yasuni National Park, Ecuador. Journal of Tropical Ecology 17:1. Nabe-Nielsen, J. 2002. Growth and mortality rates of the liana Machaerium cuspidatum in relation to light and topographic position. Biotropica 34:319. Negrelle, R. R. B. 1995. Sprouting after uprooting of canopy trees in the Atlantic rain forest of Brazil. Biotropica 27:448. Negreros-Castillo, P., and R. B. Hall. 2000. Sprouting capability of 17 tropical tree species after overstory removal in Quintana Roo, Mexico. Forest Ecology and Management 126:399. Nelson, G. C., V. Harris, and S. W. Stone. 1999. Spatial Econometric Analysis and Project Evaluation: Modeling Land Use Change in the Darin. Inter-American Development Bank, Washington, D.C. Nepstad, D. C., A. Verissimo, A. Alencar, C. Nobre, E. Lima, P. Lefebvre, P. Schlesinger, C. Potter, P. Moutinho, E. Mendoza, M. Cochrane, and V. Brooks. 1999. Large-scale impoverishment of Amazonian forests by logging and fire. Nature 398:505. OEA. 1978. Repblica de Panam Proyecto de Desarrollo Integrado de la Regin Oriental de Panam Darin. Organizacin de los Estados Americanos. Accessed April, 2004. http://www.oas.org/usde/publications/Unit/oea30s/begin.htm#Contents. O'Brien, S. T., S. P. Hubbell, P. Spiro, R. Condit, and R. B. Foster. 1995. Diameter, height, crown and age relationships in eight neotropical tree species. Ecology 76:1926. Oliver, C. D., and B. C. Larson. 1996. Forest Stand Dynamics, Update Edition edition. John Wiley & Sons, Inc, New York. Pacala, S. W., C. D. Canham, J. Saponara, J. A. Silander, R. K. Kobe, and E. Ribbens. 1996. Forest models defined by field measurements: Estimation, error analysis and dynamics. Ecological Monographs 66:1.

PAGE 171

156 Paciorek, C. J., R. Condit, S. P. Hubbell, and R. B. Foster. 2000. The demographics of resprouting in tree and shrub species of a moist tropical forest. Journal of Ecology 88:765. Panfil, S. N., and R. E. Gullison. 1998. Short term impacts of experimental timber harvest intensity on forest structure and composition in the Chimanes Forest, Bolivia. Forest Ecology and Management 102:235. Parren, M., and F. Bongers. 2001. Does climber cutting reduce felling damage in southern Cameroon? Forest Ecology and Management 141:175. Parrotta, J. A., and O. H. Knowles. 1999. Restoration of tropical moist forests on bauxite-mined lands in the Brazilian Amazon. Restoration Ecology 7:103. Pearce, D., F. E. Putz, and J. K. Vanclay. 2003. Sustainable forestry in the tropics: panacea or folly? Forest Ecology and Management 172:229. Pereira, R., J. Zweede, G. P. Asner, and M. Keller. 2002. Forest canopy damage and recovery in reduced-impact and conventional selective logging in eastern Para, Brazil. Forest Ecology and Management 168:77. Prez-Salicrup, D. R. 2001. Effect of liana cutting on tree regeneration in a liana forest in Amazonian Bolivia. Ecology 82:389. Prez-Salicrup, D. R., and M. G. Barker. 2000. Effect of liana cutting on water potential and growth of adult Senna multijuga (Caesalpinioideae) trees in a Bolivian tropical forest. Oecologia 124:469. Prez-Salicrup, D. R., V. L. Sork, and F. E. Putz. 2001. Lianas and trees in a liana forest of Amazonian Bolivia. Biotropica 33:34. Philip, M. S. 1994. Measuring Trees and Forests, 2nd edition. CAB International, Oxon, UK. Phillips, P. D., T. E. Brash, I. Yasman, P. Subagyo, and P. R. van Gardingen. 2003. An individual-based spatially explicit tree growth model for forests in East Kalimantan (Indonesian Borneo). Ecological Modelling 159:1. Pinard, M. A., and F. E. Putz. 1996. Retaining forest biomass by reducing logging damage. Biotropica 28:278. Pinard, M. A., F. E. Putz, and J. C. Licona. 1999. Tree mortality and vine proliferation following a wildfire in a subhumid tropical forest in eastern Bolivia. Forest Ecology and Management 116:247. Pinedo-Vasquez, M., D. J. Zarin, K. Coffey, C. Padoch, and F. Rabelo. 2001. Post-boom logging in Amazonia. Human Ecology 29:219-239.

PAGE 172

157 Pittier, H., and C. D. Mell. 1931. The cativo or Prioria-tree. Pages 26 in A Century of Trees of Panama. Pizano SA. 1995. Proyecto de aprovechamiento integral sostenible con silvicultura comunitaria para los bosques del area Domingodo-Truando en el Choco. Pizano, SA, Bogota. Poore, D., et al. 1998. No forest without management. Tropical Forest Update 8. Porter, D. M. 1973. The vegetation of Panama: A review. Pages 167 in A. Graham, editor. Vegetation and Vegetational History of Northern Latin America. Elsevier Scientific Publishing Company, New York. Putz, F. E. 1983. Liana biomass and leaf area of a tierra firme forest in the Rio Negro Basin, Venezuela. Biotropica 15:185. Putz, F. E. 1984. The natural history of lianas on Barro Colorado Island, Panama. Ecology 65:1713. Putz, F. E. 1990. Liana stem diameter growth and mortality rates on Barro Colorado Island, Panama. Biotropica 22:103. Putz, F. E. 1991. Silvicultural effects of lianas. Pages 493 in F. E. Putz and H. A. Mooney, editors. The Biology of Vines. University Press, Cambridge. Putz, F. E. 2000. Economics of home grown forestry. Ecological Economics 32:9. Putz, F. E., and N. V. L. Brokaw. 1989. Sprouting of broken trees on Barro Colorado Island, Panama. Ecology 70:508. Putz, F. E., and P. Chai. 1987. Ecological studies of lianas in Lambir National Park, Sarawak, Malaysia. Journal of Ecology 75:523. Putz, F. E., D. P. Dykstra, and R. Heinrich. 2000a. Why poor logging practices persist in the tropics. Conservation Biology 14:951. Putz, F. E., K. H. Redford, J. G. Robinson, R. Fimbel, and G. M. Blate. 2000b. Biodiversity Conservation in the Context of Tropical Forest Management. Wildlife Conservation Society, Washington, D.C. Raintree, J. B. 1991. Socioeconomic Attributes of Trees and Tree Planting Practices. Food and Agriculture Organization, Rome. Record, S. J. 1925. Introductory note. Tropical Woods 1:1. Reid, J. W., and R. E. Rice. 1997. Assessing natural forest management as a tool for tropical forest conservation. Ambio 26:382.

PAGE 173

158 Repetto, R. 1988. The Forest for the Trees? Government Policies and the Misuse of Forest Resources. World Resources Institute, Washington, D.C. Repblica de Panam. 1978. Proyecto de Desarrollo Integrado de la Regin Oriental de Panama Darin. Organizacin de Estados Americanos, Washington, D.C. Repblica de Panam. 1995. Situacin Fsica Meterologa. Contralora General de la Repblica, Direccin de Estadstica y Censo, Panam. Repblica de Panam. 2003. Gua Tcnica para la Reforestacin en Panam. Servicio Nacional de Desarrollo y Administracin Forestal, Direccin Nacional de Patrimonio Natural, Autoridad Nacional del Ambiente, Panam. Rice, R. E., R. E. Gullison, and J. W. Reid. 1997. Can sustainable management save tropical forests? Scientific American 276:44. Rice, R. E., C. A. Sugal, S. M. Ratay, and G. A. Fonseca. 2001. Sustainable forest management: A review of conventional wisdom. Advances in Applied Biodiversity Science, No. 3, p. 1. Rijks, M. H., E. J. Malta, and R. J. Zagt. 1998. Regeneration through sprout formation in Chlorocardium rodiei (Lauraceae) in Guyana. Journal of Tropical Ecology 14:463475. Rimbach, A. 1932. The forests of Ecuador. Tropical Woods 31:1. Romero, C. and G. Andrade. 2004. International conservation organizations and the fate of local tropical forest conservation initiatives. Conservation Biology 18:578. Romero, E. 1982. Tratamiento profilctico de trozas en el bosque hmedo tropical. Universidad Distrital Francisco Jos de Caldas, Bogot. Romero, M., A., M. Antonio, and V. Dalia. 1999. La industria forestal en Panam. INRENARE, Panama. Ronnback, P., and J. H. Primavera. 2000. Illuminating the need for ecological knowledge in economic valuation of mangroves under different management regimes a critique. Ecological Economics 35:135. Ruitenbeek, J., and C. Cartier. 1998. Rational exploitations: Economic criteria & indicators for sustainable management of tropical forests. Occasional Paper No.17, Center for International Forestry Research, Bogor, Indonesia. Saint-Paul, U., J. Zuanon, M. A. V. Correa, M. Garcia, N. N. Fabre, U. Berger, and W. J. Junk. 2000. Fish communities in central Amazonian whiteand blackwater floodplains. Environmental Biology of Fishes 57:235.

PAGE 174

159 Saleska, S. R., S. D. Miller, D. M. Matross, M. L. Goulden, S. C. Wofsy, H. R. da Rocha, P. B. de Camargo, P. Crill, B. C. Daube, H. C. de Freitas, L. Hutyra, M. Keller, V. Kirchhoff, M. Menton, J. W. Munger, E. H. Pyle, A. H. Rice, and H. Silva. 2003. Carbon in Amazon forests: Unexpected seasonal fluxes and disturbance-induced losses. Science 302:1554. Schmieg, K. 1927. Notes on cabinet woods. Tropical Woods 9:1. Schnitzer, S. A., and F. Bongers. 2002. The ecology of lianas and their role in forests. Trends in Ecology & Evolution 17:223. Schnitzer, S. A., J. W. Dalling, and W. P. Carson. 2000. The impact of lianas on tree regeneration in tropical forest canopy gaps: evidence for an alternative pathway of gap-phase regeneration. Journal of Ecology 88:655. Sheil, D., and D. Burslem. 2003. Disturbing hypotheses in tropical forests. Trends in Ecology & Evolution 18:18. Sheil, D., and R. M. May. 1996. Mortality and recruitment rate evaluations in heterogeneous tropical forests. Journal of Ecology 84:91. Sherman, P. M. 2002. Effects of land crabs on seedling densities and distributions in a mainland neotropical rain forest. Journal of Tropical Ecology 18:67-89. Sierra, R., M. Tirado, and W. Palacios. 2003. Forest-cover change from laborand capital-intensive commercial logging in the Southern Choco rainforests. The Professional Geographer 55:477. Silver, W. L., R. Ostertag, and A. E. Lugo. 2000. The potential for carbon sequestration through reforestation of abandoned tropical agricultural and pasture lands. Restoration Ecology 8:394. Sist, P., R. Fimbel, D. Sheil, R. Nasi, and M. Chevallier. 2003. Towards sustainable management of mixed dipterocarp forests of Southeast Asia: moving beyond minimum diameter cutting limits. Environmental Conservation 30:364. Sist, P., and N. Nguyen-The. 2002. Logging damage and the subsequent dynamics of a dipterocarp forest in East Kalimantan (1990). Forest Ecology and Management 165:85. Sist, P., T. Nolan, J. G. Bertault, and D. Dykstra. 1998. Harvesting intensity versus sustainability in Indonesia. Forest Ecology and Management 108:251. Sitoe, A., B. Finegan, and M. Camacho. 1999. Modelling Pentaclethra forest an important Central American lowland rain forest type for timber production. Pages 143 in C. Kleinn and M. Khl, editors. Long-Term Observations and Research in Forestry. CATIE, Turrialba, Costa Rica.

PAGE 175

160 Stevens, G. C. 1987. Lianas as structural parasites the Bursera simaruba example. Ecology 68:77. Stumpf, K. A. 1993. The estimation of forest vegetation cover descriptions using a vertical densitometer. in J. I. a. B. W. Groups, editor. Society of American Foresters National Convention, Indianapolis, Indiana. Tamayo Velez, M. P. 1991. Ecofisiologa y anatoma de las semillas y plantulas de cativo (Prioria copaifera Griseb.). Ingeniero Forestal. Universidad Nacional de Colombia, Medelln. Tapia, I. E. 1999. Estudio de los Factores Edficos en las Parcelas del Proyecto Manejo de Cativales y Productos No Maderables. Autoridad Nacional del Ambiente, Direccin Nacional de Cuencas Hidrogrficas, Panam. TechnoServe-Panam. 1996. Auto-Diagnsticos Comunitarios en Comunidades Indgenas Ember-Wounaan en Darin. Informe Final Contrato F.96A, TechnoServe, Panam. Torrealba, P. 1996. Las Polticas de Aprovechamiento Forestal en Darin, Panam. Groupe de Recherche et Dechanges Technologiques (GRET), Fundacin para el Desarrollo Econmico y Social de Centroamerica (FUNDESCA), Consejo Centroamericano de Bosques y reas Protegidas (CCAB-AP), Centro de Estudios y Accin Social de Panama (CEASPA), Panama City. Torti, S. D., P. D. Coley, and D. P. Janos. 1997. Vesicular-arbuscular mycorrhizae in two tropical monodominant trees. Journal of Tropical Ecology 13:623. Torti, S. D., P. D. Coley, and T. A. Kursar. 2001. Causes and consequences of monodominance in tropical lowland forests. American Naturalist 157:141-153' Tosi, J. A. 1976. Informe sobre la zonificacin ecolgica preliminar de la regin del Darin en la Repblica de Colombia. 178 p. OEA, IICA, Costa Rica. Universidad Nacional de Colombia. 1984. Estudio integral de la madera para construccin, segunda fase. Laboratorio de Productos Forestales, Departamento de Recursos Forestales, Medelln. van Beukering, P. J. H., H. S. J. Cesar, and M. A. Janssen. 2003. Economic valuation of the Leuser National Park on Sumatra, Indonesia. Ecological Economics 44:43. van Mantgem, P. J., N. L. Stephenson, L. S. Mutch, V. G. Johnson, A. M. Esperanza, and D. J. Parsons. 2003. Growth rate predicts mortality of Abies concolor in both burned and unburned stands. Canadian Journal of Forest Research 33:1029. Vargas, C. 2003. Desparecen los bosques, y lo que queda es misera. in El Panam Amrica, Panam, June 18.

PAGE 176

161 Veiman, C. S. 1982. Plan piloto para manejo forestal de los terrenos de J.A.P.D.E.V.A. en Costa Rica. Maestra. Centro Agronmico Tropical de Investigaciones y Enseanza, Turrialba. Verissimo, A., M. A. Cochrane, C. Souza, and R. Salomao. 2002. Priority areas for establishing national forests in the Brazilian Amazon. Conservation Ecology 6: Art. No. 4. Vidal, E., J. Johns, J. J. Gerwing, P. Barreto, and C. Uhl. 1997. Vine management for reduced-impact logging in eastern Amazonia. Forest Ecology and Management 98:105. Webb, E. L. 1997. Canopy removal and residual stand damage during controlled selective logging in lowland swamp forest of northeast Costa Rica. Forest Ecology and Management 95:117. Whigham, D. 1984. The influence of vines on the growth of Liquidambar styraciflua L (sweetgum). Canadian Journal of Forest Research 14:37. Whitford, H. N. 1921. Opportunities for American foresters in the tropics. Journal of Forestry 19:156. Whitmore, T. C. 1999. Arguments on the forest frontier. Biodiversity and Conservation 8:865. Wightman, K. E., T. Shear, B. Goldfarb, and J. Haggar. 2001. Nursery and field establishment techniques to improve seedling growth of three Costa Rican hardwoods. New Forests 22:75. Wilshusen, P. R., S. R. Brechin, C. L. Fortwangler, and P. C. West. 2002. Reinventing a square wheel: Critique of a resurgent "protection paradigm'' in international biodiversity conservation. Society & Natural Resources 15:17. Wishnie, M. H., J. Deago, E. Mariscal, and A. Sautu. 2002a. The efficient control of Saccharum spontaneum (Graminae) in mixed plantations of six native species of tree and teak (Tectona grandis) in the Panama Canal Watershed, Republic of Panama: 2nd Annual Report. Working Paper ECO-03-03-En, Proyecto de Reforestacin con Especies Nativas (PRORENA), Yale University, Smithsonian Tropical Research Institute, Panama City. Wishnie, M. H., J. Deago, A. Sautu, and E. Mariscal. 2002b. Viability of three native tree species for reforestation in riparian area within the Panama Canal Watershed, Republic of Panama: 2nd Annual Report. Working Paper ECO-04-03-En, Proyecto de Reforestacin con Especies Nativas (PRORENA), Yale University, Smithsonian Tropical Research Institute, Panama City.

PAGE 177

162 Wyckoff, P. H., and J. S. Clark. 2000. Predicting tree mortality from diameter growth: a comparison of maximum likelihood and Bayesian approaches. Canadian Journal of Forest Research 30:156. Yamada, T., Y. Kumagawa, and E. Suzuki. 2001. Adaptive significance of vegetative sprouting for a tropical canopy tree, Scaphium longiflorum (Sterculiaceae), in a peat swamp forest in Central Kalimantan. Ecological Research 16:641.

PAGE 178

BIOGRAPHICAL SKETCH William Grauel received a Bachelor of Science degree in range-forest management in 1982 and a Master of Science degree in forest management in 1994 from Colorado State University, in Fort Collins. He was the scientific coordinator of an applied forest ecology project in Panama, a research associate for the Organization of Tropical Studies in Costa Rica, and served three years as an agroforestry extensionist in Paraguay with the Peace Corps. While living in the Rocky Mountains he worked as a forester with the Bureau of Indian Affairs in Dulce, New Mexico, and as a forest technician for the Forest Service in Saratoga, Wyoming. Mr. Grauel was born and raised in Lorain, Ohio, and attended high school in Amherst, Ohio. 163


Permanent Link: http://ufdc.ufl.edu/UFE0004366/00001

Material Information

Title: Ecology and Management of Wetland Forests Dominated by Prioria copaifera in Darien, Panama
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: UFE0004366:00001

Permanent Link: http://ufdc.ufl.edu/UFE0004366/00001

Material Information

Title: Ecology and Management of Wetland Forests Dominated by Prioria copaifera in Darien, Panama
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: UFE0004366:00001


This item has the following downloads:


Full Text










ECOLOGY AND MANAGEMENT OF WETLAND FORESTS DOMINATED BY
Prioria copaifera INT DARIEN, PANAMA
















By

WILLIAM THOMAS GRAUEL


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


2004

































Copyright 2004

by

William Thomas Grauel































Dedicated to the memory of Jack Westoby, who knew that forestry is more about people
than about trees.
















ACKNOWLEDGMENTS

I express sincere gratitude to my committee chair, Jack Putz, for sharing many

insights on ecology, forestry, and writing. I thank my committee members, Eric Jokela

and Daniel Zarin of the School of Forest Resources and Conservation, Benj amin Bolker

of the Department of Zoology, and Thomas Kursar from the Department of Biology at

the University of Utah, for their time and guidance, and I am very grateful to Dr. Stephen

Humphrey of the School of Natural Resources and Environment for providing funding at

the University of Florida.

I particularly thank Claudia Romero for providing me with much of the Colombian

literature on these forests. Along with Claudia I am very thankful for friendship and

moral support from Geoff Blate, Clea Paz, and Kevin Gould, who, as fellow graduate

students, understand.

