USA National Phenology Network

Development, Rationale, and Vetting of the USA-NPN Plant Species List, 2008

DRAFT

October 2008

Species and Protocols Working Group

1 Table of Contents

Table of Contents...... 2 Contributing Species and Protocols Working Group Members ...... 3 Introduction...... 4 Development of the USA-NPN Plant Species Lists, Beta 2008...... 4 Beaubien & Schwartz 2003 ...... 4 Betancourt 2006 Native and Non-Native Species by Ecoregion...... 6 Species & Protocols Working Group October 2006...... 7 Betancourt & Thomas 2007...... 9 Project BudBurst (PBB) species list ...... 10 Species & Protocols Working Group August 2007 ...... 12 Species & Protocols Working Group October 2007...... 18 National Vetting...... 19 USA-NPN Species List, Beta 2008 ...... 20 Justifications for each plant functional types...... 20 Agriculture & Phenology...... 20 Phenology and Invasive Plants ...... 24 Phenology and Allergens ...... 26 Deciduous Trees, Remote Sensing, and Fall Color Tourism...... 27 Phenology and Coniferous trees ...... 30 Showy and/or Urban Species...... 31 Herbaceous Species ...... 32 The Importance of a Clonal Backbone for the NPN...... 33 Appendix List...... 41 Appendix 1. Beaubein & Schwartz 2003 list of 17 species by ecoregion...... 41 Appendix 2. Beaubein & Schwartz list of 36 species...... 41 Appendix 3. Betancourt 2006 list of native species by ecoregion...... 41 Appendix 4. Betancourt 2006 list of non-native species by Ecoregion...... 41 Appendix 5. McNulty evaluation of trees in Appendix 4 list for SE USA ...... 41 Appendix 6. Master list of 1,258 species compiled in 2006...... 41 Appendix 7. Anderson list of 95 species, 2006 ...... 41 Appendix 8. McKee list of 46 species, 2006...... 41 Appendix 9. McKee list of 29 native species, 2006 ...... 41 Appendix 10. McKee list of non-native species by Ecoregion, 2006 ...... 41 Appendix 11. Thomas & Betancourt compilation of lists, 2007 ...... 41 Appendix 12. Thomas & Betancourt prioritization of lists, 2007 ...... 41 Appendix 13. Project BudBurst species list, 2007 ...... 41 Appendix 14. SPWG 2007 list of proposed calibration and focal species ...... 41 Appendix 15. Cover letter for SPWG 2007 list, version 2 vetting...... 41 Appendix 16. SPWG 2007 list, version 2...... 41 Appendix 17. Summary of responses to review of SPWG 2007 list, version 2...... 41 Appendix 18. USA-NPN Species List, Beta 2008 ...... 41 Appendix 19. Select phenophase information...... 41

2

Contributing Species and Protocols Working Group Members

October 2006: Julio Betancourt, David Breshears, Kathy Goodin, Art McKee, Craig Anderson, Michael Loik, Bruce Jones, Bob Szaro, and Gretchen Meyer

August 2007: Stephen Baenziger, Julio Betancourt, Dave Breshears, Kjell Bolmgren, Ellen Denny, Pauline Drobney, Abe Miller-Rushing, Carol Spurrier, Robert Szaro, and Jack Williams

October 2007: Jake Weltzin, Julio Betancourt, Mark Losleben, Kathryn Thomas and Art McKee with Ellen Denny, Mark Schwartz and Abe Miller-Rushing remotely

3

Introduction

The purpose of the USA-NPN is to provide phenological information used to understand the timing of life cycle events and their role in the biosphere. It will establish a nationwide network of phenological observations with simple and effective means to input, report, and utilize these observations, including the resources to provide information for a wide range of decisions made routinely by individual citizens and by the Nation as a whole.

U.S. Geological Survey and the University of Arizona cooperatively established the USA-NPN National Coordinating Office (NCO) in Tucson in 2007. Prior to implementation of the NCO, two NPN Implementation Planning Workshops guided development of the NPN. National Science Foundation Research Coordination Network (RCN) was established to support the NPN. (Schwarz) The Plant Phenology Program of the NPN was initiated through the NPN in the fall of 2008.

The first Implementation Planning Workshop in August 2005 developed draft objectives and short-term, mid-term, and long-term products for the evolving NPN identified the need to target plant species for phenological monitoring that are:

“of [regional] importance and whose phenology is of socioeconomic and scientific importance in understanding the ecosystem consequences of phenological dynamics and its regional variability, and to improve efficiency and reduce uncertainty of remote sensing application. (http://www.uwm.edu/Dept/Geography/npn/meetings/2006/towards_a_usa-npn.pdf).

In order to achieve this objective, the NPN required a listing of recommended plant species for phenological monitoring that was thoroughly documented and vetted. This report documents the development of the recommended plant species list for the NPN. In the summer of 2008, the NPN initiated its first public phenology recommendations of the resulting list through National Phenology Network web site at http://www.usanpn.org/

Development of the USA-NPN Plant Species Lists, Beta 2008

Beaubien & Schwartz 2003

In 2003, Mark Schwartz contracted with Elizabeth Beaubien, U. Alberta/PlantWatch Canada, used five criteria (see below) to develop a list of 17 native target species (Appendix 1) and annotated with applicable criteria for each (see below). The biogeographic unit of analysis was Bailey's 11 Ecoregions, with a map available on the prototype NPN web page (http://www.uwm.edu/Dept/Geography/npn/native.html, see Figure 1). This initial list was later expanded to 36 target native species (Appendix 2).

4

Figure 1. The map of ecoregions used for the Beaubien & Schwartz 2003 listing of species. The ecoregions included are: (1) Warm continental; (2) Hot continental; (3) Subtropical; (4) Pacific marine; (5) Prairie; (6) Mediterranean; (7) tropical/Subtropical Steppe (8) tropical/Subtropical Desert; (9) Temperate Steppe; (10) Temperate desert; (11) Tundra and Subarctic.

Beaubien & Schwartz used five criteria for selecting species:

1) Widespread distribution and dominance; easy to find near human habitation;

2) Diverse variety of life forms and functional types, (trees/shrubs/herbaceous annuals and perennials that produce flowers every year), with preference for monoecious species (having both male and female flowers on same plant);

3) Distinctiveness: ease of recognition of both the target plant and its phenophase/growth stages (no similar-looking species or subspecies, with different flowering times);

5

4) Flowering characteristics. Clear and sharp phenophases or growth stages, with early spring bloom time is preferable and bloom timing largely driven by temperature accumulation; flowers stay open once blooms begins (they don't close at night or in cloudy or wet weather) and the lower buds are not attractive to herbivores.

5) Horticultural cultivars: species has no horticultural cultivars developed that are similar in appearance and hard to distinguish from wild specimens.

Betancourt 2006 Native and Non-Native Species by Ecoregion

(Appendix 3-5)

In March 2006, Julio Betancourt used similar criteria as Beaubien and Schwartz to develop two expanded lists by Bailey Ecoregions, one for native species and another for problematic, invasive species. An additional criterion was species that occurred in 2 or more of the 11 Ecoregions. The effort was mostly subjective and driven by perusal of the literature, rather than quantitative use of digital floral databases to ensure coverage and dominance.

This effort produced lists of native species by Ecoregion (1-11) (Appendix 3): (1) 72 spp.; (2) 127; (3) 75; (4) 38; (5) 36; (6) 42; (7) 52; (8) 42; (9) 49; (10) 24; (11) 49.

A similar exercise was done for non-native species (Appendix 4), which yielded lists by Ecoregion (1-11). (1) 7; (2) 9; (3) 13; (4) 5; (5) 15; (6) 4; (7) 11; (8) 7; (9) 10; (10) 12; (11) 5. The non-native species list was annotated with specific impacts. There was little regard in the non-native list for whether phenological observations for each species would actually help predict or manage invasions and their impacts. This criterion would apply, however, to many of the species in this list.

Another exercise involved the help of Steve McNulty with the Forest Service's Southern Global Change Program. Betancourt asked Steve to use the Forest & Inventory Analysis (FIA) database to evaluate the appropriateness of the subjective list for the SE U.S. Steve looked at Betancourt's list of non-native species for Bailey Ecoregion 3 and cross-referenced it with a query of the SE FIA database. Appendix 5 shows two types of tree distribution. The first show the number of times a species falls within either the top 10 or 20 most dominant species within a state (based on the number of stems counted in all FIA plots within the state). Therefore, 13 would be the highest value. Only Red maple was within the top 20 most dominant species for all SE states, and in 9 of the top 10 for 12 states, etc. The second breakdown shows the top 10 and top 20 most dominant species within each state. Of course, distribution is more important than dominance for a phenological study, but these analysis should help to determine how difficult it may be to find good spatial representation for the phenological network, particularly for some applications such as ground-truthing of remote sensing products.

6 Species & Protocols Working Group October 2006

At the 2nd NPN Implementation Planning Workshop in October 2006, the Species & Protocol Working Group (SPWG) developed criteria for selecting target species. A general goal was to select less than 100 species ‘widespread, dominant and sensitive to temperature, plus species of interest’. Additional notes on this goal and other goals set by the group are in the unpublished notes from that workgroup.

Criteria for selecting target species

Criteria for selecting target species will be varied, and the reasons for selection will be nuanced. Groups interested in large-scale process (e.g., biogeochemical cycling, disturbance ecology, ecohydrology, vegetation modeling, and ground truthing of remote sensing) will naturally focus on widespread and dominant plant species. Conservation biologists focused on alpine floras will want to target at risk plants or host plants for at risk organisms, while phenology of non-native species is essential to most efforts to manage plant invasions via herbicides or biocontrol. Also, NPN will be constrained, at least initially, by the participating networks. These include diverse networks such as LTER, AmeriFlux, NEON, Organization of Biological Field Stations, National Park Service Inventory & Monitoring, USDA FS Experimental Forests, USFWS National Wildlife Refuges, etc. Obviously, NPN has to include species that occur at their sites and that interest scientists, interpretive staff, and land managers.

