0121d52b642d2c124c15735f4eb5b2cd remotesensing-18-01588-v2.pdf 1308c57342cb4c45a135c7a11d38607b925fe11c remotesensing-18-01588-v2.pdf c8d7c92daa4cecb70b785180b4740966ed4155f0eb58467d3bd79dbef55eb0cf remotesensing-18-01588-v2.pdf Title: Assessing the Effect of Long-Term Soil Warming on Subarctic Grasslands Using High-Resolution Multispectral Drone Images Subject: Rising temperatures, driven by global climate change, are profoundly altering high-latitude ecosystems, influencing vegetation phenology and productivity. However, understanding the long-term, nuanced responses of these ecosystems remains a critical challenge. Soil warming experiments have served as useful tools for understanding these shifts. However, many of these studies have relied on a single measure, predominantly the Normalized Difference Vegetation (NDVI), measured at a single level of warming. This approach often fails to separate structural greening from underlying physiological responses. To address these gaps, this study provided a comprehensive snapshot assessment of growing season vegetation dynamics in a subarctic grassland ecosystem in Iceland that had been exposed to continuous geothermal soil warming for over 60 years. Using high-resolution multispectral drone imagery, twelve different vegetation indices (VIs) were derived to assess not only greenness but also physiological stress and photosynthetic efficiency across a range of mean annual soil temperatures (MATs). Using linear regression and redundancy analysis (RDA), the responses of these indices to warming and their relationships with other environmental drivers, such as standing biomass and plant nutrient concentrations (nitrogen and phosphorus), were analyzed. The results revealed significant positive linear relationships between most of the indices and MATs across the 5 to 11 °C range. This indicated that higher MATs led to increased biomass and structural growth, without revealing any significant thresholds or tipping points in vegetation response within the observed warming range. However, the Photochemical Reflectance (PRI) showed a significant negative relationship with warming, suggesting a decoupling between structural greening and photosynthetic light-use efficiency. Furthermore, RDA results indicated that, while most of the VIs were primarily driven by biomass, the decline in PRI was likely a compounding effect of physical canopy self-shading and plant phosphorus constraints. Ultimately, this study demonstrated that, while these subarctic grasslands exhibited local evidence of “Arctic greening” under further warming, multispectral drone remote sensing could detect underlying physiological adjustments and nutrient constraints that traditional greenness indices might overlook, providing a more nuanced understanding of ecosystem response. Keywords: soil warming; subarctic grasslands; vegetation indices; climate change; remote sensing; multispectral images; drone Author: Amir Hamedpour, Ruth P. Tchana Wandji, Bjarni D. Sigurdsson, Asra Salimi, Iolanda Filella and Josep Peñuelas Creator: LaTeX with hyperref Producer: pdfTeX-1.40.25; modified using OpenPDF 1.4.2 CreationDate: Thu May 21 08:03:42 2026 CEST ModDate: Thu May 21 08:37:23 2026 CEST Custom Metadata: yes Metadata Stream: no Tagged: no UserProperties: no Suspects: no Form: none JavaScript: no Pages: 22 Encrypted: no Page size: 595.276 x 841.89 pts (A4) Page rot: 0 File size: 20194908 bytes Optimized: no PDF version: 