e3d46f23629592aee2f0671b2f864c63 pmc_34822376.pdf ff7ca37e51f3961595ffe9cca62d19a4c4fbfaa3 pmc_34822376.pdf 12abaa610a2ddc4716a17797dc244f363c73ddaa8ec38ce4bf9f562e42db8140 pmc_34822376.pdf Title: Tree Species and Epiphyte Taxa Determine the “Metabolomic niche” of Canopy Suspended Soils in a Species-Rich Lowland Tropical Rainforest Subject: Tropical forests are biodiversity hotspots, but it is not well understood how this diversity is structured and maintained. One hypothesis rests on the generation of a range of metabolic niches, with varied composition, supporting a high species diversity. Characterizing soil metabolomes can reveal fine-scale differences in composition and potentially help explain variation across these habitats. In particular, little is known about canopy soils, which are unique habitats that are likely to be sources of additional biodiversity and biogeochemical cycling in tropical forests. We studied the effects of diverse tree species and epiphytes on soil metabolomic profiles of forest floor and canopy suspended soils in a French Guianese rainforest. We found that the metabolomic profiles of canopy suspended soils were distinct from those of forest floor soils, differing between epiphyte-associated and non-epiphyte suspended soils, and the metabolomic profiles of suspended soils varied with host tree species, regardless of association with epiphyte. Thus, tree species is a key driver of rainforest suspended soil metabolomics. We found greater abundance of metabolites in suspended soils, particularly in groups associated with plants, such as phenolic compounds, and with metabolic pathways related to amino acids, nucleotides, and energy metabolism, due to the greater relative proportion of tree and epiphyte organic material derived from litter and root exudates, indicating a strong legacy of parent biological material. Our study provides evidence for the role of tree and epiphyte species in canopy soil metabolomic composition and in maintaining the high levels of soil metabolome diversity in this tropical rainforest. It is likely that a wide array of canopy microsite-level environmental conditions, which reflect interactions between trees and epiphytes, increase the microscale diversity in suspended soil metabolomes. Keywords: bacteria; canopy soils; epiphyte; French Guiana; metabolomics Author: Albert Gargallo-Garriga, Jordi Sardans, Abdulwahed Fahad Alrefaei, Karel Klem, Lucia Fuchslueger, Irene Ramírez-Rojas, Julian Donald, Celine Leroy, Leandro Van Langenhove, Erik Verbruggen, Ivan A. Janssens, Otmar Urban and Josep Peñuelas Creator: LaTeX with hyperref Producer: pdfTeX-1.40.21 CreationDate: Thu Oct 21 12:43:45 2021 CEST ModDate: Thu Oct 21 13:59:53 2021 CEST Tagged: no UserProperties: no Suspects: no Form: none JavaScript: no Pages: 16 Encrypted: no Page size: 595.276 x 841.89 pts (A4) Page rot: 0 File size: 9185050 bytes Optimized: no PDF version: 1.7 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- ACKGZT+VnURWPalladioL Type 1 Custom yes yes no 10 0 EZKMGU+URWPalladioL-Ital Type 1 Custom yes yes no 15 0 PNLRTO+URWPalladioL-Bold Type 1 Custom yes yes no 20 0 DOZPIO+URWPalladioL-BoldItal Type 