Selection Criteria determined by SPWG 2006

1. Ecoregions/biomes 2. Breadth of distribution – widespread 3. Dominance 4. Ease of identification 5. Ease of observation of phenophase 6. Ecological amplitude 7. Disease/pest susceptibility 8. Functional categories – wind and animal (insects and vertebrates), pollinators, methods of dispersal 9. Annuals, perennials, long-lived species 10. Deciduous or evergreen 11. Plants for which there are questions about changing synchrony/asynchrony between plant and animal hosts 12. Masting Species 13. Resilience to disturbance/change 14. Economic importance – horticultural, agricultural, medicinal 15. Iconic species – touristic value 16. Invasive species that could be better managed with phenological data and models (Julio use NatureServe web) 17. Garden and cloned species (Kay Havens will be following this up in the southern states) 18. Conservation concern and/or rarity 19. Ease of remote sensing – ability to validate signals, species codominants

7 20. Species at edge of their geographic and elevational range 21. Overlap with spp. measured or likely to be measured in other countries 22. List of spp ‘at risk’ from climate change 23. Regional importance (exercise could be done by region) 24. Separate Spp./Phenophases that are sensitive to temperature vs. photoperiod 25. Suite of species that allow continuous phenological observations throughout year 26. NIDIS- National Integrated Drought Information System- Drought Indicators

With the help of Kathy Goodin and her co-workers at Nature Serve, The SPWG 2006 applied many of those criteria to the NatureServe (Nature Explorer) plant database for the U.S. (http://www.natureserve.org/explorer/). The NatureServe database was queried by frequency of occurrence by: 1) number of states; 2) number of provinces; 3) number of plant association or gname; and ( 4) GRank = global ranking for conservation significance).

This produced a list of 1258 species that served as the basis for subsequent exercises (Appendix 6). Complete fields in this dataset are the number of times a plant occurred in a NVC association name (gname) and the number of provinces in which the plant occurs. Common name, origin, Grank, and number of states of occurrence are also included as fields but the information is not complete for all species.

We used Grank a different way than was intended, focusing on G5 species, or species that are actually quite widespread, abundant and therefore not threatened. We did this not to ignore conservation issues but to actually get lists of truly dominant species that characterize major ecosystems. Grank is based on the total number of known, extant populations worldwide, and to what degree they are threatened by destruction. Criteria also include securely-protected populations, size of populations, and the ability of the species to persist. The GRanks are as follows:

G1 — Extremely rare: Usually 5 or fewer occurrences in the overall range or very few remaining individuals; or because of some factor(s) making it especially vulnerable to extinction.; G2 — Very rare: Usually between 5 and 20 occurrences in the overall range or with many individuals in fewer occurrences; or because of some factor(s) making it vulnerable to extinction; G3 — Rare to uncommon: Usually between 20 and 100 occurrences; may have fewer occurrences, but with a large number of individuals in some populations; may be susceptible to large-scale disturbances; G4 — Common: Usually more than 100 occurrences; usually not susceptible to immediate threats; G5 — Very common: Demonstrably secure under present conditions. If the NPN decides to target use of phenological observations to evaluate threatened species, the Grank could be used in the conventional way to select species.

8 A number of spreadsheets were created by pairing down the list to ~100-150 species. This was done by group collaboration, and then individually by Craig Anderson (Appendix 7) and by Art McKee (Appendix 8). Craig Anderson ranked species by number of provinces the species occurred in, and he added some charismatic species. Appendix 7 provides Craig's list of 1258 species used in at least 2 association gnames, with frequency of use and distribution by province based on association distribution. Craig paired down this list to 95 species (see second worksheet in his Excel file). Art McKee did a similar sort, where he combined frequency of occurrence across states, gname (associations), and the Grank. Art's list of 46 species played a prominent role in the third-generation selection process attempted by the August 2007 S&P Group (Appendix 8). Art also did some quick sorting by Bailey's Ecoregion using a combination of files to come up with a shorter list of 29 species (Appendix 9) and non-native species (Appendix 10).

Three additional issues were discussed concerning selection of species: What is the appropriate taxonomic unit of analysis (species, variety, race)? Are there cloned indicator spp. for subtropics? Should there be overlap with species measured or likely to be measured in other countries?

Betancourt & Thomas 2007

The Nature Serve list of 1258 species (Appendix 6) was used as the base evaluation list. Species that occurred on any of the Beaubien & Schwartz (Appendix 1 and 2), Betancourt (Appendix 3 and 4), Anderson (Appendix 7), McKee (Appendix 8 and 9) or Project Budburst (see below) lists were added to the base evaluation list. A field was added for each of the above databases and a ‘vote’ for that species by the base list developer was indicated. Several iterations of the list were developed.

Species list 1: all NatureServe list (n=1258) (Appendix 6)

Species list 2: all Species list 1 + Beaubien/Schwartz (B/S), Betancourt (B), Anderson (A), and McKee (M) lists (n = 1680). Note that we used the initial Beaubien/Schwartz list and not the one that appears on the prototype NPN web page. This was an oversight that could affect some of the priority rankings if the objective was to maintain all of the website native species on the final target list.

Species list 3: All species 2 list + Project Budburst Species. List subsequently edited using following criteria: All species noted that had 10 or more provinces or any ‘votes’ by B/S, B, A, or M or >= 30 states (where number of states known) or >= 40 associations (n=302). The species list 3 was edited to fill in US Origin, common name, lifeform, Family, # states using PLANTS (http://plants.usda.gov/). Single genera occurring on species list 2 were eliminated. Where Goodin list had undetermined origin the PLANTS origin was used.

Species list 4: The species were prioritized. 1 Vote assigned for every time on one of B/S, B, A, or M lists. N = 138.

Priority 1: All Project Bud Burst list, All species with 4 or 3 ‘votes’,

9 Priority 2: All species on list with 2 votes Priority 3: All species on list with 1 vote.

Species list 4 includes a number of genera pairs.

Species list 5: Additional species from the NPN list were added. The list was edited to update species names to USDA PLANTS taxonomy. The NatureServe Explorer site was used to list the number of states each species occurred in. States where the species occurs as exotic or where the species are extirpated were not included. (Appendix 11)

Issues noted by Thomas & Betancourt

• The current list contains several examples of multiple species in one genera. Need to check geographic spread and ecological distinction. • The prioritized list has 46 priority 1, 52 priority 2, and 40 priority 3. All Project Budburst are priority one; if they are not included in the priority 1 count there are 28 species in priority one. • The list needs to be analyzed to evaluate spread throughout ecoregions/provinces or states.

Project BudBurst (PBB) species list

Project Budburst developed a species list and associated protocols subsequent to the October 2006 NPN Meeting in Milwaukee. The Education Working Group from this meeting (August 2007 RCN USA NPN Meeting) includes a more comprehensive report from the meeting. Additional information on Project Budburst is at http://www.windows.ucar.edu/citizen_science/budburst/. The species list use in Project Budburst is presented in Appendix 13.

Associated species protocols and key phenophases were identified for each species, as illustrated in the figure below.

10

11 Species & Protocols Working Group August 2007

To initiate that action the first meeting (August 27-29, 2007) of the RCN chartered an ad-hoc Species Protocols Working Group (SPWG) as follows:

1. Draft native plant species (with appropriate protocols) lists (10-12 pages) organized by ecoregion (to cover all areas of the continental USA) for use at USA-NPN Tier 3 (backbone) and Tier 4 (Intensive) observation sites. Include a 1 page summary for use in the USA-NPN Strategic Plan.

The draft list hould include three levels: a. Basic-about 6 species appropriate for each region that all sites will be strongly urged to observe to maximize species commonalities among all regions. b. Extended-6-12 additional species appropriate for each region that will support comparisons within each ecoregion c. Advanced-additional species that are important for one or more collaborating research sites. This work will require consultation with the Collaboration Group.

2. Draft Continental Indicator plant plan (5 pages or less) that recommends additional cloned plants (beyond lilacs) to be used to provide full continental coverage. Dogwood (Cornus florida) has been suggested for the SE USA and Ocotillo (Fouquieria splendens) for the SW USA, but their suitability needs to be explored, as well as if either will work in the areas of the Pacific NW that do not receive sufficient chilling to support the cloned lilacs.

3. Draft Agricultural plant species plan (5 pages or less) that recommends an initial list of agricultural species and protocols that could be included in USA-NPN observations, and the regions where they would be appropriate

4. Draft Observation Scope and Prospect plan (5 pages or less) that describes the plans and timetables for incorporating observations beyond plants into USA-NPN (birds, insects, mammals etc.), and end-of-season observations. This work will require consultation with the Collaboration Group.

The group proceeded by reviewing the charge above, revisiting previous related NPN reports and analyses (described below), and considering the species lists and associated protocols developed by Project Budburst, which were developed following the most recent NPN report on species and protocols (from October 2006).

Procedure for prioritizing plant species selection

See Appendix 14 for final target list of calibration and focal species

12 Further development of NPN should build on the approach adopted by Project BudBurst, in which there was a recommended set of species to observe AND/OR observers could add information on other species of interest. The key advantage of developing a prioritization for species observations is to encourage a larger number of observations that could span a larger part of the country. A key disadvantage to developing a prioritization is that potential observers (at any skill level) that do not find a species in that list that is of particular interest to them might be discouraged from participating. The Group sought to develop an approach that would have the advantage associated with a prioritization scheme while avoiding the disadvantage of discouraging participation focused on other species. This balance between depth and breadth is important at all levels of NPN, from Citizen Scientists that might have a particular plant species in their yard or school that they wish to study to professional ecological researchers working at intensively instrumented sites such at Long Term Ecological Research (LTER) sites (http://www.lternet.edu/) or at Ameriflux sites (http://public.ornl.gov/ameriflux/) where towers are located for measuring carbon dioxide fluxes.

The Group discussed the pros and cons of different prioritization schemes. We identified key criteria that could be important in prioritizing which species to focus on. These included:

• Widespread distribution • Associated with dominant plant cover type • Have an important biological influence on other species • Readily detectable and observable plants and phenophases • Accessible for schools and in urban areas • Pollinator or wind-borne seed dispersal • Diversity of plant functional types • Diversity of temporal responses in phenophases • Overstory and understory relationships • Synchrony between plants and pollinators • Potential responses to changes in fall as well as spring • Existing related long-term sets, including herbarium collections • Key influences on applied areas of interest such as recreation and tourism (fall foliage), medicine (allergens), agriculture, and natural resources management

Most of these criteria had been previously identified in previous NPN meetings. The key challenges were to determine how long the list of prioritized species should be; whether or not to develop a tiered list (beyond “preferred” and “other” categories); how to weight the criteria above in developing the list.

Options were discussed for species lists that ranged from as few as ~10 to as many as ~100. If only a few species are selected, they may not occur at enough of the locations. If too many species are selected, participants could also be discouraged from participating in that the task appears too daunting. In addition, the resulting data might be too spatially dispersed to be effective in addressing issues of concern.