In Panama I received support and guidance from scientists and administrators at the

Smithsonian Tropical Research Institute, including Ira Rubinoff, Egbert Leigh, Leopoldo

Le6n, Stanley Heckadon, Rick Condit, Jim Dalling, Elena Lombardo, and Raineldo

Urriola. Manuel Rodes and Jose Solis administered the project and allowed me and my

co-workers to concentrate on the enormous amount of fieldwork. Among the many

people who assisted me in the swamps of Darien, I deeply thank Ricardo Pineda, Ivan

Cabrera of La Chunga, and especially Delfin Jaramillo of Tucuti for their staunch and

reliable fieldwork. Thanks to the personnel of the Panamanian Environmental Authority

in La Palma and Panama City for all their work in keeping the project moving. John










Leigh of the International Tropical Timber Organization agreed to allow me to use

proj ect data for this dissertation and provided timely direction during the four-year

proj ect.

Special thanks to Sarah Dalle for helping me keep faith in the Macondo world that

is Darien, and a thank you to Julie Velasquez-Runk for sharing the burden and for finding

stuff.

Data for this study were collected as part of a proj ect funded by the International

Tropical Timber Organization (ITTO). The Panamanian Environmental Authority

(Autoridad Nacional del Ambiente, ANAM) and the Smithsonian Tropical Research

Institute (STRI) provided logistical support.





















TABLE OF CONTENTS


page


ACKNOWLEDGMENT S .............. .................... iv


LI ST OF T ABLE S ............_...... .............. ix...


LI ST OF FIGURE S .............. .................... xi


LIST OF OBJECT S ..............._ ..............xiii........ ......


AB STRAC T ................ .............. xiv


CHAPTER


1 LITERATURE REVIEW OF Prioria copalfera ....._._._ ........___ ........_.......1


Introducti on ........._ ....... ...............1.....
Distribution ........._ ....... ...............1....

Species Description .............. ...............3.....
Phenology .............. ...............4.....
Ecology ........._..... ....... ...............4....
Wood Uses and Properties............... ...............
Diseases and Insects .............. ...............10....
Y ield .......... _..... .......... ...............11....
Growth and Mortality ........._. ........_. ...............14...
Natural Regeneration ........._.. ........_. ...............16...
Artificial Regeneration .............. ...............17....
Conclusion ........._ ........_. ...............17....


2 EFFECTS OF LIANAS ON GROWTH AND REGENERATION OF Prioria

copaifera INT DARIEN, PANAMA .............. ...............19....

Introducti on ................. ...............19.................

Study Site ................. ...............22.................
M ethods .............. ...............23....
Re sults ................ ...............26.................
Discussion ................. ...............28.................


3 GROWTH AND SURVIVAL OF Prioria copaifera SEEDLINGS PLANTED
ALONG A HABITAT GRADIENT INT A PANAMANIAN SWAMP .....................40












Introducti on ............. ...... ._ ...............40...

Study Site ............ _. .... ...............42....
M ethods .............. ...............43....
Re sults............. ...... ._ ...............45...
Discussion ............. ...... ._ ...............47...


4 STRUCTURE, COMPOSITION, AND DYNAMICS OF Prioria copalfera-
DOMINATED SWAMP FORESTS IN DARIEN, PANAMA............... ................54


Introducti on ............. ...... ._ ...............54...

Study Sites .............. ...............57....
Principal Sites ............. ...... ._ ...............58...
Secondary Sites .............. ...............60....
M ethod s .............. ...............61....
Plot Descriptions .............. ...............61....
Sampling and Analyses .............. ...............61....
Re sults.............. .... ..._ .. ... .. .. ... ...........6
Tree Species Diversity and Stand Structure .............. ...............65....
Cativo Growth, Mortality, and Recruitment................ .............6
Growth of other Tree Species............... ... ...............6
Growth-dependent Mortality of Cativo Trees .............. ..... ............... 6
Cativo Regeneration .............. ...............70....
Discussion ................. ...............70.................


5 GROWTH AND YIELD PROJECTIONS OF Prioria copaifera FROM FOUR
SWAMP FORESTS IN DARIEN, PANAMA ................. .............................91


Introducti on ................. ...............91.................

Study Sites .............. ...............94....
M ethods .............. ...............95....
Scenarios............... ...............9
Re sults ................ ...............100................
Discussion ................. ...............102................


6 GEOGRAPHICAL, ECOLOGICAL, SOCIAL, AND SILVICULTURAL
CONTEXTS FOR CATIVO (Prioria copaifera) SWAMP CONSERVATION
IN THE DARIEN OF PANAMA ................. ...............115........... ...


Introducti on ................. ........... ...............115......
Timber Harvesting in Darien ................. ...............119...............
Forest Conservation Perspectives ................. ...............123................
Conclusions............... ..............12


APPENDIX


A MODELING METHODOLOGY USED IN CHAPTER 5 ............. ...................133












B SOURCE DATA FOR CHAPTERS 4 AND 5............... ...............139...


LIST OF REFERENCES ............. ...... ._ ...............141..


BIOGRAPHICAL SKETCH ............. ...... ...............163...






































































V111

















LIST OF TABLES


Table pg

1-1. Cativo wood properties ................. ...............10........... ...

2-1. Mean (+1 SE) annual diameter growth (mm) of Prioria copalfera one, two, and
two to four years after liana cutting. ............. ...............34.....

3-1. Mean canopy openness at 1.3 m above the ground as estimated with a spherical
densiometer, arranged by seedling age and habitat. ................ ..................5

3-2. Initial mean seedling height and diameter (at 20 cm above the ground; standard
errors noted parenthetically)............... ............5

3-3. Mean annual growth rates. ........._.._.. ...._... ...............51..

4-1. Total plot area measured for different minimum tree diameters and number
of tree species found. ........._.._.. ...._... ...............78...

4-2. Species diversity indices and relative dominance of cativo (Prioria copaifera).....78

4-3. Stem density and basal area of all species (above) and cativo only (below). ..........78

4-4. Incidence (%) of prostrate, inclined, broken stems and sprouts from prostrate
trunks................ ...............79

4-5. Forest-wide annual treefall and tree incline rates (i.e., partial uprooting) for small
(above) and large (below) trees for four sites............... ...............79.

4-6. Mean annual diameter growth (mm/year) of cativo trees of three stem types,
based on 1997-2001, 1998-2001, or 1997-2000 census periods. ..........................80

4-7. Mean annual diameter growth (mm/year) of cativo trees. ................. ........._......81

4-8. Ingrowth by stem type. Percentage of recruited individuals from broken stems,
undamaged stems, or sprouts from prostrate and inclined trees. ............. ................82

4-9. Cativo annual recruitment and mortality rates (%) by stem diameter class for four
census periods. ............. ...............83.....

4-10. Annual mortality rates (%) of cativo trees by stem type and stature
for four census periods. ............. ...............84.....










4-11i. Mean annual growth (mm/year) of cativo trees that were alive at the end of the
study and those that died during the study for which there was one or more
years of growth data ................ .............. .................. ...............85

4-12. Abundance of cativo trees < 1 cm dbh by height class............_._. .........._._. ...86

4-13. Annual mortality rates (%) of cativo trees by height class for the period
November 1998 November 1999. ............. ...............86.....

4-14. Mean annual height (cm/year) and diameter (mm/yr) growth rates for
cativo trees < 1 cm dbh, (sample sizes noted parenthetically). ............... ...............86

5-1. Characteristics of 4 cativo-dominated forests in Darien, Panama. ........................ 109

5-2. Cativo volume (m3 ha- ) and volume increment (m3 ha-l yr- ) of four cativo-
dominated forests in Darien, Panama ................. ...............110........... ...

5-3. Total volume yield after 65 years of growth and harvest simulations at three
different cutting cycles for three riverine swamp forests in Darien, Panama. .......110

5- 4. Percentage total volume reduction due to logging-induced damage for three
riverine forests at three cutting cycles ................. ...............110..............

A-1. Parameter estimates for the four sites .............. ...............134....

















LIST OF FIGURES


Figure pg

1-1. Distribution of Prioria copalfera........___ .... .._.. ...............2..

2-1. Diameter distributions of ascending lianas in six 25 x 25 m plots in heavily
infested riverine Prioria copaifera forest degraded by repeated entry logging ........35

2-2. Mean (f 1 SE) density of Prioria copalfera regeneration (< 1 cm dbh) in areas
of high (N = 10) and low (N = 6) liana densities. ................ ......... ...............36

2-3. Mean (f 1 SE) Prioria copaifera seedling recruitment censused two years after
liana cutting in three control and three treatment plots. .............. .....................3

2-4. Mean (f 1 SE) annual Prioria copaifera diameter growth based on five annual
censuses of all trees > 4 cm dbh in three control plots and three plots in which
all lianas were cut at the beginning of the study. ............. ...... ............... 3

2-5. Mean (f 1 SE) annual Prioria copaifera diameter growth of cativo based on
five annual censuses according to liana infestation level in control and treatment
pl ots. ............. ...............3 9....

3-1. Mean diameters and heights (+ 1 SE) of planted Prioria copaifera (cativo)
seedlings. .............. ...............52....

3-2. Percent seedling survival, beginning with the first census (November 1997) after
planting (September 1997). ................ ...............53................

4-1. Principal study sites. a) Casarete, b) Sambu, c) Juanacati, and d) Naranzati. ..........59

4-2. Stem types: a) prostrate and inclined and b) vertical sprouts from a fallen stem. ....63

4-3. Darien Province, Panama showing principal sites (S= Sambu, N=Naranzati,
C= Casarete, J= Juanacati) and seven secondary sites. .............. ....................8

4-4. Annual diameter growth (mm/year) ................. ........... ......... ................88

5-1. Growth trajectories of cativo at four sites in Darien, Panama starting
at 10 cm dbh. ........._._ .... ._ ...............111.











5-2. Cativo volume projoooooooooections for three previously logged riverine forests in Darien,
Panam a. ................ ...............112......... ......

5-3. Year 2000 dbh for trees as they attain 60 cm dbh during harvest simulation
of a 20-year cutting cycle. ........._.__...... .__ ...............113..

5-4. Cativo volume projections for an unlogged inland swamp forest
in Darien, Panama. ........... ...............114.....

A-1. Quadratic regression curves fit to relative growth data for the four sites...............134

















LIST OF OBJECTS


Object pg

1 Comma delimited variable Cativo data (as a text file) ........_.._.. ... ......_.._.. .....139

2 Cativo data (as an Excel spreadsheet) ........._._.. ....__.. ...._.._._..........3
















Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

ECOLOGY AND MANAGEMENT OF WETLAND FORESTS DOMINATED BY
Prioria copaifera INT DARIEN, PANAMA

By

William Thomas Grauel

August 2004

Chair: Francis E. Putz
Major Department: Natural Resources and Environment

The state of knowledge of Neotropical swamp forests dominated by Prioria

copaifera (cativo) is reviewed based on the available literature. Results of two

silvicultural experiments (liana-cutting and reforestation) are reported, forest-stand

dynamics are described, and growth and yield proj sections are presented.

Permanent sample plots at four sites in three watersheds in Darien, Panama were

installed in 1997 and 1998. Woody plants > 1 cm diameter at breast height (dbh, 1.3 m

above the ground) were tagged, mapped and measured, and censused annually to gather

demographic information on growth, recruitment, and mortality. Cativo regeneration

(trees < 1 cm dbh) was also censused at short intervals to monitor seedling dynamics.

Cativo growth was studied at an additional six sites along an inundation gradient that

varied in water salinity and hydroperiod.

Cativo growth was slow to moderate at sites flooded by tidal waters, moderate at

inland swamps flooded for long periods, and moderate to fast at riverine sites flooded by









fresh water; mean annual diameter growth ranged from 0.05 > 0.8 cm/year. Mortality

and recruitment rates of cativo varied widely among sites and years as well, with

mortality exceeding recruitment at two sites for two of the five years of monitoring. At

one slow growing site, trees that died during the study grew significantly slower prior to

death than trees that lived, demonstrating the occurrence of species-level growth-

dependent mortality for the first time in tropical forest.

Unlogged inland swamps contained very large standing volumes in trees > 60 cm

dbh (up to 180 m3/hectare). Riverine forests that were repeatedly logged during 1950-

2000 contained few large trees, but showed potential for future timber harvests in the

form of abundant regeneration and large standing volumes in trees 40-60 cm dbh. Sixty

year volume proj sections suggest that higher dbh cutting limits and longer cutting cycles

reduce residual damage, and can produce high timber yields for inland swamps and

riverine forests, respectively.

Volatile world timber markets and log shortages may be reducing incentives for

cativo logging (and thus swamp forest conservation) in the face of large development

projects and increased colonization in Darien, Panama. Cativo swamp forests have

important hydrological and carbon-sequestering values that should be incorporated in

land-use decisions.















CHAPTER 1
LITERATURE REVIEW OF Prioria copalfera

Introduction

Unlike the vast maj ority of tropical tree species, much is known about the ecology

of the swamp species Prioria copaifera Griseb. (hereafter cativo). Its potential

commercial value was recognized in the 1920s, but thousands of hectares of cativo-

dominated forests had already been cleared to make way for the banana boom of the early

twentieth century. By the end of the century much had been converted to agriculture and

of the remainder, most had been cutover and degraded.

Distribution

Cativo is found from Nicaragua to Colombia, and is also present in Jamaica (Figure

1-1, Holdridge 1970). Although cativo was included in lists of the tree species of the

coastal region of Ecuador by Rimbach (1932) and Acosta Solis (1947) its presence in that

country is not confirmed. Barbour (1952) reported that the commercial range includes

the Atlantic coast of Central America from Nicaragua to Panama, the watershed of the

Bayano River, the rivers flowing into the Gulf of Darien (Golfo de San Miguel), and the

area around the mouth of the Atrato River in northwest Colombia. Cooper (1928)

referred to enormous stands of cativo in the Valle Estrella of Costa Rica and the Laguna

de Chiriqui in Panama. In 1987, it was estimated that cativo forests covered 49,000 ha in

Panama, with the maj or concentration in the easternmost Province of Darien but

including eastern Panama Province ( INRENARE, Instituto Nacional de Recursos

Naturales Renovables 1987). The same report noted 4000 ha of cativo on Coiba Island










and 17,000 ha in the Chucunaque River watershed; 9000 ha in the combined watersheds

of the Tuira, Balsas, and Marea Rivers; and an additional 4000 ha in adj acent swampy

areas. By 1999, the National Ministry of the Environment in Panama estimated an area

of only 15,000 ha of cativo in the country (ANAM 1999a).

I.JITED STATES











Figure ~ ~ ~ ~ ~ A~s~ 1-.Dsrbto fPira cpafea.
In ColmbiaLinare reo(98 esrbdctv' anea nldnh

waesed fthe Aratoic an en iesinteUub eio nte otwsto h

conr.Acodn oEsoa n Vsuz(98) aio sas oudi h
Deplartet fAtouaadCoo oi(96 ttdta aiowsntfudo










Esoa ndClmba Vasquez (18)rieport98)dsed cativo intelo-y ng aras aroludnd the vlaeo







Santa Marta along the Nechi River and referred to a sample of cativo in the Gabriel

Gutierrez "Medel" Herbarium of the Agronomy Department at the National University in

Medellin collected in the Department of Magdalena in 1949, all of which are on the









Pacific. Colombia contained 363,000 ha of cativo forest (Linares Prieto 1987b, c,

Gonzalez Perez. et al. 1991). Subsequently, the area was estimated at 173,000 ha in 1978

(Linares Prieto 1987b); and in the late 1980s estimates ranged from 60,000 ha (Linares

Prieto 1987c), to 90,000 ha (Linares Prieto 1987b), tol63,000 ha (Linares Prieto 1988).

In Costa Rica, the areas of cativo-dominated forests on the Pacific (Allen 1956) as well as

on the Atlantic side of the country (Bethel 1976) have been severely reduced (Veiman

1982) and today the species is listed as threatened in the country (Jimenez Madrigal

1995).

Species Description

Prioria copaifera is the only species in the genus, and is in the subfamily

Caesalipinioideae of the Fabaceae. Average adult height is 25-30 m, with a dbh

(diameter at breast height, 1.3 m) from 45 tol00 cm (INRENARE 1987a), although the

trees reach 180 dbh (Grauel, unpublished data) and 40 m in height (Del Valle 1972). The

branches of mature trees are somewhat arched and the foliage is distributed uniformly in

a round, thick crown (Escobar and Vasquez 1987). The bark is smooth, brownish-gray to

gray in color, and with abundant lenticels found in continuous horizontal bands

(Echavarria A. and Varon P. 1988). Cativo has no buttresses, and its compound leaves

generally have four opposite leaflets, elliptic-lanceolate in shape, with a swollen petiole.

The leaflets are 16 cm long and 8 cm wide, asymmetric, rounded at the base and with an

acuminate apex. The tiny, white flowers develop in panicles (Mufioz Valencia 1966).

The flowers have 10 stamens and no petals, but the sepals resemble petals (Gentry 1996).

The fruit is a flat indehiscent pod, with one side slightly convex and the opposite side

concave, 10 cm long, 7 cm wide, 3 cm thick, woody, with a single seed (Mahecha Vega

et al. 1984). Seeds are large (mean fresh weight = 48 g, Lopez 2001, to 96 g, Dalling et









al. 1997) and dispersed by water. In addition to the common name "cativo", P. copaifera

has been known as cautivo, kartiva, trementino, floresa, tabasara, amanza muj er, camibar,

murano, and Spanish walnut (Schmieg 1927, Anonymous 1933, Harrar 1941, Hess et al.

1950, Escobar and Vasquez 1987).

Phenology

Del Valle [cited in Escobar and Vasquez (1987)] noted that cativo leaf-drop is

uniform throughout the year and the species is evergreen. Cooper (1928) found that the

species flowers generally in March and April and fruits in October and November, but

also found a tree with flowers and immature fruits in February. Linares Prieto (1988)

reported that flowering typically begins in June and peaks in August and September.

Fruiting then begins in September and October and peaks in April and May. In

Colombia, seedling recruitment peaks at the beginning of the rainy season from April to

June (Linares and Martinez Higuera 1991). Although cativo produces seeds twice a year,

large seed crops seem to be produced once every 2 years (Pizano SA 1995, Grauel in

review 2004a).

Ecology

Holdridge (1964) reported that throughout its range from eastern Nicaragua into

northwestern South America, cativo is found in the Tropical Moist, Wet, and Rain Forest

lifezones. According to Tosi (1976), the cativo forests of Colombia are found in the

Tropical Moist Forest and Tropical Wet Forest lifezones of the Holdridge classification

system; annual precipitation ranges from 2000 to 8000 mm with average annual

temperature of 24o to 28o C (Gonzalez Perez et al. 1991), while in Darien, Panama, cativo

is found in the Tropical Moist Forest lifezone (Holdridge and Budowski 1956).

Cativo is found in four distinct habitats:










* On the Atlantic coast of Costa Rica, Panama, and Colombia, it is found just inland
from mangrove forests where salt water does not intrude.

* Along the many rivers of southern Central America and northwest Colombia,
cativo is found in alluvial valleys that are flooded periodically, generally in the
ramny season.

* Away from maj or rivers cativo is found in low-lying areas that are inundated for
extended periods, up to the entire 9-month rainy season in Darien, Panama.