1 Custom yes yes no 24 0 YTHWDY+URWPalladioL-Roma Type 1 Custom yes yes no 28 0 WLENKM+VnURWPalladioL-Bold Type 1 Custom yes yes no 32 0 ZZPUEX+ArialMT Type 1C WinAnsi yes yes no 41 0 TWGUDY+TimesNewRomanPS-BoldItalicMT Type 1C WinAnsi yes yes no 44 0 MCBCCI+SourceSansRoman-Regular Type 1C Custom yes yes no 66 0 MCBCCM+SourceSansRoman.613wght Type 1C Custom yes yes no 70 0 MCBCCO+SourceSansRoman-Bold Type 1C Custom yes yes no 74 0 YFFKLH+CMSY10 Type 1 Builtin yes yes no 89 0 ZXDKIP+EURM10 Type 1 Builtin yes yes no 97 0 Jhove (Rel. 1.22.1, 2019-04-17) Date: 2022-03-22 02:33:44 CET RepresentationInformation: pmc_34822376.pdf ReportingModule: PDF-hul, Rel. 1.12.1 (2019-04-17) LastModified: 2022-01-10 08:12:55 CET Size: 9185050 Format: PDF Version: 1.7 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf PDFMetadata: Objects: 452 FreeObjects: 13 IncrementalUpdates: 0 DocumentCatalog: PageLayout: SinglePage PageMode: UseNone Outlines: Item: Title: Introduction Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@7b69c6ba Item: Title: Results Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@46daef40 Item: Title: Discussion Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@12f41634 Children: Item: Title: Metabolome of Forest Floor and Canopy Soils Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@13c27452 Item: Title: Metabolic Profile Differences among Canopy Soils Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@262b2c86 Item: Title: Global Overview of the High Diversity of Metabolomic Profile in the Studied Soils Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@371a67ec Item: Title: Materials and Methods Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@5ed828d Children: Item: Title: Study Area Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@50d0686 Item: Title: Soil Sampling and Analysis Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@7a3d45bd Item: Title: Metabolite Extraction and Analysis Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@1e7c7811 Item: Title: Metabolite Identification and Quantification Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@77ec78b9 Item: Title: Statistical Analysis Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@1a3869f4 Item: Title: Conclusions Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@a38d7a3 Item: Title: References Destination: edu.harvard.hul.ois.jhove.module.pdf.PdfSimpleObject@77f99a05 Info: Title: Tree Species and Epiphyte Taxa Determine the “Metabolomic niche” of Canopy Suspended Soils in a Species-Rich Lowland Tropical Rainforest Author: Albert Gargallo-Garriga, Jordi Sardans, Abdulwahed Fahad Alrefaei, Karel Klem, Lucia Fuchslueger, Irene Ramírez-Rojas, Julian Donald, Celine Leroy, Leandro Van Langenhove, Erik Verbruggen, Ivan A. Janssens, Otmar Urban and Josep Peñuelas Subject: Tropical forests are biodiversity hotspots, but it is not well understood how this diversity is structured and maintained. One hypothesis rests on the generation of a range of metabolic niches, with varied composition, supporting a high species diversity. Characterizing soil metabolomes can reveal fine-scale differences in composition and potentially help