1.7 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- GWGSQX+URWPalladioL-Bold Type 1 Custom yes yes yes 154 0 ZPJUOQ+URWPalladioL-Roma Type 1 Custom yes yes yes 155 0 YSNVNJ+URWPalladioL-Ital Type 1 Custom yes yes yes 156 0 AXWEOY+CMSY10 Type 1 Builtin yes yes yes 283 0 XPMKRN+CMEX10 Type 1 Builtin yes yes yes 350 0 GIGFZE+CMR10 Type 1 Builtin yes yes yes 351 0 ZKGCPY+PazoMath-Italic Type 1 Builtin yes yes yes 371 0 SYFPBV+CMMI10 Type 1 Builtin yes yes yes 372 0 WBVKDJ+URWPalladioL-BoldItal Type 1 Custom yes yes yes 419 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-06-03 03:55:51 CEST RepresentationInformation: remotesensing-18-01588-v2.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-06-02 14:16:01 CEST Size: 20194908 Format: PDF Version: 1.7 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf PDFMetadata: Objects: 671 FreeObjects: 1 IncrementalUpdates: 0 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: true CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseNone Outlines: Item: Title: Introduction Item: Title: Materials and Methods Children: Item: Title: Study Site Item: Title: Data Collection Item: Title: Remote Sensing Indices Item: Title: Measured Ecosystem and Environmental Factors Item: Title: Statistical Analysis Children: Item: Title: Data Aggregation and Pre-Processing Item: Title: Regression Analysis Item: Title: Multivariate Ordination (PCA and RDA) Item: Title: Results Children: Item: Title: Similarity of Vegetation Indices Item: Title: Response of VIs to Soil Warming Item: Title: Redundancy Analysis (RDA) Children: Item: Title: Group 1 (NDVI, SAVI, MSAVI, CIG, GARI) Item: Title: Group 2 (CCI, WDRVI, EVI, NDRE, MCARI) Item: Title: Group 3 (PRI, PSRI) Item: Title: Discussion Children: Item: Title: Response of VIs to MATs Item: Title: Linearity of Vegetation Response to MATs Item: Title: Other Environmental Drivers Item: Title: Methodological Considerations Item: Title: Conclusions Item: Title: Appendix A Item: Title: References Info: Title: Assessing the Effect of Long-Term Soil Warming on Subarctic Grasslands Using High-Resolution Multispectral Drone Images Author: Amir Hamedpour, Ruth P. Tchana Wandji, Bjarni D. Sigurdsson, Asra Salimi, Iolanda Filella and Josep Peñuelas Subject: Rising temperatures, driven by global climate change, are profoundly altering high-latitude ecosystems, influencing vegetation phenology and productivity. However, understanding the long-term, nuanced responses of these ecosystems remains a critical challenge. Soil warming experiments have served as useful tools for understanding these shifts. However, many of these studies have relied on a single measure, predominantly the Normalized Difference Vegetation (NDVI), measured at a single level of warming. This approach often fails to separate structural greening from underlying physiological responses. To address these gaps, this study provided a comprehensive snapshot assessment of growing season vegetation dynamics in a subarctic grassland ecosystem in Iceland that had been exposed to continuous geothermal soil warming for over 60 years. Using high-resolution multispectral drone imagery, twelve different vegetation indices (VIs) were derived to assess not only greenness but also physiological stress and photosynthetic efficiency across a range of mean annual soil temperatures (MATs). Using linear regression and redundancy analysis (RDA), the responses of these indices to warming and their