* Cativo is also found in upland forests, but never as abundant and dominant as in
flooded habitats. Although Consultores Ambientales LTDA (1995) declared that
cativo cannot germinate or develop in well-drained soil, it is a common component
of the upland, mixed-species forest on Barro Colorado Island in Panama (Condit et
al. 1993b, 1995a, Sheil and Burslem 2003).

Colombian researchers have classified several types of"cativales" based on

landscape position and duration of inundation. Linares Prieto (1988) used landscape

position to classify cativo forests as low, medium, and high alluvial plains. The National

Corporation for Forest Research and Promotion (CONIF, Corporaci6n Nacional de

Investigaci6n y Fomento Forestal, cited in Echavarria A. and Varon P. 1988) classified

different types of cativo forest according to length of inundation as greater than 6 months,

3 to 6 months, and less than 3 months. Gonzalez Perez et al. (1991), referring to the

Institute Geografico Agustin Codazzi cited in G6mez (1990), defined the inundation

periods as permanent, 6 to 8 months, 3 to 6 months, and less than three months. In a

study of early natural cativo regeneration, Martinez Higuera (1989) defined their two

study sites as low alluvial plain frequently flooded and low alluvial plain infrequently

flooded.

Linares Prieto (1987b) offered various criteria for the classification of cativo-

dominated forests:

* three types of forest distinguished by landscape position (plain, terrace, and alluvial
fans.










* six types of cativales defined by the combination of life zone and landscape
position.

* number of cativo stems per hectare; median cativo basal area, average distance
between trees, number of trees of other species, the number of other species apart
from cativo, median basal area of other species than cativo, and median total basal
area per hectare.

* phytosociology of Prioria in terms of abundance, frequency, dominance,
importance, and productivity.

* common associates being Cynonsetra spp, Pterocarpus officinalis, Gustavia spp,
Carapa guianensis, Anacardium excelsunt, Escinveilera spp, nuanamo
(Myristicaceae), Castilla elastica, and Lecythis spp., the number of species
increasing with elevation and with better drained soils.


In a study that used life zone and landscape position as classification criteria, Escobar and

Vasquez (1987) proposed nine types of cativo forest but concluded that their

mathematical analysis failed to support the categories.

A defining ecological characteristic of cativo-dominated forests is their

monodominance or low species diversity within stands. Maximum homogeneity is found

in the riverine forests inundated by high Pacific Ocean tides in Darien, Panama. In these

sites, cativo can comprise more than 95% of the stems > 1 cm dbh and the forest contains

less than ten woody species per hectare (Grauel and Kursar 1999, Grauel and Putz 2004).

Along the Marea River in Darien, cativo had a relative abundance of 91% in a 10 x 1000

m transect that included five other tree species and one palm (2 10 cm dbh, Mayo

Melendez 1965). Away from the influence of tides, floristic diversity increases. Linares

Prieto (1988), citing the thesis of Escobar and Vasquez (1987), reported the relative basal

area dominance of cativo as 50 to 92%. In Colombia, Gonzalez Perez et al. (1991) found

that cativo-dominated forests contain approximately 60 tree species, 15 of which

comprise 95% of the individuals and are in the families Fabaceae, Bonabacaceae, and









Sterculiaceae, although the minimum diameter of the study was not stated. In cativo-

dominated forests in the area of Domingodo-Truando in the Colombian Department of

Choco, Consultores Ambientales LTDA (1995) found 86 tree species per hectare (>10

cm dbh). In Darien, Panama, Golley et al. (1975) found 44 species of trees per hectare in

a cativo-dominated forest along the Chucunaque River. Elsewhere in Darien, Holdridge

(1964) found Carapa guianensis to be the only other large tree associated with cativo,

although he noted that Pterocalpus officinalis was restricted to the edges of small streams

in the same forest.

The dominance of cativo in seasonally flooded habitats was suggested as being

attributed to the better competitive ability conferred by ectomycorrhizae (EM) compared

to vesicular-arbuscular mycorrhizae (VAM, Connell and Lowman 1989); but Torti et al.

(1997) showed the existence of VAM in cativo. Lopez and Kursar (1999) demonstrated

that 3 tierra firme species survived inundation as well as "flood-tolerant" cativo, and

suggested that a cycle of inundation and drought-induced water stress may better explain

patterns of tree diversity than inundation alone.

An inventory of animal diversity in a cativo forest in Colombia reported 24 species

of mammals, 16 species of birds (principally in the families Sittacidae, Cracidae, and

Ramphastidae), 6 species of fish, and 7 species of reptiles (Martinez Higuera 1989).

Several of these mammal, bird, and reptile species are becoming increasingly rare due to

hunting pressure, including Taxpirus bairdii, Mazamna amnericana reperticia, Penelope

pwrpura~scens wagler, Crax rubra, Lutra longicandis, and Caiman sclerops (Linares

Prieto 1988). Some other studies on wildlife in cativo forests include those of









MondragC~n et al. (1994), Ospina Torres (1994, 1995a, 1995b, 1996, 1997), and Orozco

Rey (1995).

Holdridge (1964) described how cativo forests alternate with monospecific forests

dominated by M~ora oleifera along the Tuira and Tucuti (Balsas) Rivers. M~ora-

dominated forests are typically found at slightly lower elevations where the effects of

brackish water from high tides are greater, but cativo trees can be found in M~ora-

dominated forests (Porter 1973). Cativo forests are characterized by a distinct

microtopography where adult trees are surrounded by mounds 20 to 30 cm in height and

5 m in diameter (Duke 1964). Duke (1964) also mentioned that fallen cativo trees were

very common. Although adult cativo trees are shallow-rooted, in a comparison of cativo-

dominated "gallery" forests with tropical moist, premontane, and mangrove forests.

Golley et al. (1969) found belowground biomass to be greatest in mangrove forests, and

similar among the other forest types. Overstory biomass, however, was notably higher in

gallery (cativo) forests, with over 100 Mg/ha dry weight in stems alone. Holdridge

(1964) estimated a leaf area index of 6. 1 in a cativo forest along the Tuira River in

Darien, Panama.

In Colombia, soil fertility of cativo forests varies from very low to moderate

(Martinez Higuera 1989) with fine to medium texture, pH from 5.1 to 6.0, and poor

drainage in general (Linares Prieto 1987c). Soils of riverine cativo forests in Panama are

composed of clay to loamy clay with pH 5.2 to 6.8 (Mariscal et al. 1999, Tapia 1999)

while soils of inland swamps are more clayey and acidic (pH 4.4-6.0, Tapia 1999). The

Colombian Institute of Hydrology and Soil Use, cited in Linares Prieto (1988) and









Martinez Higuera (1989), classified soils of cativo forests as Inceptisols (63%) and

Entisols (37%).

Wood Uses and Properties

The value of cativo wood lies in its historic abundance and accessibility, not

necessarily in its inherent properties. Early descriptions noted cativo for its cylindrical

form and general abundance (Kluge 1926, Cooper 1928) as well as its potential for

supplying raw material for architects and interior decorators (Schmieg 1927), although

Pittier and Mell (1931) considered the wood to be of little or no use.

Later, cativo was included in a series of studies before (Kynoch and Norton 193 8)

and during WWII (Harrar 1941, 1942a, b) that sought to provide technical information on

the physical and mechanical properties of foreign and domestic woods. Further research

beginning in 1947, funded by the U. S. Office of Naval Research, resulted in

recommendations of cativo for plywood, cabinetry, and furniture (Hess et al. 1950).

Similar studies of potential applications of cativo occurred later in Colombia (Hoheisel

and L6pez G. 1972, Universidad Nacional de Colombia 1984).

Several authors have mentioned the abundant resin that bleeds from freshly cut logs

and can make sawing difficult (Cooper 1928, Hess et al. 1950, Barbour 1952, Del Valle

1972). The copious resin of cativo was used by indigenous groups for such diverse uses

as repairing boats and for medicine (Cooper 1928, Duke 1986). By using high

temperatures during kiln-drying, appreciable amounts of resin can be removed from the

lumber with the additional benefit of relieving some of the internal stresses that are

caused by the presence of tension wood (Kukachka 1965).









Table 1-1. Cativo wood properties
Specific gravity.
Author .1 Shrmnkage %
(g cm)
Harrar (1941) 0.48 9.87
Hess et al. (1950) 0.40 8.9
0.41 sapwood 9.2 sapwood
Barbur (952) 0.50 heartwood 22.9 heartwood
Kukachka (1965) 0.40 8.8

Cativo wood properties have been investigated extensively (Kynoch and Norton

1938, Hernandez Hurtado 1984, Jaramillo Gallego and Velasquez Salazar 1992, Escobar

C. and Rodriguez 1993). It is moderately light in weight (Table 1-1) and although it is

relatively nondurable with respect to both fungal decay and insect attack, cativo has good

dimensional stability and was used as a base for piano keyboards for that reason

(Kukachka 1965).

Cativo veneer from Panama was marketed in Canada and the United States in 1933

(Anonymous 1933). Large-scale imports of cativo to the United States occurred in the

mid to late 1940s from Costa Rica (Hess and Record 1950). By 1952, almost 9500 m3/y

were exported from Colombia and Costa Rica to the US, the Eigure increasing to over

47,000 m3/yr by 1958 (Kukachka 1965).

While some cativo from the Caribbean side of western Panama was exported, it

eventually supplied 90% of the raw material for the domestic plywood industry and 50%

of sawn-wood production in the country (FAO, Food and Agriculture Organization

1982). The Panamanian Institute of Renewable Natural Resources recommended using

cativo for furniture, packing crates, and cabinetry (INRENARE 1987a).

Diseases and Insects

Cativo was classified as moderately to non-durable in its resistance to the white-rot

fungus Polyporus versicolor (Tramnetes versicolor) and durable to non-durable for the









brown rot fungus Poria monticola (Hess et al. 1950). Ferrer (1999 ) collected 615 fungi

associated with living and dead cativo trees in five different forests, and found 58%

Ascomycetes and 42% Basidiomycetes. Apparently, it is not known how many of these

are pathogens, saprohytes, or mutualists. In a cativo forest along the Sambu River in

Darien, Panama, Ferrer (1999) found that 27% of the Basidiomycetes belong to the genus

Phellinus, one of which is among the most important tree pathogens of temperate forests

(Phellinus weirri).

Hess et al. (1950), Barbour (1952), and Kukachka (1965) mention the susceptibility

of the boles of recently felled trees to attack by ambrosia beetles. Insects that perforate

recently cut logs belong principally to the families Scolytidae and Platypodidae, and

occasionally Brentidae and Tenebrionidae (Romero 1982). The species most commonly

found on recently cut logs, but not specific to Prioria, are Platy~pus para~lllleh Fabricius

and Xyleborus affinis Eichhoff (Estrada L6pez and G6mez Quiceno 1988). Some

protection from attack is rendered by direct sunlight, immersion of the logs in water, and

the presence of bark; one application of insecticides may prevent attack for 6 to 15 days

(Romero 1982, Estrada L6pez and G6mez Quiceno 1988).

Yield

Early research on cativo as a timber source stressed the large sizes and clear boles

of the trees. Cooper (1928) noted an average size of 60 to 90 cm dbh. Barbour (1952)

found the commercial size range of the species to be 60 to 120 cm dbh, with maximum

sizes of 150 to 180 cm. Barbour (1952) emphasized the straight form of the trunks,

without branches for 12 m (and many times branchless up to 30 m in height).

From 195 1 to 1953 Bruce Lamb studied the forests of Darien for the Panama Forest

Products Company to develop log-supply sources and determine available timber









volumes for both upland and lowland forests. Lamb estimated cativo wood volume along

the Chucunaque, Tuira, Balsas, Sambu, Congo, and Cucunati Rivers and around the

Laguna de la Pita (today called Matusagarati). Along the Balsas River, Lamb found pure

stands of cativo for a distance of 20 km and up to a km in width on each side of the river.

He estimated an average volume of 71 m3/ha and a total of 141,600m3 for the watershed

(Lamb 1953). On the Tuira River, Lamb encountered cativo forests 25 km upriver from

the Tuira' s confluence with the Balsas River up to the mouth of the Chucunaque River.

In this area of approximately 4000 ha, volumes averaged 24 m3/ha. Although Lamb did

not examine the forests upriver from the mouth of the Chucunaque, there were reports of

cativo forests up to Boca Cupe, and he estimated a total of 23,600 m3 for the Tuira

watershed. According to Lamb, the highest-quality cativo wood came from the

Chucunaque River watershed, and he estimated a total volume of 47,200 m3 along the 80-

km course of the river. Along the Sambu River, Lamb found cativo 10 km from the

river' s mouth at the confluence of the Jesus River up to the Sambu' s confluence with a

small stream called Morobichi (8 km further upriver). This cativo forest extended up to

1500 m inland from the river, and Lamb estimated an average volume of 35 m3/ha and a

total of 23,600 m3 for the Sambu watershed. Lamb estimated a total of 1,180,000 m3 foT

the entire province (Lamb 1953).

Cativo forests are known to contain large wood volumes per hectare, but

comparison of different estimates is difficult where the minimum diameter is not

specified; in addition, some estimates are for commercial volume and others for total

volume. In a previously unlogged inland swamp in the Balsas River watershed (near a

small stream called Naranzati), 96 m3/ha of commercial (> 60 cm dbh) cativo wood was









measured in 1999 (Grauel, unpublished data). Also in 1999, a 100% inventory was

carried out of a 50-ha plot along the Sambu River in Darien where cativo comprises 95%

of the species diversity. Using a form factor specifically developed for Prioria, a mean

volume of 65 m3/ha was calculated for trees > 60 cm dbh; while across the river in a

series of smaller permanent plots, mean volume totaled only 40 m3/ha for the same forest

type. The difference can probably be attributed to different management histories under

different ownership regimes: the latter being found on open-access public land that is

subj ect to frequent, low-intensity timber harvesting by local loggers; while the 50-ha plot

is located on land belonging to Embera-Wounaan indigenous communities who harvest

cativo much less frequently. When considering a minimum diameter of 40 cm dbh and

the commercial height to the lowest branch, this 50-ha plot contains 190 m3/ha (Grauel,

unpublished data). Recent volume measurements in cativo forests along the Balsas River

ranged from 20 m3/ha (> 60 cm dbh, Grauel, unpublished data) to 25 m3/ha (total volume,

Mariscal et al. 1999).

In Colombia, Linares Prieto (1987b) stated that a cativo forest contained more than

150 m3/ha in commercial wood and a mean of 80 to 100 m3/ha for trees > 52 cm dbh. In

a cativo forest with 60 tree species where cativo comprises 60% of the basal area, Linares

Prieto (1988) measured a total volume of 123 m3/ha and 46 m3/ha in trees > 40 cm dbh.

In a cativo forest with 5 other commercial tree species in Podega, Colombia Escobar

Munera (1981) calculated a mean of 36 m3/ha for all species. In another forest inventory

of trees > 49 cm dbh of 15 tree species, a mean of 7.3 individuals and 27 m3/ha, cativo

comprised 36% of the total volume (Consultores Ambientales LTDA 1995).









Growth and Mortality

As with volume estimates, growth estimates vary and depend on the methodology,

age and size of the trees, management history of the forest, and site-specific biotic,

abiotic, and climatic factors. Comparisons of growth estimates are difficult where

different field methodologies and modeling approaches are used.

Like many tropical trees, cativo produces growth rings, but no dendrochronology

based on crossdating has been performed to show that the rings are produced annually.

Using the pinning technique (Kuroda and Shimaji 1984) where wood is wounded and

subsequent growth is measured with destructive harvesting, however, McKenzie (1972)

concluded that cativo produces annual rings. I strongly suspect that cativo may produce

one or more nings per year.

Cativo diameter growth is probably influenced by many variables. Londofio

Londofio and Gonzalez Perez (1993) reported that crown area, crown position, and

Hegyi's diameter-distance competition index had significant effects on growth of cativo

in less diverse forests but not in the more diverse sites. In an unlogged cativo forest with

50 to 60 tree species, Del Valle (1979) found that maximum diameter increment was

attained by trees approximately 70 cm dbh. Two studies in Panama found maximum

diameter increment in medium-sized trees, from 20 to 50 cm dbh depending on the site.

In an upland forest in Panama, Condit et al. (1993a) measured maximum annual diameter

increments of 2 to 4 cm, while cativo from flooded forests showed maximum annual

growth rates of 1.5 to 2.0 cm, with means of 0.6 to 1.0 cm (Grauel 1999).

Several modeling approaches have been used to estimate lifetime growth

trajectories based on short-term growth rates. Del Valle (1979) used a matrix modeling

approach to produce an estimate of 98 years for a 10 cm dbh tree to reach 60 cm.









Gonzalez Perez (1995) developed a von Bertalanffy growth model for cativo, and

produced an estimate of 90 years for a 14.5 cm dbh tree to reach 60 cm. In a comparative

study of two sites where diameter structure, floristic diversity, spatial distribution, and

growth were contrasted, Gonzalez Perez et al. (1991) found growth to be three times

greater in the more diverse forests. Although the authors admit to small sample sizes,

they estimated 168 years in less diverse forests and 77 years in more diverse forests for a

10 cm dbh tree to reach 60 cm (Gonzalez Perez et al. 1991). There was no difference in

growth between diverse and cativo-dominated forests in another study in Colombia,

where Linares Prieto (1987b) estimated that a tree would reach optimum harvest size (60

cm dbh) in 55 years in less diverse forests and in 60 years in the more diverse forests. In

a different study, the same author estimated that a tree could reach 60 cm dbh in only 38

years, and suggested that this time period could be reduced with "adequate silvicultural

techniques" (Linares Prieto 1988). In a study of growth and yield potential of cativo on

an upland site in Panama, Condit et al. (1995b) estimated a period of 130 years for a stem

to grow from 1 cm dbh to 60 cm, based on mean growth using data from 1982-1985, and

180 years based on data from 1985-1990. Using the same modeling approach and based

on data from 1997-2001, I found two sites with similar inundation regimes to vary

considerably; one requiring 315 years, and the other 179 years for a 1 cm dbh stem to

grow to 60 cm. At another forest farther upriver, only 80 years would be required for a 4

cm dbh tree to reach 60 cm. For an inland swamp, based on growth data from 1997-

2000, 157 years would be required for a 1 cm dbh tree to reach 60 cm, and 186 years to

reach 80 cm.









Natural Regeneration

In a demographic study of cativo in Colombia, Montero (1996) estimated 30,490

seeds/ha were produced during the 6-month period from December to May. Montero

(1996) noted that trees with relatively low seed production tended to produce greater

numbers of established seedlings than trees that had greater seed production and

hypothesized that there was some optimal period for seedfall that resulted in higher

probability of survival and maximum recruitment rates.

Linares and Martinez Higuera (1991) found lower densities of cativo natural

regeneration in frequently inundated forests than in less frequently flooded forests,

although the tendency toward monodominance was greater where flooding was more

frequent. Linares and Martinez Higuera (1991) also found a strong correlation between

mean monthly precipitation and seedling density, and determined that 2% of the large

initial seedfall became established. Lopez (2002) followed a 1997 cohort of cativo

seedlings and found 2.5% survivorship after 3 years.

Dalling et al. (1997) found that less than 10% of cativo seeds were viable 2 months

after seedfall, and that 30% of the viable seeds had suffered damage by insects or

pathogens. Even with up to 60% of the seedmass damaged, however, there was no

reduction in the probability of germination; and seeds with up to eight insect larvae

germinated as often as seeds with no infestation (Dalling et al. 1997). Furthermore,

cativo seeds have the ability to produce an average of 2.1 additional, sequential resprouts

after the initial sprout is damaged or lost (Dalling et al. 1997).