To determine how to most effectively address these issues, the Working Group returned to previous documentation on NPN goals. In particular, we focused on a table of listing the ways in

13 which NPN seeks to provide benefits. These were outlined in the document “Towards a National Phenology Network” (October 2006):

These 6 areas can be aggregated into three more general types: 1) Scientific Research, 2) Education, and 3) Applications (including Agriculture, Tourism and Recreation, Human Health, Natural Resources, and Education). The Working Group recommends that a top tier of species (10-20) be selected based on the criteria that they would enable some successes in all six of the categories above. In addition, this top tier list should provide spatially extensive coverage complimentary to that of the lilac network and should include some species for every state and most counties in the US. The species should include a diversity of functional types and phenophases. This resulted in three categories, originally referred to as tiers but perhaps better referred to as follows (Project BudBurst and others could be consulted to evaluate if there are better terms for each category):

• (Tier 1): National Network Focal Species (the most important 10-20 species that would enable NPN to successfully achieve goals in all of the categories discussed above) • (Tier 2): National Network Integrating Species (widespread important species spanning across regions that can provide additional information at large spatial scales) • (Tier 3): Regionally and Individually Important Species (not included in the other lists; importance could be associated with dominance, endemism, conservation or other criteria).

Each of these three categories is important and enables complimentary objectives. The National Network Focal Species set provides the best means of assessing larger-scale changes. If more observations are concentrated on these species, this data set will have the most power for unraveling broad-scale phonological responses. The National Network Integrating Species set provides more in-depth coverage that still is very broad in extent. The Regionally and Individually Important Species set will be of particular interest for researchers at many sites, for land management in a species-specific context, and for Citizen Scientists with interest and access to particular plant species. The Working Group developed a list of plant functional types

14 that would need to be included to address the six categories. A focal research question was identified for each category. Then the Working Group evaluated candidate species for each plant functional type (detailed below).

The categories and associated questions for the plant functional types were:

Plant Question Number Functional of Type Species Selected* Crops Can NPN information affect and improve real-time crop decisions? 2 (3) Invasives Can NPN information affect and improve management of 2 (2) important invasive species? Allergens Can NPN information inform and improve a transition from post- 1 (8) pollen assessments to pre-pollen forecasting? Deciduous Can NPN information improve forecasting of fall foliage for key 3 (5) woody plants deciduous species of interest in tourism and recreation? Coniferous Does conifer phenology help improve predictions related to water 3 (3) woody plants stress and associated land management issues (fire, bark beetles, drought-impacts)? Herbaceous How is herbaceous productivity and associated grazing and 4 (7) plants harvesting times affected by varying climate? Showy, Urban, Do widespread and readily accessible species (e.g., schoolyards, 4 (7) and/or urban) provide similar metrics to phenological responses as native widespread species? Clonal Does controlling for genetic variation improve predictions of 2 (3) phenological responses?

These research questions are included to help ensure that the National Network Focal Species can address questions in all six categories of interest listed above. They are not intended to discourage pursuit of other related and important questions. One important cross-cutting issue relates to spatial-temporal variation in phenology and the extent to which it can be detected through remote sensing. Measuring phenology for the National Network Focal Species may partially address this issue, but the species have not been selected to optimize this objective alone. Nor were they selected as a set of species likely to be most sensitive to climate variation. Additional relevant research questions are available in previous NPN planning documents (also see the report from the Research Working Group from this meeting).

The approach developed here for key plant species can also be expanded to address other organisms, such as insects, amphibians, and birds (see related section Below).

Species selected for the National Network Focal Species are presented in the table below. Subsequent sections describe the rationale for selection of the species in each of the three categories (National Network Focal Species, National Network Integrating Species and Regionally and Individually Important Species).

15 16

We evaluated our selections with respect to the degree of national coverage on a state-by-state basis. We previously focused on ecoregions (aka. Bailey) for assessing coverage but found that data were more readily available on a state basis. In addition, we anticipate that NPN participants will enter the system through a state/county basis. In our attempt to compile a national list based on widespread and common species, we evaluated the distribution of our species sample using the documentation from Nature Serve (http://www.natureserve.org/explorer/servlet/NatureServe). Out of the 22 tier 1 species, all states except Hawaii (5) and Alaska (5) have 10 or more species. 42 states have 12 or more species in tier 1, and no state has more than 16. Summing up the tier 1 and tier 2 (104 sp.), all but two states (AK, HI) have more than 40 species or more.

Species no. by state

100

90

80

70

60 tier_2 50 tier_1 40

30 20

10

0

I A IL S A D D H Y Y AK AR CA CT DE GA I K L M M MO MT N N NM N OK PA SC TN UT VT WI W State

Species Protocols and Citizen Science Engagement Strategy (August/Sept. 2007)

Julio Betancourt, Dave Breshears, Mark Losleben and Jake Weltzin had some important post- meeting discussions about species and protocols in August and September 2007 following the RCN meeting in Milwaukee. We developed a modification of the rationale and prioritization scheme for species inclusion.

Recall that we struggled with the tradeoff between having a few species that span the country and not discouraging interest in other species. Dave B. tried to address this in his draft by changing the terms from Tier 1s to 3 to less value-laden terms. But, the main problem remained as to how to build a core set with lots of depth AND encourage people to study the species of interest.

The idea that we came up with is to encourage participants to chose and study a species of their choice, as well as a "backbone" species, which would help us extrapolate their results for them and for science.

Key Points:

Engage citizen scientists through inclusive invitation: when participants enter the main web portal for participation, we will ask them to chose one or two species for monitoring.

17 Chose your species of interest, which we term FOCAL species. The rationale for choice of species will depend on the situation and participant; for example, the species may be a visual dominant, a rare species of special interest or prior study, the focus of local management activities, an allergen, an important agricultural crop, etc.

We will develop a suite of potential focal species (the 80 species in the old tier 2 and tier 3 categories), with monitoring protocols; however, if the focal species is not on the list, the NPN will develop protocols for monitoring and add that species to the list. If the participant does not have a particular preference, they will be encouraged to select a species from the CALIBRATION list, or secondarily from the FOCAL list.

We would also like you to consider also selecting at least one other species from the CALIBRATION species list to help us calibrate (your observations at a national level and relative to longer time scales ?)

Species & Protocols Working Group October 2007

The National Science Foundation funded a Research Coordination Network (RCN) grant to NPN that supported an Implementation Planning Workshop in August 2007. It was decided at that workshop to prepare a vetted list of plant species to be included in the Network for 2008. In late October of 2007, several people assembled at the NPN offices on the University of Arizona campus to discuss and initiate the vetting process. Participants in those October discussions were Jake Weltzin, Julio Betancourt, Mark Losleben, Kathryn Thomas and Art McKee; with Ellen Denny and Mark Schwartz engaging via conference calls, and Abe Miller-Rushing participating by e-mail.

The participants reviewed the proposed list of plant species to be considered for the NPN (Appendix 14, Species and Protocols Breakout Group Report, RCN Implementation Planning Meeting, Aug 2007) and its reorganization by Jake Weltzin into three categories: Indicator/Cloned, Calibration and Regional/Focal Species. These had been ordered alphabetically by genus and species within each category. At this point in the process, it was decided to keep the list in the Indicator/Cloned category very short and focus on species that could be easily cloned and had wide geographic ranges. It was recognized that the species included in that category did not cover the southern or southwestern US very well, that species were needed for that geographic area.

It was also decided to try and keep the Calibration species list relatively short, with a target of around 20 species. The partitioning of species into Calibration or Regional/Focal lists was done utilizing input from all participants up to that point and emphasizing abundance and wide distribution as criteria for inclusion the Calibration list. Following this partitioning, the 2007 list from Project Budburst , and the prototype list from earlier NPN efforts led by Mark Schwartz were added as separate columns and the final compilation to be vetted by independent review was labeled USANPN_ProposedSpp.

18 National Vetting

A request for assistance with vetting the proposed USA-NPN species and accompanying list were mailed out in early November 2007. The cover letter is included in Appendix 15, and the proposed list (USANPN_ProposedSpp) is included in Appendix 16. This list was mailed to over 350 scientists, field research stations and network offices. In addition to the USA-NPN mailing list, Jake Weltzin, Mark Losleben and Kathryn Thomas had been compiling addresses of scientists who had expressed interest in vetting a proposed list. This compilation included agency scientists (e.g. National Park Service, US Forest Service, U S Fish and Wildlife Service) as well as university faculty. These names were added to lists from several networks: the Northeast Regional Phenology Network (Ne-RPN); the Organization of Biological Field Stations (OBFS); the Long-Term Ecological Research Network (LTER); and the National Ecological Observatory Network (NEON) with requests to forward the letter of request for help and proposed USA-NPN list to appropriate scientists in the respective networks. Responses were mailed to Mark Losleben who compiled and archived them at the national office of NPN.

The responses to this round of vetting came from a mix of individuals and groups that collectively represent over 100 scientists. Working with Mark Losleben, Art McKee summarized the feedback (Appendix 17) and met with the USA-NPN Vetting and Protocol Committee in January of 2008 to incorporate the suggested revisions and additions into a USA- NPN Species List for 2008. Members of the Committee that participated in these discussions were: Jake Weltzin, Mark Losleben, Kathryn Thomas, Theresa Crimmins, Art McKee, and, by phone, Ellen Denny, Mark Schwartz, and Bruce Wilson. Abe Miller-Rushing participated by e- mail.

Some of the changes incorporated into an NPN Beta species list for 2008 included: eliminating the Indicator/Cloned category of USANPN_ProposedSpp and moving those species into the Calibration category; deleting and adding species in both the Calibration and Regional/Focal categories, and adding two new Regional/Focal lists to ensure species representation in Hawaii and Puerto Rico. The primary factors used in deleting or adding species were the range and abundance of each species, their regional representativeness and importance and whether or not the species was already included in a sampling network. Of the species that had been considered for cloned material within the NPN, Ocotillo (Fouquiera splendens) was dropped from consideration because of the extreme phenological variation within the species evidenced over small spatial scales, and flowering dogwood (Cornus florida) was moved into the Regional/Focal species list because of the uncertainty of obtaining large amounts of clonal material. This species could well become part of a clonal network for calibration in future years.

The vetting process produced well reasoned arguments for substituting one species for another in the Calibration list and adding species in the Regional/Focal lists. For example, several scientists working in the plains suggested substituting Bouteloua gracilis (blue grama) for Bouteloua curtipendula (side-oats grama) because blue grama tends to be the more common species throughout the plains.

In addition, it had been pointed out by several during this vetting process that the proposed lists did not include species specific to either Hawaii or Puerto Rico. The vetting scientists from both

19 regions had made suggestions of species to be included, and it was clear that the species proposed were the result of careful consideration, including, in many cases, species already being monitored in other programs. Thus, the addition of two more lists in the Regional/Focal category.

USA-NPN Species List, Beta 2008

The result of all these considerations was a draft “beta” list for USA-NPN species for 2008 that was mailed to the USA-NPN Board of Directors and Project BudBurst leaders. Following their feedback, a final USA-NPN Species List, Beta 2008, was produced (Appendix 18). In early February, 2008, Jake Weltzin mailed out the USA-NPN Species Lists, Beta 2008 to the USA- NPN network. That list was to be the basis for the 2008 NPN campaign, and phenology sampling protocols were to be drafted for as many species on the lists as possible.