relationships with other environmental drivers, such as standing biomass and plant nutrient concentrations (nitrogen and phosphorus), were analyzed. The results revealed significant positive linear relationships between most of the indices and MATs across the 5 to 11 °C range. This indicated that higher MATs led to increased biomass and structural growth, without revealing any significant thresholds or tipping points in vegetation response within the observed warming range. However, the Photochemical Reflectance (PRI) showed a significant negative relationship with warming, suggesting a decoupling between structural greening and photosynthetic light-use efficiency. Furthermore, RDA results indicated that, while most of the VIs were primarily driven by biomass, the decline in PRI was likely a compounding effect of physical canopy self-shading and plant phosphorus constraints. Ultimately, this study demonstrated that, while these subarctic grasslands exhibited local evidence of “Arctic greening” under further warming, multispectral drone remote sensing could detect underlying physiological adjustments and nutrient constraints that traditional greenness indices might overlook, providing a more nuanced understanding of ecosystem response. Keywords: soil warming; subarctic grasslands; vegetation indices; climate change; remote sensing; multispectral images; drone Creator: LaTeX with hyperref Producer: pdfTeX-1.40.25; modified using OpenPDF 1.4.2 CreationDate: Thu May 21 08:03:42 CEST 2026 ModDate: Thu May 21 08:37:23 CEST 2026 ID: 0x14c1fb66fd655c3f9d0839d5285e96c1, 0xe77732f1a9584a39e6bfdc92e16ec583 Filters: FilterPipeline: FlateDecode Images: Image: NisoImageMetadata: FormatName: image/png ImageWidth: 3408 ImageHeight: 2610 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1114 ImageHeight: 639 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1084 ImageHeight: 584 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 2769 ImageHeight: 2400 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 2700 ImageHeight: 3360 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 7200 ImageHeight: 2400 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 3600 ImageHeight: 1800 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Image: NisoImageMetadata: FormatName: image/png ImageWidth: 2700 ImageHeight: 1800 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Fonts: Type1: Font: BaseFont: ZKGCPY+PazoMath-Italic FontSubset: true FirstChar: 98 LastChar: 121 FontDescriptor: FontName: ZKGCPY+PazoMath-Italic Flags: Symbolic FontBBox: -70, -277, 902, 733 FontFile: true ToUnicode: true Font: BaseFont: WBVKDJ+URWPalladioL-BoldItal FontSubset: true FirstChar: 112 LastChar: 112 FontDescriptor: FontName: WBVKDJ+URWPalladioL-BoldItal Flags: Symbolic FontBBox: -170, -300, 1073, 935 FontFile: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: SYFPBV+CMMI10 FontSubset: true FirstChar: 60 LastChar: 62 FontDescriptor: FontName: SYFPBV+CMMI10 Flags: Symbolic FontBBox: -32, -250, 1048, 750 FontFile: true ToUnicode: true Font: BaseFont: GWGSQX+URWPalladioL-Bold FontSubset: true FirstChar: 2 LastChar: 241 FontDescriptor: FontName: GWGSQX+URWPalladioL-Bold Flags: Symbolic FontBBox: -152, -301, 1000, 935 FontFile: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: ZPJUOQ+URWPalladioL-Roma FontSubset: true FirstChar: 2 LastChar: 253 FontDescriptor: FontName: ZPJUOQ+URWPalladioL-Roma Flags: Symbolic FontBBox: -166, -283, 1021, 943 