Tamayo Velez (1991) determined that a germinating seed required 48 days to

develop into a 29-cm tall seedling, and declared that height growth of cativo plants

between 30 and 150 cm in height was 50-60 cm/month. Martinez Higuera (1989)










suggested an annual growth rate of 2.4 m for trees in the same height range. In two

strongly monodominant cativo forests in Darien, Panama, density of cativo natural

regeneration (trees < 1 cm dbh) varied widely, from less than 5,000/ha at one site to over

17,000/ha at the other. Mean annual height growth for trees between 30 and 150 cm tall

varied little, from 2-5 cm, with maximum annual growth rates of 15-25 cm (Grauel, in

review 2004a).

Artificial Regeneration

In a study of artificial regeneration of cativo carried out in Uruba, Colombia,

Linares Prieto (1987a) tested five planting methods and two planting seasons, and

determined that average survival and height were greatest for bare root seedlings planted

during the dry season. After 4 years, these seedlings had reached 2.7 m in height

(Linares Prieto 1987a). Caycedo (1988), cited by Martinez Higuera (1989), measured

annual height growth in seedlings of 70 cm. Cativo seedlings of two ages were planted in

three habitats in Darien, Panama. Seedling mortality after 4 years was highest in the

natural habitat of the forest understory (89%) and lowest in partial sun on the edge of the

forest bordering a treeless marsh (54%, Grauel in review 2004b). Maximum height

growth was observed in the full sun, where seedlings of both ages grew approximately 50

cm/yr.

Conclusion

Relatively few tropical tree species have been studied as extensively as cativo.

Because of its accessibility, abundance, form, and wood properties, cativo logging has

provided livelihoods for thousands of rural Latin Americans as well as for forest

industries in North, Central, and South America. Little research has been carried out on

the importance of the ecosystem services that cativo forests provide, however.









Unfortunately, the existence of technical, silvicultural, and ecological knowledge of

cativo and the forests where it is abundant has not resulted in sustainable management.

As is frequently the case in natural resource management, technical knowledge is

insufficient when social and economic forces can influence the strength of forest policies

that determine the quality of management carried out on the ground. Nevertheless,

technical knowledge can form the foundation for research that links ecological dynamics

with the social and economic policies that affect forest management, with the aim of

promoting socioeconomic development that conserves natural ecosystems.















CHAPTER 2
EFFECTS OF LIANAS ON GROWTH AND REGENERATION OF Prioria copalfera
INT DARIEN, PANAMA

Introduction

The abundance and ecological roles of lianas in tropical forests have long attracted

the attention of tropical silviculturists (Fox 1968, Appanah and Putz 1984, Chaplin 1985,

Putz 1991, Vidal et al. 1997, Carse et al. 2000). Because lianas are a major component of

woody plant diversity and provide important food sources for wildlife, they play critical

roles in maintaining biological diversity (Nabe-Nielsen 2001, Burnham 2002, Schnitzer

and Bongers 2002). Unfortunately, where sustainable forest management is the primary

tool for forest conservation and the primary obj ective is timber production, lianas can be

a maj or impediment. Given that the likelihood of forest conversion to more profitable

land uses than forestry is enhanced if prospects for subsequent timber harvests are not

economically competitive, liana proliferation can contribute indirectly to forest loss.

The large trees that provide the timber value of a forest are more likely than smaller

trees to be infested with lianas (Putz 1984, Putz and Chai 1987, Nabe-Nielsen 2001,

Perez-Salicrup et al. 2001), and lianas can have various silvicultural implications for

forest management. During harvesting operations for example, felling of liana-laden

trees can induce excessive stand damage, because their crowns are likely to be connected

to their neighbors (Putz 1984). Avoidance of this accessory damage has frequently, but

not always (Parren and Bongers 2001), been accomplished through pre-felling liana cutting

(Fox 1968, Appanah and Putz 1984, Johns et al. 1996). An additional benefit of pre-









felling liana cutting is the post-harvest reduction in liana proliferation in logging gaps

(Alvira et al. 2004, Gerwing and Vidal 2002). Reducing post-logging liana infestations is

desirable, because lianas can seriously impede succession in gaps (Schnitzer et al. 2000)

and diminish opportunities for rapid recruitment and growth of desirable timber species.

In addition to physically impeding establishment of seedlings and saplings of tree species

in logging gaps, lianas can reduce host tree fecundity (Stevens 1987), lowering the

reproductive output of valuable timber species in forests where natural regeneration is the

only cost-effective silvicultural option for stand perpetuation. Heavy liana infestations

can also substantially reduce diameter growth of adult trees (Whigham 1984, Gerwing

2001, Clark and Clark 1990), which lowers the net present value of future timber yields

by prolonging cutting cycles.

Prioria copaifera (hereafter "cativo," Fabaceae), a canopy tree found in freshwater

wetland forests from Nicaragua to Colombia, has been exploited for timber for decades

(Barbour 1952), with little apparent concern for long-term management. Today,

commercial stands are found principally in eastern Panama and northwest Colombia.

Repeated logging of monodominant cativo stands during 40 years of exploitation testifies

to the regenerative capacity of the species. Nevertheless, large areas of cativo forest have

been converted to agricultural production or to mixed-species secondary forest and liana

tangles as a result of overharvesting. Of the original 363,000 ha of cativo in Colombia

for example, less than 90,000 ha remain (Linares Prieto 1987b). Similarly, extensive

stands of cativo were once found in western as well as eastern Panama, but today

commercial stands are found only in Darien Province. Of 30,000 ha of cativo-dominated

forest in Darien in 1987 (INRENARE 1987a), an estimated 15,000 ha remained in 1999










(ANAM 1999a). Increasingly, Panamanian foresters as well as local Darien community

members desire to promote sustainable logging of the remaining cativo forests.

The stands of almost pure cativo that are found as bands along the principal rivers

of Darien vary between 100 m and 1 km in width. Behind the forest, treeless wetlands

composed of the palms Elaeis oleifera and Oenocarpus nzapora and various lianas

i ncludi ng Dalbergia brownei, Conabre tunt sa~n buensis, Elachyptera floribunda,

Tetrapteris nzacrocalpa, AllllllllllIllllllla~nzad cathartica, Phrygan2ocydia corynabosa, Cydista

diversifolia, Snzilax spinose, Banisteriopsis spp., and Heteropteris spp. often dominate

the landscape. In the absence of silvicultural interventions other than logging, high-

statured riverine forests are likely to be converted into palm- and liana-dominated

vegetation.

Present day stand structure of many riverine cativo forests in Darien is a result of

traditional logging methods that do not employ heavy machinery (Grauel and Pineda M.

2001). Instead, logs are levered or rolled by hand towards the river on roads constructed

from 15 to 30 cm dbh (diameter at breast height, 1.3 m) cativo trunks cut and laid end to

end to form two parallel rails. In many riverine cativo forests, the combination of

removing all harvestable-size trees as well as many subcanopy individuals for rail

building has left a very discontinuous canopy and large multiple-tree gaps, which are

habitats favorable for liana proliferation.

The leguminous liana Dalbergia brownei proliferates abundantly in disturbed

cativo forests. A principal component of the treeless wetlands found behind the natural

river levees where cativo dominates, this liana uses cativo forest edges to climb into the

forest canopy. Although this species does not establish in the deep shade of the cativo









forest understory, large stems (up to 20 cm diameter) are commonly found hanging from

the 30-40 m high canopy in many of the cativo forests of the lower Balsas, Sambu, and

Tuira Rivers (Grauel and Pineda M. 2001). Areas with high liana densities seem to have

developed in large logging gaps created 20-30 years ago. Many mature cativo trees in

heavily infested areas are visibly deformed, apparently from having developed while

carrying large liana loads or from having been damaged during logging. In other areas

that have been continually and recently subj ected to small scale harvesting, D. brownei is

proliferating on the ground in large canopy gaps and appears to delay cativo regeneration.

In the present study I measured, by observation and experimental liana removal, the

effect of lianas on cativo adult stem growth as well as on seedling height growth,

recruitment, and mortality.

Study Site

The study was conducted in a riverine cativo forest along the Balsas River in

eastern Panama (8o 07' N, 77o 52' W). Mean annual precipitation at Camoganti, the

nearest town (approximately 8 km from the study site), is 2457 mm (based on

Government of Panama published reports for 1978-1982, 1984, 1986, and 1988-1994)

while rainfall measured at the study site in 1998 and 1999 totaled 2970 mm and 2758

mm, respectively. The forest is inundated periodically with rainwater during the 9-month

wet season from April to December. In addition, it is flooded twice per day for about

five days during the monthly spring tides known locally as the 'aguaj e.' The freshwater

backup caused by the Pacific spring tides affects the riverine forests as far as Camoganti,

73 km from the mouth of the Tuira River at the Gulf of San Miguel. Although at the

study site the tidal flooding is mostly the freshwater backup, soil samples show a slight

brackishness (electrical conductivity 5.0 mmhos/cm) and mangrove forests are found










only 7 km downriver from the study site. Soils at the study site are heavy clays classified

in the suborders fluvent and aquept, are acidic to slightly acidic, and poorly drained

(Tapia 1999).

The study site is on private land owned by a logger and is next to an operating

sawmill. The owner, who has been logging cativo in Darien since 1960, is currently

logging further upriver and has protected the forest where the study took place because he

values it for hunting and aesthetics, although he told us that he had harvested a few

scattered trees about ten years prior to the study. This cativo forest is composed of about

95% Prioria copaifera of all size classes (Grauel and Kursar 1999). Other tree species

include Pterocarpus officinalis, M~ora oleifera, and Caurapa guianensis. Results from a 1

ha permanent plot show 10 cativo trees per hectare > 60 cm dbh, the legal cutting limit,

but the maj ority of these were left due to bad form or hollowness. Regeneration of cativo

of all sizes is abundant.

Methods

In September 1997 six 25 x 25 m plots were installed in a line at 50-75 m intervals

in areas with intact canopies but with relatively high densities of lianas compared to the

forest overall. Each plot was subdivided into twenty-five 25-m2 Subplots to facilitate

stem mapping. Inside the plots I measured all trees > 4 cm dbh as well as the diameters

of all ascending liana stems > 1 cm at breast height. I did not attempt to differentiate

genetically distinct lianas; every stem encountered at 1.3 m above the ground was

measured. To increase the sample size for the growth analyses of cativo, additional trees

were measured up to 5 m outside of each plot, but no lianas were measured outside the

plots. Every plant was tagged and mapped and subsequent censuses were carried out in










1998, 1999, and 2001. For the growth analysis, diameter classes for trees were selected

based on relative canopy position; 15 cm dbh was used as the cutoff between canopy and

understory individuals. Due to the low canopy of the forest where lianas are abundant,

even trees 15-30 cm dbh may receive substantial direct illumination, while trees < 15 cm

dbh are generally in the understory.

During the initial measurements, each tree was classified as severely or lightly

infested by lianas. Severely infested trees had at least five individual liana stems hanging

from the crown and some stems or branches apparently deformed by lianas. Lightly

infested trees had fewer than five liana stems hanging from the crown and no visible

deformations. For the growth analyses, growth rates of liana-free trees were included in

the lightly infested category. All lianas were cut with a machete inside and up to 10 m

outside of three randomly chosen plots.

In 10 randomly chosen 25m2 Subplots in each plot, all natural regeneration of

cativo from seedlings to small trees 1 cm in diameter were counted, tagged, and measured

(height) before the vine cutting treatment and two years later. Where necessary to reduce

heteroskedasticity, seedling frequency data were natural log-transformed. Mean relative

height growth for seedlings in treated and control plots was compared with a two-sample

t-test and the difference of mean absolute height growth was tested by ANOVA using

initial height as a covariate. Mortality of these seedlings and small trees in treatment and

control plots was also compared. Two years after the initial census, the same subplots

were surveyed for new cativo regeneration. Treatment differences in mean density of

seedlings recruited per plot was compared with t tests.









For several reasons, including the observation that increases in cross sectional area

of lianas are associated with much larger increases in leaf area than in trees (Putz 1983), it

is desirable to estimate diameter growth rates of lianas. In 2001, four years after the

initial measurements, 56 Dalbergia brownei lianas in the control plots were again

measured to estimate stem diameter growth. Individuals < 6 cm dbh were measured with

dial calipers; a mean diameter was calculated from measurements of the long and short

axes. Lianas > 6 cm dbh were measured with a diameter tape. Wood density was

estimated using ten bark-free stem samples, to allow comparisons with other studies.

While growth rates of trees are often negatively correlated with wood density, this pattern

may not hold for lianas that do not produce structural wood for support.

Canopy openness above 2 m was measured immediately before and two months

after liana cutting in all plots with a vertical densitometer (Stumpf 1993). Both

measurements were made during the rainy season. This instrument proj ects a point

vertically upward that encounters either canopy or open sky at each evenly spaced sample

point along a linear transect. Canopy openness is estimated as the proportion of points of

open sky along three transects in each plot.

To compare rates of cativo growth and regeneration in heavily vine-infested areas

with forest with low liana infestation, data from the six plots of the present study were

compared with data from plots selected at random for a demographic study of cativo in

the same forest. The demographic study was based on five 20 x 20 m and five 40 x 40 m

plots established in March 1997. All trees > 10 cm dbh were tagged, mapped, and

measured (dbh), while trees > 1 cm dbh were measured in all five 20 x 20 m plots and in

five randomly chosen 20 x 20 m subplots in each of the 40 x 40 m plots. All trees were









measured annually from 1997 to 2001. In addition, all trees < 1 cm dbh in eight

randomly chosen 5 x 5 m subplots of each plot were tagged and mapped and were

measured (height only) in November 1997. This population of seedlings and saplings

was censused approximately every two months for two years whereas height was

measured annually.

Results

Two months after cutting lianas, significant but modest increases in canopy

openness were observed in the treated plots. There was no difference in the before and

after canopy coverage in the three control plots, while the three treated plots showed a

mean increase of 7% (p < 0.01) in canopy openness.

Mean annual diameter growth of Dalbergia brownei was 1.3 mm yr- (n = 56, sd =

1.4, range = -0.8 to 5.5 mm). Mean wood density of D. brownei (dry weight/fresh

volume) based on ten samples was 0.38 g cm-3 (Sd = 0.047).

Based on the mean number of stems > 4 cm from the six 25 x 25 m plots, cativo

dominated the forest with 1320 stems ha-l (sd = 212), virtually identical to nearby areas

of riverine forest with lower liana densities (133 8 stems ha l, Grauel and Kursar 1999).

Pterocarpus officinalis, the only other abundant tree species, was represented by 51 stems

ha-l > 4 cm dbh. The 35.1 m2 ha-l of cativo basal area represents 96.6% of the total basal

area of trees > 4 cm dbh.

For all cativo trees > 4 cm dbh, 71% had lianas hanging from the crowns, while

93% of mid- and upperstory trees (> 15 cm dbh) had lianas. There were 1757 ascending

liana stems ha-l > 1 cm dbh (sd = 270), with a mean liana basal area of 3.40 m2 ha-l (sd =

0.8). Of the two liana species found, Dalbergia brownei comprised 96.9% of the basal









area. The only other liana encountered, Elachyptera floribunda (Hippocrateaceae), was

mainly represented by small stems, with 75% of the ascending stems < 3 cm in diameter,

while over 80% of the D. brownei stems were > 3 cm in diameter (Figure 2-1).

Prior to liana cutting, in the six heavily liana-infested plots the mean density of

cativo seedlings and saplings (< 1 cm dbh) was 707 ha-l (sd = 1154). In contrast, in the

ten randomly located plots for the demographic study of cativo at the same site, the mean

density of cativo seedlings and saplings was 6350 ha-l (sd = 12882, Figure 2-2).

Although this was almost an order of magnitude difference and is plainly discernible in

the forest, variability was large due to the clumped distributions of seedlings, but the

difference was significant (t = 3.12, df = 14, p = 0.008).

For cativo regeneration present at the beginning of the study, relative and absolute

height growth over two years did not differ between liana-cut and control plots.

Although mean initial height for seedlings and saplings happened to be significantly

greater in the three control than in the three treatment plots, there was no difference in

initial height of only those trees that survived to produce growth records. Cativo seedling

and sapling mortality was nearly double in treated than in control plots (63% vs. 36%,

Pearson X = 6.2, p = 0.01).

Cativo seedling recruitment during two years after liana cutting was more than

three times greater in the treated than in the control subplots but, due to large variability,

was not statistically significant (t = 1.30, df = 4, p = 0.26, Figure 2-3). On a per ha basis,

over 7700 cativo seedlings recruited during two years after lianas were cut compared to

just over 2200 seedlings for the control plots.









Mean annual diameter growth of cativo trees during 1997-2001 was about twice as

rapid in the liana-cut compared to the control plots (Figure 2-4). For trees > 15 cm dbh

the difference was significant (t = 3.41, df = 4, p = 0.03), while for trees between 4 and

15 cm the difference was not statistically significant (t = 2.61, df = 4, p = 0.06).

Surprisingly, severity of liana infestation had little apparent effect (no significant

differences found) on cativo diameter growth for either control or treated plots (Figure 2-

5). The largest difference was for canopy trees in the control plots, where severely

infested trees grew slightly slower than lightly infested trees.

Discussion

Despite their abundance, liana cutting had only a slight (7%) but statistically

significant (p < 0.01) effect on canopy openness of the cativo forest, because most of the

liana foliage is displayed on the tops of tree crowns. Liana-infested cativo forests look

"feathery" at the top of the canopy, due to the abundant emergent branches of small-

leaved D. brownei searching for higher trellises. Two years after liana cutting, canopy

openness was similar for all plots, perhaps because the cativo canopies increased leaf

production after liana cutting. In contrast, Gerwing (2001) found that increases in canopy

light transmittance persisted for two years following vine cutting, and Perez-Salicrup

(2001) measured no change in canopy openness four months after vine cutting in a

Bolivian lowland forest but an increase in openness two years later. Both the Bolivian

and Brazilian studies took place in much drier forests and probably on less fertile soils

than the present study; perhaps the cativo trees were better able to take advantage of the

removal of lianas and produce foliage rapidly.

The low diameter growth rate of Dalbergia brownei is similar to the growth rate

found by Putz (1990, 1.4 mm yr ) for fifteen liana species from a tropical moist forest in









Panama. In a lowland wet forest in Ecuador the most abundant liana, Machaerium

cuspidatum, had an average annual growth rate of 1.4 mm yrl for stems between 30 and

50 mm (Nabe-Nielsen 2002). In contrast to these low diameter growth rates, a single shoot

of D. brownei was observed to grow 1.24 m in length in 71 days.

Increased mortality of cativo seedlings in liana-cut plots compared to control plots

appeared to be due to numerous large liana stems falling from the canopy, most within

the first year following cutting. In addition, floodwaters commonly move coarse woody

material around on the forest floor, which frequently results in the bending and breakage

of seedlings. This additional impact may explain why tree seedling mortality increased

significantly following liana-cutting here but not in a tropical tierra firme forest in

Bolivia with higher liana densities (Perez-Salicrup 2001).