Justifications for each plant functional types

Agriculture & Phenology

In this section, we will use agriculture in its broadest context, simply the managed ecosystem that is used to produce plants and animals for food, feed, fiber, and aesthetics, as well as to remediate the effects of humanity (e.g. the land application of urban effluent) and sustain the environment. What separates agriculture from many ecosystems is the level of its management and the importance of making timely decisions and the presence of “real-time” decision models, such as, when or if to plant, when to fertilize, when to irrigate, when or if to control pests, when to bring in pollinators or to detassel, when to graze or to stop grazing, when to harvest, and when to sell the products. A central objective of the USA-NPN is to develop phenological landmarks that may be used to monitor and anticipate phenomena that affect our ecosystems in both the short term (days to months) as would be needed for annual cropping systems and long term (years to decades) as would be needed for perennial crop systems . In addition, agricultural plants can be used to enrich the regional information developed by the NPN, as well as, greatly benefit by this information.

Specifically, the NPN can provide the biologically relevant ability to synthesize diverse seasonal weather fluctuations and climate information (the current basis of agricultural decisions) to temper the agricultural model predictions to enhance their predictive capabilities. We believe that the indicator plants (e.g. lilacs and others) that form the backbone of the NPN can be tied to agricultural plant, pest, disease, and animal growth. The advantages of using lilacs are that they are perennials similar to many agriculturally important plants (e.g. forage legumes and grasses, forestry plants, horticultural trees and shrubs) and that by being perennial they avoid the issue of when they were planted and how that affects subsequent plant development. Widely grown agricultural perennials include alfalfa and switchgrass (expected to increase as a biofuel crop).

20 Winter annuals (e.g. winter wheat, rye, barley, oats, triticale, and canola, etc.) can also be used, as their ability to overwinter, minimizes the affect of planting date on subsequent phenological measurements. Ideally, broadly grown summer annuals (e.g. corn, soybeans, vegetable crops) could be used in phenological networks once the relationship between planting date and subsequent phenology has been developed (e.g. http://www.ca.uky.edu/agc/pubs/agr/agr184/agr184.pdf ). In some cases, the well characterized genomes and genotypes of agricultural species can be used to optimize their NPN linked potential. For example, preliminary results indicate a winter barley line has been identified that lacks vernalization genes, hence can be grown as a fall sown winter barley from New York to Nebraska to Oregon and states further south. In more northern regions, it can be planted in the spring and complete its life cycle.

Table of Backbone Crop Plants:

CROPS 1 2 3 apple Almond oranges alfalfa Grapes avocados winter wheat peaches maize/corn cotton Soybean Strawberries Blueberries

For the tier one plants, apples (Malus domesticus) and alfalfa (Medicago sativa) were chosen based upon their being: 1. widely grown, 2. perennial, 3. easily identified, 4. representing widely different genetic structures, and growth and development patterns, 5. having easily identified phenological stages which affect the value of the crop, 6. have important pollinator interactions, and 7. having phenological information potentially available from commercial sources. In addition, commercial apples are clonally propagated with the additional benefits described in the importance of clonal species. Alfalfa is grown as a population and is expected to have greater environmental buffering. Its relative (M. truncatula; http://www.medicago.org/) has been chosen as a model plant for genome study and will provide a wealth of accessible genome information. To augment the tier one species, almonds (Prunis dulcis) which are highly dependent upon pollinator species, grapes (Vitus spp.), strawberries (Fragaria spp.), blueberries (Vaccinnium spp.), winter wheat (Triticum aestivum), maize (Zea mays), and soybeans (Glycine max) were chosen as tier two species. These species were chosen for the same reason as the tier one species but they lack in one or more of the six selection criterion. In addition, commercial almonds and grapes are grafted plants, blueberries are propagated in various ways, and strawberries are sold as plugs, hence are clonal plants (similar to apples). Winter wheat was chosen for this wide distribution, genomic resources, easy of phenological measurements and as a winter annual effectively removes the impact of seeding date on subsequent phenological measurements. Maize and soybeans were similarly chosen as winter wheat except that they are summer annuals; hence provide the opportunity to link data that are affected by seeding date to the NPN

21 backbone. Wheat and soybeans are inbred cultivars, whereas maize is a grown as a hybrid. The tier three plants were added to represent smaller regions that may be underrepresented by the tier 1 and 2 species. The tier 3 species include oranges (Citrus sinensis), avocados (Persea americana), peaches (Prunus persica), and cotton (Gossypium hirsutum and barbadense). Commercial oranges, avocados, and peaches are clonal plants, while cotton is planted from seed and is an inbred cultivar.

Representative Information/Protocols

Recommend using generic tree protocols that include fruit phenology for apple, almond, orange, avocado, and peach. Can a similar protocol be used for grape? I tried to attach what I can find for blueberries and strawberries (note there are cultivated versus wild species and the genetic population structure will be different). The protocols for maize, soybeans, and wheat are well described and documented. I added a generic detailed perennial grass reference that may be used for most forage grasses (see Moore et al.).

Apple information: http://en.wikipedia.org/wiki/Apple#Botanical_information

Alfalfa: http://cahe.nmsu.edu/pubs/_a/a-330.pdf , http://en.wikipedia.org/wiki/Alfalfa

Almond: http://en.wikipedia.org/wiki/Almond , http://www.botanical.com/botanical/mgmh/a/almon026.html

Grape: http://en.wikipedia.org/wiki/Grape , http://www.kingestate.com/wines/wine_info/grapedevelopment.php , http://www.practicalwinery.com/JulyAugust02/julaug02p14.htm ,

Strawberry: http://en.wikipedia.org/wiki/Strawberry ,

Blueberries: http://en.wikipedia.org/wiki/Blueberry , http://www.gnb.ca/0171/10/0171100026-e.asp ,

http://danielorzuza.files.wordpress.com/2006/07/growthstages.pdf --This is good.

Wheat: Most agricultural crops have defined multiple growth stage systems (e.g. Zadoks

[http://en.wikipedia.org/wiki/Zadoks_scale ,

http://www.usask.ca/agriculture/plantsci/winter_cereals/Winter_wheat/CHAPT10/cvchpt10.php]

22 or Feekes scale [http://ohioline.osu.edu/iwy/feekes.html] in wheat) and these can be used or simplified to fit into the NPN data acquisition. Many crops also have predictive phenological other decision based models for contolling diseases and pests using weather input data based upon archival climate data.

Corn: http://www.ag.ndsu.edu/pubs/plantsci/rowcrops/a1173/a1173w.htm

Soybean: http://www.ag.ndsu.edu/pubs/plantsci/rowcrops/a1174/a1174w.htm , http://www.oznet.ksu.edu/kansascrops/soybean_varieties.htm

Oranges: http://en.wikipedia.org/wiki/Orange_(fruit) , http://www.sunkist.com/products/growing_packing.asp#propagation

Avocados: http://www.crfg.org/pubs/ff/avocado.html , http://en.wikipedia.org/wiki/Avocado

Peach: http://en.wikipedia.org/wiki/Peach , http://classes.hortla.wsu.edu/hort310/hort%20310%20- %20documents/Fruit%20Growth%20&%20Development.pdf -- This is kind of generic for fruits of all kinds.

Cotton: http://pubs.caes.uga.edu/caespubs/pubcd/B1252.htm , http://edis.ifas.ufl.edu/AG235 , http://en.wikipedia.org/wiki/Cotton

NOT NEEDED--Canola: http://ndawn.ndsu.nodak.edu/help.html?topic=canolagdd- infohttp://ndawn.ndsu.nodak.edu/help.html?topic=canolagdd-info http://www.lethamshank.co.uk/gsosr.htm http://www.scijournals.org/cgi/collection/crop_growth_and_development?page=27

Collaborations

Due to the commercial importance of agricultural plants, there is a wealth of phenological data that may be accessible from grower networks, company, and governmental sources. For example, hybrid corn companies will have planting date, detasselling date, male plant removal,

23 harvest date with grain moisture contents (an indicator of maturity) for their diverse hybrid seed production fields. Similarly, alfalfa produces will have information on first cutting dates, as would switch grass producers. For winter cereals that are grazed, the appearance of hollow stems will be recorded as will when they are harvested for grain. Other readily available phenological information includes new crop harvest dates obtainable from grain elevators and fruit suppliers, and possibly chemical spray dates, which are based upon plant developmental stages. Potential partners include commercial seed companies, crop improvement associations (http://www.aosca.org/), crop scouting companies and organizations, crop insurance companies who have weather related crops claims (freeze, frost, heat drought, etc. damages). In addition, there may be “volunteer” networks with an agricultural bias that may be useful partners: 4H students (http://www.4husa.org/), Sustainable agricultural networks (http://attra.ncat.org/; http://www.sustainableagriculture.net/), seed savers (http://www.seedsavers.org/), native range and pasture groups.

Phenology and Invasive Plants

Invasive exotic species threaten conservation of native ecosystems, agriculture, recreation, economics, and aesthetics. These species did not evolve in concert with suites of native species within North American ecosystems, and thus they disrupt species interactions and ecosystem function, and degrade wildlife habitat. The cost of invasive species to the national economy has been estimated as high as $137 billion per year and increasing, due primarily to losses in agriculture, forestry and fisheries, as well as to the cost of clearing invasive-clogged waterways and fighting invasive-fueled fires. About 42% of the species on the Federal Threatened or Endangered species lists are at risk primarily because of invasive species.

Invasive species selected for North American phenological observation respond to environmental cues that are important for their control or for restricting range expansion. These species are listed in Table 1.

Table 1. Invasive species included in the NPN Project.

1 2 3 Scientific Name Scientific Name Scientific Name Centaurea stoebe Bromus rubens Brassica tournefortii Cirsium arvense Bromus tectorum Rhamnus cathartica Centaurea solstitialis Pennisetum ciliare Euphorbia esula Lespedeza cuneata Lonicera japonica Lythrum salicaria Myriophyllum spicatum

24 Polygonum cuspidatum Pueraria Montana Tamarix spp.

In many cases, such as in Canada thistle (Cirsium arvense), control actions such as mowing, herbicide treatment or application of biological control is critically related to development of buds, flowering, or other phenophases. In many cases, integrated pest management requires a precise understanding of the timing of various types of treatments relative to phenophase in order to most effectively control an invasive species.

The ranges of some invasive species are expanding due to changes in weather patterns. Kudzu (Pruraria montana), for example, is now surviving in areas north of its former range, in response to shorter or warmer winters. In such cases, observation of phenology throughout their North American range could provide advance warning of range expansion, and provide time to prevent spread. Prevention of invasive species is more effective and far less costly than containment and eradication.