FontFile: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: AXWEOY+CMSY10 FontSubset: true FirstChar: 0 LastChar: 20 FontDescriptor: FontName: AXWEOY+CMSY10 Flags: Symbolic FontBBox: -29, -960, 1116, 775 FontFile: true ToUnicode: true Font: BaseFont: YSNVNJ+URWPalladioL-Ital FontSubset: true FirstChar: 39 LastChar: 121 FontDescriptor: FontName: YSNVNJ+URWPalladioL-Ital Flags: Symbolic FontBBox: -170, -305, 1010, 941 FontFile: true EncodingDictionary: Differences: true ToUnicode: true Font: BaseFont: XPMKRN+CMEX10 FontSubset: true FirstChar: 0 LastChar: 112 FontDescriptor: FontName: XPMKRN+CMEX10 Flags: Symbolic FontBBox: -24, -2960, 1454, 772 FontFile: true ToUnicode: true Font: BaseFont: GIGFZE+CMR10 FontSubset: true FirstChar: 40 LastChar: 61 FontDescriptor: FontName: GIGFZE+CMR10 Flags: Symbolic FontBBox: -40, -250, 1009, 750 FontFile: true ToUnicode: true Pages: Page: Sequence: 1 Annotations: Annotation: Subtype: Link Rect: 34, 216, 124, 235 Annotation: Subtype: Link Rect: 89, 77, 152, 84 Annotation: Subtype: Link Rect: 34, 64, 120, 72 Annotation: Subtype: Link Rect: 34, 773, 185, 809 Annotation: Subtype: Link Rect: 515, 773, 560, 809 Annotation: Subtype: Link Rect: 134, 663, 145, 677 Annotation: Subtype: Link Rect: 257, 663, 268, 677 Annotation: Subtype: Link Rect: 373, 663, 384, 677 Annotation: Subtype: Link Rect: 453, 663, 464, 677 Annotation: Subtype: Link Rect: 546, 663, 557, 677 Annotation: Subtype: Link Rect: 135, 650, 146, 663 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 2 Annotations: Annotation: Subtype: Link Rect: 467, 440, 474, 449 ActionDest: 118 Annotation: Subtype: Link Rect: 475, 440, 482, 449 ActionDest: 118 Annotation: Subtype: Link Rect: 475, 412, 482, 421 ActionDest: 118 Annotation: Subtype: Link Rect: 482, 412, 489, 421 ActionDest: 118 Annotation: Subtype: Link Rect: 442, 398, 449, 407 ActionDest: 118 Annotation: Subtype: Link Rect: 450, 398, 457, 407 ActionDest: 118 Annotation: Subtype: Link Rect: 513, 357, 520, 365 ActionDest: 118 Annotation: Subtype: Link Rect: 521, 356, 528, 365 ActionDest: 118 Annotation: Subtype: Link Rect: 408, 301, 415, 310 ActionDest: 118 Annotation: Subtype: Link Rect: 416, 301, 428, 310 ActionDest: 118 Annotation: Subtype: Link Rect: 501, 273, 513, 282 ActionDest: 118 Annotation: Subtype: Link Rect: 514, 273, 526, 282 ActionDest: 118 Annotation: Subtype: Link Rect: 270, 245, 282, 254 ActionDest: 118 Annotation: Subtype: Link Rect: 285, 245, 297, 254 ActionDest: 118 Annotation: Subtype: Link Rect: 208, 203, 220, 212 ActionDest: 118 Annotation: Subtype: Link Rect: 221, 203, 233, 212 ActionDest: 118 Annotation: Subtype: Link Rect: 498, 189, 510, 198 ActionDest: 118 Annotation: Subtype: Link Rect: 513, 189, 525, 198 ActionDest: 118 Annotation: Subtype: Link Rect: 297, 161, 309, 170 ActionDest: 118 Annotation: Subtype: Link Rect: 310, 162, 322, 170 ActionDest: 118 Annotation: Subtype: Link Rect: 325, 162, 337, 170 ActionDest: 237 Annotation: Subtype: Link Rect: 328, 147, 340, 157 ActionDest: 118 Annotation: Subtype: Link Rect: 341, 147, 353, 157 ActionDest: 237 Annotation: Subtype: Link Rect: 441, 106, 453, 115 ActionDest: 237 Annotation: Subtype: Link Rect: 454, 106, 466, 115 ActionDest: 237 Annotation: Subtype: Link Rect: 408, 78, 420, 87 ActionDest: 118 Annotation: Subtype: Link Rect: 421, 78, 433, 87 ActionDest: 237 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 3 Annotations: Annotation: Subtype: Link Rect: 217, 758, 229, 767 ActionDest: 237 