explain variation across these habitats. In particular, little is known about canopy soils, which are unique habitats that are likely to be sources of additional biodiversity and biogeochemical cycling in tropical forests. We studied the effects of diverse tree species and epiphytes on soil metabolomic profiles of forest floor and canopy suspended soils in a French Guianese rainforest. We found that the metabolomic profiles of canopy suspended soils were distinct from those of forest floor soils, differing between epiphyte-associated and non-epiphyte suspended soils, and the metabolomic profiles of suspended soils varied with host tree species, regardless of association with epiphyte. Thus, tree species is a key driver of rainforest suspended soil metabolomics. We found greater abundance of metabolites in suspended soils, particularly in groups associated with plants, such as phenolic compounds, and with metabolic pathways related to amino acids, nucleotides, and energy metabolism, due to the greater relative proportion of tree and epiphyte organic material derived from litter and root exudates, indicating a strong legacy of parent biological material. Our study provides evidence for the role of tree and epiphyte species in canopy soil metabolomic composition and in maintaining the high levels of soil metabolome diversity in this tropical rainforest. It is likely that a wide array of canopy microsite-level environmental conditions, which reflect interactions between trees and epiphytes, increase the microscale diversity in suspended soil metabolomes. Keywords: bacteria; canopy soils; epiphyte; French Guiana; metabolomics Creator: LaTeX with hyperref Producer: pdfTeX-1.40.21 CreationDate: Thu Oct 21 12:43:45 CEST 2021 ModDate: Thu Oct 21 13:59:53 CEST 2021 ID: 0x9bbb1ccf65031613251bfe7c50d2aa90, 0x9bbb1ccf65031613251bfe7c50d2aa90 Filters: FilterPipeline: FlateDecode Fonts: Type1: Font: BaseFont: WLENKM+VnURWPalladioL-Bold FontSubset: true FirstChar: 189 LastChar: 189 FontDescriptor: FontName: WLENKM+VnURWPalladioL-Bold Flags: Symbolic FontBBox: -152, -301, 1000, 935 FontFile: true EncodingDictionary: Differences: true Font: BaseFont: ZXDKIP+EURM10 FontSubset: true FirstChar: 11 LastChar: 22 FontDescriptor: FontName: ZXDKIP+EURM10 Flags: Symbolic FontBBox: -32, -243, 1060, 720 FontFile: true Font: BaseFont: PNLRTO+URWPalladioL-Bold FontSubset: true FirstChar: 3 LastChar: 241 FontDescriptor: FontName: PNLRTO+URWPalladioL-Bold Flags: Symbolic FontBBox: -152, -301, 1000, 935 FontFile: true EncodingDictionary: Differences: true Font: BaseFont: DOZPIO+URWPalladioL-BoldItal FontSubset: true FirstChar: 77 LastChar: 116 FontDescriptor: FontName: DOZPIO+URWPalladioL-BoldItal Flags: Symbolic FontBBox: -170, -300, 1073, 935 FontFile: true EncodingDictionary: Differences: true Font: BaseFont: YFFKLH+CMSY10 FontSubset: true FirstChar: 0 LastChar: 48 FontDescriptor: FontName: YFFKLH+CMSY10 Flags: Symbolic FontBBox: -29, -960, 1116, 775 FontFile: true Font: BaseFont: ACKGZT+VnURWPalladioL FontSubset: true FirstChar: 160 LastChar: 243 FontDescriptor: FontName: ACKGZT+VnURWPalladioL Flags: Symbolic FontBBox: -166, -283, 1021, 943 FontFile: true EncodingDictionary: Differences: true Font: BaseFont: YTHWDY+URWPalladioL-Roma FontSubset: true FirstChar: 2 LastChar: 252 FontDescriptor: FontName: YTHWDY+URWPalladioL-Roma Flags: Symbolic FontBBox: -166, -283, 1021, 943 FontFile: true EncodingDictionary: Differences: true Font: BaseFont: EZKMGU+URWPalladioL-Ital FontSubset: true FirstChar: 2 LastChar: 122 FontDescriptor: FontName: EZKMGU+URWPalladioL-Ital Flags: Symbolic FontBBox: -170, -305, 1010, 941 FontFile: true EncodingDictionary: Differences: true Pages: Page: Sequence: 1 Annotations: Annotation: Subtype: Link Rect: 35, 759, 166, 795 Annotation: Subtype: Link Rect: 515, 759, 560, 795 Annotation: Subtype: Link Rect: 258, 641, 269, 654 Annotation: Subtype: Link Rect: 408, 641, 419, 654 Annotation: Subtype: Link Rect: 481, 641, 492, 654 Annotation: Subtype: Link Rect: 276, 616, 287, 629 Annotation: Subtype: Link Rect: 389, 616, 400, 629 Annotation: Subtype: Link Rect: 113, 358, 154, 367 Annotation: Subtype: Link Rect: 35, 346, 141, 355 Annotation: Subtype: Link Rect: 35, 162, 97, 184 Annotation: Subtype: Link Rect: 153, 89, 154, 98 Annotation: Subtype: Link Rect: 35, 77, 147, 86 Annotation: Subtype: Link Rect: 122, 17, 273, 28 Annotation: Subtype: Link Rect: 394, 17, 558, 28 Annotation: Subtype: Link Rect: 35, 482, 87, 501 Page: Sequence: 2 Annotations: Annotation: Subtype: Link Rect: 441, 721, 448, 730 ActionDest: 168 Annotation: Subtype: Link Rect: 451, 721, 458, 730 ActionDest: 168 Annotation: Subtype: Link Rect: 193, 709, 200, 718 ActionDest: 168 Annotation: Subtype: Link Rect: 201, 708, 208, 718 ActionDest: 168 Annotation: Subtype: Link Rect: 221, 696, 228, 705 ActionDest: 168 Annotation: Subtype: Link Rect: 229, 696, 236, 705 ActionDest: 168 Annotation: Subtype: Link Rect: 235, 683, 242, 692 ActionDest: 168 Annotation: Subtype: Link Rect: 374, 683, 381, 692 ActionDest: 168 Annotation: Subtype: Link Rect: 382, 683, 389, 692 ActionDest: 171 Annotation: Subtype: Link Rect: 392, 683, 404, 692 ActionDest: 171 Annotation: Subtype: Link Rect: 517, 683, 529, 692 ActionDest: 171 Annotation: Subtype: Link Rect: 249, 671, 261, 680 ActionDest: 171 Annotation: Subtype: Link Rect: 423, 671, 435, 680 ActionDest: 171 Annotation: Subtype: Link Rect: 370, 658, 381, 667 ActionDest: 171 Annotation: Subtype: Link Rect: 359, 633, 371, 642 ActionDest: 171 Annotation: Subtype: Link Rect: 258, 608, 270, 617 ActionDest: 171 Annotation: Subtype: Link Rect: 487, 570, 499, 579 ActionDest: 171 Annotation: Subtype: Link Rect: 502, 571, 514, 579 ActionDest: 171 Annotation: Subtype: Link Rect: 251, 533, 263, 542 ActionDest: 171 Annotation: Subtype: Link Rect: 532, 533, 544, 542 ActionDest: 171 Annotation: Subtype: Link Rect: 545, 533, 557, 542 ActionDest: 171 Annotation: Subtype: Link Rect: 504, 520, 516, 529 ActionDest: 171 Annotation: Subtype: Link Rect: 349, 495, 361, 504 ActionDest: 171 Annotation: Subtype: Link Rect: 265, 483, 277, 492 ActionDest: 171 Annotation: Subtype: Link Rect: 369, 457, 381, 466 ActionDest: 171 Annotation: Subtype: Link Rect: 382, 457, 394, 466 ActionDest: 171 Annotation: Subtype: Link Rect: 394, 457, 406, 466 ActionDest: 171 Annotation: Subtype: Link Rect: 448, 