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   <dataset id="HF442">
      <alternateIdentifier system="https://doi.org">doi:10.6073/pasta/b26a7d03402dac0daff0e8f3232bac1d</alternateIdentifier>
      <title>Using Phenology to Forecast Species Distributions across the Eastern United States in the 2070s</title>
      <creator>
         <individualName>
            <givenName>Shijia</givenName>
            <surName>Peng</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0002-3510-119X</userId>
      </creator>
      <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>Charles</givenName>
            <surName>Davis</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0001-8747-1101</userId>
      </creator>
      <associatedParty>
         <individualName>
            <givenName>Isaac</givenName>
            <surName>Park</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0001-5539-1641</userId>
         <role>Researcher</role>
      </associatedParty>
      <associatedParty>
         <individualName>
            <givenName>Tadeo</givenName>
            <surName>Ramirez-Parada</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0002-9900-8373</userId>
         <role>Researcher</role>
      </associatedParty>
      <associatedParty>
         <individualName>
            <givenName>Susan</givenName>
            <surName>Mazer</surName>
         </individualName>
         <role>Researcher</role>
      </associatedParty>
      <associatedParty>
         <individualName>
            <givenName>Sydne</givenName>
            <surName>Record</surName>
         </individualName>
         <userId directory="https://orcid.org">https://orcid.org/0000-0001-7293-2155</userId>
         <role>Researcher</role>
      </associatedParty>
      <pubDate>2023</pubDate>
      <language>English</language>
      <abstract>
         <section>
            <para>Studies that use species distribution models (SDMs) to document the relationship between species’ geographic range and environmental conditions rarely consider functional traits, such as phenology, that strongly affect species’ demography and fitness. Using more than 120,000 herbarium specimens representing 360 plant species across the eastern United States, we created a novel “phenology-informed” SDM that integrates dynamic phenological responses to changing climates. Compared to standard SDMs based only on abiotic variables, our phenology-informed SDMs forecast significantly lower species habitat loss, and less species turnover within communities under climate change. These results suggest that phenotypic plasticity and/or local adaptation in phenology may help many species adjust their ecological niches and persist in their habitats during periods of rapid environmental change. By modeling historical data that link phenology, climate and species distributions, our findings reveal how species’ reproductive phenology mediates their geographic distributions along environmental gradients and affect regional biodiversity patterns under future climate changes. More importantly, our newly developed model also circumvents the need for mechanistic models that explicitly link traits to occurrences for each species, and could thus facilitate the deployment of trait-based SDMs across unprecedented spatial and taxonomic scales.  </para>
         </section>
      </abstract>
      <keywordSet>
         <keyword>climate change</keyword>
         <keyword>distribution</keyword>
         <keyword>modeling</keyword>
         <keyword>phenology</keyword>
         <keyword>plant species</keyword>
         <keyword>species diversity</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=hf442</url>
         </online>
      </distribution>
      <coverage>
         <geographicCoverage>
            <geographicDescription>Eastern United States. Coordinates based on WGS84 datum.</geographicDescription>
            <boundingCoordinates>
               <westBoundingCoordinate>-87.53</westBoundingCoordinate>
               <eastBoundingCoordinate>-66.97</eastBoundingCoordinate>
               <northBoundingCoordinate>+47.31</northBoundingCoordinate>
               <southBoundingCoordinate>+24.55</southBoundingCoordinate>
               <boundingAltitudes>
                  <altitudeMinimum>2.78</altitudeMinimum>
                  <altitudeMaximum>1113.89</altitudeMaximum>
                  <altitudeUnits>meter</altitudeUnits>
               </boundingAltitudes>
            </boundingCoordinates>
         </geographicCoverage>
         <temporalCoverage>
            <rangeOfDates>
               <beginDate>
                  <calendarDate>1895</calendarDate>
               </beginDate>
               <endDate>
                  <calendarDate>2018</calendarDate>
               </endDate>
            </rangeOfDates>
         </temporalCoverage>
      </coverage>
      <maintenance>
         <description>
            <para>complete</para>
         </description>
      </maintenance>
      <contact>
         <positionName>Information Manager</positionName>
         <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>
         <electronicMailAddress>hf-im@lists.fas.harvard.edu</electronicMailAddress>
      </contact>
      <publisher>
         <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>
      </publisher>
      <methods>
         <methodStep>
            <description>
               <section>
                  <title>Species selection and phenological data collection</title>
                  <para>We used digitized specimens from two of the most comprehensive digitized regional floras in the world: the Consortium of Northeastern Herbaria (CNH, https://neherbaria.org /portal/) and the Southeast Regional Network of Expertise and Collections (SERNEC, https://sernecportal.org/portal/). These two online portals include more than six million digitized herbarium specimens from across the eastern United States. Following previously established protocols, we selected species and specimens for analysis when i) there were at least 50 unique collections of the species across space and time in the eastern United States, ii) the specimens included both an exact collection date and at least county-level location information, and iii) the specimens had easily identifiable and quantifiable buds, flowers, and fruits. Applying these criteria yielded 124,847 total herbarium specimens representing 360 species that we used in our models and analyses. These species vary in life history, growth form and native status (i.e., native or not to the eastern United States). Information on species’ growth form and native status were derived from the United States Department of Agriculture PLANTS Database (https://plants.isda.gov/). Our final database included 57 herbaceous annuals, 224 herbaceous perennials, 12 vine perennials, 67 woody species, of which 314 are native and 46 are introduced.</para>