Our information about invasive species control and exclusion is imperfect and detailed information about the phenology of individual species helps develop more effective control strategies. This will be especially important in the face of global climate changes.

Though arguments could be made for inclusion of numerous exotic and invasive species in the NPN Project, some species perhaps should be excluded because of a history of horticultural desirability. Purple loosestrife (Lythrum salicaria) has been widely grown as an ornamental and flowering of large populations have even been known to be the focus of a local celebration due to its beauty. State governments have often had great difficulty in banning its sale due to the popularity of purple loosestrife in the horticultural trade. Purple loosestrife still adorns numerous gardens in areas including highly vulnerable wetlands. It could be argued that inclusion of this species in the NPN Program would provide opportunities to educate otherwise uninformed individuals of the degree of threat due to this invasive species. However, we feel that inclusion of purple loosestrife at this phase of development of the NPN may result in the unintended consequence of encouraging people to justify keeping or even providing plants to others in order to observe phenophases for this project. For these reasons, and because there are many more invasive species than can be included in this project, we recommend that it not be included in the study at this time.

Specific phenological information by invasive species chosen for observation has been compiled below. Much of the information below was copied from Element Stewardship Abstracts compiled by the Nature Conservancy and hosted on their web site: www.tncweeds/ucdavis.edu/esadocs.htlm

Where information on phenology stages of the species was available, it is included in Appendix 19 with examples of how phenology measurements can be useful in managing these invasive species. Other information was downloaded from various sites available by searching on species names through Google.

25 Phenology and Allergens

A central objective of the USA-NPN is to develop phenological landmarks that may be used to monitor and anticipate phenomena that affect human health. In particular, allergy (hay fever) and pulmonary (asthma) problems affect millions of Americans, and there is the risk that climate change may affect the timing of pollen release, length of pollen season, and total pollen loadings in the atmosphere. Currently, atmospheric pollen monitoring is performed by networks of independent stations run by allergenicists who measure atmospheric pollen concentrations. These data are assembled by several non-profit (AAAAI) and for-profit organizations (e.g. pollen.com, weather.com) and combined with meteorological data to produce same-day and up to 4-day allergy forecasts for the US (e.g. pollen.com, weather.com).

Pollen-forecast models currently do not incorporate other relevant phenological observations- i.e. flowering time and senescence or vegetation green-up that could provide longer-lead forecasts of pollen production and allergenic risk. An important application for the NPN would be to quantify the relationship between earlier phenophases and pollen production at local to regional scales for use in pollen forecast models. For example, phenological observations for junipers could improve forecasting models that rely on day-to-day release forecasts at source areas and dispersion forecasts to downwind areas to predict allergenic risk (Levetin and Van de Water 2003). Phenological observations could also parameterize dispersion and mesoscale meteorological models now being adapted to forecast atmospheric pollen concentrations (Pasken and Pietrowitz 2005). Additionally, monitoring of species-level phenological phenomena should be able to distinguish inter-species differences in phenological sensitivity to climate change, which will improve predictions of future changes in the onset/abundance/duration of pollen load.

There are several potential opportunities for collaboration between the USA-NPN and the medical community. First, allergenicists have been collecting daily data on atmospheric pollen loadings since the 1920’s. These data are spatially sparse (dozens of sites nationally), temporally spotty (few if any continuous records prior to the mid-1990’s), and suffer from data- heterogeneity issues (measured pollen concentrations are highly sensitive to sampling design), but nonetheless offer a rare and currently unexplored source of long-term phenological observations.

Second, the NPN may be able to provide a service to the medical community by 1) collecting phenological observations for key allergenic species and 2) using this information to improve prediction accuracy of short-term pollen-forecasting models, and 3) model the human-heath impacts of phenological changes driven by climate-change scenarios. A wide variety of US trees and herbs produce allergenic pollen (Trees: elm, juniper, birch, alder, oaks, maple, ash, olive, cottonwood, mulberry, pecan, walnut, sycamore. Herbs: grass, mugwort, ragweed, chenopods, pigweed, sorrel, plantain) (www.aaaai.org/nab/index.cfm?p=common_outdoor_allergens). This list includes many USA-NPN Tier 1 and Tier 2 species.

We have selected Ambrosia (ragweed) as a Tier 1 taxon because 1) it is a well-known allergen, and has been the subject of much research and 2) its constituent species are found throughout the US. The earliest continental survey (1929-1933) for Ambrosia comprised 70 cities and ca. 13,000 samples (Durham, 1929, 1931a, 1931b, 1933, 1935). Other historical data compilations

26 include ca. 30 US sites with intermittent data from the 1960’s to 1980’s (Frenz et al. 1999) and data from the 1980’s to mid 1990’s (Frenz et al. 1995, 1999). Experiments with Ambrosia show that pollen production and the allergenicity of pollen increase with elevated CO2 (Ziska, Bazzaz REFS). Scott Isaard and others at the Computational Epidemiology and Aerobiology Lab at Penn State are currently developing a predictive model of ragweed phenological development applicable in both Europe and North America, and will be using phenological observations to calibrate the model (http://www.ceal.psu.edu/ragweed.htm). The NPN could significantly enhance the success of these efforts.

Phenological measurements for Ambrosia should concentrate on flowering phenophases. Unlike other taxa in the NPN, we here focus on the genus level because Ambrosia pollen cannot be identified.

Deciduous Trees, Remote Sensing, and Fall Color Tourism

The need to include deciduous trees in the NPN is obvious: spring green-up and fall senescence of woody deciduous trees is one of the most pervasive and easily observed phenological events in the US. From a scientific perspective, monitoring and anticipating year-to-year variations in the timing of these events is essential to understanding interannual and long-term variations in global seasonal fluxes of carbon and water between the biosphere and atmosphere (MAUNA LOA REF). Many deciduous tree species are the dominant tree species in their community and hence are primary drivers of ecological dynamics XX From an economic perspective, many towns and regions rely heavily on revenue derived from fall colors tourism. Additionally, many deciduous tree species are well suited for the educational citizen-science objectives of the NPN, because these species are aesthetically important, locally dominant, easy to identify, and/or provide easy-to-observe phenological signals.

Currently, remote sensing already offers or is developing ‘wall-to-wall’ phenological products for the US that are based on the MODIS, Landsat (AND OTHER?) sensors (Morissette, Fisher, other REFS and WEBSITES). Such products are essential for assessing biospheric responses to global change at regional to global scales and carbon cycle modeling. These products, however, require validation against on-the-ground phenological observations, so that spectrally observed events such as vegetation green-up can be cross-referenced against species-level phenological indices such as leaf onset and flowering time. This cross-referencing has been persistently confounded by 1) the limited availability of on-the-ground phenological observations in the US and 2) scale mismatches between the minimum pixel size of synoptic remote sensors (e.g. 500m for MODIS) and site-level ground observations. A central motivation for the USA-NPN is to help overcome the ground-level observational limitations to remote sensing validation.

Remote sensing validation can be improved at two scales: the intensive observation of many individuals from plant communities at landscape scales (Schwartz, Liang REFS) and the extensive sampling of many individuals from a select list of species at national scales. As described above (DAVID’S SECTION), the three-tier strategy pursued here is intended to create a high-priority list of species for nationwide extensive phenological observations (Tier 1) while recommending species suitable for intensive regional- or landscape-scale investigation (Tiers 2

27 and 3). We recognize that for the purpose of landscape-scale remote sensing validation, phenological observations from only Tier 1 species will be insufficient. Instead, the intent of the Tier 1 list is to prioritize a few widely distributed and abundant species that can be used for national-scale assessments and intersite comparisons. We further stress that the assignment of species to Tiers 1-3 is not in any way meant to preclude species of importance to a particular organization, location, or research question, and that the USA-NPN welcomes all phenological information.

In assembling the limited portfolio of woody deciduous species for the Tier 1 list, we picked species that were nationally widespread, had complementary species ranges, were dominant at local to regional scales, and had strong fall colors and/or showy flowers. Based on these criteria, we chose Acer rubrum (red maple), Populus tremuloides (quaking aspen), Cornus florida (flowering dogwood), and Prunus virginiana (chokecherry). All four are widespread (A. rubrum: 33 states, P. tremuloides: 39 states, C. florida; 31 states, P. virginiana: 42 states) (CITE: Julio’s Natureserve tables). These species all have overlapping ranges at the state and county level (Kartesz and Meacham, Synthesis of North American Flora), making it possible to cross- reference phenological phenomena. P. tremuloides, A. rubrum, and C. florida were chosen as widespread representatives of the western US (P. tremuloides), eastern deciduous forests (A. rubrum) and southeastern deciduous/mixed forests (C. florida). C. florida and P. virginiana were chosen as widespread eastern deciduous trees with showy flowers, enabling joint phenological measurements of leaf and flowering phenology. Additionally, C. florida is a clonal species being evaluated as a possible southeastern indicator species to supplement the Lilac Observation Network (which does not extend south of ca. 43ºN).

Although it is probable that some observation sites will contain none of these species, it should be possible to find Tier 2 and Tier 3 species at those sites that elsewhere co-occur with one of the Tier 1 species, allowing a cross-referencing of phenophases for other regionally and nationally important species to one or more of the Tier 1 species.

Note: The list of Tier 2 species is currently based on a one-off brainstorming session, with no serious attempt to parse it into Tier 2 and Tier 3 taxa (or to ensure completeness of the list). This list needs to be reviewed. There was some initial attempt to pick sets of congeneric species with complementary range maps (e.g. Acer spp.) which would be worth following through.

Tier 1 Tier 2 Tier 3 Acer rubrum Acer glabrum Purshia tridentata Cornus florida Acer macrophyllum Cercocarpus letifolius Populus tremuloides Acer negundo Cornus nuttallii Cercidium Acer saccharum floridum/microphyllum Amelanchier canadensis Prosopis pubescens Amelanchier utahensis Prosopis velutina Arctostaphylos uva-ursi Artemisia tridentata Betula papyrifera Cornus sericea

28 Corylus americana Corylus cornuta Corylus servicea Fagus grandifolia Fraxinus americana Fraxinus pensylvanica Liquidambar styraciflora Liriodendron tulipifera Mahonia repens Quercus alba Robinia pseudoacacia Sambucus nigra syn. canadensis Synphoricarpos albus Tilia americana Quercus macrocarpa Cercis canadensis Alnus incana Alnus rubrum Carya glabra Carya ovata Juglans nigra

29 Phenology and Coniferous trees

The rationale for including coniferous trees in the NPN National Network Focal Species set is to ensure some phenological coverage for important, extensive vegetation types. These ecosystems are particularly extensive and important in the west. Relative to ecosystems dominated by herbaceous species or deciduous tree species, coniferous forests are evergreen and therefore overall phenological changes such as ecosystem greenness are relatively stable though time. However, these ecosystems and associated management issues are tightly tied to periods of seasonal water stress. These periods of seasonal water stress can be related to key events such as fire risk and/or bark beetle outbreaks and drought-induced tree mortality. In addition, some coniferous species such as pinyon pines are particularly important biologically due to masting production of nuts. In addition, previous phenological research has quantified relationships between the lilac network and conifers. Phenological responses should be interrelated with water stress in these species and could be important in improving understanding of ecosystem processes and management in coniferous systems.

