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         packageId="knb-lter-hfr.332.4">
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   <dataset id="HF332">
      <alternateIdentifier system="https://doi.org">doi:10.6073/pasta/af85078378dc3f0b4e82d4630a78d393</alternateIdentifier>
      <title>Species Distribution Modeling of Carnivorous Plants Worldwide</title>
      <creator>
         <individualName>
            <givenName>Aaron</givenName>
            <surName>Ellison</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0003-4151-6081</userId>
      </creator>
      <creator>
         <individualName>
            <givenName>Matt</givenName>
            <surName>Fitzpatrick</surName>
         </individualName>
      </creator>
      <creator>
         <individualName>
            <givenName>Nicholas</givenName>
            <surName>Gotelli</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0002-5409-7456</userId>
      </creator>
      <pubDate>2023</pubDate>
      <language>English</language>
      <abstract>
         <section>
            <title/>
            <para>Forecasting how carnivorous plant species will respond to climatic change is a key issue in their conservation and management but presents a number of challenges. These challenges derive from interactions between the relatively simplistic statistical methods typically used to forecast species responses to climatic change, which to date have been limited mainly to species distribution models (“SDMs) and particular aspects of the ecology of carnivorous plants, including their rarity, habitat specialization, and limited dispersal ability. The small ranges and oftentimes low local abundance of carnivorous plants provide few occurrence records, which increase the potential for poorly or over-fitted SDMs and misspecification of relationships with their “optimal” environments. The unique habitats in which carnivorous plants often grow also are difficult to characterize using the basic temperature and precipitation data that often undergird SDMs. Rather, habitats in which carnivorous plants are common often are decoupled from broader climatic patterns (e.g., many retain high soil moisture even during seasonal drought) and may be associated with frequent disturbance. Last, dispersal limitation also may constrain range shifts of carnivorous plants as the climate changes. These three issues raise two related questions that are critical for understanding and forecasting the future of carnivorous plants. First, to what extent are current carnivorous plants distributions constrained by climate; and second, how readily, if at all, might carnivorous plants disperse to colonize new habitat as it becomes climatically suitable?</para>
         </section>
         <section>
            <title/>
            <para>We estimated the vulnerability of carnivorous plants to climatic change in light of challenges identified with SDMs in general and their particular application to these unique species. We combined two approaches: “ensembles of small models”, which attempt to deal with the challenges of fitting SDMs for data-limited species; and “bioclimatic velocity”, which is an estimate of how fast a species would have to migrate to track its climatic niche, to provide initial assessments of the vulnerability of carnivorous plants to climatic change. We also explored, for one carnivorous plant (Sarracenia purpurea) how to additionally incorporate demographic complexity into its SDM.</para>
         </section>
      </abstract>
      <keywordSet>
         <keyword>carnivorous plants</keyword>
         <keyword>climate change</keyword>
         <keyword>demography</keyword>
         <keyword>dispersal</keyword>
         <keyword>distribution</keyword>
         <keyword>modeling</keyword>
         <keywordThesaurus>LTER controlled vocabulary</keywordThesaurus>
      </keywordSet>
      <keywordSet>
         <keyword>populations</keyword>
         <keywordThesaurus>LTER core area</keywordThesaurus>
      </keywordSet>
      <keywordSet>
         <keyword>Harvard Forest</keyword>
         <keyword>HFR</keyword>
         <keyword>LTER</keyword>
         <keyword>USA</keyword>
         <keywordThesaurus>HFR default</keywordThesaurus>
      </keywordSet>
      <intellectualRights>
         <section>
            <para>This dataset is released to the public under Creative Commons CC0 1.0 (No Rights Reserved). Please keep the dataset creators informed of any plans to use the dataset. Consultation with the original investigators is strongly encouraged. Publications and data products that make use of the dataset should include proper acknowledgement.</para>
         </section>
      </intellectualRights>
      <licensed>
         <licenseName>Creative Commons Zero v1.0 Universal</licenseName>
         <url>https://spdx.org/licenses/CC0-1.0.html</url>
         <identifier>CC0-1.0</identifier>
      </licensed>
      <distribution>
         <online>
            <url function="information">https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf332</url>
         </online>
      </distribution>
      <coverage>
         <geographicCoverage>
            <geographicDescription>Global. Coordinates based on WGS84 datum.</geographicDescription>
            <boundingCoordinates>
               <westBoundingCoordinate>-180</westBoundingCoordinate>
               <eastBoundingCoordinate>+180</eastBoundingCoordinate>
               <northBoundingCoordinate>+90</northBoundingCoordinate>
               <southBoundingCoordinate>-90</southBoundingCoordinate>
            </boundingCoordinates>
         </geographicCoverage>
         <taxonomicCoverage>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Aldrovanda</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Brocchinia</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Byblis</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Catopsis</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Cephalotus</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Darlingtonia</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Dionaea</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Drosera</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Drosophyllum</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Genlisea</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Heliamphora</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Ibicella</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Nepenthes</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Pinguicula</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Sarracenia</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Sarracenia</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>purpurea</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Triphyophyllum</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
            <taxonomicClassification>
               <taxonRankName>genus</taxonRankName>
               <taxonRankValue>Utricularia</taxonRankValue>
               <taxonomicClassification>
                  <taxonRankName>species</taxonRankName>
                  <taxonRankValue>sp.</taxonRankValue>
               </taxonomicClassification>