               </section>
               <section>
                  <title>Phenological data extraction</title>
                  <para>Crowdsourcers hired through Amazon’s Mechanical Turk service (MTurk; https://www.mturk.com/) counted the number of each type of structure (i.e., buds, flowers, and fruits) present on the specimen using the CrowdCurio platform. Each specimen was scored independently by at least three crowdsourcers. To estimate the reliability of each crowdsourcer and their data, each 10-image set scored by one person included nine unique images and one duplicate image randomly selected from the other nine. The nine images plus the duplicate image were presented to each crowdsourcer in a random order. We then calculated a reliability score for each crowdsourcer based on the data for each 10-image set. The absolute differences between the two duplicate specimens in counts of buds, flowers, or fruits, were separately divided by the total counts for each phenological state, and then subtracted from 1.</para>
                  <para>The reliability score ranged from 0 (unreliable) to 1 (reliable). Specimen observations scored by crowdsourcers with a reliability score of zero were excluded from the analysis; if an individual received a reliability score equal to zero for one phenological phase, the reliability scores for that same individual’s counts of the other phenological states were also set to zero (and similarly excluded from the analyses). Our final dataset included a total of 393,610 phenological observations spanning 124 years (1895–2018).</para>
                  <para>We calculated the median number of buds, flowers, and fruits from the multiple phenological observations for each specimen. We then estimated the day of year (DOY) of “peak” budding, flowering, and fruiting if their buds, open flowers, or fruits separately represented ≥ 50% of the total number of reproductive structures. Two species each were excluded from the “peak budding” category (Primula mistassinica and Sarracenia rubra) and the “peak fruiting” category (Calopogon barbatus and Narcissus poeticus) because there were no specimens of these species at these developmental stages. Of the original 124,847 herbarium specimens, 29,094 specimens were scored as being at peak budding (representing 358 species), 55,767 specimens at peak flowering (representing 360 species), and 37,071 specimens at peak fruiting (representing 358 species).</para>
               </section>
               <section>
                  <title>Species distribution data</title>
                  <para>County-level species’ distribution data were obtained from the Biota of North America Program’s (BONAP; http://www.bonap.org/) North American Plant Atlas (NAPA), which included 19,039 taxa from 227 families (as of November 2022). These data are available as binary occurrences (i.e., presence/absence) for 3,067 counties in the USA, excluding Alaska and Hawai’i. We supplemented the species’ distribution records from BONAP using additional data from the GBIF database (https://www.gbif.org) and all available specimen records in the CNH and SERNEC database. Most (≈ 70%) of these additional data points (i.e., GBIF+NH+SERNEC) represented specimens collected between 1960 and 2000.</para>
                  <para>Since the median area of all counties across the eastern United States is 1221 km2, we divided the contiguous eastern United States into 40 × 40-km (1600-km2) grid cells using the Mercator projection (EPSG 3857). After removing incomplete grid cells (those less than 800 km2) within national borders and coasts, a total of 1,158 grid cells remained. We then overlaid grid cells with county-layer species-occurrence data. A species was deemed “present” in a grid cell if more than half of the cell’s area was covered by the county-level distribution of that species. If a grid cell included parts of several counties in which a species occurred, the total coverage of this grid cell was defined as the sum of all intersected parts of those counties.</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. Other funding: NSF grants DEB-1754584 and DEB-2242804.</funding>
      </project>
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         <entityDescription>growth form</entityDescription>
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                  <url function="download">https://harvardforest.fas.harvard.edu/data/p44/hf442/hf442-01-growth-form.csv</url>
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         </physical>
         <attributeList>
            <attribute>
               <attributeName>Species</attributeName>
               <attributeDefinition>genus and species</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <textDomain>
                           <definition>identifier</definition>
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            <attribute>
               <attributeName>Growth_Form</attributeName>
               <attributeDefinition>growth form</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <enumeratedDomain>
                           <codeDefinition>
                              <code>Herbaceous annual</code>
                              <definition>herbaceous annual</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>Herbaceous perennial</code>
                              <definition>herbaceous perennial</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>Vine perennial</code>
                              <definition>vine perennial</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>Woody</code>
                              <definition>woody</definition>
                           </codeDefinition>
                        </enumeratedDomain>
                     </nonNumericDomain>
                  </nominal>
               </measurementScale>
            </attribute>
            <attribute>
               <attributeName>Status</attributeName>
               <attributeDefinition>status</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <enumeratedDomain>
                           <codeDefinition>
                              <code>Introduced</code>
                              <definition>introduced</definition>
                           </codeDefinition>
                           <codeDefinition>
                              <code>Native</code>
                              <definition>native</definition>
                           </codeDefinition>
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            </attribute>
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         <numberOfRecords>360</numberOfRecords>
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               <online>
                  <url function="download">https://harvardforest.fas.harvard.edu/data/p44/hf442/hf442-02-grid40km.csv</url>
               </online>