We selected Pinus ponderosa, ponderosa pine, for one of the conifer study species in the National Network Focal Species because it was the pine species found in the most states (15). It is also important in terms of forest management, recreation, fire risk and restoration, and bark beetle outbreaks and associated drought-induced water stress. Many of the large recent, severe wildfires in the west were at least partially in ponderosa pine forests. Extensive restoration efforts are being focused on ponderosa pine, including restoration at the wildland-urban interface. This species is located mostly in the western US. To extend coverage east, we also selected Juniperus virginia (redcedar) for National Network Focal Species set. This species has extensive range across the coterminus US (43 states). Its range includes most of the species ranges for Pinus ponderosa and Pinus edulis/monophylla (see below) and extends well beyond.

In addition, we selected a pair of pinyon pine species: Pinus edulis and Pinus monophylla, which are co-dominant species of pinyon-juniper woodlands with other species of junipers. Pinyon juniper woodlands are one of the most prevelant vegetation types in the US and span much of the west. Distributions of Pinus edulis and Pinus monophylla overlap somewhat but together cover most the US pinyon-juniper woodlands. Most importantly, pinyon pines, through pine nut production associated with masting events, have enormous influence on other organisms in the community, including jays and rodents. They also indirectly affect disease spread related to Hantavirus. Pine nuts are also gathered, harvested and sold in communities thoughout the Southwest. Masting behavior is a fundamental phenological event that is poorly understood but very ecologically and socially important.

Table XX: List of coniferous species, grouped into Tiers 1, 2, and 3. The assignment of taxa to Tiers 2 and 3 is not finalized.

30

1 2 3 Pinus ponderosa Pinus palustris Tsuga canadensis Juniperus virginiana Larix laricina Tsuga mertensiana Tsuga Pinus edulis/monophylla Picea engelmanii heterophylla Picea glauca Picea mariana Pseudotsuga menziesii Pinus strobus Abies lasiocarpa Abies balsamifera Abies concolor Abies grandis Juniperus asheii Pinus taeda

Showy and/or Urban Species

For the purposes of Citizen Science outreach, it is important to include some key showy and/or urban species. Project BudBurst has already been under way for 1 year and in that time has identified several popular species for Citizen Scientists. It is important to note that many of these observations are within in urban areas and perhaps at or near schools or homes. For these taxa, the educational mission is of particularly important. These taxa can still be tied to important applied questions such as urban heat island effects, as well as the growing interest in urban ecology. In addition, the observations on the selected showy and/or urban species can be evaluated against observations of clonal species (e.g. lilacs) or native species to broaden the phenological knowledge base. For this category, taxa that were most popular in Project BudBurst were selected for the National Network Focal Species set. The four selected species were Prunus virginiana (chokecherry), Taraxacum officinale (dendelion), Fosythia sp., and Syringa vulgaris (Lilac – non clonal). Additional information on all of these species is available at the Project BudBurst website (http://www.windows.ucar.edu/citizen_science/budburst/)

Table XX: List of Showy species. No species have been selected yet for Tiers 2 and 3. [This may be a good place to insert other taxa from Project Budburst.]

1 2 3 Prunus virginiana Taraxacum officinale Forsythia spp. Syringa vulgaris Cornus florida* Fouquiera splendens*

31 Syringa chinensis*

*These species were selected as representatives of other PFT’s (e.g. Cornus florida is a primary representative of the Deciduous PFT) but have secondary value as showy species.

Herbaceous Species

Native herbaceous species are included in the NPN Project because they are the dominant vascular plants in grassland or tundra ecosytems. In forest or savanna ecosystems, herbaceous plants exist as important understory elements that provide complexity to the system.

Herbaceous species selected for national monitoring were broadly distributed nationally and include big bluestem (Andropogon gerardii), side oats grama (Bouteloua curtipendula), and switch grass (Panicum virgatum.) These species are present in many states and collectively are included in the majority of grassland types in North America. Seed production of local ecotype and of cultivars and selections of these species are important in prairie ecological restoration, site rehabilitation, rain garden, and landscaping projects. These species are also important for forage and for bio-fuels production projects, and in carbon sequestration studies.

Flowering, seed set and other phenophases are important for the above listed economic conservation, and landscaping purposes. In addition, shifts in phenophases in these species may be sensitive indicators of global climate change trends, especially in areas where these they are abundant and widespread. In such areas, remote sensing in tandum with ground observations, could be a powerful tool in indicating greenup or scenescens, and thus providing convenient indications of climate trends.

Though we recognized usefulness in observation of these species for reasons stated, we also note some potential complications in using these species, especially big bluestem and switch grass. Cultivars and named selections that have been commercially produced and broadly distributed often exhibit very different phenologies than their counterparts existing in local remnant natural communities or plantings from remnant natural communities that are in close proximity to their seed source. For this reason, it will be important to indicate from which situation observations are derived.

It was difficult to identify native forbs with a broad range that would be useful in the first category. Common species widely distributed were the characteristic that we were most looking for in this category. Species selected are listed in Table 2. Very little was known or discussed about how valuable phonological measurements for these species might be. With three important genera we looked for two or three species that could cover the entire country. This is why there are three columbines, three evening-primroses, two claytonias and two trout lilies. Having the three columbine species together substitute for one species added to the first group.

Because of time limitation many of the species on list two were not sufficiently discussed in the group after being suggested. Asclepius tuberose, Heracleum maximum, Caltha palustris, Impatiens capensis, Mertensia virginica, and Podophyllum peltatum were not discussed, nor

32 were the merits of California poppy or fireweed in the third tier. Wild strawberry is widely distributed and was later added because it will compliment data collection on agricultural strawberries.

1 2 3 Scientific Name Scientific Name Scientific Name Andropogon gerardii Aquilegia caerulea Eschscholtzia californica Bouteloua curtipendula Aquilegia canadensis Panicum virgatum Aquilegia formosa Arisaema triphyllum Asclepius tuberosa Caltha palustris Claytonia lanceolata Claytonia virginica Dodecatheon spp. Erythronium albidum Erythronium americanum Fragaria virginiana Heracleum maximum Impatiens capensis Linnea borealis Mertensia virginica Oenothera biennis Oenothera caespitosa Oenothera speciosa Podophyllum peltatum

Table 2. Herbaceous species recommended for use in the NPN Project.

The Importance of a Clonal Backbone for the NPN

An important goal of the NPN is to measure phenology as an indicator of climate variation nationally. All phenological field measurements are phenotypic measurements, in which the phenotype is determined by the organism’s genotype, environment, and the interaction between the genotype with the environment. Many of the tier one plants are clonally propagated plants because they have a consistent or constant genotype, hence their phenotype in diverse environments will reflect the effect of the environment and the interaction of the genotype and the environment, thus allowing more precise measurements of climate variation. The interaction of the genotype with the environment, while confounded with any single clone, can be indirectly measured by taking phenological measurements using multiple species within the environment

33 and comparing these measurements across species and environments. Hence we have proposed more than one species for measurement in the NPN environments.

In addition to clonal plants, highly inbred cultivars (e.g. winter wheat, soybeans, and cotton cultivars) would have a constant genotype due to their homozygosity/homogeneity, as would the hybrid progeny of inbred lines (e.g. single cross hybrid maize). If the clonal or inbred cultivar, or hybrid changes among locations as would be expected in commercial production fields due to differing adaptation requirements (e.g. winter vs. spring wheat, or the phenologically based maturity groupings of soybeans), the genotypes will change. However commercial cultivars and hybrids are often well described in public databases (e.g. plant patents; Certificates of Plant Variety Protection, http://www.ams.usda.gov/science/PVPO/CertificatesDB.htm; or commercial literature). Having a constant clonal lilac throughout all of the network where it is able to be grown is critical to truly take advantage of the benefits a constant genotype as a phenological landmark. Many of the native plants will have poorly known genetic/genotypic population structures, but can be tied to the lilac clones or other backbone plant species.

Useful references:

http://www.ca.uky.edu/agc/pubs/agr/agr184/agr184.pdf Soybean maturity group affecting first flowering date when planted at different planting dates.

http://beaumont.tamu.edu/eLibrary/Reports/2003%20EOY%20Alternate%20Crops%20Report/Avoiding% 20Soybean%20UGC.%201.49.pdf Soybean maturity groups and insect interactions.

http://www.oznet.ksu.edu/kansascrops/soybean_varieties.htm Soybean maturity groups, also protocols for growth

References

Agency Technical Work Group State Of New Mexico. 2005. Potential Effects of Climate Change on New Mexico. New Mexico Environment Department, Albuquerque, NM. Available online at: http://www.nmenv.state.nm.us/aqb/cc/Potential_Effects_Climate_Change_NM.pdf. Ahas, R. 1999. Long-term phyto-, ornitho- and ichthyophenological time-series analyses in Estonia. Int. J. Biometeorol. 42, 119−123. Askeyev, O.V., · T.H. Sparks, · I.V. Askeyev and· P. Tryjanowski. 2005. Is earlier spring migration of Tatarstan warblers expected under climate warming? J Ornithol 146:200- 205. Avirona, S., P. Kindlmann and F. Burela. 2007. Conservation of butterfly populations in dynamic landscapes: The role of farming practices and landscape mosaic. Ecological Modelling 205(1-2):Pages 135-145. Beaumont, L.J., Mcallan, I.A. and L. Hughes. 2006. A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds. Global Change Biology 12(7):1339–1354.