Annotation: Subtype: Link Rect: 229, 758, 241, 767 ActionDest: 237 Annotation: Subtype: Link Rect: 512, 717, 524, 726 ActionDest: 237 Annotation: Subtype: Link Rect: 527, 717, 539, 726 ActionDest: 237 Annotation: Subtype: Link Rect: 482, 689, 494, 698 ActionDest: 237 Annotation: Subtype: Link Rect: 497, 689, 509, 698 ActionDest: 237 Annotation: Subtype: Link Rect: 493, 675, 505, 684 ActionDest: 237 Annotation: Subtype: Link Rect: 506, 675, 518, 684 ActionDest: 237 Annotation: Subtype: Link Rect: 541, 343, 548, 356 ActionDest: 24 Annotation: Subtype: Link Rect: 262, 288, 269, 300 ActionDest: 24 Annotation: Subtype: Link Rect: 280, 246, 287, 258 ActionDest: 24 Annotation: Subtype: Link Rect: 405, 221, 417, 230 ActionDest: 237 Annotation: Subtype: Link Rect: 544, 193, 556, 202 ActionDest: 237 Annotation: Subtype: Link Rect: 463, 165, 475, 174 ActionDest: 237 Annotation: Subtype: Link Rect: 519, 109, 531, 118 ActionDest: 237 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 4 Annotations: Annotation: Subtype: Link Rect: 289, 728, 296, 740 ActionDest: 24 Annotation: Subtype: Link Rect: 285, 700, 292, 712 ActionDest: 169 Annotation: Subtype: Link Rect: 196, 616, 211, 628 ActionDest: 113 Annotation: Subtype: Link Rect: 484, 616, 499, 628 ActionDest: 113 Annotation: Subtype: Link Rect: 543, 188, 550, 200 ActionDest: 24 Annotation: Subtype: Link Rect: 337, 135, 349, 144 ActionDest: 237 Annotation: Subtype: Link Rect: 349, 134, 361, 144 ActionDest: 294 Annotation: Subtype: Link Rect: 290, 104, 297, 116 ActionDest: 169 Annotation: Subtype: Link Rect: 199, 76, 206, 88 ActionDest: 169 Annotation: Subtype: Link Rect: 414, 76, 421, 88 ActionDest: 29 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 5 Annotations: Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 6 Annotations: Annotation: Subtype: Link Rect: 348, 385, 360, 394 ActionDest: 294 Annotation: Subtype: Link Rect: 361, 385, 373, 394 ActionDest: 294 Annotation: Subtype: Link Rect: 218, 343, 230, 352 ActionDest: 294 Annotation: Subtype: Link Rect: 233, 343, 244, 352 ActionDest: 294 Annotation: Subtype: Link Rect: 254, 232, 266, 241 ActionDest: 294 Annotation: Subtype: Link Rect: 462, 204, 474, 213 ActionDest: 294 Annotation: Subtype: Link Rect: 531, 176, 543, 185 ActionDest: 294 Annotation: Subtype: Link Rect: 544, 176, 556, 185 ActionDest: 294 Annotation: Subtype: Link Rect: 506, 148, 518, 157 ActionDest: 294 Annotation: Subtype: Link Rect: 282, 107, 294, 116 ActionDest: 294 Annotation: Subtype: Link Rect: 461, 79, 473, 88 ActionDest: 294 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 7 Annotations: Annotation: Subtype: Link Rect: 395, 758, 407, 767 ActionDest: 294 Annotation: Subtype: Link Rect: 307, 730, 319, 740 ActionDest: 294 Annotation: Subtype: Link Rect: 367, 689, 379, 698 ActionDest: 237 Annotation: Subtype: Link Rect: 380, 689, 392, 698 ActionDest: 294 Annotation: Subtype: Link Rect: 388, 661, 400, 670 ActionDest: 294 Annotation: Subtype: Link Rect: 353, 633, 365, 642 ActionDest: 294 Annotation: Subtype: Link Rect: 324, 616, 331, 628 ActionDest: 352 Annotation: Subtype: Link Rect: 528, 500, 538, 507 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 474, 538, 482 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 441, 538, 449 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 401, 538, 408 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 372, 538, 380 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 342, 538, 349 