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Combined Report Field Assessments to Assist with Status Updates and Plant Conservation Status Updates II

Prepared For: Wild Resources Conservation Program Grants #013486 and #14513.

Prepared By: Western Conservancy / Pennsylvania Natural Heritage Program

In partnership with: Cleveland Museum of Natural History Carnegie Museum of Natural History Morris Arboretum of the University of Pennsylvania

March 17, 2017

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Table of Contents Introduction ...... 5 Methods ...... 5 Results ...... 6 Table 1. Summary of work completed for each taxon...... 8 Accounts ...... 14 Small-leaved White-snakeroot – aromatica (L.) Spach ...... 15 spp. (Tim Block, due December 15h 2016) ...... 17 Toothcup – Ammannia coccinea L...... 21 Whorled Milkweed – Asclepias verticillata L...... 23 Screw-stem — paniculata (Michx.) Muhl...... 26 Spike Sedge – Carex ormostachya Wieg...... 30 Short’s Sedge – Carex shortiana Dewey ...... 33 Indian-paintbrush – Castilleja coccinea Mutis ...... 35 Short’s Slender Chervil – procumbens (L.) Crantz var. shortii Torr. & Gray ...... 37 Strawberry Goosefoot – Chenopodium capitatum (L.) Asch...... 39 Smartweed Dodder – Cuscuta polygonorum Englemann ...... 41 Laurentian Bladder laurentiana (Weath.) Blasdell ...... 45 Heller’s Witchgrass – Dichanthelium oligosanthes (Schult.) Gould ...... 47 Log Fern – celsa (W.Palmer) Small ...... 50 Clinton’s Fern – (D.C. Eaton) Dowell ...... 52 Schweinitz’s Water-weed – Elodea schweinitzii (J.E. Planchon) Caspary ...... 55 Slender Wheat Grass – trachycaulus (Link) Gould ex Shinners ...... 57 White Trout-lily – Erythronium albidum Nutt...... 59 Godfrey’s – Eupatorium godfreyanum Cronquist ...... 61 Threepetal Bedstraw – Galium trifidum L...... 63 Hawkweed – Hieracium umbellatum L...... 65 Virginia Sweetspire, Virginia-willow, Tassel-white – Itea virginica L...... 67 Grass Rush – Juncus biflorus Elliott ...... 69 Fetter-bush — Leucothoe racemosa (L.) Gray ...... 72 Red Mulberry – Morus rubra Linneaus ...... 74 Sourwood – Oxydendrum arboretum (L.) DC...... 80 Stiff Cowbane – rigidior (L.) Raf...... 82 Yellow-fringed Orchid — Platanthera ciliaris (L.) Lindl...... 84 Bluegrass – paludigena Fern & Wieg...... 87

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Racemed Milkwort – Polygala polygama Walter ...... 91 Lion’s Foot – Prenanthes serpentaria Pursh ...... 93 Swamp chestnut oak – Quercus michauxii Nuttall...... 95 Water-plantain Spearwort – Ranunculus ambigens S. Wats...... 97 Long-stalked Crowfoot – Ranunculus hederaceus L...... 99 Swamp Currant — Ribes lacustre (Pers.) Poir...... 101 Meadow Rose – Rosa blanda Aiton ...... 104 Bristly Blackberry – setosus Bigelow ...... 106 Blue Willow – Salix myricoides Muhl...... 109 Bog Willow — Salix pedicellaris Pursh ...... 112 Blue-eyed Grass – Sisyrinchium montanum Greene ...... 115 Purple Meadow-rue – Thalictrum dasycarpum Fisch. & Avé-Lall ...... 117 Nodding L...... 119 Declined Trillium – Raf...... 122 Appalachian Blue Violet – appalachiensis Henry ...... 127 Netted Chain Fern – Woodwardia areolata (L.) T.Moore ...... 130 Appendix A: Annotated Species List ...... 132 Appendix B: PNHP Field Surveys 2014-2016 for All Plant Status Update Projects Taxa...... 138

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Introduction The purpose of these projects is to provide improved documentation of proposed legal status for plant taxa identified as high priority by DCNR. The official status of in Pennsylvania as approved by legislative action under Chapter 45 of the Pennsylvania Code has not been updated since 1992. However, the Technical Committee (VPTC) of the Pennsylvania Biological Survey (PABS) has been updating the status of Pennsylvania’s rarest species for the entire period of time the act has been in effect. DCNR has reviewed the numerous species whose status has been proposed to change by the VPTC, and identified some which require additional work to document and justify the proposed status. A list of 56 high priority taxa was provided at the start of the 2014 WRCP project Plant Status Updates Collaborative, and work on these taxa has continued through the 2015 WRCP project Plant Status Updates II. These taxa require some combination of additional fieldwork and / or documentation of existing knowledge about the status of the species, and some cases are also complicated by taxonomic problems or questions around the native status of Pennsylvania populations.

The Plant Status Updates projects brought together major institutions with botanical expertise in Pennsylvania to evaluate these taxa and determine how best to resolve any information gaps. Project partners were the Pennsylvania Natural Heritage Program, Morris Arboretum at the University of Pennsylvania, the Cleveland Museum of Natural History, and the Carnegie Museum of Natural History.

Methods

Expert Review of Taxa The list of taxa provided by DCNR was initially reviewed by a group of experts from the partner institutions including Dr. Timothy Block (Morris Arboretum), Jim Bissell (Cleveland MNH), Steven P. Grund (PNHP), John Kunsman (PNHP), Jessica McPherson (PNHP), Dr. Cynthia Morton (Carnegie MNH), and Loree Speedy (Carnegie MNH). The group briefly summarized the state of existing knowledge for each taxon, and decided whether additional fieldwork was needed to assign legal status and provide appropriate justification. The taxa that needed additional fieldwork were each assigned a lead investigator to coordinate work and write summary justifications.

Review of Historical Records PNHP provided all investigators with exports of all Biotics records for the taxa for which they were designated as leads. Lead investigators were responsible for reviewing all historic records and annotating each according to their assessment of whether there is still a reasonable chance of relocating the occurrence. Historical rescords with annotation are listed in an accompanying document. Individual records were not reviewed for some species that had a large number of specimens collected in SE Pennsylvania, because a number of factors make historical specimens an unreliable and inefficient method of locating populations in that region. These factors include vague location data in combination with very extensive landscape alteration, as well as many instances of repeat collection from the same locations by multiple people without clear label annotations (Block and Kunsman, personal communication). For taxa where this is relevant, it is noted in Table 1.