Enhanced seedling recruitment in liana-cut plots (Figure 2-3) more than

compensated for increased mortality of the initially scarce regeneration in these heavily

liana-infested areas. Large cativo seed crops were produced in 1997 and 1999, and the

germinated seedlings from the May-June 1999 seedfall were captured in the plot census

in November 1999 before dry season (January-April) mortality occurred. Three possible

mechanisms for this increased seedling recruitment following liana removal include: 1)

reduced seed movement out of the liana-cut plots; 2) enhanced seedling survival; and, 3)

increased seed production.

Reduced seed movement could result because cativo seeds are large (mean fresh wt

= 48 g, Lopez 2001) and mainly dispersed by water. Seeds that fall and that are not

partially buried in the heavy wet soil may be transported by the monthly spring tides or










the periodic flooding caused by wet season rains. Fallen liana stems could have acted as

small dams inhibiting cativo seed movement during inundation.

If seed production and retention as well as capture of water dispersed seeds were

equal in all plots, high early mortality rates in the control plots would explain the

difference in seedling densities between control and treatment plots. But most cativo

seedling mortality occurs during the short dry season from January to April (Lopez 2002),

and the plots were censused after seedfall but before the onset of the dry season of the

second year after liana cutting.

Cativo trees newly liberated from lianas may have produced more seeds. A

possible mechanism that could help explain an increased production of seeds by cativo is

that lianas interfere with cativo flower or seed production, either physically or through

competition for light. D. brownei produces long recurved spines that wrap around small

diameter objects that they encounter, such as flower-bearing branches. A drain on host

resources caused by the constriction of vascular elements in branches or twigs was

proposed by Stevens (1987) to explain the negative effect of lianas on the fecundity of

Bursera simaruba trees in Costa Rica, and a similar mechanism may be at work in the

cativo/Dalbergia canopy.

Another possible mechanism for the proposed increased seed production in the

treated plots is the removal of belowground competition after the death of the lianas,

resulting in increased nutrient availability. Increased water availability may also have

been a factor, since seasonally flooded cativo forests can experience severe short-term

annual droughts. Putz (1991) suggests that because lianas do not need to produce large

diameter structural roots, root systems of lianas may be more efficient in water and









nutrient uptake. Lianas were also experimentally shown to be effective belowground

competitors in a study with north temperate vine species (Dillenburg et al. 1993).

Although liana cutting resulted in a notable positive increase in Prioria copalfera

stem diameter growth, lianas may not be solely responsible for low forest-wide growth

rates, as proposed by Gerwing (2001) for a seasonal Amazonian forest in Brasil. In the

riverine cativo swamps in Panama discussed in this study, mean annual growth of trees in

the control plots was not significantly different than similar sized trees in the nearby

permanent plots where lianas were generally less abundant (data not shown).

Mean annual growth of cativo trees of all sizes in both treatment and control plots

was greatest during the second year following liana cutting and then declined (Table 2-

1). Although the liana leaves fell during the first two months after cutting and the

hanging stems fell within the first year following treatment, this pattern of increased

growth followed by decline is probably not due to a "fertilizer effect" from the fallen

liana material; forest-wide growth rates for cativo, based on permanent plots at this site

and three others in different watersheds in Darien Province, showed the same pattern,

suggesting a correlation with climate.

The observation that the growth rates of liana-infested control plot trees > 15 cm

dbh did not differ from growth rates of trees of the same dbh in adj acent permanent plots

with few lianas (data not shown) could be attributed to the lower overall canopy height in

the area of the liana-cutting plots. With fewer large trees in the liana-abundant areas of

the forest, trees that otherwise would be subcanopy individuals may receive more light

than a similar sized tree in adj acent non-liana forest. This interpretation suggests that

these liana-abundant areas are old canopy gaps in the process of recovery, similar to the









"vine-dominated disclimax" of Whigham (1984) or the "stalled gap" of Schnitzer et al.

(2000).

The profound negative effect of lianas on cativo growth and reproduction probably

results from a combination of aboveground and belowground influences. Lianas were

estimated to occupy 3 1% of the forest canopy surface area during the wet season in a

seasonally dry tierra firme forest in central Panama (Avalos and Mulkey 1999) and liana

leaves might significantly reduce light availability for cativo leaves. Belowground

competition for nutrients and water in the dry season could also be a factor. Given that

rooting depth in cativo forests is limited by the high water table (Lopez 2002),

belowground competition for water may be severe during the dry season because root

systems die back during wet season flooding (Lopez 2002). In a semi-deciduous lowland

forest in Bolivia, Perez-Salicrup and Barker (2000) found significantly less negative

water potentials in Senna multifuga trees where lianas were cut as well as increased tree

diameter growth. In contrast, in the same forest after liana cutting, Barker and Perez-

Salicrup (2000) found no difference in water status of mahogany trees with and without

lianas and concluded that lianas and trees had access to different sources of water due to

different rooting depths.

Two issues pertinent to considerations of liana cutting as a silvicultural tool are cost

of implementation and biodiversity impacts. Based on the experimental plots, it would

require 16 person-hours to treat one hectare of forest. Although the experiment took

place on private land, the maj ority of these degraded forests are on state land managed by

the Panamanian Environmental Ministry (ANAM). Currently, ANAM is working with

several communities in Darien to develop a partnership whereby forest areas are









identified for potential timber production and legal tenure is transferred to local

community groups. The government intends to train community members in mapping,

inventory, and other management activities; liana cutting could be one of the

recommended silvicultural treatments.

Conservation of biodiversity is a critical issue in our study area, especially because

of the presence of Darien National Park and the current improvement of the Pan-

American highway that will doubtlessly increase colonization rates and forest clearing.

Liana-cutting in seasonally flooded cativo forests is not expected to have severe effects

on species diversity for several reasons. Tree species diversity is extremely low in

tidally-flooded cativo forests (Grauel and Kursar 1999) and only one species of liana

seems to have proliferated excessively as a result of several decades of uncontrolled

timber exploitation. Because this species, Dalbergia brownei, is also a common

component of nearby treeless wetlands, attempts at controlling its proliferation in cativo

forests suited for timber production probably will never eliminate it. Promoting

sustainable management of cativo forests for timber production could actually serve as a

biodiversity conservation tool. By providing local communities with viable economic

activities in the areas surrounding Darien National Park, pressure to exploit the natural

resources of the park could be reduced.






34


Table 2-1. Mean (+1 SE) annual diameter growth (mm) of Prioria copalfera one, two,
and two to four years after liana cutting.
1997-1998 1998-1999 1999-2001


Liana
Treatment Infestation


Control ih
severe

.light
Lianas Cut
severe


Mean Annual
N Increment (mm)

171 1.5 (0.2)
204 1.6 (0.2)

166 2.2 (0.2)
180 3.3 (0.3)


Mean Annual
N Increment (mm)

168 2.2 (0.3)
196 3.1 (0.3)

164 4.1 (0.4)
172 6.6 (0.4)


Mean Annual
N Increment (mm)

152 1.6 (0.2)
181 1.8 (0.2)

144 2.7 (0.3)
154 4.5 (0.3)

















600

0 Elachyptera floribunda
5001 1 Dalberqia browlnei




(n 300





100







0.6
O Elachyptera floribunda
0.51 g Dalberqia browvlei

.c~ 0.4


S0.3


S0.2






1-2 2-3 3-4 4-5 5-6 6-7 7-8 8-9 9-10 10-11 11-12 >12

Diameter Class (cm)


Figure 2-1. Diameter distributions of ascending lianas in six 25 x 25 m plots in heavily
infested riverine Prioria copaifera forest degraded by repeated entry logging.
a) Number of stems per hectare. b) Basal area in square meters per hectare.













10000 I


8000-


v, 6000--


.c 4000--


2000-



High Low
Liana Density



Figure 2-2. Mean (-E 1 SE) density of Prioria copalfera regeneration (< 1 cm dbh) in
areas of high (N = 10) and low (N = 6) liana densities.













301 I i, 12000


25- 10000 (D


~f20- __, 8000


S15- 6000 v,


S10- 4000


v, 5- 2000


Control Vines Cut
Treatment



Figure 2-3. Mean (-E 1 SE) Prioria copaifera seedling recruitment censused two years
after liana cutting in three control and three treatment plots.



















S5-
C3
S4-




I I I Treatment
<( 1 -
o Control
01 m Lianas Cut
4-15 >1
DBH (cm)



Figure 2-4. Mean (-E 1 SE) annual Prioria copaifera diameter growth based on five
annual censuses of all trees > 4 cm dbh in three control plots and three plots
in which all lianas were cut at the beginning of the study.



















2 6 Control Lianas cut







Infestation Level
o Light
0 m Severe
4-15 >15 4 -15 >15
DBH (cm) DBH (cm)




Figure 2-5. Mean (-E 1 SE) annual Prioria copaifera diameter growth of cativo based on
five annual censuses according to liana infestation level in control and
treatment plots.















CHAPTER 3
GROWTH AND SURVIVAL OF Prioria copaifera SEEDLINGS PLANTED ALONG
A HABITAT GRADIENT IN A PANAMANIAN SWAMP

Introduction

Until relatively recently, native tropical species have been little utilized for

reforestation programs (Evans 1992). For plantation forestry in the past, a general lack of

information led to a reliance on few familiar exotic species with high growth rates.

Concern over the loss of biodiversity as well as recognition of other production systems

besides monospecific plantations has increased attention on native species for

reforestation (e.g., Butterfield 1995, Haggar et al. 1998). Furthermore, much effort has

now been expended in acquiring knowledge about growth and mortality rates, as well as

propagation techniques, of native tree species in the tropics, often with the specific aim of

identifying promising candidates for reforestation (Condit 1995, Foroughbakhch et al.

2001, Wightman et al. 2001, Moulaert et al. 2002).

Reforestation with native tropical species may be carried out for timber production

(Keenan et al. 1999), site restoration (Parrotta and Knowles 1999, Engel and Parrotta

2001, Montagnini 2001), fuelwood production (Kataki and Konwer 2002), carbon

sequestration (Silver et al. 2000), biodiversity conservation (Blakesley et al. 2002a,

Blakesley et al. 2002b), or other reasons. For a given end use, species choice should not

be based soley on site characteristics or growth rates because the purposes for which trees

are planted may vary not only with the type of tree but also with the type of 'user'

(Raintree 1991). Certain landowners may be more amenable to reforestation with native










species where opportunity costs for labor are low or motivation consists of a wider range

of benefits than only financial profitability (Putz 2000).

In this study, a valuable native timber tree, Prioria copaifera (hereafter 'cativo'),

was planted to evaluate growth and mortality rates in different habitats in swamp forests

in eastern Panama. I hypothesized that both mortality and growth would be greatest in

the high-light environment and lowest in the forest understory where cativo natural

regeneration was abundant.

Cativo is a large Caesalpinoid timber tree that has been harvested commercially for

decades (Anonymous 1933, Hess and Record 1950, Kukachka 1965). Originally

distributed from Nicaragua to Colombia, logging and conversion to banana plantations

have severely reduced the area of cativo forests in Costa Rica and western Panama

(Veiman 1982, Jimenez Madrigal 1995). Today, commercial stands are restricted to

eastern Panama and northwest Colombia. Cativo is known principally from single

species or monodominant stands in swamp and riparian habitats, but is also found, less

frequently, in upland soils where it sometimes dominates (Condit et al. 1993b).

In Darien, Panama, the area of cativo-dominated forests has been reduced to less

than 15,000 ha from an original coverage of 60,000 ha (Grauel and Pineda M. 2001).

Many riverine cativo forests are adj acent to extensive marshes composed of palms, dense

liana tangles, and occasional "vine towers" where trees still stand from the remnant

forest. In the early 1950s Lamb (1953) described a belt of commercial cativo forests

along the Balsas River that averaged 1km in depth on each bank of the river, but in 2000

the belt was only 100m deep in some areas. Decades of logging has left many of these

cativo forests badly liana-infested and otherwise severely degraded (Grauel and Putz









2004). Hence, an important obj ective of the study was to determine the feasibility of

restoring treeless marshes to cativo forests.

Study Site

The study was conducted in a forest on private land along the banks of a tributary

of the Tuira River, the Balsas River, in Darien Province in eastern Panama (8o 07' N, 77o

52' W), 48 km from the mouth of the Tuira River at the Gulf of San Miguel. The

landowner has been logging cativo in Darien for forty years, and although he has cut only

a few scattered trees at the study site within the last ten years, it is likely that this forest

was subjected to an initial selective cutting around thirty years ago. In an early survey of

cativo-dominated forests Barbour (1952) noted that commercial-sized trees ranged from

60 to 120 cm in dbh (diameter at breast height, 1.3 m), with occasional specimens of 150

to 180 cm dbh. Today, few large trees are found in these easily accessible riverine forests

(Grauel and Kursar 1999).

The arboreal component of the study site is composed of 95% cativo (basal area

and stems ha l) of all sizes (Grauel and Kursar 1999). Other tree species that occur in the

stands include Pterocarpus officinalis, M~ora oleifera, and Carapa guianensis. Marshes

are found adj acent to the cativo forest and are composed of scattered palms (Elaeis

oleifera), liana tangles, and occasional "vine towers", suggesting that the tall forest has

been displaced. The lianas of the marsh, principally Dalbergia brownei, climb the tree

canopy at the well-defined edge of the forest where there is abundant light, but seldom

colonize the understory of the high-statured forest where little light penetrates.

Mean annual precipitation at Camoganti, the nearest town (approximately 8 km

from the study site), is 2457 mm (based on Government of Panama published reports for

1978-1982, 1984, 1986, and 1988-1994) while rainfall measured at the study site in 1998









and 1999 totaled 2970 mm and 2758 mm, respectively. The study site is subjected to

periodic flooding from rain events during the 9-month wet season (April-December) and

is also flooded when river water is backed up by high monthly tides. The slightly

brackish soils at the study site (electrical conductivity 5.0 mmhos/cm) are heavy clays

classified in the suborders Fluvents and Aquepts, are acidic to slightly acidic, and are

poorly drained (Tapia 1999).

Methods

In August and September 1997, cativo wildings of two ages were dug up in the

forest and planted in three different habitats with distinct light environments. First year

seedlings, seeds that matured in May-June of 1997, were identified by having an attached

seed and little woody tissue. Individuals referred to as "older seedlings" had no attached

seed and had woody stems. These latter seedlings may have been 3 years or older given

that cativo seedlings can survive for a number of years with little or no growth (Lopez

2002). These latter seedlings were likely 2-4 years old.

I tested the effects of 3 levels of canopy cover on two ages of seedlings in four sites

(= blocks) separated by 50-200 m along forest-marsh interfaces. In each of four blocks,

side-by-side pairs of 7 x 7 m plots were installed in the shade of the forest, on the edge

between the forest and treeless marsh, and in the marsh in full sunlight. Site preparation

in the marsh involved extensive liana cutting to establish plots and facilitate access but

only moderate liana cutting in the edge habitat where liana density was not as abundant

due to the partial shade of the adj acent forest canopy. Seedlings were carefully dug up

and bare root planted at 1 x 1 m spacing. For each plot pair in a given light environment,

seedling age was randomly assigned and 49 seedlings of a given age were planted in each

plot. Therefore, for each habitat seedling age combination, 196 seedlings were planted









for a total of 1176 seedlings. In the first plot that was planted, the seedlings (older,

shade) were transplanted with entire blocks of soil to protect the roots. This practice

proved to be excessively laborious and was abandoned. All subsequent results exclude

these 49 seedlings that had high survivorship but little height growth at the end of the

study .

Canopy openness is defined as the proportion of the sky hemisphere that is not

obscured by vegetation when viewed from a single point (Jennings et al. 1999). At the

center of each plot, canopy openness at 1.3 m above the ground was measured with a

spherical densiometer (Lemmon 1957) by averaging four readings taken in the cardinal

directions.

After planting in August and September 1997, seedlings were tagged, each stem

was marked at 20 cm above the ground, and two perpendicular diameter measurements

were taken with Vernier calipers and averaged. Total height of each seedling was also

measured. Both height and diameter were recorded at approximately six month intervals

for two years; Einal measurements were made after an additional 20 months had elapsed

(July 2001). Repeated measures analysis of covariance was used with either initial height

or diameter as the covariate to test the significance of seedling age (between-subj ects),

habitat (within-subjects), and their interaction. In addition, final mean and maximum

height and diameter were each compared between seedling ages within habitats. Height

and diameter data were natural log-transformed to comply with the ANOVA assumption

of normality. For tests on seedling sizes after the initial measurements, Geisser-

Greenhouse adjusted p-values were used because the assumption of homogeneity of

variances of treatment-differences was not met (Maxwell and Delaney 1999).









Seedling survival was censused Hyve times in the first two years (1997-1999) with a

Einal tally in July 2001. Survival among the three habitats was compared for each

seedling age with one-way ANOVA, then Bonferroni post-hoc tests were used to

determine differences among means. All data were arcsine square root-transformed to

reduce the unequal variances found in a few plots.

Annual rates of absolute and relative growth in height and diameter were calculated

for one year, two years, and four years after planting. Growth was first examined for

each seedling age by comparing performance among the three habitats with ANOVA and

Bonferroni comparisons. Younger and older seedling growth was then compared with t-

tests for each habitat. For growth over the entire four years of the study, as well as Einal

seedling size, comparisons could only be made between the edge and sun habitats

because of high seedling mortality in the shade.

Results

Canopy openness among the three habitats ranged from 10-85% (Table 3-1). The

high variability for the readings in the edge environment resulted from one reading

capturing the forest canopy, one capturing the open sky of the marsh, and the other two

including both canopy and sky.

Younger and older seedlings differed significantly in both height and diameter at

the time of planting (Table 3-2). Older seedlings were twice as large in diameter and

about 25% taller than first year seedlings.

The repeated measures ANCOVA revealed that only habitat, not seedling age, had

significant effects on height and diameter (both G-G adjusted p < 0.001) during the six

measurements, and there was no age-habitat interaction (Figure 3-1). Maximum heights

for both seedling ages were found in the full sun, where mean maximum height among









the four blocks for younger seedlings was 382 cm and for older seedlings 350 cm.

Within the edge and sun habitats, however, there was no significant difference in

maximum or mean height attained between seedling ages at the end of the four year

study. Because of high mortality of first-year seedlings in the shade, no comparisons

could be made regarding final size attained in that habitat.

The planted seedlings were also largest in diameter in the full sun habitat. The only

significant difference in all comparisons of diameter and DBH, however, was found in

mean stem diameter, where older seedlings were slightly larger than younger seedlings in

the sun (38.6 mm vs 32.2 mm, t = 2.5, df = 6, p = 0.047).

For the edge and sun habitats, 88% 99% of total four year mortality occurred in

the first seven months after planting (Figure 3-2). Seedlings initially survived better in

the shade but at the end of the study mortality was highest for both seedling ages in the

shaded understory. Results from the ANOVAs showed that survival was consistently

highest for both seedling ages in the edge habitat, but the Bonferroni tests revealed no

significant differences in survival at the end of the study between the edge and full sun

habitats for either seedling age. Furthermore, for the older seedlings there was no

significant difference between survival in the shade and full sun habitats. Variation

among plots was fairly modest (Figure 3-2).