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 308, 538, 316 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 275, 538, 283 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 241, 538, 249 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 203, 538, 211 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 165, 538, 173 ActionDest: 294 Annotation: Subtype: Link Rect: 528, 127, 538, 135 ActionDest: 294 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 8 Annotations: Annotation: Subtype: Link Rect: 529, 255, 541, 264 ActionDest: 294 Annotation: Subtype: Link Rect: 342, 241, 354, 250 ActionDest: 294 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 9 Annotations: Annotation: Subtype: Link Rect: 468, 204, 475, 216 ActionDest: 64 Annotation: Subtype: Link Rect: 387, 190, 394, 202 ActionDest: 64 Annotation: Subtype: Link Rect: 515, 78, 522, 90 ActionDest: 64 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 10 Annotations: Annotation: Subtype: Link Rect: 233, 271, 245, 280 ActionDest: 294 Annotation: Subtype: Link Rect: 223, 227, 230, 239 ActionDest: 363 Annotation: Subtype: Link Rect: 456, 171, 463, 183 ActionDest: 363 Annotation: Subtype: Link Rect: 492, 171, 499, 183 ActionDest: 68 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 11 Annotations: Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 12 Annotations: Annotation: Subtype: Link Rect: 365, 730, 372, 742 ActionDest: 363 Annotation: Subtype: Link Rect: 199, 500, 206, 512 ActionDest: 68 Annotation: Subtype: Link Rect: 473, 486, 480, 498 ActionDest: 64 Annotation: Subtype: Link Rect: 343, 119, 350, 131 ActionDest: 68 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 13 Annotations: Annotation: Subtype: Link Rect: 502, 725, 509, 737 ActionDest: 68 Annotation: Subtype: Link Rect: 409, 517, 416, 529 ActionDest: 68 Annotation: Subtype: Link Rect: 544, 338, 551, 350 ActionDest: 68 Annotation: Subtype: Link Rect: 396, 130, 408, 139 ActionDest: 118 Annotation: Subtype: Link Rect: 409, 130, 421, 139 ActionDest: 237 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 14 Annotations: Annotation: Subtype: Link Rect: 322, 673, 329, 686 ActionDest: 363 Annotation: Subtype: Link Rect: 549, 662, 556, 671 ActionDest: 118 Annotation: Subtype: Link Rect: 314, 648, 321, 657 ActionDest: 118 Annotation: Subtype: Link Rect: 322, 648, 334, 657 ActionDest: 118 Annotation: Subtype: Link Rect: 334, 648, 346, 657 ActionDest: 237 Annotation: Subtype: Link Rect: 529, 620, 541, 629 ActionDest: 294 Annotation: Subtype: Link Rect: 544, 620, 556, 629 ActionDest: 294 Annotation: Subtype: Link Rect: 366, 481, 378, 490 ActionDest: 294 Annotation: Subtype: Link Rect: 378, 481, 390, 490 ActionDest: 294 Annotation: Subtype: Link Rect: 324, 467, 336, 476 ActionDest: 118 Annotation: Subtype: Link Rect: 339, 467, 351, 476 ActionDest: 118 Annotation: Subtype: Link Rect: 343, 426, 355, 435 ActionDest: 237 Annotation: Subtype: Link Rect: 257, 395, 264, 409 ActionDest: 68 Annotation: Subtype: Link Rect: 257, 286, 269, 295 ActionDest: 294 Annotation: Subtype: Link Rect: 192, 272, 204, 281 ActionDest: 294 Annotation: Subtype: Link Rect: 204, 272, 216, 281 ActionDest: 294 Annotation: Subtype: Link Rect: 262, 175, 273, 184 ActionDest: 294 Annotation: Subtype: Link Rect: 350, 147, 362, 156 ActionDest: 294 Annotation: Subtype: Link Rect: 415, 105, 427, 114 ActionDest: 294 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 15 Annotations: Annotation: Subtype: Link Rect: 