445, 460, 454 ActionDest: 171 Annotation: Subtype: Link Rect: 461, 432, 473, 441 ActionDest: 171 Annotation: Subtype: Link Rect: 473, 432, 485, 441 ActionDest: 171 Annotation: Subtype: Link Rect: 494, 420, 506, 429 ActionDest: 171 Annotation: Subtype: Link Rect: 393, 382, 405, 391 ActionDest: 171 Annotation: Subtype: Link Rect: 406, 382, 418, 391 ActionDest: 171 Annotation: Subtype: Link Rect: 418, 382, 430, 391 ActionDest: 171 Annotation: Subtype: Link Rect: 531, 370, 543, 379 ActionDest: 171 Annotation: Subtype: Link Rect: 544, 369, 556, 379 ActionDest: 171 Annotation: Subtype: Link Rect: 531, 344, 543, 353 ActionDest: 174 Annotation: Subtype: Link Rect: 544, 344, 556, 353 ActionDest: 174 Annotation: Subtype: Link Rect: 307, 282, 319, 291 ActionDest: 174 Annotation: Subtype: Link Rect: 304, 244, 316, 253 ActionDest: 174 Annotation: Subtype: Link Rect: 237, 232, 249, 241 ActionDest: 174 Annotation: Subtype: Link Rect: 499, 206, 511, 215 ActionDest: 174 Annotation: Subtype: Link Rect: 412, 181, 424, 190 ActionDest: 174 Annotation: Subtype: Link Rect: 378, 169, 389, 178 ActionDest: 174 Annotation: Subtype: Link Rect: 390, 169, 402, 178 ActionDest: 174 Annotation: Subtype: Link Rect: 217, 131, 229, 140 ActionDest: 174 Annotation: Subtype: Link Rect: 229, 131, 241, 140 ActionDest: 174 Annotation: Subtype: Link Rect: 242, 131, 254, 140 ActionDest: 174 Annotation: Subtype: Link Rect: 543, 118, 555, 128 ActionDest: 174 Annotation: Subtype: Link Rect: 306, 93, 318, 102 ActionDest: 174 Annotation: Subtype: Link Rect: 319, 93, 331, 102 ActionDest: 174 Annotation: Subtype: Link Rect: 471, 94, 483, 102 ActionDest: 171 Page: Sequence: 3 Annotations: Annotation: Subtype: Link Rect: 203, 735, 215, 744 ActionDest: 171 Annotation: Subtype: Link Rect: 215, 735, 227, 744 ActionDest: 174 Annotation: Subtype: Link Rect: 228, 735, 240, 744 ActionDest: 174 Annotation: Subtype: Link Rect: 358, 697, 370, 706 ActionDest: 171 Annotation: Subtype: Link Rect: 318, 609, 330, 618 ActionDest: 174 Annotation: Subtype: Link Rect: 498, 609, 510, 618 ActionDest: 171 Annotation: Subtype: Link Rect: 492, 557, 499, 568 ActionDest: 86 Annotation: Subtype: Link Rect: 519, 557, 526, 568 ActionDest: 86 Annotation: Subtype: Link Rect: 329, 531, 336, 543 ActionDest: 93 Annotation: Subtype: Link Rect: 434, 531, 441, 543 ActionDest: 108 Annotation: Subtype: Link Rect: 299, 518, 306, 530 ActionDest: 108 Annotation: Subtype: Link Rect: 326, 518, 333, 530 ActionDest: 126 Annotation: Subtype: Link Rect: 266, 418, 273, 430 ActionDest: 93 Annotation: Subtype: Link Rect: 285, 393, 292, 405 ActionDest: 108 Page: Sequence: 4 Page: Sequence: 5 Annotations: Annotation: Subtype: Link Rect: 349, 434, 355, 445 ActionDest: 86 Annotation: Subtype: Link Rect: 214, 420, 221, 432 ActionDest: 93 Annotation: Subtype: Link Rect: 472, 64, 479, 77 ActionDest: 93 Page: Sequence: 6 Annotations: Annotation: Subtype: Link Rect: 517, 691, 524, 703 ActionDest: 126 Annotation: Subtype: Link Rect: 544, 691, 551, 703 ActionDest: 144 Page: Sequence: 7 Annotations: Annotation: Subtype: Link Rect: 496, 411, 508, 420 ActionDest: 174 Annotation: Subtype: Link Rect: 465, 348, 477, 357 ActionDest: 174 