34 Berthold, P., W.Fiedler and Møller, A.P. 2003. Report of the European Science Foundation workshop entitled “Bird migration and climate change”. Available online at: www.ifv.terramare.de/ESF/WS_Konstanz0303.pdf. Berteaux, D., M. M. Humphries, C. J. Krebs, M. Lima, A. G. McAdam, N. Pettorelli, D. Réale, T. Saitoh, E. Tkadlec, R. B. Weladji, and N. Chr. Stenseth. 2006. Constraints to projecting the effects of climate change on mammals. Climate Research 32:151–158. Bezzel, E. & W. Jetz (1995): Delay of the Autumn Migratory Period in the Blackcap (Sylvia- Atricapilla) 1966-1993 - a Reaction to Global Warming? – Journal of Ornithology. 136: 83-87. Blaustein, A.R., B. Han, B. Fasy, J. Romansic, E.A. Scheessele, R.G. Anthony, A. Marco, D.P. Chivers, L.K. Belden, J.M. Kiesecker, T. Garcia, M. Lizana and L. B. Kats. 2004. Variable breeding phenology affects the exposure of amphibian embryos to ultraviolet radiation and optical characteristics of natural waters protect amphibians from uv-b in the u.s. pacific northwest: comment. Ecology 85(6):1747–1754. Both, C. & Visser, M. E. 2001. Adjustment to climate change is constrained by arrival date in a long-distance migrant bird.Nature 411:296–298. Bradley, N. L., Leopold, A. C., Ross, J. & Huffaker, W. 1999. Phenological changes reflect climate change in Wisconsin. Proc. Natl Acad. Sci. USA 96, 9701−9704. Brooker, R. 2005. Climate Change And Biodiversity In Europe: A Review Of Impacts And Policy. available online at: http://www.nbu.ac.uk/biota/Archive_climatechange/ReviewV1.pdf. Brown, J. L., Li, S. H. & Bhagabati, N. 1999. Long-term trend toward earlier breeding in an American bird: a response to global warming? Proc. Natl. Acad. Sci. USA 96:5565–5569. Buckley, L.B. & W. Jetz (2007): Environmental and historical constraints on global patterns of amphibian richness. Proceedings of the Royal Society B: Biological Sciences. Campbell Grant E.H., R.E. Jung and K.C. Rice. 2005. Stream Salamander Species Richness and Abundance in Relation to Environmental Factors in Shenandoah National Park, Virginia. Am. Midl. Nat. 153:348–356. Canelas, M.A.S. and Bertoluci, J. 2007. Anurans of the Serra do Caraça, southeastern Brazil: species composition and phenological patterns of calling activity. Iheringia, Sér. Zool. 97(1):21-26. Chalmers, R.J., and Loftin, C.S., 2006, Hemidactylium scutatum (four-toed salamander): morphology/phenology: Herpetological Review 37:69-71. Cook, D.G., P.C. Trenhamb, and P.T. Northen. 2006. Demography And Breeding Phenology Of The California Tiger Salamander (Ambystoma Californiense) In An Urban Landscape. Northwestern Naturalist 87(3):215–224. Corn, P.S., 2005, Climate change and amphibians: Animal Biodiversity and Conservation 28(1):57–65. Corn, P.S., and Muths, E., 2002, Variable breeding phenology affects the exposure of amphibian embryos to ultraviolet radiation: Ecology 83(11):2958–2963. Corn, P.S., and Muths, E., 2004, Variable breeding phenology affects the exposure of amphibian embryos to ultraviolet radiation – Reply: Ecology 85(6):1759–1763. Corn, P.S., Battaglin, W.A., Hossack, B.R., Muths, E., Olson, D.H., Patla, D.A., Pilliod, D.S., Vredenburg, V.T., 2004, Amphibians and snow – Estimated changes in breeding phenology in the mountains of the western United States: Presentation at Ecological Society of America Annual Meeting, Portland, Ore.

35 Cotton, P.A. 2003. Avian migration phenology and global climate change. Proc Natl Acad Sci U S A. 100(21):12219–12222. Crick, H. Q. P., Dudley, C., Glue, D. E. & Thomson, D. L. 1997. UK birds are laying eggs earlier. Nature 388, 526. Dennis, B., and W.P. Kemp. 1988. Further statistical inference methods for a stochastic model of insect phenology. Environmental Entomology 17:887-893. Dennis, B., W.P. Kemp, and R.C. Beckwith. 1986. A stochastic model of insect phenology: estimation and testing. Environmental Entomology 15:540-546. Donnelly M. A. 1989. Reproductive phenology and age structure of Dendrobates pumilio in northeastern Costa Rica. Journal of Herpetology. 23:362–367. Donnelly M. A. 1999. Reproductive phenology of Eleutherodactylus brandsfordii in northeastern Costa Rica. Journal of Herpetology. 33:624–631. Fitzgerald, L.A., F.B. Cruz and G. Perotti. 1999. Phenology of a Lizard Assemblage in the Dry Chaco of Argentina. Journal of Herpetology 33(4):526-535. Forchhammer, M. C., Post, E. & Stenseth, N. C. 1998. Breeding phenology and climate. Nature 391, 29−30. Forister, M. L., Shapiro, A. M. 2003. Climatic trends and advancing spring flight of butterflies in lowland California, Global Change Biology 9:1130-1135. Frederiksen, M., Harris, M.P., Daunt, F., Rothery, P. & Wanless, S. 2004. Scale-dependent climate signals drive breeding phenology of three seabird species. Global Change Biology 10: 1214-1221. Gibbs, J.P. and A.R. Breisch. 2001. Climate Warming and Calling Phenology of Frogs near Ithaca, New York, 1900-1999. Conservation Biology 15(4):1175-1178. Green, R.E., M. Harley, M. Spalding and C. Zöckler. 1999. Impacts of climate change on wildlife. Published by the RSPB on behalf of English Nature, WWF-UK, UNEP World Conservation Monoitoring Centre and the Royal Society for the Protection of Birds. Available online at: http://www.english-nature.org.uk/pubs/publication/PDF/ImpactsCChange.pdf. Greenberg, R. 1981. The abundance and seasonality of forest canopy birds on Barro Colorado Island, Panama. Biotropica 13:241-252. Hartel, T. 2006. Aspects of breeding activity of Rana dalmatina and Rana temporaria reproducing in a seminatural pond. North-Western Journal of Zoology 1:5-13. Holbrook, S. J., Schmitt, R. J. & Stephens, J. S.Jr. 1997. Changes in an assemblage of temperate reef fishes associated with a climatic shift. Ecol. Appl. 7, 1299−1310. Jakob C., Poizat G., Veith M., Seitz A., and Crivelli A.J. 2003. Breeding phenology and larval distribution of amphibians in a Mediterranean pond network with unpredictable hydrology. Hydrobiologia 499(1-3):51-61. Jonzén, N., A. Lindén, T. Ergon, E. Knudsen, J. O. Vik, D. Rubolini, D. Piacentini et al. 2006. Rapid Advance of Spring Arrival Dates in Long-Distance Migratory Birds. Science 312:1959 – 1961. Kemp, W.P., and B. Dennis. 1992. Toward a general model of rangeland grasshopper (Orthoptera: Acrididae) phenology in the steppe region of Montana. Environmental Entomology 20:1504-1515. Kemp, W.P., B. Dennis, and R.C. Beckwith. 1986. A stochastic phenology model for the western spruce budworm (Lepidoptera: Tortricidae). Environmental Entomology 15:547-554. Kordges, Thomas. 2002. Notizen zur Phänologie und Larvalökologie von Amphibien in Hochgebirgslagen der Alpen [Some notes on phenology and larval ecology of

36 amphibians in mountainous regions of the Alps]. Zeitschrift für Feldherpetologie. 9(119- 123. [[email protected]] Kordges, Thomas and Burkhard Thiesmeier. 2000. Zur Phänologie und Biometrie metamorphosierter Teich- und Bergmolche (Triturus vulgaris und T. alpestris) in einem Abgrabungskomplex in Wuppertal (Nordrhein-Westfalen) [On phenology and biometry of metamorphosed smooth and alpine newts (Triturus vulgaris and T. alpestris) in a pit complex in Wuppertal (Northrhine-Westphalia)]. Zeitschrift für Feldherpetologie. 7(1/2):203-210. [[email protected]] Kneitz, Stephan. 1999. Zur Jahresphänologie adulter Gras- (Rana temporaria) und Springfrösche (Rana dalmatina) an Laichgewässern im Drachenfelser Ländchen südwestlich von Bonn [Annual phenology of adult common frogs (rana temporaria) and agile frogs (Rana dalmatina) at breeding sites in the Drachenfelser Ländschen southwest of Bonn]. Zeitschrift für Feldherpetologie. 6:159-185. Inouye, D. W., Barr, B., Armitage, K. B. & Inouye, B. D. 2000. Climate change is affecting altitudinal migrants and hibernating species. Proc. Natl. Acad. Sci. USA 97: 1630–1633. La Sorte, F.A. and F. R. Thompson III. 2007. Poleward Shifts In Winter Ranges Of North American Birds. Ecology 88:1803–1812. Marra, P. P., Hobson, K. A. & Holmes, R. T. 1998 Linking winter and summer events in a migratory bird by using stable-carbon isotopes. Science 282:, 1884–1886. Marra, P.P., Francis, C., Mulvihill, R. & Moore, F. 2005. The influence of climate on the timing and rate of spring bird migration. Oecologia 142, 307-315. Parmesan, C. 2006. Ecological and Evolutionary Responses to Recent Climate Change. Annual Review of Ecology, Evolution, and Systematics 37:637-669. Mcauley, D.G. and J.R. Longcore. Nesting Phenology and Success of Ring-Necked Ducks in East-Central Maine. J. Field Ornithol. 60(1):112-119. Morin, P.J., S.P. Lawler and E.A. Johnson. 1990. Ecology and breeding phenology of larval Hyla andersonii: the disadvantage of breeding late. Ecology 71:1590–1598. Owen, R. D. 1996. A comparison of breeding phenology and microhabitat requirements among three species of chorus frog from east central Florida. Page 28 (abstract) in Proceedings of the 39th Annual Meeting of the Society for the Study of Amphibians and Reptiles, 24– 29 July 1996, University of Kansas, Lawrence, Kansas, USA. Owen, R. D. 1996. Breeding phenology and microhabitat use among three chorus frog species (Pseudacris) in east-central Florida. M.S. Thesis, University Central Florida, Orlando, Florida, USA. 256pp. Parmesan, C. and J. Matthews. 2006. Biological Impacts of Climate Change. Pages 333-374 in Martha J. Groom, Gary K. Meffe, and C. Ronald Carroll, Principles of Conservation Biology, Third Edition. Sinauer Associates, Inc. Stamford, Connecticut. Parmesan, C. T.L. Root, and M.R. Wil. 2000. Impacts of Extreme Weather and Climate on Terrestrial Biota. Bulletin of the American Meteorological Society 81:443-450. Parmesan C. and G. Yohe. 2003. A globally coherent fingerprint of climate change impacts across natural systems. Nature 421(6918):37-42. Parmesan, C. et al. 1999. Poleward shifts in geographical ranges of butterfly species associated with regional warming. Nature 399, 579−583. Paton, P., Stevens, S. and L. Longo. 2000. Seasonal Phenology of Amphibian Breeding and Recruitment at a Pond in Rhode Island. Northeastern Naturalist 7(3):255-269.