331, 730, 343, 740 ActionDest: 294 Annotation: Subtype: Link Rect: 544, 661, 556, 670 ActionDest: 294 Annotation: Subtype: Link Rect: 426, 633, 438, 642 ActionDest: 237 Annotation: Subtype: Link Rect: 260, 495, 272, 504 ActionDest: 294 Annotation: Subtype: Link Rect: 387, 467, 399, 476 ActionDest: 484 Annotation: Subtype: Link Rect: 403, 439, 415, 448 ActionDest: 237 Annotation: Subtype: Link Rect: 298, 367, 313, 379 ActionDest: 91 Annotation: Subtype: Link Rect: 421, 342, 433, 351 ActionDest: 484 Annotation: Subtype: Link Rect: 498, 300, 510, 309 ActionDest: 118 Annotation: Subtype: Link Rect: 292, 216, 304, 226 ActionDest: 118 Annotation: Subtype: Link Rect: 305, 216, 317, 226 ActionDest: 237 Annotation: Subtype: Link Rect: 544, 175, 556, 184 ActionDest: 237 Annotation: Subtype: Link Rect: 370, 133, 382, 142 ActionDest: 118 Annotation: Subtype: Link Rect: 382, 133, 394, 142 ActionDest: 237 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 16 Annotations: Annotation: Subtype: Link Rect: 197, 683, 204, 695 ActionDest: 68 Annotation: Subtype: Link Rect: 506, 672, 518, 681 ActionDest: 294 Annotation: Subtype: Link Rect: 518, 672, 530, 681 ActionDest: 294 Annotation: Subtype: Link Rect: 241, 547, 253, 556 ActionDest: 484 Annotation: Subtype: Link Rect: 334, 546, 346, 556 ActionDest: 484 Annotation: Subtype: Link Rect: 219, 463, 231, 472 ActionDest: 294 Annotation: Subtype: Link Rect: 231, 463, 243, 472 ActionDest: 484 Annotation: Subtype: Link Rect: 326, 279, 341, 291 ActionDest: 91 Annotation: Subtype: Link Rect: 505, 254, 517, 263 ActionDest: 294 Annotation: Subtype: Link Rect: 399, 172, 411, 181 ActionDest: 237 Annotation: Subtype: Link Rect: 412, 172, 424, 181 ActionDest: 237 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 17 Annotations: Annotation: Subtype: Link Rect: 471, 758, 483, 767 ActionDest: 118 Annotation: Subtype: Link Rect: 484, 758, 496, 767 ActionDest: 237 Annotation: Subtype: Link Rect: 297, 744, 309, 754 ActionDest: 484 Annotation: Subtype: Link Rect: 310, 744, 322, 754 ActionDest: 484 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 18 Annotations: Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 19 Annotations: Annotation: Subtype: Link Rect: 59, 716, 96, 726 Annotation: Subtype: Link Rect: 102, 716, 138, 726 Annotation: Subtype: Link Rect: 230, 689, 268, 700 Annotation: Subtype: Link Rect: 359, 651, 396, 662 Annotation: Subtype: Link Rect: 402, 651, 439, 662 Annotation: Subtype: Link Rect: 146, 626, 183, 636 Annotation: Subtype: Link Rect: 187, 601, 224, 611 Annotation: Subtype: Link Rect: 445, 574, 482, 585 Annotation: Subtype: Link Rect: 354, 550, 391, 560 Annotation: Subtype: Link Rect: 423, 523, 461, 534 Annotation: Subtype: Link Rect: 411, 498, 448, 509 Annotation: Subtype: Link Rect: 59, 460, 96, 470 Annotation: Subtype: Link Rect: 467, 434, 504, 445 Annotation: Subtype: Link Rect: 510, 434, 546, 445 Annotation: Subtype: Link Rect: 276, 396, 313, 407 Annotation: Subtype: Link Rect: 337, 370, 375, 381 Annotation: Subtype: Link Rect: 472, 345, 510, 356 Annotation: Subtype: Link Rect: 264, 319, 301, 330 Annotation: Subtype: Link Rect: 59, 282, 96, 292 Annotation: Subtype: Link Rect: 385, 255, 423, 266 Annotation: Subtype: Link Rect: 148, 218, 185, 228 Annotation: Subtype: Link Rect: 191, 218, 227, 228 Annotation: Subtype: Link Rect: 59, 179, 96, 190 Annotation: Subtype: Link Rect: 102, 179, 138, 190 Annotation: Subtype: Link Rect: 458, 153, 496, 164 Annotation: Subtype: Link Rect: 392, 115, 430, 126 Annotation: Subtype: Link Rect: 436, 115, 472, 126 Annotation: Subtype: Link Rect: 403, 89, 440, 100 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 20 Annotations: Annotation: Subtype: Link Rect: 104, 731, 142, 742 Annotation: Subtype: Link Rect: 181, 693, 218, 703 Annotation: Subtype: Link Rect: 59, 655, 96, 665 Annotation: Subtype: Link Rect: 135, 629, 173, 639 Annotation: Subtype: Link Rect: 135, 591, 173, 601 Annotation: Subtype: Link Rect: 406, 565, 443, 576 Annotation: Subtype: Link Rect: 59, 527, 96, 537 Annotation: Subtype: Link Rect: 118, 489, 155, 499 Annotation: Subtype: Link Rect: 318, 462, 356, 473 Annotation: Subtype: Link Rect: 279, 438, 316, 448 Annotation: Subtype: Link Rect: 323, 438, 359, 448 Annotation: Subtype: Link Rect: 169, 412, 207, 422 Annotation: Subtype: Link Rect: 367, 373, 404, 384 Annotation: Subtype: Link Rect: 253, 347, 291, 358 Annotation: Subtype: Link Rect: 297, 347, 333, 358 Annotation: Subtype: Link Rect: 392, 309, 429, 320 Annotation: Subtype: Link Rect: 520, 284, 557, 295 Annotation: Subtype: Link Rect: 329, 258, 366, 269 Annotation: Subtype: Link Rect: 311, 220, 348, 231 Annotation: Subtype: Link Rect: 226, 194, 263, 205 Annotation: Subtype: Link Rect: 309, 169, 346, 180 Annotation: Subtype: Link Rect: 87, 143, 165, 154 Annotation: Subtype: Link Rect: 337, 105, 375, 116 Annotation: Subtype: Link Rect: 370, 79, 408, 90 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 21 Annotations: Annotation: Subtype: Link Rect: 95, 731, 132, 742 Annotation: Subtype: Link Rect: 425, 718, 463, 729 Annotation: Subtype: Link Rect: 316, 679, 354, 690 Annotation: Subtype: Link Rect: 208, 655, 245, 665 Annotation: Subtype: Link Rect: 462, 628, 499, 639 Annotation: Subtype: Link Rect: 319, 603, 356, 614 Annotation: Subtype: Link Rect: 217, 578, 254, 588 Annotation: Subtype: Link Rect: 59, 553, 96, 563 Annotation: Subtype: Link Rect: 189, 526, 227, 537 Annotation: Subtype: Link Rect: 233, 526, 269, 537 Annotation: Subtype: Link Rect: 108, 502, 146, 512 Annotation: Subtype: Link Rect: 421, 488, 458, 499 Annotation: Subtype: Link Rect: 59, 450, 96, 461 Annotation: Subtype: Link Rect: 102, 450, 138, 461 Annotation: Subtype: Link Rect: 273, 425, 310, 435 Annotation: Subtype: Link Rect: 59, 387, 96, 397 Annotation: Subtype: Link Rect: 331, 360, 369, 371 Annotation: Subtype: Link Rect: 116, 323, 153, 333 Annotation: Subtype: Link Rect: 409, 284, 446, 295 Annotation: Subtype: Link Rect: 59, 259, 96, 269 Annotation: Subtype: Link Rect: 356, 220, 393, 231 Annotation: Subtype: Link Rect: 428, 194, 466, 205 Annotation: Subtype: Link Rect: 337, 169, 374, 180 Annotation: Subtype: Link Rect: 117, 144, 155, 154 Annotation: Subtype: Link Rect: 487, 92, 524, 103 Annotation: Subtype: Link Rect: 193, 67, 230, 78 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Page: Sequence: 22 Annotations: Annotation: Subtype: Link Rect: 59, 731, 96, 742 Annotation: Subtype: Link Rect: 108, 706, 146, 716 Annotation: Subtype: Link Rect: 258, 667, 295, 678 Annotation: Subtype: Link Rect: 461, 641, 498, 652 Annotation: Subtype: Link Rect: 491, 616, 528, 627 Annotation: Subtype: Link Rect: 468, 590, 505, 601 Annotation: Subtype: Link Rect: 429, 27, 560, 37 Checksum: d1336346 Type: CRC32 Checksum: 0121d52b642d2c124c15735f4eb5b2cd Type: MD5 Checksum: 1308c57342cb4c45a135c7a11d38607b925fe11c Type: SHA-1 Checksum: c8d7c92daa4cecb70b785180b4740966ed4155f0eb58467d3bd79dbef55eb0cf Type: SHA-256