            </taxonomicClassification>
         </taxonomicCoverage>
      </coverage>
      <maintenance>
         <description>
            <para>complete</para>
         </description>
      </maintenance>
      <contact scope="document">
         <positionName>Information Manager</positionName>
         <organizationName>Harvard Forest</organizationName>
         <address scope="document">
            <deliveryPoint>324 North Main Street</deliveryPoint>
            <city>Petersham</city>
            <administrativeArea>MA</administrativeArea>
            <postalCode>01366</postalCode>
            <country>USA</country>
         </address>
         <phone phonetype="voice">(978) 724-3302</phone>
         <electronicMailAddress>hf-im@lists.fas.harvard.edu</electronicMailAddress>
      </contact>
      <publisher scope="document">
         <organizationName>Harvard Forest</organizationName>
         <address scope="document">
            <deliveryPoint>324 North Main Street</deliveryPoint>
            <city>Petersham</city>
            <administrativeArea>MA</administrativeArea>
            <postalCode>01366</postalCode>
            <country>USA</country>
         </address>
         <phone phonetype="voice">(978) 724-3302</phone>
         <phone phonetype="fax">(978) 724-3595</phone>
         <onlineUrl>https://harvardforest.fas.harvard.edu</onlineUrl>
      </publisher>
      <methods>
         <methodStep>
            <description>
               <section>
                  <title/>
                  <para>For our initial SDMs of carnivorous plants worldwide, we assembled a comprehensive database of occurrence records of carnivorous from multiple sources, including the Global Biodiversity Information Facility (GBIF), the Australia Virtual Herbarium, and numerous regional herbaria. We applied four quality control measures to the occurrence records before using them for modeling: [1] updated and standardized species names to the latest taxonomy and to identify hybrids, which were removed from further consideration; [2] species records without geographic (e.g., latitude–longitude) coordinates but with collection location information were georeferenced using GEOLocate 2.0; [3] removed occurrence records with [a] low spatial precision; [b] whose coordinates were far outside the known range; [c] whose coordinates matched the centroids of political boundaries; [d] or whose coordinates fell within botanical gardens or museums; [4] removed all spatial replicates (i.e., records for the same species with identical coordinates). The archived dataset contains those occurrence records that fell within the known native distribution of each species and which had sufficient spatial precision given the resolution of the climate data (WorldClim)— maximum temperature of the warmest month, mean temperature of the coldest quarter, annual precipitation, and precipitation seasonality—at 2.5 arc-minute spatial resolution.</para>
               </section>
               <section>
                  <title/>
                  <para>For scenarios of future climates, we used decadal averages of these same four bioclimatic variables for 2050 from 32 future climate simulations statistically downscaled to 2.5 arc-minute spatial resolution. The simulations included output from numerous general circulation models and one representative concentration pathway (RCP 8.5) developed as part of the IPCC AR5 and available from the Research Program on Climate Change, Agriculture and Food Security.1 28.5.3</para>
               </section>
               <section>
                  <title/>
                  <para>We modeled habitat suitability for carnivorous plants using MaxEnt version 3.3.3E as implemented in the dismo package in R. We fit models using the default values for all settings. We fit MaxEnt models (default values for all settings and optimized for model complexity) for any species with at least 10 spatially unique occurrence records and constrained selection of background data (points selected at random from the study region and also used to inform model fitting) to those areas within any terrestrial ecoregion in which the species has been observed and immediately adjacent ecoregions, as defined using the World Wildlife Fund’s definitions of terrestrial ecoregions. We partitioned the occurrence data for each species randomly 10 times into calibration (70%) and evaluation (30%) datasets, and models were run on each of the 10 resulting datasets. The multiple models for each species resulting from different random splits of the occurrence data into training and test partitions were combined into a single ensemble by weighted averaging.</para>
               </section>
               <section>
                  <title/>
                  <para>For more detailed SDMs of Sarracenia purpurea, we augmented the distribution records in hf332-01-carnivorous-plant-data.csv with additional records collected by Hannah Buckley (HF193) and data collected by Aaron Ellison and Nick Gotelli throughout New England (HF159).</para>
               </section>
            </description>
         </methodStep>
      </methods>
      <project>
         <title>Harvard Forest Long-Term Ecological Research</title>
         <personnel>
            <organizationName>Harvard Forest</organizationName>
            <address>
               <deliveryPoint>324 North Main Street</deliveryPoint>
               <city>Petersham</city>
               <administrativeArea>MA</administrativeArea>
               <postalCode>01366</postalCode>
               <country>USA</country>
            </address>
            <phone phonetype="voice">(978) 724-3302</phone>
            <phone phonetype="fax">(978) 724-3595</phone>
            <onlineUrl>https://harvardforest.fas.harvard.edu</onlineUrl>
            <userId directory="https://ror.org">https://ror.org/059cpzx98</userId>
            <role>pointOfContact</role>
         </personnel>
         <abstract>The Harvard Forest Long-Term Ecological Research (LTER) program examines ecological dynamics in the New England region resulting from natural disturbances, environmental change, and human impacts.</abstract>
         <funding>National Science Foundation LTER grants: DEB-8811764, DEB-9411975, DEB-0080592, DEB-0620443, DEB-1237491, DEB-1832210.</funding>
      </project>
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         <entityName>hf332-01-carnivorous-plant-data.csv</entityName>
         <entityDescription>all carnivorous plant records</entityDescription>
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            <objectName>hf332-01-carnivorous-plant-data.csv</objectName>
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            </dataFormat>
            <distribution>
               <online>
                  <url function="download">https://harvardforest.fas.harvard.edu/data/p33/hf332/hf332-01-carnivorous-plant-data.csv</url>
               </online>
            </distribution>
         </physical>
         <attributeList>
            <attribute id="1571756687277">
               <attributeName>id</attributeName>
               <attributeDefinition>unique record identifier</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <textDomain>