Fieldwork Potential fieldwork sites were identified by a combination of historical record review and use of other sources (aerial photographs, geology maps, personal experience, etc.) to identify potential habitat for de

5 novo surveys. Experts partitioned fieldwork regionally according to the location of their home office; in cases where the state range of taxa extended beyond the region of the lead investigator, experts in other regions identified field sites and completed the field visits. Jim Bissell did field surveys in the NW and to a lesser extent the NC region; Tim Block did field surveys in the SE region; Loree Speedy did field surveys in the western 2/3 of the state; PNHP staff worked throughout the state. Field surveys were conducted according to standard PNHP protocols. Results of all field visits have been submitted to PNHP data management for processing and entry into Biotics.

Documentation of Legal Status Justifications The DCNR’s standards for documentation of legal status justification continued to evolve during the course the Plant Status I and II projects, and therefore slightly different documentation is provided for taxa addressed in the two project rounds. The NatureServe rank calculator spreadsheet and a written summary justification were completed for all taxa where legal status could be assigned. For those taxa addressed in Plant Status I, a spreadsheet provided by the DCNR to summarize characteristics of the species was also completed. Where sufficient information is not currently available about a taxon to reliably assign it a legal status, a written summary of the current state of knowledge and the most critical knowledge gaps was provided.

The Plant Status II project continued fieldwork for taxa where more work was still needed after Plant Status I, and also provided rank calculator spreadsheets and written summary justifications for the taxa on the DCNR’s list that did not require additional fieldwork. The taxa that did not require fieldwork were divided between PNHP staff and Morris Arboretum based on staff expertise and taxon range within the state. All documentation was reviewed by multiple experts as noted in the individual materials.

Review by Vascular Plant Technical Committee / Rare Plant Forum In cases where the work completed as part of Plant Status I and II indicated that a change should be made in the current PABS-recommended status, changes were proposed at the Rare Plant Forum and considered by the VPTC. Results are summarized in table I. Proposed status changes for a few taxa will be considered at the 2017 rare plant forum based on work completed in 2016.

Results

Table 1 summarizes the work completed for each taxon on the list initially provided by DCNR. Deliverables (underlined headings) are described below; accompanying documents are listed in bold.

Annotated list of Taxa: Appendix A: Annotated Species List shows the expert annotations of the list initially provided by DCNR. This information is more fully developed in the status justification summaries included in this report, but the annotations are included because they are listed specifically as a deliverable.

Status Justifications: • Written summaries are included in this document for each taxon. The written summaries are designed as stand-alone documents, with references listed below each summary rather than at the conclusion of this report. • The NatureServe Rank Calculator spreadsheet results are summarized in Table 1, and the full worksheets are provided in an accompanying document, element_rank_estimator_PNHP_Master_v3.185.xlsm

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• The spreadsheet provided by DCNR is submitted as an accompanying document, DCNR_Worksheet_Justifications-combined.xlsx

Annotated historical records for taxa requiring fieldwork are provided in an accompanying document, historical_record_review_deliverable.xlsx

Documentation of Field Surveys Appendix B lists the PNHP field surveys completed during the course of the Plant Status Updates I and II projects for all taxa on the list provided by DCNR.

Summaries of Carnegie Museum surveys and Cleveland Museum surveys are submitted separately in the following documents: • Cleveland Museum Plant Status I Report.docx • Cleveland Museum Plant Status II Report.docx • Carnegie Museum Plant Status Surveys 2014.xlsx • Loree Speedy 2016 Plant Status Surveys.pdf (this document shows the field survey codes and the locations for 36 unsuccessful surveys for conducted in the spring of 2016).

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Table 1. Summary of work completed for each taxon.

CurrentProposed StatusFieldwork StatusStatusRank summary DCNRCalculator spreadsheetReview Historics VPTC/RPF Review needed? Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? Ageratina N PT X X X Considered at 2017 Rare Plant Forum; Historical locations are of limited use for this species aromatica proposed status changed PR->PT due to its early successional habitat. Amelanchier N PE Summary of conservation needs for the canadensis genus Amelanchier in PA, rectification of current PA with new FNA taxonomy. Individual taxa not assessed. Amelanchier TU PE (as above) humilis Amelanchier TU PE (as above) obovalis Amelanchier TU PE (as above) sanguinea Ammannia PE W X Considered at 2017 Rare Plant Forum; Historical locations are of limited use for this species coccinea L proposed status changed PT->WL due to its early successional habitat. Asclepias N PT X X S2 X X 5 historical locations are in biotics, and these were verticillata reviewed for viability. There are additional museum specimen records along the Delaware River and extreme SE PA (about 40 occurrences in total) that have not been entered into biotics because they are judged to have minimal value for relocation.*

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Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? Astragalus N PE X Considered at 2014 RPF; proposed status See proposal materials for evaluation of historical canadensis changed from TU to PE. Proposal materials records. 26 of the 27 historics are pre-1950, and this available as status justification. species requires open, early successional habitat; it is unlikely that habitat remains suitable at historical locations after such a long period of time. This species also occurs on limestone, and limestone habitats have been converted at higher rates than other geology types in Pennsylvania, hence it is likely many have been lost to habitat conversion. Bartonia N PR X X X Potentially review Historical records not reviewed individually, because paniculata this again they are not considered an important source for eventually. potential extant populations of this species; there are relatively few, and all historical records are in SE PA*, and new populations are being found in NE and NC regions. Carex N T X X S1 X X Proposed PT->PE at RPF 2015, but VPTC ormostachya U determined TU. Proposed status changed TU->PT at 2017 Rare Plant Forum.

Carex N PR X X X shortiana Castilleja TU PT X S2 Individual records not reviewed, considered unlikely coccinea to persist due to successional nature of species

Chaerophyllum N PE X X S1S2 X X PABS-recommended status changed from No - current There is only one historical specimen, and the site procumbens TU to PE at 2013 Fall VPTC meeting. recommended was visited (Bedford County); habitat now converted. var. shortii status of PE supported in summary sheet. Chenopodium TU PE X SH No - this species Individual records not reviewed, considered unlikely capitatum should be PH, and to persist due to successional nature of species from a practical perspective the current rec of PE is a reasonable surrogate (or PX). 9

Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? N T Not addressed due to time limitations. No - more work dilatata U needed on Crataegus Crataegus N T Not addressed due to time limitations. No - more work pennsylvanica U needed on Crataegus Cuscuta corylii TU PX X PABS-recommended status changed Considered unlikely to persist due to successional PT->PX at 2014 RPF. Proposal materials nature of species available as justification. Cuscuta TU PT X S1 Individual records not reviewed, considered unlikely polygonorum to persist due to successional nature of species Cystopteris TU PE X X S1 X laurentiana Dichanthelium N PT X X S2 X X Yes- review 9 historical locations are in biotics, and these were oligosanthes varieties and reviewed for viability. There are additional museum assign status (PE specimen records along the Delaware River and for var. extreme SE PA (about 40 occurrences in total) that scribnerianum, PT have not been entered into biotics because they are for var. judged to have minimal value for relocation.* oligosanthes). Species rank OK. Dryopteris N PE X X S1 X celsa Dryopteris N PT X X S2 X clintoniana Elodea PX T X N/ Determined that there is not reliable No historics to review. schweinitzii U A evidence this taxon exists: PABS recommended status changed from TU to not present in Pennsylvania in 2017. Elymus N T X X Summary of current information, no Eventually - not trachycaulus U status recommended. now; wait for more data to come in and revisit in a couple of years. Erythronium X X S3 X X albidum Eupatorium N PT X S2 Individual records not reviewed, considered unlikely godfreyanum to persist due to successional nature of species 10

Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? Galium N PR X X S3 X trifidum Hieracium N PE X S1 Not a fieldwork species, historical records not umbellatum reviewed. Itea virginica PX PE X S1 Not a fieldwork species, historical records not reviewed. Juncus biflorus TU PR X X S3 X X PABS-recommended status changed Western and central records reviewed; not SE.* PT->PR at 2015 RPF/VPTC meeting Leucothoe TU PR X X S3 X PABS-recommended status changed Individual historical records not reviewed, located in racemosa PT->PR at 2015 RPF/VPTC meeting. extreme SE PA* Morus rubra N PT X X S2 Eastern historical records not reviewed individually*

Oxydendrum TU PR X X S4 X arboreum Oxypolis TU PR X X S3 X PABS-recommended status changed Reviewed western and central records, not eastern rigidior PT->PR at 2014 RPF/VPTC meeting ones.* Parthenium TU PE Not yet addressed due to time constraints. integrifolium Platanthera TU PT X S2 Not a fieldwork species, historical records not ciliaris reviewed. Poa PT PR X X S2S3 X paludigena Polygala TU PE X X S1 X polygama Prenanthes N PT X X No - wait a few serpentaria years for more data, then reassess. Quercus N PE X X S1 X No historical records to review. michauxii Ranunculus N T X X S1 X X PABS-recommended status changed Did not review individually; as a group, the historical ambigens U TU->PT at 2017 RPF/VPTC meeting locations are old - ca 40 percent from 1900 or previous and ca 84 percent from 1950 or previous. Surveys at locations of some historical records were unsuccessful. Most have not been surveyed, but their age and accessibility argue against the likelihood of confirming very many of them.

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Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? Ranunculus PX PE X Not at this time; hederaceus further work needed to determine nativity. Ribes lacustre TU PE X X X Rosa blanda N T Summary, no status can be assigned at Historical records in vague or disturbed sites, of U this time. questionable value for relocating extant populations. Rubus setosus TU PE X X S2 X Yes - but wait a few years to see if we find more; PABS status is PE, summary sheet recommends PT Salix N PE X X S1 X myricoides Salix N PE X S1 No historical records to review. pedicellaris Sisyrinchium N T Summary, no status can be assigned at Historical records not likely to remain due to early montanum U this time. successional nature of species. N PT X PABS-recommended status changed Annotated historical records included in proposal. borealis TU->PT in 2014. Thalictrum N T X SH Herbarium specimens reviewed to One historical location, addressed in summary sheet. dasycarpum U accurately ID historical records. PABS- recommended status changed TU->PX at 2017 RPF/VPTC meeting Trillium N PT X S2 Not needed - historical collections mainly from SE cernuum PA*, better use of time to do de novo surveys. Trillium N PT X X Addressed under erectum L.× Trillium flexipes flexipes Trillium TU PT X X S2 X X flexipes Viola PT PR X X X X VPTC recommended status change TU->PR appalachiensis in 2014. Viola renifolia TU PE X X S1 X X

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Scientific CS PS FW SS RC DS RH Other work VPTC/RPF Review Historical Record Review Notes Name needed? Woodwardia N PT X X S3 X PABS-recommended status changed Individual historical records not reviewed because areolata PT->PR at 2015 RPF/VPTC meeting not likely to be a significant source of extant populations; most are located in extreme SE PA* *Historical records from specimens collected in SE PA are judged to be of little value for locating extant populations, because this region has seen a very high rate of landscape change and a great increase in invasive plants in recent decades, and also because of a pattern of frequent repetitive collections from the same populations in combination with vague labelling of historical collections.

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Species Accounts

The following pages contain stand-alone accounts for each taxon. In most cases these provide documentation and rationale to justify the recommended legal status. In a few cases they do not recommend a legal status at this time, but outline what is currently known and make recommendations for gathering any additional information needed in order to be able to assign a status.

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Small-leaved White-snakeroot – Ageratina aromatica (L.) Spach Author: Dr. Timothy Block, Morris Arboretum, Jessica McPherson, PNHP.

Introduction Small-leaved white-snakeroot (Ageratina aromatica) is a plant of mostly dry, often sandy, low nutrient soils on open sites and is distinguished from the more common A. altissima by habitat type (A. altissima on moist, wooded sites) and smaller with shorter petioles. Specific identification can be problematic due to the occurrence of hybrids between these species where their ranges overlap, including Pennsylvania. Hybrids possess characteristics intermediate between the parent species.

Proposed PA Status Small-leaved white-snakeroot is proposed to be classified as Pennsylvania Threatened (PT) due to a low number of populations known in the state and a possible decline and range contraction within the state. The PABS Vascular Plant Technical Committee reviewed the status in 2017 and recommended PT at that time.

Recent field and herbarium work has highlighted the need for this species to receive more attention.

Habitat All of Pennsylvania’s known extant sites for this species are on serpentine barrens in the southeastern part of the state. This species was once much more widespread, occurring on dry, open sites as far west as Bedford County and as far north as Lackawanna County.

North American Distribution and Global / Regional Conservation Concerns Small-leaved white-snakeroot is widely distributed in the southeastern ; its range extends from Louisiana east to Florida and north to , but it appears to become more restricted to the coast from Pennsylvania northwards (Kartesz 2003), and it is rare in all states in this portion of its range. Natureserve ranks small-leaved white-snakeroot as Vulnerable (S3) in Pennsylvania and Critically Imperiled (S1) in , , Massachusetts, and West Virginia, and Imperiled (S2) in . Its global rank is given as secure (S5), as the species is considered more common in the southeastern U.S. There is, however, some nomenclatural confusion as this species is treated to varietal level by some authorities and not by others.