            </distribution>
         </physical>
         <attributeList>
            <attribute>
               <attributeName>Gridcode</attributeName>
               <attributeDefinition>grid code</attributeDefinition>
               <measurementScale>
                  <nominal>
                     <nonNumericDomain>
                        <textDomain>
                           <definition>identifier</definition>
                        </textDomain>
                     </nonNumericDomain>
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            </attribute>
            <attribute>
               <attributeName>longitude</attributeName>
               <attributeDefinition>longitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>dimensionless</standardUnit>
                     </unit>
                     <precision>1e-05</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute>
               <attributeName>latitude</attributeName>
               <attributeDefinition>latitude</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>dimensionless</standardUnit>
                     </unit>
                     <precision>1e-05</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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         </attributeList>
         <numberOfRecords>1158</numberOfRecords>
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         <entityName>hf442-03-access-cm2.csv</entityName>
         <entityDescription>future climate data, ACCESS-CM2 model</entityDescription>
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                  <url function="download">https://harvardforest.fas.harvard.edu/data/p44/hf442/hf442-03-access-cm2.csv</url>
               </online>
            </distribution>
         </physical>
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            <attribute>
               <attributeName>Gridcode</attributeName>
               <attributeDefinition>grid code</attributeDefinition>
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                     <nonNumericDomain>
                        <textDomain>
                           <definition>identifier</definition>
                        </textDomain>
                     </nonNumericDomain>
                  </nominal>
               </measurementScale>
            </attribute>
            <attribute>
               <attributeName>bio1</attributeName>
               <attributeDefinition>mean annual temperature</attributeDefinition>
               <measurementScale>
                  <interval>
                     <unit>
                        <standardUnit>celsius</standardUnit>
                     </unit>
                     <precision>0.001</precision>
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               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute>
               <attributeName>bio2</attributeName>
               <attributeDefinition>mean diurnal range</attributeDefinition>
               <measurementScale>
                  <interval>
                     <unit>
                        <standardUnit>celsius</standardUnit>
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                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
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               <attributeName>bio3</attributeName>
               <attributeDefinition>isothermality</attributeDefinition>
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                  <ratio>
                     <unit>
                        <standardUnit>dimensionless</standardUnit>
                     </unit>
                     <precision>0.001</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </ratio>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute>
               <attributeName>bio4</attributeName>
               <attributeDefinition>temperature seasonality</attributeDefinition>
               <measurementScale>
                  <ratio>
                     <unit>
                        <standardUnit>dimensionless</standardUnit>
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               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute>
               <attributeName>bio5</attributeName>
               <attributeDefinition>maximum temperature of warmest month</attributeDefinition>
               <measurementScale>
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                     <unit>
                        <standardUnit>celsius</standardUnit>
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                     <numericDomain>
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               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
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               <attributeName>bio6</attributeName>
               <attributeDefinition>maximum temperature of coldest month</attributeDefinition>
               <measurementScale>
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                     <unit>
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                     </unit>
                     <precision>0.001</precision>
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                  </interval>
               </measurementScale>
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                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
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               <attributeName>bio7</attributeName>
               <attributeDefinition>temperature annual range</attributeDefinition>
               <measurementScale>
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                        <standardUnit>celsius</standardUnit>
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                        <numberType>real</numberType>
                     </numericDomain>
                  </interval>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
            </attribute>
            <attribute>
               <attributeName>bio8</attributeName>
               <attributeDefinition>mean temperature of wettest quarter</attributeDefinition>
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                        <standardUnit>celsius</standardUnit>
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               </measurementScale>
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               </measurementScale>
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                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
               </missingValueCode>
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               <attributeName>bio10</attributeName>
               <attributeDefinition>mean temperature of warmest quarter</attributeDefinition>
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                        <standardUnit>celsius</standardUnit>