Maximum mean annual height and diameter growth for both seedling ages occurred

in the sun habitat during 1999-2001, when seedlings grew about 1 cm in diameter (at 20

cm stem height) and from 80-90 cm in height (Table 3-3). For each seedling age,

absolute and relative height and diameter growth were significantly different in most

habitat comparisons. The few exceptions occurred when growth in the edge habitat was









similar to growth in either the full sun or the shade habitat for a few growth intervals. In

comparisons of seedling ages within each habitat, annual relative height growth was

never significantly different for any growth period. During the 12 months after planting,

younger seedlings grew relatively faster in diameter than older seedlings in the shade and

edge habitats. In the second year after planting, younger seedlings' relative diameter

growth was faster only in the shade. Annual relative diameter growth rates for the period

1997-2001 were significantly greater for younger seedlings than older seedlings in both

the edge and sun habitats.

Discussion

This experiment was motivated by the continuing decrease in the area of cativo-

dominated swamp forests due to degradation from repeated entry logging. Planting

cativo wildings in the understory of the forest provided a baseline for comparison of

growth and survival with wildings planted in two habitats that might be considered when

the restoration of degraded swamps is a land management obj ective. There is a tradeoff

in terms of growth and survival and the labor requirements of site preparation and

maintenance. Initial clearing of dense, extensive liana tangles by hand proved to be much

more difficult than clearing areas in the edge habitat, and periodic cleaning was also

much easier in the edge habitat because lianas did not resprout as vigorously there.

Cativo has often been described as shade tolerant (Linares Prieto 1987a, 1988,

Tamayo Velez 1991) or as shade tolerant early in life but requiring high light levels for

further development (Linares Prieto et al. 1997). Younger seedlings had much higher

mortality in the shade of the forest understory than in the high light habitat, while older

seedlings had similar mortality rates in these two habitats, suggesting that seedling

establishment requires more than a single year.









Mortality rates observed in this study were generally higher than in a cativo

reforestation experiment in Colombia that examined the effects of different types of

planting techniques and timing of planting (Linares Prieto 1987a). That study occurred

on abandoned agricultural soils while the study site in Darien is less suited for agriculture

due to a slight brackishness in the soil and long hydroperiods (Tapia 1999).

Although mortality was highest in the shaded understory of the forest, seedling

deaths were not soley attributable to shade or root competition. During periodic

inundations, fallen branches float around and damage small cativo seedlings. Several

planted seedlings in the understory were found bent over completely by coarse woody

debris. Liana tangles in open habitats, in contrast, prevent this woody debris from

moving. Another agent of mortality present in the forest but not likely in liana-infested

areas was trampling by people, particularly hunters.

These findings add to the already highly variable published growth rates of cativo

seedlings and saplings. Maximum mean annual height growth after four years in the

present study was observed in the high light environment, where younger and older

seedlings grew 48 and 54 cm yr- respectively. These growth rates are similar but

predictably lower than those from a reforestation experiment in Colombia on less acidic

soils and in high light, where Linares Prieto (1987) reported annual rates of height growth

ranging from 67-76 cm yr- In that study, slow plantation height growth was attributed

to a lack of suitable mycorrhizae, infertile soils, or poor plantation management. Still,

plantation growth greatly exceeds seedling growth in natural forest. In natural cativo-

dominated forest in Darien, Panama, a large population (~ 1300 trees) of natural

regeneration, from newly germinated seedlings to small saplings 150 cm tall, revealed









essentially zero mean height growth for trees between 60 cm and 150 cm tall. Mean

annual height growth of seedlings less than 30 cm tall was 15 cm yrl and for seedlings

between 30 and 60 cm tall mean annual height growth was 5 cm yrl (Grauel 1999).

Cativo's fastest height growth seems to occur after germination until seed reserves are

exhausted (Tamayo Velez 1991).

In certain cases cativo may have the desirable socioeconomic and biophysical

attributes that would make it the opportune choice for reforestation. Small landholders or

cooperatives may have more reasons than just timber production for reforestation.

During the course of this study the author was asked by a member of a small-scale

loggers' cooperative about the feasibility of restoring degraded swamps by planting

cativo. A comment was also made regarding the lack of wildlife in the increasingly

extensive teak plantations in Panama, implying that native species attract wildlife. Cativo

seeds are consumed by agoutis (Dasyprocta aguti) and its leaves by land crabs (possibly

Gecarcinus spp), a species that also is harvested by local people.

Cativo is the principal raw material for the domestic plywood industry in Panama

but is also used locally for furniture and construction; its value lies in its relative

abundance and accessibility. Native timber species such as Tabebuia rosea, Dalbergia

retusa, Astronium graveolens, and Pachira quinata are currently being planted on a

commercial scale in Panama (Mariscal et al. 2002, Wishnie et al. 2002a, Wishnie et al.

2002b), but it remains to be seen if cativo can compete with other, more valuable

hardwoods, both native and exotic, for planting on upland soils. This study demonstrates

that cativo is particularly suited to reforesting severely degraded sites that are unsuited

for agriculture due to flooding and that previously were in cativo.










Table 3-1. Mean canopy openness at 1.3 m above the ground as estimated with a
spherical densiometer, arranged by seedling age and habitat.
Standard
Seedling Age Habitat % Openness
Deviation
shade 10.7 0.7
Younger edge 38.7 12.3
sun 83.9 4.1
shade 11.1 1.6
Older edge 27.8 13.4
sun 85.2 9.2


Table 3-2. Initial mean seedling height and diameter (at 20 cm above the ground;
standard errors noted parenthetically). All differences between seedling ages
are significant (p < 0.01) based on t-tests. N = 4 plots in blocks 2-4 and N = 3
plots in block 1.
Age Younger Older Younger Older
Block Height (cm) Diameter (mm)
1 41.4 (0.75) 56.9 (1.56) 2.7 (0.05) 5.6 (0.25)
2 40.0 (0.82) 55.1 (1.94) 2.6 (0.08) 5.8 (0.41)
3 43.3 (0.39) 60.5 (2.60) 2.7 (0.02) 5.9 (0.21)
4 50.4 (2.73) 65.8 (1.33) 3.0 (0.19) 6.5 (0.45)













Table 3-3. Mean annual growth rates. Units of absolute growth are millimeters for
diameter and centimeters for height. Comparisons are among habitats within
each seedling age for different time periods throughout the study. All
comparisons are significantly different (P < 0.05) except where noted, "a"
indicates no difference between edge and sun, and "b" indicates no difference
between edge and shade.
Diameter Growth Year
1997-1998 1998-1999 1999-2001 1997-2001
Seedling
Age Habitat Relative Absolute Relative Absolute Relative Absolute Relative Absolute
Sun 1.27 4.0 1.11 8.0 0.62 9.6 2.44 7.5
Younger Edge 0.83" 2.6ab 0.54 3.0b 0.28b 3.1 1.03 3.3b
Shade 0.27 0.7 0.14 0.6 0.04 0.1 0.05 0.2
Sun 0.54 3.5 0.98 9.7 0.51 10.4 1.30 8.3
Older Edge 0.33 1.9 0.46 3.7b 0.24 3.2 0.53 3.1
Shade 0.02 0.1 0.04 0.2 0.05 0.3 0.05 0.3



Height Growth Year
1997-1998 1998-1999 1999-2001 1997-2001
Seedling
Age Habitat Relative Absolute Relative Absolute Relative Absolute Relative Absolute


Sun
Younger Edge
Shade
Sun
Older Edge
Shade


0.36
0.15b
-0.01
0.25
0.09b
-0.03


18.2
7.3a"b
-0.6
15.8
4.9
-2.2


0.45
0.18
-0.10
0.48
0.25
0.01


31.4
11.6
-5.5
39.4
17.4
0.4


0.80
0.27b
0.04
0.74
0.23
-0.02


79.7
22.0b
1.8
87.2
22.5b
-1.4


0.99
0.32b
0.02
0.87
0.26
-0.01


48.4
16.0b
0.8
54.0
16.1
-0.07














Seedling Diameter Sun


Seedling Height Sun


40


E 30


20


10


-*- Younger Seedlings
-o Older Seedlings


Seedling Diameter Edge


Seedling Height Edge


Seedling Diameter Shade


Seedling Height Shade


I I I


E
E
U 6
a,
E
m
O


Aug 97 Apr 98 Nov 98 May 99Nov 99


Jul01 Aug 97 Apr 98 Nov 98May 99Nov 99


Jul 01


Figure 3-1. Mean diameters and heights (+ 1 SE) of planted Prioria copaifera (cativo)
seedlings. Repeated measures analysis showed significant differences in size
among habitats but not between seedling ages when initial size (at time of
planting) was included as a covariate.















Younger Seedlings Sun


Older Seedlings Sun


Plot

v-- F2
-o F3
~-- E4


8,
v
~----~ -o~--


40



20


Younger Seedlings Edge

so \
4\

60 i



40 -0 -0- -


Older Seedlings Edge


Plot


9?


a- C1

-o C3
O-- C4


-o- D1
-v-- C2


Younger Seedlings Shade


Older Seedlings Shade

Plot

-v- A2
--o B3
S-0- B4


\ "


Plot


v-- B2
o- A3
O- A4


Nov 97Apr 98 Nov 98 May 99 Nov 99 Jul01 Nov 97Apr 98 Nov 98 May 99 Nov 99


Jul 01


Date Date

Figure 3-2. Percent seedling survival, beginning with the first census (November 1997)
after planting (September 1997). Each point pertains to percent survival of
the original 49 seedlings planted. Plot letters refer to pairs within habitats (A
and B shade, C and D edge, E and F sun), plot numbers refer to blocks.















CHAPTER 4
STRUCTURE, COMPOSITION, AND DYNAMICS OF Prioria copalfera-
DOMINATED SWAMP FORESTS IN DARIEN, PANAMA

Introduction

Unplanned selective timber harvesting over time results in a pattern of chronic

disturbance that strongly shapes forest structure and composition (Kittredge et al. 2003).

Timber harvesting is a discrete event that, alone, does not necessarily lead to forest

degradation. But where logging is poorly done or is too frequent, forests may become

susceptible to fires and liana infestations (Nepstad et al. 1999, Pinard et al. 1999,

Gerwing 2002). Given the unfortunate commonness of this disturbance regime, an

understanding of forest stand development in response to chronic degradation is critical

to the pursuit of sustainable forest management because the diverse values of forests

depend largely on forest structure and species composition (Oliver and Larson 1996).

The disturbance history of a site substantially influences present day forest stand

structure and productivity. Unfortunately, detailed site histories are usually unavailable

and evidence of past disturbances may not be obvious. Given that stand structure alone is

insufficient to indicate population trends in natural forests (Condit et al. 1998),

understanding patterns of recovery in degraded forests requires demographic information

as well. Without knowledge of how species, cohorts, and even individual trees respond

to disturbances such as logging, predictions cannot be made regarding the likely

responses of forest stands to further management interventions.









Neotropical swamp forests dominated by Prioria copaifera, a Caesalpinoid legume,

were long ago noted for their theoretical ease of management due to their low diversity

and the ability of this species to regenerate naturally (Barbour 1952, Holdridge 1964).

These traits have apparently allowed Prioria-dominated forests to be especially resilient

to repeated-entry logging. While potentially easy to manage, managers of Prioria-

dominated forests suffer from a lack of demographic information. In more diverse forests

where trees of most species are scarce, predictions are often based on small sample sizes,

although recent innovations such as large plots (Condit et al. 1999) and landscape-scale

sampling (Clark and Clark 1994, 1999) are addressing this limitation. But with few

exceptions (but see Favrichon 1998, Finegan and Camacho 1999, Fredericksen and

Mostacedo 2000), most information has been derived from unlogged forest preserves that

bear little resemblance to the actual working forests where harvesting occurs. And rarely

have studies of a particular forest type been replicated at different sites.

Although forest stand structure can reveal clues about disturbance regimes and past

uses, simple size distributions of stems often obscure underlying dynamic processes. In

particular, resprouting from snapped or partially uprooted trees can greatly influence

demographic parameters such as growth, recruitment, and mortality. Although

resprouting is increasingly recognized as an important component of forest dynamics,

many forest dynamic models omit this mechanism of regeneration and consequently

overestimate forest recovery rates (Paciorek et al. 2000).

Foresters typically focus on growth rates and on the adequacy of regeneration,

overlooking the importance of mortality rates of target species in the development of

management plans, often because they lack data. Although correlations have been made









between mortality and stand density (Lugo and Scatena 1996), light environment (Davies

2001), forest fragmentation (Mesquita et al. 1999), and climate (Condit et al. 1995b, Aiba

and Kitayama 2002), the estimation of a given species' mortality rate is usually difficult

due to small sample sizes and limited study periods. Although my study is based on only

Hyve annual censuses, I had the advantage of fairly large sample sizes that permitted

examination of mortality by stem type (fallen, inclined, resprout from erect stem, stem

sprout from prostrate or inclined stem) as well as evidence of growth-dependent

mortality. Although growth-dependent mortality has been reported for temperate forests

using growth estimates derived from tree rings (Kobe et al. 1995, Pacala et al. 1996,

Kobe and Coates 1997, Wyckoff and Clark 2000, Caspersen and Kobe 2001, Lin et al.

2001, Bigler and Bugmann 2003, van Mantgem et al. 2003), a frequent lack of tree rings,

inadequate sample sizes, and insufficient numbers of censuses have precluded

investigations of growth-dependent mortality in tropical forests (but see Finegan and

Camacho 1999 for a stand-level analysis).

Studying forests dominated by Prioria copaifera allowed collection of a large

dataset for a single, commercially important species and its main associates. In this paper

I describe the stand structure of five P. copaifera dominated forests in eastern Panama

and report on stand dynamics of four of those five sites based on Hyve years of monitoring

data; I also supply tree growth data from an additional six sites. All trees were described

on the basis of evident lean, breakage, and resprouting. Because P. copaifera sprouts

from erect broken stems as well as from inclined and partially uprooted trees, I

differentiate between these sprout types in presentations of data on growth, recruitment,

and mortality rates.










Distributed from Nicaragua to Colombia and also found in Jamaica, Prioria

copaifera (hereafter "cativo") was recognized as a potential source of commercial timber

in the first half of the twentieth century (Kluge 1926, Schmieg 1927, Anonymous 1933).

During and after WWII, interest in cativo wood increased (Harrar 1941, 1942a, b, Hess

and Record 1950, Hess et al. 1950, Barbour 1952). Much of the cativo forest in Costa

Rica was exploited to the point of current scarcity (Veiman 1982, Jimenez Madrigal

1995), while in the more remote parts of Panama and Colombia cativo forests were and

continue to be subj ected to silvicultural experimentation and intensive timber harvesting

(Lamb 1953, Mayo Melendez 1965, Christiansen 1980, FAO 1982, INRENARE 1987,

Linares Prieto 1988, FAO 1990, Alvarado Q. et al. 1996, CONIF 1997, Mariscal et al.

1999, Grauel and Pineda M. 2001). Although limited in extent, wetland forests

dominated by cativo are valued as sources of timber because of their large commercial

volumes (Golley et al. 1969, Grauel and Pineda M. 2001) and their ready accessibility by

niver.

Study Sites

Permanent plots were installed in 1997 at four sites in three watersheds in Darien,

Panama (Figure 4-1). Small populations of cativo trees at an additional six sites

provided additional information about growth of this species under a range of inundation

regimes. Eight of the ten sites were previously logged at various intensities and

frequencies, most recently three years before this study began. Plots were also installed

at a remote site near the Colombian border in Darien National Park where there was no

evidence of cativo logging.









Principal Sites

Casarete is located along the banks of the Balsas River (8o 07' 1 1-14" N, 77o 52' 19-

47"Wi), 19 km upriver from its confluence with the Tuira River and 48 km from the

Tuira' s mouth at the Gulf of San Miguel (Figure 4-1). Soils are heavy clays classified in

the suborders Fluvent and Aquept, are acidic to slightly acidic (pH 5.2-6.5), poorly

drained, and slightly brackish (electrical conductivity 5.0 mmhos/cm, Tapia 1999);

mangrove forests are found only 7 km downriver. Rainfall measured at the study site in

1998 and 1999 was 2970 mm and 2758 mm, respectively. Mean annual rainfall at

Camoganti, the nearest town, (8.5 km from the study site), is 2457 mm (Reputblica de

Panama 1995). Because the forest owner values the area for hunting and aesthetics he

protected it from logging for approximately 25 years (but did harvest a few large trees

approximately ten years before the study).

The Sambu River site (8o 03' 49"-04' 06" N, 78o 13' 16-33" W) at the mouth of the

small Chunga River, is 17 km upstream from the Sambu's mouth at the Gulf of San

Miguel. This site is approximately 4.5 km from Boca de Sabalo, also on the Sambu

River, and 9.5 km from the Wounaan village of Taimati, on the coast of the Gulf of San

Miguel, where mean annual precipitation is 1342 mm and 1592 mm, respectively

(Reputblica de Panama 1995). Soils are similar in texture but slightly less acidic than

Casarete (pH 5.7-6.8), and although it is nearer to the ocean, there was no evidence of

salinity (Tapia 1999). The forest is located on open-access, public land and was

repeatedly logged, the most recent entry three years before the study began.

Juanacati is 65 km from the mouth of the Tuira River near the town of El Real (8o

04' 38-49" N, 77o 46' 37-48" W). Mean annual rainfall at El Real (5 km from the site) is

2096 mm (Reputblica de Panama 1995). Although Juanacati, like the previous riverine










sites, is flooded by monthly spring tides, no soil salinity is evident and soil pH is higher

than the other sites (pH 6.2-6.3). There is no evidence of recent logging, but anecdotal

evidence and the forest' s proximity to El Real suggest that the site was repeatedly logged

in the past. Indeed, L.R. Holdridge noted the presence of stumps only a few kilometers

downriver from this site in 1962 (Holdridge 1964).

The fourth site is an inland location near a small intermittent stream called

Naranzati (8. _02' 58"-4' 26" N, 770 -55' 40"-58' 02" W), approximately 7 km west of the

town of Camoganti. Unlike the three riverine study sites, this forest is flooded for the

entire nine month wet season (April to December). Due to its inaccessibility, this site and

other inland swamps were subj ect to logging only recently as riverine cativo forests

became increasingly depleted of large trees.


Figure 4-1. Principal study sites. a) Casarete, b) Sambu, c) Juanacati, and d) Naranzati.









Secondary Sites

To compare cativo growth in different landscape positions and flooding regimes, I

chose three sites adj acent to mangrove forests and subj ect to brackish water inundation.

Bajo Grande (80 22' 23" N, 780 09' 24-31" W) is an area near La Palma on the coast of the

Gulf of San Miguel; the study site is a few hundred meters inland from the coast, behind

the coastal Rhizophora mangle forests. The other two tidal sites are slightly upriver from

the transition zone of red mangrove to cativo forest, one along the Tuira River (8o 10' 13-

33" N, 77o 50' 18-23" W) and the other along the Balsas River (8o 09' 03-17" N, 77o 53' 04-

10" W). All three tidal sites are monodominant, uneven-aged cativo stands.

Additionally, I chose three sites that are unaffected by brackish water for

monitoring cativo growth. Two of these freshwater sites are along the Balsas River,

between the principal Balsas River site (Casarete) and the community of Camoganti

(Bongales: 8o 05' 30" N, 770 51' 35" W and Limon: 8o 04' 13" N, 770-53' 16" W). Although

these sites are occasionally inundated by high tides, the floodwaters are comprised of

freshwater tidal backup and are not brackish. The final freshwater site is an inland

swamp far up the Amarraderro River, a small tributary of the Balsas River (8o 00' 48-55"

N, 77o 49' 38-41" W). All six secondary sites, except possibly the latter inland swamp,

were subjected to intermittent logging during the last decades of the 20th Century. A

seventh site is found near the headwaters of the Balsas River inside Darien National Park

near the Colombian border (7o 34' 23"-35' 32" N, 77o 47' 02-11" W), 130 km upriver from

the mouth of the Tuira River. This site was probably never logged for cativo due to its

remoteness, a condition that limited my access to a single visit.