Extant Locations There are currently 7 known extant sites in Pennsylvania; 4 in Chester County, 1 in Delaware County, 1 on the Northampton/Carbon county border, and 1 in Fulton County. Three (3) sites (Fulton, Chester, Delaware) have populations of 50 or fewer individuals, which are small enough for there to be some concern about long term viability, especially the Fulton County site. Three (3) sites in Chester County have larger populations with good viability; 2 sites with several hundred plants, and 1 site with a population of 5000-10,000 individuals. The Northampton/Carbon County site does not have census data recorded. This plant’s major habitat at the present time, serpentine soils in SE Pennsylvania, has been fairly comprehensively surveyed, so few additional populations are likely to be documented at sites of this nature.

Historical Locations and Potential for Relocation Many historic locations have been rechecked without success. However, due to the early successional nature of the species’ habitat, and the likelihood of successional regrowth in the intervening years since historical specimens were collected, relocation of plants at the same locations as historical collections

15 are viewed as unlikely. The southeastern collections in particular are viewed as unreliable sources for present-day leads because of vagueness in locational data, redundancy in collection, and high rate of habitat conversion in that region. It appears that the species may have historically been collected from generic early successional habitats, but today it is known only from serpentine habitats. Because no information about population sizes, and very little habitat data, is typically provided with historical specimens, little is known about the sites where collections appear to have been made outside of serpentine habitat in the past, or how many plants may have been present at the time.

The reasons for the decline of small-leaved white-snakeroot in Pennsylvania may be related to woody plant succession and/or habitat loss.

Threats The primary threat to small-leaved white-snakeroot in Pennsylvania is probably woody plant succession. However, most serpentine barrens sites are protected as preserves by conservation organizations and mostly actively managed to maintain open habitat, thus helping to ensure the long-term survival of the species.

Status Justification Pennsylvania Threatened is the most appropriate status for this species, because of its restriction to serpentine habitats, and the small number of known sites despite fairly comprehensive surveys of habitat in the appropriate region of the state. While the total state population is on the larger end of what is typical for this status, the unevenness of population size between sites (one very large population, a couple of good-sized populations, and three very small populations) indicate that many of the known extant sites still have potential viability concerns. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and . Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Amelanchier spp. Author: Dr. Timothy Block, Morris Arboretum. Editor: Jessica McPherson, PNHP. Editors’ note: this account has been somewhat extensively edited to include expanded explanation of the technical complexity of the genus, and also to incorporate information from the most recent treatment of Amelanchier published in the volume of the New Manual of the Vascular Plants of Northeastern United States and Adjacent Canada (Campbell and Burgess 2017). A follow-up publication to this treatment is set to be published in 2018, and this account should be revisited at that point.

Amelanchier is a genus in which the frequency of asexual reproduction, polyploidy, and sexual reproduction across ploidy levels complicate the application of the traditional species concepts of reproductive isolation in combination with morphological distinction. Because understanding of the ploidy levels within the genus, whether particular taxa commonly reproduce sexually or asexually, and which mechanisms of genetic exchange commonly occur between and within ploidy levels have only been elucidated relatively recently, and because differences of opinion exist on how best to apply the species concept in this context, taxonomy has been revised and re-revised, in sometimes contradictory ways. Historically, various authors have introduced substantial differences of opinion as to the number of species to include in a taxonomic treatment, as well as varying circumscriptions of species included. In eastern , Amelanchier is an actively evolving genus. Distinct taxa do exist, but they are not always completely isolated from other taxa; exchanges of genetic material are occurring, and intermediate individuals that cannot be classified to existing taxa are common. Because of these complications, it is not possible to create a traditional taxonomy that will resolve individuals to a taxon concept cleanly in all cases. However, recent research shows promise in unravelling the complexities and creating a taxonomic framework that, while non-traditional, reflects the reproductive systems that are occurring and accommodates the existence of intermediates.

In addition to reproductive complexity and the existence of intermediate individuals, the subtlety of morphological distinctions in the genus creates difficulties for understanding taxon distribution. Campbell, et al (2014) note that relatively few morphological characters in Amelanchier are taxonomically informative. They state that:

“Identification from herbarium specimens is often inconclusive. Identification is best undertaken in the field, with visits during flowering and fruiting seasons, and observations of habitat, habit, presence of congeners, and flowering time relative to sympatric congeners.”

When the herbarium specimen record cannot be relied upon, it is difficult to obtain a clear picture of the distribution of various taxa within the state. Ongoing research by Campbell and Burgess is providing some elucidation of patterns, but field identification of individual plants as they are discovered remains problematic.

The taxonomic approach that Campbell and Burgess have taken in recent research is to recognize species first among diploids, and then separately for polyploids using different criteria; and to also recognize species complexes as containers for intermediate individuals. Most taxonomists in the past have included both diploids and morphologically similar polyploid plants that are triploid or tetraploid in the same species concept (see Table 1 for ploidy levels of Pennsylvania species).

Diploids of an Amelanchier species can be somewhat morphologically diverse, but all qualify as good species under one or more species concepts when considered separately from their conspecific

17 polyploids (Burgess, et al, 2015). Burgess, et al also note that the distributions of diploid species populations are generally allopatric, and even where sympatric, don’t tend to produce large numbers of hybrids (see distribution maps in Burgess et al. 2015; BONAP distributions reflect multi-ploidal species concepts). A diploid treatment of eastern species has been released, and treatment of polyploids is scheduled to be released within the next year; we recommend waiting on this treatment for further action on this taxon.

According to Burgess, “while Amelanchier polyploids are almost exclusively apomictic, diploids are almost exclusively sexual.” However, importantly, tetraploids and triploids can sexually reproduce with each other and with diploids at low frequency. Once established, this combination apomixis and low frequency of sex can lead to rapid morphological diversification, intergradation, and the formation of “hybrid swarms.”

Burgess, et al. further note that eastern Pennsylvania is a location of some diploid species sympatry and an exceptionally high number of triploid occurrences. Most notable are sympatry of Amelanchier arborea with A. bartramiana and A. humilis, and many hybrids among these species have been reported. Eastern Pennsylvania is also known to be a hotspot for Amelanchier polyploids.