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                     <precision>0.001</precision>
                     <numericDomain>
                        <numberType>real</numberType>
                     </numericDomain>
                  </interval>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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            </attribute>
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               <attributeName>bio11</attributeName>
               <attributeDefinition>mean temperature of coldest quarter</attributeDefinition>
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                        <numberType>real</numberType>
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                  </interval>
               </measurementScale>
               <missingValueCode>
                  <code>NA</code>
                  <codeExplanation>missing value</codeExplanation>
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            </attribute>
            <attribute>
               <attributeName>bio12</attributeName>
               <attributeDefinition>annual precipitation</attributeDefinition>
               <measurementScale>
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            <attribute>
               <attributeName>Dicentra_canadensis</attributeName>
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            <attribute>
               <attributeName>Dicentra_cucullaria</attributeName>
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            </attribute>
            <attribute>
               <attributeName>Erythronium_americanum</attributeName>
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            </attribute>
            <attribute>
               <attributeName>Erythronium_umbilicatum</attributeName>
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            <attribute>
               <attributeName>Halesia_carolina</attributeName>
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            <attribute>
               <attributeName>Halesia_diptera</attributeName>
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            <attribute>
               <attributeName>Halesia_tetraptera</attributeName>
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            <attribute>
               <attributeName>Helenium_amarum</attributeName>
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            <attribute>
               <attributeName>Helenium_autumnale</attributeName>
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            </attribute>
            <attribute>
               <attributeName>Helenium_flexuosum</attributeName>
               <attributeDefinition>presence or absence of this species</attributeDefinition>
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            <attribute>
               <attributeName>Helenium_pinnatifidum</attributeName>
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            <attribute>
               <attributeName>Helenium_vernale</attributeName>
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            <attribute>
               <attributeName>Phlox_drummondii</attributeName>
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            <attribute>
               <attributeName>Phlox_glaberrima</attributeName>
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            <attribute>
               <attributeName>Phlox_nivalis</attributeName>
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            <attribute>
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            <attribute>
               <attributeName>Geum_macrophyllum</attributeName>
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               <attributeName>Geum_peckii</attributeName>
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            <attribute>
               <attributeName>Geum_virginianum</attributeName>
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            <attribute>
               <attributeName>Rhizophora_mangle</attributeName>
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            <attribute>
               <attributeName>Rubus_arenicola</attributeName>
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            <attribute>
               <attributeName>Rubus_argutus</attributeName>
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            <attribute>
               <attributeName>Rubus_chamaemorus</attributeName>
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               <attributeName>Rubus_cuneifolius</attributeName>
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            <url name="Eastern Massachusetts Flowering Phenology 1852-2013">https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=HF258</url>
            <url name="Widespread Sampling Biases in Herbaria Revealed from Large-Scale Digitization 1656-2016">https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf296</url>
            <url name="Assessing Plant Phenological Patterns in the Eastern United States Over the Last 120 Years">https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf309</url>
            <url name="Assessing Plant Phenological Character Displacement Across the Eastern United States Since 1895">https://harvardforest.fas.harvard.edu/exist/apps/datasets/showData.html?id=hf335</url>
         </additionalLinks>
      </metadata>
   </additionalMetadata>
   <additionalMetadata>
      <metadata>
         <unitList>
            <unit id="centigramPerCubicCentimeter"
                  multiplierToSI="centigram per cubic centimeter"
                  name="centigramPerCubicCentimeter"
                  parentSI="kilogramPerMeterCubed"
                  unitType="massDensity">
               <description>centigram per cubic centimeter</description>
            </unit>
         </unitList>
      </metadata>
   </additionalMetadata>
   <additionalMetadata>
      <metadata>
         <unitList>
            <unit id="cubicCentimeterPerCubicDecimeter"
                  multiplierToSI="cubic centimeter per cubic decimeter"
                  name="cubicCentimeterPerCubicDecimeter"
                  parentSI="meter"
                  unitType="volumePerVolume">
               <description>cubic centimeter per cubic decimeter</description>
            </unit>
         </unitList>
      </metadata>
   </additionalMetadata>
   <additionalMetadata>
      <metadata>
         <unitList>
            <unit id="gramsPerKilogram"
                  multiplierToSI="0.001"
                  name="gramsPerKilogram"
                  parentSI="gramsPerGram"
                  unitType="massPerMass">
               <description>grams per kilogram</description>
            </unit>
         </unitList>
      </metadata>
   </additionalMetadata>
</eml:eml>