Methods

Plot Descriptions

Because of the low tree species diversity of cativo-dominated forests, I was able to

use small plots and still gather sufficient demographic information for cativo and several

common associates. To capture landscape heterogeneity, and because some cativo

forests are limited to narrow bands along rivers, many small plots were installed at each

site instead of single large plots. Casarete has Hyve 40 x 40 m and Hyve 20 x 20 m plots,

Sambu has Hyve 40 x 40 m plots, and at Naranzati there are Hyve 40 x 40 m and three 40 x

20 m plots. At Juanacati, all trees > 4 cm dbh were measured in six 50 x 50 m plots. At

the other three sites, all trees > 10 cm dbh were measured in all plots and trees > 1 cm

dbh were measured in a randomly chosen 20 x 20 m quadrant of each plot, or in the entire

plot if it was 20 x 20 m (Table 4-1).

Most plots were installed in early to mid-1997, but the work was interrupted at

Naranzati and Juanacati and was completed in early 1998. All trees were tagged and

mapped at the time of plot installation and all plots were subdivided into 5 x 5 m subplots

to facilitate mapping. Trees > 7 cm dbh were measured with a diameter tape to the

nearest millimeter while smaller trees were measured with calipers, with two

perpendicular measurements being averaged.

Sampling and Analyses

To increase the sample size for large cativo trees, additional trees outside the

permanent plots were tagged and measured at Casarete and Naranzati. This approach

was not feasible at Juanacati and Sambu due to the overall scarcity of large cativo trees.










Three of the eight plots at Naranzati were mistakenly logged two months before the

1999 census. Analyses after that time were done using data from only the unlogged

plots.

In late 1997 and early 1998 all cativo stems < 1 cm dbh in 80 randomly chosen 5 x

5 m subplots within the larger plots at Casarete and 28 subplots at Sambu were tagged,

mapped, and measured (height); dbh was also measured for those saplings > 1.5 m tall.

These two populations of seedlings as well as new cativo recruits at these sites were

tagged, measured, and mapped approximately every two months for two years and

marked seedlings were measured annually.

The permanent plots were censused annually until 2001, except the Naranzati site

which was last censused in 2000. At the secondary sites trees 2 20 cm dbh were

measured in 1997, 1998, and 1999, except the Amarradero site, which was logged after

the 1998 census. Inter-census intervals were always nearly annual to minimize seasonal

effects on growth.

Methods of censusing and measuring trees as well as the approach to data checking

generally followed Condit (1998). During each census, in addition to recording each

tree's status (alive, dead, recruit), diameter (trees > 1 cm dbh), and height (trees < 1 cm

dbh), codes were assigned to denote if a stem was prostrate, inclined > 450 from vertical

but not lying on the ground, broken above or below the point of measurement (POM),

resprouting from a broken stem, or sprouting from a prostrate or inclined stem. I report

the incidences of these stem types, but for growth analyses I exclude fallen stems and

combine erect and inclined stems. I refer to stems that show evidence of previous

breakage and subsequent resprouting as broken stems. Although I separately coded trees










that had broken depending on if the break was above or below the POM, in this paper I

combine the two because there were only a few cases where a substantial recorded loss in

diameter was the result of stem breakage and subsequent resprouting. Living fallen stems

were measured if possible but were always tagged and mapped because they served as

hosts for vertical sprouts. References to prostrate stems refer to uprooted, not snapped,

stems. I measured all vertical sprouts from prostrate stems and from inclined stems if

they emerged from < 1.3 m from the ground.












Figure 4-2. Stem types: a) prostrate and inclined and b) vertical sprouts from a fallen
stem.

Two professional foresters performed all the ~ 22,000 dbh measurements in the

permanent plots. One forester measured trees only in 1997 and 1998, while the other (the

author) measured half the trees in 1997 and 1998 and all trees from 1999-2001, overall

measuring 80% of the trees in the study. I was also responsible for all error checking and

database management.

All data were independently entered into a computerized database by two different

people immediately upon return from the Hield. Any discrepancies between the dbh

measurements that could not be corrected in the office were noted, and follow-up field

trips a few weeks after the principal census were carried out to remeasure trees or

confirm codes.










Within sites, I first examined variability among plots of each year' s annual cativo

growth of erect, broken, and vertical sprouts with ANOVA and Tukey post-hoc tests.

Data were natural log-transformed if variances were substantially unequal. For these

comparisons, the longest annual growth record available was used. Of 3065 cativo trees

in the four principal sites, 80% of the annual growth records were for the period 1997-

2001, 14% were for 1998-2001, and 6% were for 1997-2000. For cativo, growth was

then compared among normal stems, broken stems, and sprouts from inclined/prostrate

stems within five diameter classes using ANOVA and Bonferroni post-hoc tests. To

explore inter-annual growth of cativo I used Hyve diameter classes and I combined all

stem types but excluded those that were prostrate on the ground. I also report annual

diameter growth of cativo' s three principal arboreal associates.

I report annual recruitment and mortality rates for cativo at the four principal sites

using four diameter classes for trees > 1 cm dbh and Hyve height classes for smaller trees

at the two sites where cativo regeneration was studied. In each census I used the totals of

stems from the previous census, that is, I do not calculate demographic parameters using

only the originally-tagged 1997 population (see Sheil and May 1996). The monitored

large trees outside the plots at Casarete and Naranzati were included in calculations of

mortality rates. Furthermore, in the few cases where trees were not found in a particular

year' s census, those trees were subtracted from the previous year' s total number of trees

to exclude them from calculations of recruitment rates, instead of assuming the trees had

died.

I evaluated the extent to which cativo mortality varied with recent growth rates.

With Hyve censuses at most sites, I was able to compare annual growth for a maximum of









three years between trees that were alive at the end of the study and those that died during

the study. I used two size classes (< 10 and > 10 cm dbh) and performed one-tailed t-

tests on populations with at least Hyve dead stems to test the hypothesis that slower

growing trees suffered a higher probability of mortality.

I report species diversity using Fisher' s alpha and the Shannon-Weiner (S-W)

index in two ways, first with only all stems > 4 cm dbh so that all four sites can be

compared. Then I also calculated the S-W index for the three sites with minimum dbh of

1 cm. Voucher specimens were collected from the four principal sites in 2000;

unidentified species were not separated into morphospecies; I noted the number of

unidentified species at each site and considered all unknown trees as a single species in

the diversity calculations.

Results

Tree Species Diversity and Stand Structure

Although cativo dominated all four principal sites, species diversity varied

substantially. The riverine sites on the Sambu and Balsas (Casarete site) Rivers showed

the greatest cativo dominance, with cativo comprising 95 and 96% of the stems of all size

classes, respectively (Table 4-2). Diversity indices grouped the four sites into pairs, with

Casarete and Sambu being strongly monodominant and Juanacati and Naranzati being

relatively more diverse. Only seven tree species were tallied at each of the former two

sites, while 48 and 54 species were identified at Juanacati and Naranzati, respectively.

For trees > 10 cm dbh at the five sites with plots, cativo comprised from 46-96% of the

stems and from 33-96% of the basal area (Table 4-3). Pterocarpus officinalis was

cativo' s principal associate common to all four sites, making up 2-10% of the stems.

Other common overstory trees at Juanacati were Pentaclethra macroloba (10.2%),










Calrapaguianensis (7.2%), Licania platyipus (3.3%), and M~ora oleifera (1.3%). At the

Naranzati inland swamp, cativo tended to dominate the overstory with P. officinalis_and

P. nzacroloba, but understory associates included Andira inernzis_(1.3%), Escinveilera

integrifolia (4.7%), Gustavia nana (2.0%), and Brownea rosa-de-nzonte (2.5%). The

palms Oenocarpus nzapora and Astrocalyunt standlyanunt were common understory

species at the two more diverse sites as well, comprising 2-8% of the stems.

Stand density and basal area varied considerably among sites (Table 4-3). Stand

density of trees > 10 cm dbh ranged from 328 trees/ha (Darien NP) to 757 trees/ha

(Casarete). Stand basal area varied less markedly, but most cativo basal area was

represented by trees > 60 cm dbh at the inland swamp (Naranzati) and in Darien NP, and

by stems 10-60 cm dbh in the riverine forests.

Cativo made up ~ 95% of stand basal area at the riverine Sambu and Casarete sites

and ~ 83% at the inland Naranzati swamp (Table 4-3). The Juanacati site was visually

dominated by 14 huge, emergent M~ora oleifera trees per hectare that made up 21% of

stand basal area, the same percentage as Pterocarpus officinalis, while cativo comprised

33% of basal area and Pentaclethra nzacroloba 14%.

For all species, substantially more large prostrate and severely inclined trees were

found at the three riverine sites than at the inland swamp or the remote Darien NP site.

No prostrate, living trees were noted at either Naranzati or Darien NP. In contrast, large

prostrate stems were fairly common at the three riverine sites, where 1-5% of live stems

>10 cm dbh were on the ground (Table 4-4). Large inclined stems were also more

prevalent in the riverine forests, where 4-8% of all stems were partially uprooted and

leaning > 450









The rates at which trees partially uprooted and fell to the ground or leaned > 450

were generally higher at all sites for trees > 10 cm dbh than for smaller trees (Table 4-5).

Consistent with the proportion of live fallen and inclined stems recorded at the beginning

of the study, rates of falling and inclination were higher at the riverine sites than the

inland swamp, but rates varied greatly among years within sites.

At the time of plot installation a modest proportion of stems showed signs of

previous stem breakage at most sites. Where trees < 10 cm dbh were measured, between

6-11% were broken, and approximately 3-6% of large stems had suffered but recovered

from stem breakage. No broken and resprouted stems were noted in the plots at Darien

NP.

Sprouts from prostrate and inclined stems were much more prevalent in the riverine

forests than inland swamps. Less than 2% of small stems (< 10 cm dbh) at the inland

Naranzati swamp consisted of these sprouts, and no larger sprouts were found. In

contrast, 6-17% of smaller stems were sprouts at the riverine sites. Among the riverine

sites, Casarete stood out by having more large than small stems classified as sprouts,

where 12% of all stems > 10 cm dbh were sprouts from prostrate or inclined parent trees.

The 23.5 m long stem of a 33 cm dbh cativo that partially uprooted and fell to the ground

at Casarete between the 1999 and 2000 censuses produced 175 vertical sprouts > 1.5 m

tall by the 2001 census. Fifteen of these shoots were inside the plot (within 5.7 m from

the root system of the parent tree) and were tallied as recruits in 2001, with mean and

maximum diameters of 1.7 and 3.2 cm dbh, respectively.

Cativo Growth, Mortality, and Recruitment

Relative rates of cativo growth of three stem types (undamaged,

broken/resprouting, or sprouting from inclined and prostrate trunks) are unique to each









site. Sprouts grew faster than either undamaged or broken stems at Casarete for stems

<40 cm dbh, while at Sambu the smallest sprouts grew slower than undamaged stems and

only sprouts 4-10 cm dbh from inclined or prostrate stems grew faster than undamaged

stems of the same size. Undamaged stems grew faster than broken stems at Juanacati but

growth rates of normal stems and sprouts from prostrate/inclined trunks were similar

(Table 4-6).

Cativo annual diameter growth varied considerably among both sites and years, but

a few patterns were apparent. Growth was slowest, with some exceptions, during the El

Nifio year of 1997-1998 and fastest for the following census period (1998-1999). For

trees < 10 cm dbh, the Casarete and Naranzati sites were both characterized by very slow

growth rates, while stems of this size class grew significantly faster in almost all years at

Sambu and Juanacati (Table 4-7). Mean diameter growth rates within the larger size

classes (2 10 cm dbh) varied notably among the 9 sites (10 sites for 1997), ranging from

only 0.5 mm to > 8 mm per year.

Sprouts from prostrate or inclined stems made up a significant portion of forest-

wide recruitment in many cases (Table 4-8). About 50% of saplings entering the 1 cm

dbh size class at Casarete were sprouts of this type. Also at Casarete, these sprouts

consistently comprised substantial portions of the recruitment into the 10 cm dbh size

class.

Recruitment and mortality rates for cativo show similar inter-site patterns to those

observed for growth. Mortality equaled or exceeded recruitment at Casarete and

Naranzati for trees <10 cm dbh in the first two years of the study (Table 4-9). In general,










recruitment of cativo at Sambu and Juanacati greatly surpassed mortality for all size

classes and years.

Mortality rates of different cativo stem types varied among sites, but in general,

sprouts from prostrate/inclined trunks and undamaged stems showed the highest mortality

rates at Casarete and Sambu (Table 4-10). No sprouts from prostrate/inclined trunks died

at either Juanacati or Naranzati; at these two sites either broken or prostrate stems had

mortality rates that approached or occasionally exceeded those observed for undamaged

stems.

Growth of other Tree Species

Pterocarpus officinalis was the fastest growing tree species at all four sites, with

mean annual diameter growth of 9-1 1 mm for trees > 10 cm dbh in three of the four

forests (Figure 4-2b). The other two most common associates of cativo, Calrapa

guianensis and Pentaclethra macroloba had mean annual growth of large trees > 10 cm

dbh that approached 5 mm at the two more diverse sites where they were found.

Growth-dependent Mortality of Cativo Trees

Surviving trees grew significantly faster (p < 0.05) than trees that died for nine of

ten t-test comparisons (Table 4-11). The non-significant case revealed significance

when the size class was further divided. For 1997-1998 growth, survivors at Casarete

<10 cm dbh grew equally slowly as those trees that subsequently died, but when these

smaller trees were analyzed as two size classes, surviving trees between 1-4 cm dbh grew

significantly faster than those trees that died (mean 0. 1 vs. -0. 1 mm, df = 90, t = 3.1, p =

0.002), while trees 4-10 cm dbh in both groups had essentially zero growth.









Cativo Regeneration

Regeneration was much more abundant at the more recently logged Sambu River

site than at Casarete (Table 4-12). Small seedlings (< 30 cm tall) were not abundant at

either site, suggesting that they grow rapidly after germination. Seedlings 30-60 cm tall

were most abundant at Casarete, while at Sambu seedlings 60-90 cm tall were the most

common. Due to a large seedfall in April and May 1999, annual cativo seedling

recruitment rates based on the period November 1998 November 1999 for Casarete and

Sambu were 136.6% and 29. 1%, respectively. Annual mortality for the same period was

similarly high for seedlings < 30 cm tall but differed substantially for taller trees between

the two sites (Table 4-13). Mean annual height growth was generally < 5 cm at each site

(Table 4-14). Exceptions were the relatively small number of seedlings < 30 cm tall at

both sites and saplings > 90 cm tall at Sambu, both of which grew rapidly.

Discussion

Forests dominated by cativo in Darien, Panama vary substantially in structure,

species composition, and stand dynamics. Flooding regimes and management histories

appear to be the main determinants of present-day structure and dynamics. Slight soil

salinity, in particular, seems to favor cativo dominance (Mayo Melendez 1965). Cativo

forests near mangrove forests exhibit almost total dominance by cativo, whereas inland

swamps and riverine forests that escape tidal flooding with brackish water contain

relatively high tree species diversity.

Cativo dominance can probably be attributed to a variety of mechanisms. Flood

tolerance alone is an insufficient explanation because other flood tolerant species are

generally rare in cativo forests (Lopez and Kursar 1999). Cativo's root system dies back

to a much lesser extent than other flood tolerant species and gives the species competitive









advantage in seasonally flooded forests that are subj ect to short but severe annual

droughts (Lopez 2002). High leaf area index, as shown by Holdridge (1964), may

modify the understory environment to be more favorable for cativo seedling survival than

for other species. Such a modification was attributed to Gilbertiodendron dewevrei, a

tropical tree species that forms monodominant stands in West Africa. Although the crab

species common to riverine cativo forests have not been studied in Darien, the land crab

Gecarcinus quadratus was shown to affect species diversity in a coastal Costa Rican

forest by selective seedling consumption (Sherman 2002).

The degree of cativo dominance varies widely in Panama. Cativo is a locally

common species in the 50 ha forest dynamics plot on Barro Colorado Island, with a mean

of 27-29 trees hal > 1 cm dbh and a maximum of 223 (Condit et al. 1993b). The three

sites in Darien for which I have comparable data have densities of cativo 5-15 times

greater than on BCI. Similar to cativo swamps in Darien, the relative basal area

dominance of cativo in Colombia is 50-92% (Escobar and Vasquez 1987). Anecdotal

evidence suggests that Caurapa guianensis, a species that produces wood of a similar

quality as mahogany, was probably much more common in the past in some cativo

forests and L.R. Holdridge (1964) identified C. guianensis as cativo's sole associate in a

1962 transect very near the Juanacati site on the Tuira River.

The histories of timber harvesting of these forests undoubtedly influence present

day stand structure and dynamics, but details on logging frequencies and intensities are

largely unknown. It is likely that all the riverine forests were repeatedly logged since the

1950s (Reputblica de Panama 1978). Although I characterized the inland swamps and the









Darien NP site as intact forests, it is possible that even these remote forests were logged

for mahogany several decades ago.

Stand structure analyses of cativo forests reveal high timber volumes or at least the

potential for high volume production. The forest at Darien NP stands apart from all other

sites with the lowest density of stems but the largest stand basal area. The other sites

(except Juanacati) contain higher basal areas than most other tropical forests (Leigh

1999). This is notable because the riverine cativo forests of Darien were identified for

their timber potential in the 1950s (Lamb 1953) and L.R. Holdridge noted stumps from

harvesting activities in the early 1960s near the Juanacati site (Holdridge 1964). The

history of logging is undoubtedly responsible for the paucity of trees >60 cm dbh (the

legal cutting limit), but these riverine forests still contain a similar number of stems >10

cm dbh as most other lowland tropical forests (Leigh 1999).

High stem density at Casarete may be a result of the site' s management history.

Having been protected by its owner from the repeated logging that typically occurs on

open access state land, this forest may have passed through a period of enhanced

recruitment after the first wave(s) of logging in the 1950s and 1960s. Increased

recruitment and growth of undamaged residual trees after logging is a well-documented

phenomenon (e.g. Magnusson et al. 1999, Parrotta et al. 2002). The Casarete forest 40

years ago may have been similar to the present-day Sambu forest that was recently

logged and exhibits high seedling and sapling densities, fast sapling growth, and low

sapling mortality.

Sprouting is a well-recognized regeneration strategy in tropical forest (Putz and

Brokaw 1989, Rijks et al. 1998, Gavin and Peart 1999, Kammesheidt 1999, Negreros-









Castillo and Hall 2000, Yamada et al. 2001) but sprout density varies from being

common (Paciorek et al. 2000) to absent in mature forests (Kammesheidt 1998).

Vegetative sprouts from various sources may be the principal colonizers of gaps (Putz

and Brokaw 1989, Negrelle 1995), but mortality rates of resprouted broken stems are

generally higher than non-sprouts (Guariguata 1998, Paciorek et al. 2000, Ickes et al.

2003). In cativo dominated forests it is important to distinguish between resprouts

originating from erect, broken trees and those that emerge from fallen and inclined trees.

Sprouts from fallen or inclined trunks are frequently much more common, often grow

more rapidly than trees originating from seed but have the highest mortality rates of any

stem type, including trees that have uprooted and are lying on the ground.