A treatment of polyploid taxa forthcoming in 2018 will assign species status to polyploids with geographic ranges and levels of morphological distinction that are comparable to that of the recognized diploid species, while species complexes will be retained to accommodate intermediate individuals and “microspecies” (polyploids which have spread to some extent, but only have a small geographic range). Their research shows that morphological distinctions do tend to exist on average between diploids and polyploids (Campbell and Burgess 2017); however, the key written for the Amelanchier treatment in New Manual of Vascular Plants of Northeastern United States and Adjacent Canada based on this work includes considerable overlap in characters, such that in some cases it may not be possible to determine whether an individual is a member of a recognized species or a .

Campbell, et al (2014) note that morphologically distinct phenotypes of polyploid Amelanchier could be considered as taxonomically distinct entities if widespread enough, or that, if less widespread or less distinct, could be considered as “microspecies.”

The Flora of North America treatment of Amelanchier was published in 2014 (Campbell, et al, 2014), and reconciliation of the taxonomic treatment of the species of Amelanchier as presented in Rhoads and Block (2010) with recent treatment of the genus in Flora of North America for Pennsylvania is relatively simple and is presented in table 1. The Campbell, et al treatment will be adopted in future editions of The Plants of Pennsylvania until more research leads to a clearer picture of the genus.

Editor’s postscript: reconciliation of the taxa with the NE flora treatment, along with listing of ploidy level, has also been added to table 1.

Table 1. Taxonomic correlation (* denotes taxonomic adjustment; * for ploidy level indicates plants at that level are self-incompatible and reproduce sexually)

Rhoads and Block, Flora of North Ploidy NE Flora Manual (in State PBS G- S- 2010 America, vol. 9, press) status status rank rank 2014

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Amelanchier Amelanchier 2x, 4x A. arborea (2x), A. N N G5 S5 arborea (F. arborea (F. Michx.) canadensis complex Michx.) Fernald Fernald (4x) A. bartramiana A. bartramiana 2x*, 3x, 4x A. bartramiana (2x), PE PE G5 S1 (Tausch) M. Roem. (Tausch) M. Roem. A. bartramiana complex (3x, 4x) *A. canadensis (L.) A. canadensis (L.) 2x*, 4x A. canadensis var. N PE G5 S1 Medik. Medik. var. canadensis (2x), canadensis A. canadensis complex (4x) A. humilis A. humilis 2x, 3x, 4x A. alnifolia (2x), TU PE G5 S1 Wiegand Wiegand polyploids morphologically similar (3x, 4x) A. laevis Wiegand A. laevis Wiegand 2x, 4x A. laevis (2x), N N G5 S5 A. canadensis complex (4x) *A. obovalis A. canadensis (L.) 2x*, 3x (1 A. canadensis var. TU PE G4G S1 (Michx.) Ashe Medik. var. specimen) obovalis (2x), 5 obovalis (Michx.) A. canadensis BSP complex (3x) A. sanguinea A. sanguinea 2x, 3x, 4x A. sanguinea TU PE G5 S2 (Pursh) DC. (Pursh) DC. complex *A. stolonifera A. spicata (Lam.) K. 3x, 4x A. lucida complex N N G5 SNR Wiegand Koch

Conservation of groups like Amelanchier, of which individuals often resist resolution to the species level due to a high rate of evolutionary flux, is not a straightforward matter. The mindset of protecting species alone is misplaced here. Rather, we must adopt the philosophy and practice of protecting evolutionary process and potential. While I realize that this doesn’t mesh with existing law and regulations, it is, for the reasons given above, the only biologically supportable path forward. Toward this end, below are a few recommendations that will help protect the future of this very complex genus.

Conservation recommendations: 1. Protect all populations of rare species taxonomic elements that appear to be rare, to the best of our abilities to identify them using morphological characters. 2. Protect all putative “hybrid” individuals suspected to be of rare species parentage, as these plants could represent sources of novel speciation events (see the “triploid bridge hypothesis” of Burgess, et al, 2014). 3. Protect suspected diploid populations of all rare species, at least until ploidy level can be determined. 4. Protect distinct phenotype populations, both those covering small areas (potential microspecies) or larger areas (potential taxonomically recognizable) of all rare species. 5. Promote funding for continued systematic research, including morphometric, cytometric, and genetic studies of Amelanchier in Pennsylvania, a known “hotspot” for diversity.

Modifications of these recommendations, and perhaps others, should be applied when considering other biologically similar genera. Such genera include (but not limited to) Crataegus and Rubus in Rosaceae, and Hieracium, Antennaria, Symphyotrichum, and others in .

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Editor’s postscript: The treatment of Amelanchier in the New Manual of Vascular Plants of Northeastern United States and Adjacent Canada has also recently been released, which recognizes diploid species and puts the polyploids into species complexes. A future paper by the same authors in Systematic Botany to be released in 2018 will address the polyploid taxa, assigning species status to those with sufficient geographic range and morphological distinction (Burgess, personal communication). We recommend revisiting the Pennsylvania taxonomy upon the release of this treatment. The completion of this work may also give new insights into the distribution of our taxa and which ones are rare in the state. The approach of recognizing species for diploids and for widespread, morphologically distinct polyploids also suggests a focus for state conservation efforts on these units.

Literature cited:

Burgess, M.B., K.R. Cushman, E.T. Doucette, N. Talent, C.T. Frye, and C.S., Campbell, 2014. Effects of apomixis and polyploidy on diversification and geographic distribution in Amelanchier (Rosaceae). American Journal of Botany 101(8): 1375-1384.

Burgess, M.B., K.R. Cushman, E.T. Doucette, C.T. Frye, and C.S. Campbell, 2015. Understanding diploid diversity: a first step in unraveling polyploid, apomictic complexity in Amelanchier. American Journal of Botany 102(12): 2041-2057.

Campbell, C.S., M.B. Burgess, K.R. Cushman, E.T. Doucette, A.C. Dibble, and C.T. Frye, 2014. Amelanchier in Flora of North America North of Mexico, volume 9, pg. 646-661. Oxford University Press, New York.

Campbell, C.S. and Michael B. Burgess. 2017. “Amelanchier” in Rosaceae, the Rose Family. Genus treatment in echapter of New Manual of Vascular Plants of Northeastern United States and Adjacent Canada. Ed. Robert F.C. Naczi. NYBG Press, ebook,

Rhoads, A.F. and T.A. Block, 2010. The Plants of Pennsylvania: An Illustrated Manual, 2nd edition. University of Pennsylvania Press, Philadelphia.

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Toothcup – Ammannia coccinea L. Author: Dr. Timothy Block, Morris Arboretum

Introduction Like the closely related Rotala ramosior (also commonly called toothcup), Ammannia coccinea is a true annual. The most obvious distinguishing field characteristic is the presence of clustered in the axils in Ammannia coccinea, while the flowers are solitary in the leaf axils in Rotala.