Annotation: Subtype: Link Rect: 225, 273, 237, 282 ActionDest: 171 Annotation: Subtype: Link Rect: 319, 223, 331, 232 ActionDest: 174 Annotation: Subtype: Link Rect: 462, 172, 474, 181 ActionDest: 174 Annotation: Subtype: Link Rect: 223, 135, 235, 144 ActionDest: 174 Annotation: Subtype: Link Rect: 332, 122, 344, 131 ActionDest: 174 Annotation: Subtype: Link Rect: 524, 97, 536, 106 ActionDest: 171 Annotation: Subtype: Link Rect: 522, 72, 534, 81 ActionDest: 174 Page: Sequence: 8 Annotations: Annotation: Subtype: Link Rect: 244, 735, 255, 744 ActionDest: 171 Annotation: Subtype: Link Rect: 358, 722, 365, 731 ActionDest: 171 Annotation: Subtype: Link Rect: 366, 722, 378, 731 ActionDest: 171 Annotation: Subtype: Link Rect: 378, 722, 390, 731 ActionDest: 174 Annotation: Subtype: Link Rect: 427, 697, 439, 706 ActionDest: 174 Annotation: Subtype: Link Rect: 439, 697, 451, 706 ActionDest: 174 Annotation: Subtype: Link Rect: 236, 659, 248, 669 ActionDest: 174 Annotation: Subtype: Link Rect: 324, 659, 336, 669 ActionDest: 174 Annotation: Subtype: Link Rect: 222, 609, 234, 618 ActionDest: 174 Annotation: Subtype: Link Rect: 235, 609, 247, 618 ActionDest: 177 Annotation: Subtype: Link Rect: 516, 609, 528, 618 ActionDest: 177 Annotation: Subtype: Link Rect: 336, 597, 348, 606 ActionDest: 171 Annotation: Subtype: Link Rect: 349, 597, 361, 606 ActionDest: 177 Annotation: Subtype: Link Rect: 361, 597, 373, 606 ActionDest: 177 Annotation: Subtype: Link Rect: 487, 597, 498, 606 ActionDest: 177 Annotation: Subtype: Link Rect: 543, 597, 555, 606 ActionDest: 177 Annotation: Subtype: Link Rect: 353, 584, 365, 593 ActionDest: 177 Annotation: Subtype: Link Rect: 368, 584, 379, 593 ActionDest: 177 Annotation: Subtype: Link Rect: 497, 508, 509, 518 ActionDest: 174 Annotation: Subtype: Link Rect: 287, 446, 299, 455 ActionDest: 177 Annotation: Subtype: Link Rect: 317, 433, 329, 442 ActionDest: 174 Annotation: Subtype: Link Rect: 330, 433, 342, 442 ActionDest: 177 Annotation: Subtype: Link Rect: 342, 433, 354, 442 ActionDest: 177 Annotation: Subtype: Link Rect: 288, 408, 300, 417 ActionDest: 177 Annotation: Subtype: Link Rect: 303, 408, 315, 417 ActionDest: 177 Annotation: Subtype: Link Rect: 384, 370, 396, 380 ActionDest: 171 Annotation: Subtype: Link Rect: 373, 333, 385, 342 ActionDest: 177 Annotation: Subtype: Link Rect: 408, 270, 420, 279 ActionDest: 174 Annotation: Subtype: Link Rect: 295, 107, 302, 116 ActionDest: 171 Annotation: Subtype: Link Rect: 302, 107, 314, 116 ActionDest: 171 Annotation: Subtype: Link Rect: 315, 107, 327, 116 ActionDest: 171 Annotation: Subtype: Link Rect: 327, 106, 339, 116 ActionDest: 177 Annotation: Subtype: Link Rect: 419, 69, 431, 78 ActionDest: 171 Annotation: Subtype: Link Rect: 432, 69, 444, 78 ActionDest: 174 Annotation: Subtype: Link Rect: 444, 69, 456, 78 ActionDest: 174 Annotation: Subtype: Link Rect: 457, 69, 469, 78 ActionDest: 174 Page: Sequence: 9 Annotations: Annotation: Subtype: Link Rect: 375, 735, 387, 744 ActionDest: 174 Annotation: Subtype: Link Rect: 387, 735, 399, 744 ActionDest: 174 Annotation: Subtype: Link Rect: 325, 570, 337, 579 ActionDest: 177 Annotation: Subtype: Link Rect: 509, 533, 521, 542 