The tree recruitment assemblage in newly formed treefall gaps in cativo forests

may not be dominated by true seedlings. In some cativo forests the principal gap

colonists were not newly germinated seedlings or established shade tolerant saplings;

instead, regeneration was dominated by sprouts from inclined or prostrate stems. For

example, at least half the recruitment of 1 cm dbh stems every year at Casarete was

comprised of sprouts from fallen stems, which also grew faster than their conventionally

rooted counterparts. Sprouts from prostrate stems develop their own root systems

composed of roots that emerge from the bottom of the parent stem. At Casarete, these

sprouts continued their superior growth rates at least into the subcanopy after which it

was difficult to determine their mode of regeneration.

The three riverine sites share a somewhat similar history of logging as well as a

higher proportion of live prostrate and inclined trees than the inland swamps that have not

been logged. Logging and inclined or prostrate trees may not necessarily be related,









however. Riverine cativo forests were characterized by a large number of fallen trees at

about the time that widespread commercial logging was beginning in Darien (Duke 1964,

Holdridge 1964), but recent logging has been sporadic due to scarcity of commercial-

sized trees. Cativo sawnwood and plywood production during the late 1990s was only a

quarter of its peak in the late 1960s (Romero M. et al. 1999). I conclude that fallen and

inclined cativo trees may be a common feature of riverine forests due to their shallow

root systems and saturated soils, with logging being a lesser factor.

Growth rates of the various stem types at different sites may vary with forest

structure but may also be limited by different factors depending on stem type. Although

sprouts from inclined/prostrate trunks at Sambu and Casarete grew at similar rates,

undamaged stems and broken stems at Sambu grew four to five times faster than their

counterparts at Casarete, presumably because the recent logging at Sambu left a more

open canopy.

Growth of all cativo stem types increased with increasing distance from tidal

influences but decreased with increasing hydroperiods. In general, cativo trees in the

inland swamps that are flooded continuously during the rainy season grew more slowly

than in the riverine forests, and the riverine sites further upriver (Juanacati, Bongales,

Lim6n) showed faster mean cativo growth than downriver sites.

Diameter increment of canopy trees varied by site and year, but the patterns of

variability differed among sites. Canopy tree growth at Casarete varied up to 2-fold, with

1997 as the slowest growing year. At the other three principal sites, canopy tree growth

varied less, and the census period that spanned the 1997-1998 El Nifio was not always

the slowest growing year. These findings stand in contrast to the strong reductions in tree










growth in Costa Rica during the 1997-1998 El Nifio, which were negatively correlated

with daily minimum temperatures (Clark et al. 2003).

The fastest growing tree species in my sample plots was Pterocarpus officinalis,

which is not considered a timber tree. Cativo's two other most common associates,

Calrapa guianensis and Pentaclethra macroloba, are harvested in both Panama and Costa

Rica (Webb 1997, Sitoe et al. 1999). C. guianensis in particular is valued by Darien

loggers, while P. macroloba produces less valuable wood. C. guianensis is more

abundant in riverine forests flooded only with freshwater, and has moderate growth rates

and relatively high densities in some Darien cativo forests. This species may have been

locally extirpated in some riverine forests but could be reintroduced by seed scattering as

was recommended by Webb (1997) for logging gaps in swamp forest in Costa Rica.

In the absence of spatially-explicit data that allow for the development of

competition indices and the construction of distance-dependent forest dynamic models,

higher survival of faster growing trees (growth-dependent mortality) should be taken into

account when projecting growth traj ectories. When mean growth of a large population of

small trees is low, modeling lifetime growth based on mean growth may result in

unrealistically long traj ectories (Grauel and Kursar 1999), especially if a large proportion

of slow growing trees die before reaching commercial size.

I noted heavy seedfall during plot installation in April 1997, but the regeneration

study began in September 1997 at one site and May 1998 at the other, so I measured

recruitment in 1998-2000 at one site and 1999-2000 at the other. Although cativo

produces some seeds twice a year, large seedfalls seem to occur once every two years










(Pizano SA 1995), and I also measured a large pulse of seedling recruitment from July to

November 1999, two years after the large observed seedfall in April-May 1997.

When stand structures and dynamics vary so markedly among forests, it seems

inadvisable to extrapolate results widely. Witness the difference in cativo dynamics

between these Darien cativo forests and the population of cativo on Barro Colorado

Island. With much lower mortality rates and generally higher mean and maximum

growth rates for cativo on BCI, use of their data for management tasks such as timber

harvest scheduling or yield proj sections would justify over-harvesting of most Darien

cativo forests. Mortality, more than growth, was shown to be a pivotal factor in the

simulated sustainability of cativo harvest potential based on BCI data (Condit et al.

1995a), but annual mortality rates of Darien cativo forests varied greatly and were

sometimes much higher than on BCI.

A critical aspect for understanding present day forest structure and dynamics is the

history of use that has resulted in what are now degraded forests. Increasingly, these

degraded forests will be a source for wood and non-wood products as the area of intact,

mature tropical forest declines. For example, the forest most recently logged in this study

(Sambu) is one of the most dynamic and resilient, with relatively high growth and

recruitment rates, low mortality, and sufficient densities of advance regeneration to

theoretically provide additional timber harvests. Forest history, although it may only be

inferred, can yield insights into today's forest and perhaps help to guide management

direction.

This study highlights the importance of examining stand development patterns at a

variety of sites, even when a single "forest type" is identified based on species






77


composition and landscape position. Only where variability is recognized can it be

considered when making forest management decisions. Anthropogenic disturbance may

have been the primary factor in shaping the structure and function of many cativo-

dominated forests in Darien, but the persistence of these forests after decades of

harvesting attests to their resilience and should serve as inducement for better

management.











Table 4-1. Total plot area measured for different minimum tree diameters and number of
tree species found.
Plot Area (ha) Number of Tree Species
Site >1 cm >4 cm >10 cm >1 cm >4 cm >10 cm % unidentified
Casarete 0.4 0.4 1.0 8 7 6 < 0.1
Sambu 0.2 0.2 0.8 7 5 5 0.0
Juanacati -1.5 1.5 48 24 0.4
Naranzati 0.32 0.32 0.96 54 42 28 1.5


Table 4-2. Species diversity indices and relative dominance of cativo (Prioria
copaifera).
DBH > 1cm
Cativo
Site Fisher's a S-W Evenness Simpson dominance
Casarete 1.10 0.20 0.09 0.93 0.96
Sambu 1.00 0.22 0.11 0.91 0.95
Naranzati 12.37 2.10 0.53 0.32 0.55

DBH > 4cm
Cativo
Site Fisher's a S-W Evenness Simpson dominance
Casarete 1.01 0.22 0.10 0.92 0.96
Sambu 0.76 0.28 0.16 0.87 0.93
Juanacati 9.52 2.09 0.54 0.25 0.47
Naranzati 7.19 1.81 0.52 0.36 0.59


Table 4-3. Stem density and basal


area of all species (above) and cativo only (below).
Stems/ha BA/ha (nr)
>10 cm >10 cm
757 43.3
484 48.9
426 31.4
339 47.1
328 71.1
Cativo Cativo Cativo Cativo
Stems/ha BA/ha (nr') Stems/ha BA/ha (m )
>10 cm >10 cm >60 cm >60 cm
727 41.5 8 2.5
463 39.9 12 4.4
195 10.3 3 0.9
240 39.3 51 29.2
160 48.8 58 45.4


Stems/ha
1-4 cm
1460
2095

1675

Cativo
Stems/ha
1-4 cm
1415
2030

732


Stems/ha
4-10 cm
705
785
500
1057

Cativo
Stems/ha
4-10 cm
668
690
229
457


Site
Casarete
Sambu
Juanacati
Naranzati
Darien NP




Casarete
Sambu
Juanacati
Naranzati
Darien NP











Table 4-4. Incidence (%) of prostrate, inclined, broken stems and sprouts from prostrate
trunks. Other stems showed no signs of earlier breakage and presumably
regenerated from seed.
Sprouts from
Inclined or
Prostrate Stems Inclined Stems Broken Stems Prostrate Stems
<10 >10 >10 >10 >10
Site cm cm <10 cm cm <10 cm cm <10 cm cm
Casarete 0.0 5.0 3.0 7.7 8.2 4.5 8.3 12.0
Sambu 0.0 3.1 1.9 7.2 6.1 4.0 16.7 1.6
Juanacati 2.1 1.1 4.2 4.1 10.9 5.6 5.9 0.7
Naranzati 0.0 0.0 3.5 0.9 9.7 3.1 1.8 0.0
Darien NP 0.0 0.0 0.0 0.0


Table 4-5. Forest-wide annual treefall and tree incline rates (i.e., partial uprooting) for
small (above) and large (below) trees for four sites.
< 10 cm dbh 1997-98 1998-99 1999-2000 2000-01
Incline Fall Incline Fall Incline Fall Incline Fall
Casarete 0.40 0 0 0 3.22 0.57 0.23 0.23
Sambu 0.17 0 0 0 0.13 0.13 0.15 0
Juanacati 1.48 0.37 0.97 0 1.18 0.34 0.49 0
Naranzati 0.87 0 0.55 0 1.16 0.93 -


> 10 cm dbh

Casarete
Sambu
Juanacati
Naranzati


1997-
Incline
0.56
1.43
0.88
0


98
Fall
0
0
1.32
0


1998-99
Incline Fall
0.26 0
1.08 0.81
0.73 0.36
0.29 0.58


1999-
Incline
2.38
1.87
0.53
0.33


2000
Fall
1.68
0.42
1.24
0.33


2000-01
Incline Fall
0.46 0.58
0.46 0
0.26 0.38






























Sprouts from
prostrate/
DBH cm Undamaged Broken DBH cm Undamaged Broken inclined trunks
1-4 0.9" (92) 2.7" (8)-
4-10 1.2" (79) 2.6" (5) -4-10 3.0" (288) 2.0b (34) 2.7ab (15)
10-20 2.7" (61) 1.7" (5) -10-20 5.3" (164) 2.8" (9) 5.7" (3)
20-40 4.7 (43) --20-40 8.5" (70) 2.5b 7
S40 4.5 (140) 40 7.1 (32)--


Table 4-6. Mean annual diameter growth (mm/year) of cativo trees of three stem types,
based on 1997-2001, 1998-2001, or 1997-2000 census periods. Within
diameter classes, different letters denote significant differences (p<0.01)
among stem types, (sample sizes noted parenthetically).


Casarete


Sambu


Sprouts
from
prostrate/
inclined
trnmks
0.9b (25)
1.2" (14)
4.0b (63)
5.4b (19)


Sprouts from
prostrate/
inclined trnmks
1.0b (66)
4.1b (20)
6.60 (6)


DBH cm Undamaged
1-4 0.35" (427)
4-10 0.7" (205)
10-20 1.80 (288)
20-40 3.5a (204)
S40 2.6 (120)
Naranzati


Broken
0.3" (33)
0.9" (20)
1.6" (22)
1.2" (4)
0.6 (3)


DBH cm
1-4
4-10
10-20
20-40
> 40
Juanacati


Undamag~
1.5a (301)
2.00 (117)
4.60 (140)
4.2a (101)
2.8 (103)


ed Broken
1.5ab (24)


6.5" (6)
2.2" (4)
0.8 (2)










Table 4-7. Mean annual diameter growth (mm/year) of cativo trees. Statistical
comparisons are vertical, within diameter classes and among sites; different
letters denote significant differences (p<0.01; sample sizes noted
parenthetically). All stem types except prostrate are grouped.
1997-98 1-4 cm 4-10 cm 10-20 cm 20-40 cm > 40 cm
Casarete 0.1b (485) 0.0b (242) 1.2b (374) 2.3C (230) 1.7b (123)
Sambu 1.8a (396) 2.2a (138) 4.4a (153) 4.3b (108) 3.3ab (107)
Naranzati 0.3b (64) 0.2b (52) 1.0b (37) 3.0b,c,d (20) 3.5a (23)
Juanacati -3.1a (118) 4.7a (53) 5.4a.b (26) 7.5a (9)
Bajo Grande --2.0a,b (9) 1.7c~d (70) 1.9b (48)
Rio Amarradero .7a,b (11) 0.5c~d (51) 2.0b (92)
Rio Balsas1 .5d (96) -0.5b (21)
Bongales --3.1a,b (11) 5.8a.b (62) 5.9a (34)
Limon --5.0a,b (4) 8.4a (36) 8.1la (8)
Rio Tuira --3.1ab (11) 2.0cd (151) 1.2b (38)
1998-99 1-4 cm 4-10 cm 10-20 cm 20-40 cm 2 40 cm
Casarete 0.5b (491) 0.9b (237) 2.5C (376) 4.3C (227) 3.5C (126)
Sambu 1.6a (410) 3.7a (148) 6.4b (156) 5.6bc (105) 3.90 (104)
Naranzati 0.6b (185) 1.2b (117) 2.4C (97) 4.2C (67) 3.5C (176)
Juanacati -4.5a (332) 8.2a (179) 10.5a.b (78) 9.4a.b (31)
Bajo Grande --3.7abc (8) 8.0b (68) 11.0a (49)
Rio Balsas1 3.2" (98) 4.1b (20)
Bongales --4.5a,b,c (9) 8.7a.b (57) 7.2b (36)
Limon --6.8a,b,c (3) 12.0a (34) 12.7a (10)
Rio Tuira --4.6a,b,c (11) 7.2b (144) 5.7b.c (39)
1999-00 1-4 cm 4-10 cm 10-20 cm 20-40 cm 2 40 cm
Casarete 0.5b (483) 1.1b (238) 1.7b (372) 5.0b (226) 3.2C (128)
Sambu 1.3a (427) 2.6a"b (162) 4.2a.b (157) 3.8b (110) 2.3" (104)
Naranzati 1.6a (100) 1.7b (83) 2.6b (66) 5.4b (43) 5.1b (139)
Juanacati -3.0a (338) 5.1a (190) 8.4a (81) 7.8a (39)
2000-01 1-4 cm 4-10 cm 10-20 cm 20-40 cm 2 40 cm
Casarete 0.8a (479) 1.0b (243) 2.1b (355) 3.6b (224) 2.0b (126)
Sambu 1.1a (459) 1.9a (170) 4.0a (169) 3.3b (116) 1.6b (107)
Juanacati -1.3b (332) 2.6b (197) 5.1a (83) 5.2a (40)















Table 4-8. Ingrowth by stem type. Percentage of recruited individuals from broken stems, undamaged stems, or sprouts from
prostrate and inclined trees.


1997-98


1998-99


1999-00


Sprouts from
DBH inclined/prostrate
cm stems
50
1-4 5.5


Sprouts from
inclined/prostrate
stems
63
11
50
11
7
6
0
38
30
0
0
0
0
0
0


Sprouts from
inclined/prostrate
stems
53
0
0
0
21
25
0
20
0
0
0
0
0
0
0


Site
Casarete
Sambu
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati


Broken
0
5.5
0
0
0
50
0
0
0
0
0
0
0
0
0


Undamaged
50
89
100
100
71
50
100
50
100
100
100
100
100
100


Broken
6
4
0
0
0
0
0
0
0
0
0
0
0
0
0


Undamaged
31
85
50
89
93
94
100
62
70
100
0
100
100
100
100


Broken
33
0
20
0
0
6
0
0
0
0
0
0
0
0
0


Undamaged
14
100
80
100
79
69
100
80
100
100
100


4-10



10-
40



> 40











Table 4-8. Continued


2000-01


Sprouts from
DBH inclined/prostrate
cm stems
89
1-4 61


Site
Casarete
Sambu
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati


Broken
2
6




5

-
0
0

0
-
0


Undamaged
9
33


75
89
45


43
86
80


50
100
100


4-10




10-40


Table 4-9. Cativo annual recruitment and mortality rates (%) by stem diameter class for
four census periods.


Recruitment Rates


Mortality Rates
-1998- 1999- 2000-
99 00 01
4.6 2.8 2.1
1.6 1.7 0.6
4.3 3.0-
2.8 0.4 0.7
0 0.5 0
0.4 0 0.7


DBH 1997-
cm 98
1.2
1-4 8.4
3.9


1998-
99
3.0
11.6
1.7
3.6
10.2
5.5
2.1


1999-
00
3.2
3.0
4.8
3.1
6.9
5.2
8.1


2000-
01
10.7
3.7


1997-
98
8.3
1.9
3.9


Site
Casarete
Sambu
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati


4-10


0.3
2.1
10-40
4.8
2.4
0.8
1.7
> 40
7.3
0
















Table 4-10. Annual mortality rates (%) of cativo trees by stem type and stature for four census periods.


1997-98


Prostrate
0.21
0
0.81
0
0.23
0
0
0

1999-2000


1998-99
Sprouts from
inclined/prostrate
stems Pro:
1.16
0.57
0 0


0.13 0
0 0




2000-01
Sprouts from
inclined/prostrate
stems Pro:
0.70
0.45


0 0


Sprouts from
inclined/prostrate
stems
2.51
1.06



0.23


DBH
cm Broken
0.83
0.18
<10
0
3.32
0
0
>10
0
0





Broken
0.29

<10 0
0
0.54


Inclined
0.83
0
0
0
0.46
0.25
0
0


Undamaged
3.56
0.18
1.62
0
0.23
0
0
0


Broken
0.13
0.19
0
1.42

0.13
0
0
0.33


state
0
0
.35
0
.26
.56
0
0


Inclined
0.39
0
0
0
0.13
0
0
0


Undamaged
2.33
0.38
0.70
1.89
0.40
0.28
0
0


Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Nammnati






Casarete
Sambu
Juanacati
Naranzati
Casarete
Sambu
Juanacati
Naranzati


Sprouts from
inclined/prostrate
stems
0.58
0.98


Prostrate
0
0
0


Inclined
0
0
0
0
0.29
0
0
0


Undamaged
1.16
0.42
0
1.63
0.44
0
1.39
0.75


Broken



0.48

-
0
0


state






.12
0
0


Inclined
0.35
0
0


0.35
0
0.49


Undamaged
0.59
0
0.24











Table 4-11. Mean annual growth (mm/year) of cativo trees that were alive at the end of
the study and those that died during the study for which there was one or more
years of growth data.


1997-98 growth
Alive 2001 Dead 2001 df
0.04 (683) -0.09 (44) 65
1.7 (712) 0.3 (17) 25
1.9 (519) 0.5 (14) 16

0.2 (107) 0.6 (9) 9


Site DBH cm
1-10
Casarete
S10
1-10
Sambu
> 10
1-10
Naranzati
> 10
4-10
Juanacati
> 10



Site DBH cm
1-10
Casarete
S10
1-10
Sambu
> 10
1-10
Naranzati
> 10
4-10
Juanacati
S10



Site DBH cm
1-10
Casarete
S10
1-10
Sambu
> 10
1-10
Naranzati
> 10
4-10
Juanacati
> 10


0.05
<0.001
0.002

0.32






p
<0.001
0.01
<0.001


1998-99 growth
Dead 2001 df
0.2 (21) 33
1.6 (13) 13
.03 (12) 14


Alive 2001
0.6 (707)
3.3 (718)
2.2 (545)


8.7 (294) 3.6 (6) 5 2.5 0.025


1999-00 growth
Dead 2001 df
-0.1(10) 11
-0.8 (7) 15


Alive 2001
0.7 (711)
3.6 (722)


t p
4.9 < 0.001
17.2 < 0.001