Ammannia coccinea has been considered by some authors (Gleason and Cronquist, 1991) to be a species of hybrid origin, with A. auriculata and A. robusta as parents; it is diploid, as are the parent species.

The above noted species also share a propensity for wet, sandy, open, often highly disturbed habitats, and co-occur in some places over their shared range.

Proposed PA Status Ammannia coccinea should not be considered for conservation concern in Pennsylvania. Neither should this species be considered native to Pennsylvania in the strict sense. While native further south, its earliest occurrence in PA was of dubious origin at the Philadelphia Navy Yard in 1866. No collection records exist for this species from that time until 1967 when it was collected by Edgar Wherry in a disturbed park setting near Shawmont in Philadelphia County.

Ammannia coccinea is considered non-native in and Delaware where it is abundant. It’s also listed as exotic in Hawaii.

Habitat As noted above, Ammannia coccinea is a species of disturbed, wet, open areas where competition from other vegetation is limited.

North American Distribution and Global/Regional Conservation Concerns Ammannia coccinea is widely distributed across the southeastern U.S. and as far west as California, and south into Mexico and tropical America.

This species is of no conservation concern throughout most of its range. It is, however, currently listed as S2 in PA, WV, and NC, S3 in VA, and S4 in IA, KY, and GA.

Pennsylvania Distribution Pennsylvania is at the northern edge of this species’ range. Most collections of Ammannia coccinea are from SE PA, but at least two recent collections are from SW PA, occurring on disturbed stream banks. This species might be expected any place where appropriate habitat exists.

Extant Locations At least 14 locations exist in Pennsylvania that were collected from within the last 25 years. Population sizes range from a few to thousands of individuals per population.

Historical Locations and Potential for Relocation Relatively few truly historical locations exist for this species and attempting to relocate those would probably be irrelevant.

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Threats Assuming, for some reason, this species was to be considered of conservation concern, the only substantial threat to occurrence sites would be increasing competition from taller vegetation.

Status Justification As Ammannia coccinea is an abundant, often weedy, plant of highly disturbed habitats throughout its range, and was not present as a naturalized part of the Pennsylvania flora until the 1960s (perhaps later), it seems unreasonable to consider this species as strictly native to Pennsylvania, let alone to consider it as a species of special concern.

This species’ occurrence and distribution in Pennsylvania is likely the result of human mediated events rather than more natural causes.

References • Gleason, H.A. and A. Cronquist, 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd edition. New York Botanical Garden, New York.

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Whorled Milkweed – Asclepias verticillata L. Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP

Introduction Whorled milkweed is an herbaceous perennial characterized by its milky sap, slender stems, narrowly- linear whorled leaves, white or greenish-white flowers grouped in an umbel-like cluster, and pod-like fruit containing seeds that have a tuft of hairs that facilitate dispersal. It can often be identified in the vegetative condition because of the milky sap and leaf arrangement and shape.

Proposed PA Status This species was proposed and accepted as Pennsylvania Threatened at the 2013 Rare Plant Forum/PABS Vascular Plant Technical Committee meeting. It was previously on the Watch list. Whorled milkweed fits the criteria of the PT status because of its generally low population numbers and restricted “barrens” type habitat, which usually has early successional conditions and often requires active management by fire, mowing, or scraping in order to eliminate or keep in check competing plants, particularly woody exotics or aggressive native species. At a number of sites, active management has not been available, and the long-term viability of the whorled milkweed is therefore jeopardized.

North American Distribution and Global / Regional Conservation Concerns On a global basis, the species is considered to be globally secure (G5) and is distributed from southern New England to the Gulf Coast and west to the prairie states, and is by far most frequent in the latter (Kartesz 2003). In the Mid-Atlantic portion of its range, which includes Pennsylvania and adjacent states, it is considered to be Threatened (New Jersey, Pennsylvania), Rare (, New York, West Virginia), or Historical (Delaware).

Habitat and Pennsylvania Distribution Whorled milkweed grows in well drained, sunny conditions, and has a definite preference for calcareous substrates such as serpentine, limestone, diabase, and certain types of shale. It is a very characteristic, although usually not abundant, component of the flora of serpentine grasslands. To the north and west of the serpentine region, the plant inhabits limestone and diabase glades, shale barrens, open woodlands, “prairies,” rock outcroppings, and even a few human-caused disturbances, such as grazing areas associated with agriculture, if the proper substrate is present. The species apparently does not thrive in acidic conditions and so is seemingly absent from habitats, such as scrub oak barrens, that might superficially appear to have potential for it.

Extant Locations The current number of extant locations in Pennsylvania, based primarily on PNHP field forms and other recent observations from the last 30 years, is conservatively estimated in the 20 to 26 (30) range. Most of the locations and the majority of individual plants are at serpentine barrens. The larger, more accessible barrens with numerous actively managed grasslands, such as Nottingham and New Texas, probably support hundreds of stems of whorled milkweed, and the species seems secure at these locations. The populations at the smaller, less accessible and less managed barrens, most of which have not been visited by PNHP in recent years, probably only contain a fraction or less, maybe much less, of that number of stems. Even if the smaller barrens were to have active management, the populations of whorled milkweed would probably remain relatively small because these sites are limited in area and because it is the nature of the species to be relatively infrequent. Overall, the serpentine barrens, particularly the larger ones, appear to be the stronghold of this species in the state, and the species will probably always be present.

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Much less is known about the current status of whorled milkweed at the relatively few extant sites occurring outside of the serpentine region. The species was confirmed growing in a diabase grassland in Adams County in 2014, but the other non-serpentine sites known to PNHP have not been visited in recent years or have been incompletely surveyed. These sites tend to be small in area and in the number of stems present, receive minimal if any active management, have major invasive plant problems, and are located on often inaccessible private land. In addition to the documented sites, there is potential for finding new occurrences of the species in the limestone and diabase regions, but it seems unlikely that there will be any significant number. If attempting the conserve the overall range of the species in the state is important, then these non-serpentine locations can’t be ignored, and this increases the degree of threat.

Historical Locations and Potential for Relocation There appear to be 50-100 historical locations, depending on how an occurrence is defined, of whorled milkweed in Pennsylvania. Most of these herbarium specimens were collected in Chester, Delaware, and Lancaster counties, because of the presence of serpentine and the zealous collecting of specimens by Philadelphia area botanists from the late 1800s to the early 1900s. There are records from ca 15 counties to the north and west of the above three counties. The ultimate value in taking the time and expense to attempt to visit each of the historical sites and confirm the presence of whorled milkweed is questionable. Because of the age of the records, subsequent habitat changes or degradation, and the great increase in invasive plants in recent decades, it appears likely that very few of these historical sites will be found to still support the species and so add to the number of extant locations.