                           <definition>unique record identifier</definition>
                        </textDomain>
                     </nonNumericDomain>
                  </nominal>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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            </attribute>
            <attribute id="1571756687293">
               <attributeName>family</attributeName>
               <attributeDefinition>plant family</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <textDomain>
                           <definition>plant family</definition>
                        </textDomain>
                     </nonNumericDomain>
                  </nominal>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756687308">
               <attributeName>genus</attributeName>
               <attributeDefinition>plant genus</attributeDefinition>
               <measurementScale>
                  <nominal>
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                        <textDomain>
                           <definition>plant genus</definition>
                        </textDomain>
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                  <codeExplanation>missing value</codeExplanation>
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               <attributeName>species</attributeName>
               <attributeDefinition>plant species</attributeDefinition>
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                        <textDomain>
                           <definition>plant species</definition>
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                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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               <attributeName>study.name</attributeName>
               <attributeDefinition>study name</attributeDefinition>
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                           <definition>study name</definition>
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               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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            </attribute>
            <attribute id="1571756687355">
               <attributeName>range</attributeName>
               <attributeDefinition>inside or outside of known range of the taxon</attributeDefinition>
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                        <enumeratedDomain>
                           <codeDefinition>
                              <code>inside</code>
                              <definition>inside of known range</definition>
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                           <codeDefinition>
                              <code>outside</code>
                              <definition>outside of known range</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>unknown</code>
                              <definition>unknown</definition>
                           </codeDefinition>
                        </enumeratedDomain>
                     </nonNumericDomain>
                  </nominal>
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               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756687371">
               <attributeName>num.precision</attributeName>
               <attributeDefinition>spatial precision of location</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>meter</standardUnit>
                     </unit>
                     <precision>0.01</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756687386">
               <attributeName>longitude</attributeName>
               <attributeDefinition>longitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>degree</standardUnit>
                     </unit>
                     <precision>0.0000001</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756687402">
               <attributeName>latitude</attributeName>
               <attributeDefinition>latitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>degree</standardUnit>
                     </unit>
                     <precision>0.00000001</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
         </attributeList>
         <numberOfRecords>92617</numberOfRecords>
      </dataTable>
      <dataTable id="hf332-02">
         <entityName>hf332-02-sp-data.csv</entityName>
         <entityDescription>Sarracenia purpurea records only</entityDescription>
         <physical>
            <objectName>hf332-02-sp-data.csv</objectName>
            <size unit="byte">13851</size>
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               <textFormat>
                  <numHeaderLines>1</numHeaderLines>
                  <recordDelimiter>\r\n</recordDelimiter>
                  <attributeOrientation>column</attributeOrientation>
                  <simpleDelimited>
                     <fieldDelimiter>,</fieldDelimiter>
                  </simpleDelimited>
               </textFormat>
            </dataFormat>
            <distribution>
               <online>
                  <url function="download">https://harvardforest.fas.harvard.edu/data/p33/hf332/hf332-02-sp-data.csv</url>
               </online>
            </distribution>
         </physical>
         <attributeList>
            <attribute id="1571756848233">
               <attributeName>source</attributeName>
               <attributeDefinition>source</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <enumeratedDomain>
                           <codeDefinition>
                              <code>Buckley</code>
                              <definition>from HF193</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>GBIF</code>
                              <definition>from hf332-01-carnivorous-plant-data.csv</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>MAPP</code>
                              <definition>from HF159</definition>
                           </codeDefinition>
                        </enumeratedDomain>
                     </nonNumericDomain>
                  </nominal>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756848248">
               <attributeName>lat</attributeName>
               <attributeDefinition>latitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>degree</standardUnit>
                     </unit>
                     <precision>0.01</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute id="1571756848264">
               <attributeName>long</attributeName>
               <attributeDefinition>longitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>degree</standardUnit>
                     </unit>
                     <precision>0.01</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
         </attributeList>
         <numberOfRecords>738</numberOfRecords>
      </dataTable>
      <otherEntity id="hf332-03">
         <entityName>hf332-03-dist-modeling-function.txt</entityName>
         <entityDescription>distribution modeling function R script</entityDescription>
         <physical>
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