37 Paton, P.W.C, W.B. Crouch. 2002. Using the phenology of pond-breeding amphibians to develop conservation strategies. Conservation Biology 16(1):194-204. Post, E. & Stenseth, N. C. 1999. Climatic variability, plant phenology, and northern ungulates. Ecology 80, 1322−1339. Price, J.T. and P.L. Glick. 2002. The Birdwatcher's Guide to Global Warming. National Wildlife Federation and American Bird Conservancy. Available online at: http://www.abcbirds.org/climatechange/statepage.htm. Price, J.T., T.L. Root, K.R. Hall, G. Masters, L. Curran, W. Fraser, M. Hutchins, and N. Myers. 2000. Review of the potential impacts of climate change on wildlife in ecosystems. Originally prepared as supplemental information for Working Group II, Section 5.4, Third Assessment Report, Intergovernmental Panel on Climate Change. Available online at: http://www.abcbirds.org/climatechange/ecosystem.pdf. Price, J.T. and T.L. Root, 2000. Focus: Effects of climate change on bird distributions and migration patterns. Pages 65-68 in P.J. Sousounis and J.M. Bisanz (eds.) Preparing for a Changing Climate: The Potential Consequences of Climate Variability and Change. University of Michigan, Atmospheric, Oceanic, and Space Sciences Dept., Ann Arbor, Michigan. Price, J. 2000. Climate change and bird distributions in the Mid-Atlantic Region. Page 32 in A. Fisher et al., (eds). Preparing for a Changing Climate: Mid-Atlantic Overview for the National Assessment on the Potential Consequences of Climate Change for the United States. Mid-Atlantic Regional Assessment Report, The Pennsylvania State University, University Park, PA. Price, J. 2000. Modeling the potential impacts of climate change on the summer distributions of Ohio's non-game birds. Ohio Birds and Natural History 2(1): 32-39. Price, J. 2000. Modeling the potential impacts of climate change on the summer distributions of Massachusetts' passerines. Bird Observer 28(4): 224-230. Price, J. 2000. Modeling the potential impacts of climate change on the summer distributions of Michigan's nongame birds. Michigan Birds and Natural History 7(1): 3-13. Price, J. 2000. Modeling the potential impacts of climate change on the summer distribution of Colorado's nongame birds. Journal of the Colorado Field Ornithologists 34:160-167. Price, J. 2000. Climate change and Minnesota birds. Minnesota Birding 37(2): 14-15. Ptaszyk, J., Kosicki, J., Sparks, T.H., and P. Tryjanowski. 2003. Changes in the timing and pattern of arrival of the White Stork (Ciconia ciconia) in western Poland. Journal für Ornithologie 144 (3):323–329. Pearson, S. F., and S. Rohwer. 1998. Influence of breeding phenology and clutch size on hybridization between Hermit and Townsend's Warblers. Auk 115:739–745. Peterson, C.R., and M.E. Dorcas. 1994. Automated data acquisition. pp. 47-57. In Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Heyer, W.R., M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster (eds.). Smithsonian Institution Press, Washington, D.C. Richter-Boix, A., Llorente, G.A. and A. Montori. 2006. Breeding phenology of an amphibian community in a Mediterranean area. Amphibia-Reptilia 27(4):549-559. Riley SPD, Bradley Shaffer H, Randal Voss S, Fitzpatrick BM. 2003. Hybridization between a Rare, Native Tiger Salamander (Ambystoma Californiense) and Its Introduced Congener. Ecological Applications 13(5):1263–1275.

38 Root, T. L., Price, J. T., Hall, K. R., Schneider, S. H., Rosenzweig, C. & Pounds, J. A. 2003. Fingerprints of global warming on wild animals and plants. Nature 421:, 57–60. Roy, D. B. & Sparks, T. H. 2000. Phenology of British butterflies and climate change. Glob. Change Biol. 6, 407−416. Salvador, A. & carrascal, L. M. 1990. Reproductive phenology and temporal patterns of mate access in Mediterranean anurans. Journal of Herpetology 24(4):438-441. Schaefer, H.C., Jetz, W. & K. Böhning-Gaese (in press): Impact of climate change on migratory birds: community reassembly versus adaptation. Global Ecology and Biogeography. in press Scharf, W.C. and G.W. Shugart. 1984. Distribution and phenology of nesting Forster’s Terns in eastern Lake Huron and Lake St. Clair. Wilson Bull. 96(2):309-313. Sæther, B.-E., Tufto, J., Engen, S., Jerstad, K., Røstad, O. W. & Skåtan, J. E. 2000. Population dynamical consequences of climate change for a small temperate songbird. Science 287: 854–856. Sillett, T. S., Holmes, R. T. & Sherry, T. W. 2000. Impacts of a global climate cycle on population dynamics of a migratory songbird. Science 288:, 2040–2042. Sinsch, Ulrich, Verena Lang, Raffaela Wiemer and Simone Wirtz. 2003. Dynamik einer Kammmolch-Metapopulation (Triturus cristatus) auf militärischen Übungsgelände (Schmittenhöhe, Koblenz): 1. Phänologie, Wettereinfluss und Ortstreue [Dynamics of a crested newt metapopulation (Triturus cristatus) at a military training area (Schmittenhöhe, Koblenz): 1. Phenology, climate impact and pond fidelity]. Zeitschrift für Feldherpetologie.10(2):193-210. [sinsch@uni-koblenz-de] Sinsch, Ulrich, Verena Lang and Raffaela Wiemer. 2003. Dynamik einer Kammmolch- Metapopulation (Triturus cristatus) auf militärischen Übungsgelände (Schmittenhöhe, Koblenz): 2. Saisonale Variation der Bestände in zwei Laichgewässern [Dynamics of a crested newt metapopulation (Triturus cristatus) at a military training area (Schmittenhöhe, Koblenz): 2. Seasonal variation of population size in two breeding ponds]. Zeitschrift für Feldherpetologie. 10(2):211-227. Sinsch, Ulrich, Verena Lang and Raffaela Wiemer. 2003. Dynamik einer Kammmolch- Metapopulation (Triturus cristatus) auf militärischen Übungsgelände (Schmittenhöhe, Koblenz): 3. Altersstruktur [Dynamics of a crested newt metapopulation (Triturus cristatus) at a military training area (Schmittenhöhe, Koblenz): 3. Age structure]. Zeitschrift für Feldherpetologie. 10(2):229-244. Sparks, T., Heyen, H., Braslavska, O. & Lehikoinen, E. 1999. Are European birds migrating earlier? BTO News 223, 8−9. Stefanescu, C., J. Penuelas, and I. Filella. 2003. Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin. Global Change Biology 9:1494–1506. Thorne, J. H., O'Brien, J., Forister, M. L., Shapiro, A. M. 2006. Building Phenological Models from Presence/Absence Data for a Butterfly Fauna, Ecological Applications, 16(5):1842– 1853 van Noordwijk, A. J. 2003. Climate change: The earlier bird. Nature 422:29. Vega, J. H. & Rappole, J. H. 1994. Composition and phenology of an avian community in the Rio Grande Plain of Texas. Wilson Bulletin 106, 366-380. Vignoli, L., M.A. Bologna and L. Luiselli. 2007. Seasonal patterns of activity and community structure in an amphibian assemblage at a pond network with variable hydrology. Acta Oecologica 31(2):185-192.

39 Vinogradov, A.E. and O.V.Olga V. Anatskaya. 2004. Phenological resonance and quantum life history. Journal of Theoretical Biology 228 (3):417-420. Visser, M.E. and C. Both. 2005. Shifts in phenology due to global climate change: the need for a yardstick. Proc Biol Sci. 272(1581):2561–2569. Visser, M. E. & Holleman, L. J. M. 2001. Warmer springs disrupt the synchrony of oak and winter moth phenology. Proc. R. Soc. Lond. B 268, 289−294. Visser, M. E., van Noordwijk, A. J., Tinbergen, J. M. & Lessels, C. M. 1998. Warmer springs lead to mistimed reproduction in great tits (Parus major). Proc. R. Soc. Lond. B 265, 1867−1870. Walther, G. R., Post, E., Convey, P., Menzel, A., Parmesan, C., Beebee, T. J. C., Fromentin, J. M., Hoegh-Guldberg, O. & Bairlein, F. 2002. Ecological responses to recent climate change. Nature 416:389–395. Watling, J.I., J. H. Waddle, D. Kizirian, and M.A. Donnelly. 2005. Reproductive Phenology of Three Lizard Species in Costa Rica, with Comments on Seasonal Reproduction of Neotropical Lizards. Journal of Herpetology 39(3):341–348. Weiss, S.B. and A.D. Weiss. 1998. Landscape-Level Phenology of a Threatened Butterfly: A GIS-Based Modeling Approach. Ecosystems 1(3):299-309. Willi, Y., J.V. Buskirk, and A. A. Hoffmann. 2006. Limits to the Adaptive Potential of Small Populations. Annual Review of Ecology, Evolution, and Systematics. 37:433-458. Young, B. E., S. N. Stuart, J. S. Chanson, N. A. Cox, and T. M. Boucher. 2004. Disappearing Jewels: The Status of New World Amphibians. NatureServe, Arlington, Virginia. Available online at: http://www.natureserve.org/publications/disappearingjewels.jsp

40 Appendix List – Appendices are provided in SPWG_Appendices.zip

Appendix 1. Beaubein & Schwartz 2003 list of 17 species by ecoregion

Appendix 2. Beaubein & Schwartz list of 36 species

Appendix 3. Betancourt 2006 list of native species by ecoregion

Appendix 4. Betancourt 2006 list of non-native species by Ecoregion

Appendix 5. McNulty evaluation of trees in Appendix 4 list for SE USA

Appendix 6. Master list of 1,258 species compiled in 2006

Appendix 7. Anderson list of 95 species, 2006

Appendix 8. McKee list of 46 species, 2006

Appendix 9. McKee list of 29 native species, 2006

Appendix 10. McKee list of non-native species by Ecoregion, 2006

Appendix 11. Thomas & Betancourt compilation of lists, 2007

Appendix 12. Thomas & Betancourt prioritization of lists, 2007

Appendix 13. Project BudBurst species list, 2007

Appendix 14. SPWG 2007 list of proposed calibration and focal species

Appendix 15. Cover letter for SPWG 2007 list, version 2 vetting

Appendix 16. SPWG 2007 list, version 2

Appendix 17. Summary of responses to review of SPWG 2007 list, version 2

Appendix 18. USA-NPN Species List, Beta 2008

Appendix 19. Select phenophase information

Appendix 20. Summary notes of SPWG 2007 RCN meeting

41

42