Threats One of the most significant threats faced by this species is woody plant encroachment on the early- successional habitats it requires; without active management open, herbaceous communities on serpentine, diabase, or limestone in Pennsylvania typically revert to shrub and tree cover, conditions in which the whorled milkweed declines and may eventually be lost. Another major threat is competition from invasive species. Currently, very few of the sites hosting this species are receiving any management, either to maintain early successional conditions or to control invasive species. Few sites are currently protected, and land ownership and use patterns in the regions inhabited by whorled milkweed create risk that some existing sites may be lost to development or mining.

Status Justification The PT status appears to be justified for whorled milkweed, because of the moderate number of occurrences, the generally low population numbers, the requirement of active management to maintain the necessary early successional habitat, the threat of invasive species, and the potential contraction of the historical state range. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Screw-stem — Bartonia paniculata (Michx.) Muhl. Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP

Introduction Screw-stem is a very slender and delicate annual that may grow to over one foot in height, but is often much smaller and sometimes only a couple of inches. The leaves are reduced to minute scales and mostly occur alternately on the stem. The flowers, which appear in late summer to fall, are generally whitish or yellowish, with pointed and untoothed petals, and occur at the end of relatively long stalks. The fruit is a capsule with many minute seeds. The entire plant frequently has a yellowish coloration.

This species has a superficial resemblance to Bartonia virginica, which generally has opposite leaves, more oblong and usually toothed petals, shorter stalks, and is more likely to be found in dry habitats.

Proposed PA Status Although screw-stem does not currently have a Pennsylvania legal status assigned to it, it has been assigned a recommended status of Pennsylvania Rare (PR) by the PABS Vascular Plant Technical Committee. A few years ago the PABS-recommended status of the species was Pennsylvania Threatened (PT), but this status was adjusted because dozens of new locations had been documented since the late 1990s, which indicated that the species was obviously more frequent and widespread than had been believed and no serious threats were evident in regard to extirpation.

North American Distribution and Global / Regional Conservation Concerns Screw-stem has a NatureServe global rank of Secure (G5).

The global distribution extends from east into Newfoundland and south into Texas and Florida. In this large range, the species tends to be found mostly near the Atlantic and Gulf coasts and then northward in the Mississippi River lowlands, although the plant does occur in the Appalachians (Kartesz 2003).

Screw-stem has been documented in all six states that are contiguous to Pennsylvania. Five of the states have assigned a rarity rank to the species - Delaware (Imperiled S2), Maryland (Vulnerable S3), New York (Critically Imperiled S1), Ohio (Critically Imperiled S1), and West Virginia (Critically Imperiled S1). The species has not been assigned a rarity status in New Jersey, presumably because it is not of conservation concern.

Pennsylvania Distribution Screw-stem has a NatureServe state rank of Vulnerable (S3), which along with the VPTC-recommended state status was recently changed (from S2 to S3) due to the large number of “new” locations that have been documented in the past fifteen years.

Prior to the late 1990s, screw-stem was considered mostly a plant of the southeastern counties, as one might expect from a generally coastal species, and as is shown in the Atlas of Rhoads & Klein (1993). In the last fifteen years, however, fourteen counties have been added to the state range to the west and northwest, including such distant counties as Cameron, Clearfield, Lycoming, and Tioga. It is unclear if screw-stem had always been present in these counties and had just been overlooked by botanists, or if it has markedly expanded its range in recent years. There may be some reality in both explanations, although the former reason is easy to understand because of the diminutive size of the plant and

26 relatively brief time of the year when it is visible. It wouldn’t be surprising to eventually document this species as occurring in most counties of Pennsylvania.

Habitat Screw-stem grows in damp to wet places, including stream margins in forests, swamps, thickets, seepages, and in the northern counties in mossy/boggy wetlands of various sorts, including abandoned and active beaver meadows. The species can be opportunistic, as long as the substrate is suitable, and has been found growing on human-created sites such as utility line rights-of-way, old forest roads, and tree plantations. It is usually rooted in and other bryophytes, but has been found on damp/wet sand, muck, and woodland soils. The species appears to thrive in both open and shaded situations. Overall, the species occurs in a rather wide range of wetland habitats, which include certain types of human-impacted areas.

The habitats given on Pennsylvania herbarium specimens (collected primarily in the southeastern counties) of screw-stem from the last 150 years include “seepy woods; moist woods; mossy woods; wet woods; low damp woods; swamp thicket; marshy depression; open meadow; sphagnous border of pond; small wooded stream; woodland bog; swamp; wet ground in woods; wooded sphagnous area; boggy swale; in moss on old woods road; moist overgrown evergreen plantation.”

Extant locations Numbers At present, 63 extant locations in 24 counties have been documented by PNHP, with about 75% situated on land owned by state natural resource agencies, municipalities, or land conservancies. Approximately one-third of these populations have not been visited within the last 10 years.

As mentioned previously, the number of extant locations has increased greatly since the late 1990s. Probably the main reason for this marked increase was the realization that this species occurs well to the north and northwest of its previously-known range, and that it is often an inhabitant of the open boggy (often beaver impacted) wetlands that are frequent in some parts of the northern counties of Pennsylvania. Prior to this, based on the records from the southeastern counties, screw-stem was considered more of a species to be found in wet places in wooded situations. Another factor in the increase of locations was learning that the species has a relatively restricted window for botanical surveys in late summer and fall, so searches for the species must be targeted to this period of time.

Population sizes The population sizes at the 63 extant locations vary from one plant to over 500 plants, although about one-half of the occurrences have 50 plants or fewer. The accuracy of these numbers must be judged by the time of year of the survey, the inconspicuous nature of the plant, the degree of effort of the survey, and particularly by the fact that the plant is an annual, so population numbers can vary considerably from year to year. On average, based on current information, the more northern occurrences tend to have the largest populations, but this may be anecdotal.

Potential for additional EOs The trend of documenting increasing numbers of de novo extant populations for screw-stem, especially in northern Pennsylvania, seems likely to continue because of recent experience in knowing where to search for the species, the large numbers of potential locations to be surveyed (especially on public land), and the variety of habitats that can be utilized by the species. A quantitative estimate of how many additional de novo populations might be present is speculation.

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Historical locations As mentioned previously, it is very notable that this species has been documented to be much more widespread than its historical distribution would have suggested.