ActionDest: 177 Annotation: Subtype: Link Rect: 522, 533, 534, 542 ActionDest: 177 Annotation: Subtype: Link Rect: 257, 508, 269, 517 ActionDest: 177 Annotation: Subtype: Link Rect: 270, 508, 282, 517 ActionDest: 177 Annotation: Subtype: Link Rect: 224, 482, 236, 492 ActionDest: 177 Annotation: Subtype: Link Rect: 438, 482, 450, 492 ActionDest: 177 Annotation: Subtype: Link Rect: 327, 457, 339, 466 ActionDest: 177 Annotation: Subtype: Link Rect: 286, 417, 293, 429 ActionDest: 164 Annotation: Subtype: Link Rect: 383, 294, 395, 303 ActionDest: 180 Annotation: Subtype: Link Rect: 449, 168, 461, 177 ActionDest: 180 Page: Sequence: 10 Page: Sequence: 11 Annotations: Annotation: Subtype: Link Rect: 431, 312, 443, 321 ActionDest: 180 Annotation: Subtype: Link Rect: 443, 312, 455, 321 ActionDest: 180 Annotation: Subtype: Link Rect: 379, 224, 391, 233 ActionDest: 180 Annotation: Subtype: Link Rect: 544, 161, 556, 170 ActionDest: 180 Annotation: Subtype: Link Rect: 453, 58, 559, 70 Page: Sequence: 12 Annotations: Annotation: Subtype: Link Rect: 424, 570, 560, 581 Annotation: Subtype: Link Rect: 166, 558, 282, 569 Annotation: Subtype: Link Rect: 429, 215, 466, 226 Annotation: Subtype: Link Rect: 472, 215, 508, 226 Annotation: Subtype: Link Rect: 520, 192, 557, 203 Annotation: Subtype: Link Rect: 59, 181, 95, 192 Annotation: Subtype: Link Rect: 520, 169, 557, 180 Annotation: Subtype: Link Rect: 59, 158, 95, 169 Annotation: Subtype: Link Rect: 172, 135, 209, 146 Annotation: Subtype: Link Rect: 215, 135, 251, 146 Annotation: Subtype: Link Rect: 316, 112, 353, 123 Annotation: Subtype: Link Rect: 254, 89, 292, 100 Annotation: Subtype: Link Rect: 298, 89, 334, 100 Page: Sequence: 13 Annotations: Annotation: Subtype: Link Rect: 95, 710, 132, 720 Annotation: Subtype: Link Rect: 162, 686, 199, 697 Annotation: Subtype: Link Rect: 277, 663, 314, 674 Annotation: Subtype: Link Rect: 401, 640, 439, 651 Annotation: Subtype: Link Rect: 369, 617, 406, 628 Annotation: Subtype: Link Rect: 59, 584, 97, 594 Annotation: Subtype: Link Rect: 280, 560, 318, 571 Annotation: Subtype: Link Rect: 488, 537, 525, 548 Annotation: Subtype: Link Rect: 152, 515, 189, 525 Annotation: Subtype: Link Rect: 165, 492, 203, 502 Annotation: Subtype: Link Rect: 334, 468, 371, 479 Annotation: Subtype: Link Rect: 520, 433, 557, 444 Annotation: Subtype: Link Rect: 59, 423, 95, 433 Annotation: Subtype: Link Rect: 163, 400, 201, 410 Annotation: Subtype: Link Rect: 328, 364, 366, 375 Annotation: Subtype: Link Rect: 271, 341, 308, 352 Annotation: Subtype: Link Rect: 422, 318, 459, 329 Annotation: Subtype: Link Rect: 331, 295, 368, 306 Annotation: Subtype: Link Rect: 283, 261, 321, 272 Annotation: Subtype: Link Rect: 95, 227, 133, 237 Annotation: Subtype: Link Rect: 478, 215, 515, 226 Annotation: Subtype: Link Rect: 226, 192, 264, 203 Annotation: Subtype: Link Rect: 320, 169, 357, 180 Annotation: Subtype: Link Rect: 520, 157, 557, 168 Annotation: Subtype: Link Rect: 510, 146, 548, 157 Annotation: Subtype: Link Rect: 446, 123, 484, 134 Annotation: Subtype: Link Rect: 519, 111, 557, 122 Annotation: Subtype: Link Rect: 519, 100, 557, 111 Annotation: Subtype: Link Rect: 59, 89, 95, 99 Annotation: Subtype: Link Rect: 195, 66, 232, 76 Page: Sequence: 14 Annotations: Annotation: Subtype: Link Rect: 212, 722, 249, 732 Annotation: Subtype: Link Rect: 278, 698, 315, 709 Annotation: Subtype: Link Rect: 476, 675, 513, 686 Annotation: Subtype: Link Rect: 185, 652, 222, 663 Annotation: Subtype: Link Rect: 483, 640, 521, 651 Annotation: Subtype: Link Rect: 471, 629, 509, 640 Annotation: Subtype: Link Rect: 415, 594, 453, 605 Annotation: Subtype: Link Rect: 512, 571, 550, 582 Annotation: Subtype: Link Rect: 432, 548, 469, 559 Annotation: Subtype: Link Rect: 91, 503, 128, 513 Annotation: Subtype: Link Rect: 263, 445, 300, 456 Annotation: Subtype: Link Rect: 519, 422, 557, 433 Annotation: Subtype: Link Rect: 434, 399, 472, 410 Annotation: Subtype: Link Rect: 441, 376, 478, 387 Annotation: Subtype: Link Rect: 250, 330, 288, 341 Annotation: Subtype: Link Rect: 137, 295, 174, 306 Annotation: Subtype: Link Rect: 317, 272, 354, 283 Annotation: Subtype: Link Rect: 373, 249, 410, 260 Annotation: Subtype: Link Rect: 173, 226, 210, 237 Annotation: Subtype: Link Rect: 113, 204, 150, 214 Annotation: Subtype: Link Rect: 174, 180, 212, 191 Annotation: Subtype: Link Rect: 136, 158, 173, 168 Annotation: Subtype: Link Rect: 104, 123, 141, 134 Annotation: Subtype: Link Rect: 492, 100, 530, 111 Annotation: Subtype: Link Rect: 288, 77, 326, 88 Page: Sequence: 15 Annotations: Annotation: Subtype: Link Rect: 131, 722, 169, 732 Annotation: Subtype: Link Rect: 175, 722, 211, 732 Annotation: Subtype: Link Rect: 152, 699, 189, 709 Annotation: Subtype: Link Rect: 418, 675, 456, 686 Annotation: Subtype: Link Rect: 59, 641, 97, 651 Annotation: Subtype: Link Rect: 152, 618, 189, 628 Annotation: Subtype: Link Rect: 320, 594, 357, 605 Annotation: Subtype: Link Rect: 353, 572, 391, 582 Annotation: Subtype: Link Rect: 415, 548, 453, 559 Annotation: Subtype: Link Rect: 59, 515, 97, 525 Annotation: Subtype: Link Rect: 297, 491, 335, 502 Annotation: Subtype: Link Rect: 136, 469, 173, 479 Annotation: Subtype: Link Rect: 293, 445, 331, 456 Annotation: Subtype: Link Rect: 337, 445, 373, 456 Annotation: Subtype: Link Rect: 209, 410, 246, 421 Annotation: Subtype: Link Rect: 253, 410, 289, 421 Annotation: Subtype: Link Rect: 520, 387, 557, 398 Annotation: Subtype: Link Rect: 512, 364, 550, 375 Annotation: Subtype: Link Rect: 339, 341, 377, 352 Annotation: Subtype: Link Rect: 265, 318, 302, 329 Annotation: Subtype: Link Rect: 294, 295, 332, 306 Annotation: Subtype: Link Rect: 520, 272, 557, 283 Annotation: Subtype: Link Rect: 172, 249, 209, 260 Annotation: Subtype: Link Rect: 184, 192, 221, 203 Annotation: Subtype: Link Rect: 363, 169, 400, 180 Annotation: Subtype: Link Rect: 59, 146, 97, 157 Annotation: Subtype: Link Rect: 188, 112, 225, 122 Annotation: Subtype: Link Rect: 344, 66, 381, 76 Page: Sequence: 16 Annotations: Annotation: Subtype: Link Rect: 489, 721, 526, 732 Annotation: Subtype: Link Rect: 202, 687, 240, 697 Annotation: Subtype: Link Rect: 297, 663, 335, 674 Annotation: Subtype: Link Rect: 265, 640, 303, 651 Checksum: 59306068 Type: CRC32 Checksum: e3d46f23629592aee2f0671b2f864c63 Type: MD5 Checksum: ff7ca37e51f3961595ffe9cca62d19a4c4fbfaa3 Type: SHA-1