Numbers Based on herbarium specimens, an estimated 28 historical locations of screw-stem in 11 southeastern counties have not been searched by PNHP for the possibility of having extant populations. The dates of these records range from the 1890s to the 1950s, with most of them being in the 1920 to 1940 period. It is completely unclear what the size of the populations may have been at these sites at the time of the collection. Many of these locations have not yet been recorded in Biotics.

Potential for extant EOs Because of the age of these collections and the region of the state involved (the southeast), it seems likely that there has been some degree of habitat loss and degradation at some of the historical locations. Given their location in the state, the generally vague directions, the age of the collections, as well as the effort required contacting landowners for access, it appears unlikely that many of these historical locations would be worth the effort to survey for extant populations. In fact, de novo surveys in southeastern Pennsylvania, particularly on public lands, watershed properties, and nature preserves, might offer a greater potential for discovering extant populations than searching historical locations.

Threats Overall, the level of threat to screw-stem at the extant locations seems relatively low, due to the general protections given to wetlands, the number of locations on public land or having owners sympathetic to conservation, and the lesser (relatively speaking, of course) threat from invasive species at many of the populations as compared to the threat to more mesic habitats. The threats appear particularly minimal at most of the northern populations because of the remote locations, public ownership, the number of potential sites, and the lesser amount of exotic species found there as contrasted with southeastern Pennsylvania.

Status justification In the opinion of this contributor, screw-stem does not appear to require a status of Endangered or Threatened because of the increasing number of extant occurrences, its recently-expanded distribution in the state, the number of occurrences (83%) on public land, the large number of potential locations remaining to be searched, the variety of habitats that can be occupied by the species, the realization that population numbers will fluctuate greatly from year to year because of the annual life cycle of the species, and the general environmental protection given to wetlands.

It can be debated if screw-stem even qualifies for the status of Pennsylvania Rare, and perhaps should be delisted from the state rare plant list. We believe the status of Pennsylvania Rare is appropriate, since that status can be used to give the habitat an added degree of protection and can give conservation attention to local populations. This species will be considered at the 2017 Rare Plant Forum.

The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of (TBD RPF 2017) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

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References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Rhoads, Ann Fowler and William McKinley Klein, Jr. 1993. The Vascular Flora of Pennsylvania: annotated checklist and atlas. American Philosophical Society, Philadelphia, PA.

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Spike Sedge – Carex ormostachya Wieg. Author: Steven P. Grund, PNHP

Introduction Spike sedge is a member of Section Laxiflorae in the genus Carex. Sedges are grass-like plants, and the genus Carex is the largest in Pennsylvania, with about 180 species (Rhoads and Klein 1993). Carex ormostachya is distinguished from other members of Section Laxiflorae in our region by the combination of red leaf bases, granular but not serrulate sheath angles, and slightly bent perigynium beaks (Voss and Reznicek 2012).

Proposed PA Status The Pennsylvania Biological Survey recommended Pennsylvania Threatened for Carex ormostachya in 2017 because the species is known to be extant in Pennsylvania at only two sites. Although the difficulty of finding and correctly identifying the species leaves some uncertainty as to how many undiscovered populations might exist, Pennsylvania Threatened is considered appropriate because a significant effort to relocate the species in areas with appropriate habitat in the vicinity of historical collection sites has resulted in the discovery of no additional extant sites.

North American Distribution and Global / Regional Conservation Concerns Carex ormostachya ranges from Ontario (to at least 50.7° N) and Quebec, NE to above the tension zone. New England and eastern New York, sporadically through Pennsylvania in the Ridge and Valley, and disjunct at a few stations in the mountains of Virginia and West Virginia (Brouillet et al 2017, Bryson and Naczi 2002, Kartesz 2017, NatureServe 2015). There is also a dubious record from western (Kartesz 2017). The Global rank is G4. The isolated occurrences in West Virginia and Virginia lead to S1 statuses in those states (NatureServe 2015). North Carolina is not included in the range of the species by NatureServe; we have located two specimens purported to represent Carex ormostachya collected in North Carolina (Southwest Environmental Information Network 2017). One of those specimens has been determined to represent another species (R.F.C. Naczi, pers. comm.). The other has not been examined by a specialist to our knowledge, and we consider the identification suspect. If it turns out to be correctly identified, presumably it will become S1 in North Carolina as well. Carex ormostachya is also considered to be of conservation concern at the northeast edge of its range: S1 in Nova Scotia; S3 in Quebec and New Brunswick.

Pennsylvania Distribution Reports of Carex ormostachya from Erie, Crawford, Bedford, and Fulton Counties (Rhoads and Klein 1993, Kartesz 2013) were based on misidentified specimens of other species (S.P. Grund, pers. obs., J.K. Bissell, unpubl. data). Most of the remaining specimens were have annotations by H.A. Wahl or C.T. Bryson, both highly regarded experts in the genus Carex. The scattered pattern of collections, one or two each from Wayne, McKean, Centre, Columbia, Pike, Tioga, Cameron, Lycoming, Clinton, Blair, and Monroe, is typical both for a species at the edge of its range, and of a species that is difficult to identify.

Habitat Carex ormostachya grows in mesic to dry-mesic forests and openings, often in sandy soils, and often in moderately disturbed habitats such as along logging roads (Voss and Reznicek 2012, Robert F.C. Naczi pers. comm.).

Extant Locations

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One extant location was discovered by R.F.C. Naczi in 1994 on Barclay Mountain in Bradford County. The size of the population is not known, but was not large enough for L.M. Speedy, C.M. Bowers, and S.P. Grund to relocate on a very rainy day. The second known extant population, in Blair County, is based on two specimens collected in 2003 during a survey of Bell’s Gap Run, a forested valley where there was a historical collection of Carex ormostachya recorded, and where other northern-affinity species have also been documented. The specimens have been confirmed as Carex ormostachya by R.F.C. Naczi. No population data was recorded due to uncertain identification in the field; however, we have revisited the locality several times without successfully confirming the occurrence, so we believe that any population of Carex ormostachya at that site is likely small.

Historical Locations and Potential for Relocation Some of the collection localities are too vague for targeted surveys (e.g., “Rocky mountains, Clinton County”). We were refused permission to survey at a few of the collection sites. We have conducted surveys to attempt to rediscover six of the fourteen historic collection sites. We spent multiple days at a few of these sites. The survey sites were chosen as the most likely to be rediscovered considering the specificity of the location information and the likelihood of appropriate habitat remaining in the vicinity. We found no Carex ormostachya during these surveys. Potential habitat for this species is abundant in Pennsylvania, but our position at the edge of the range of the species may mean that these occurrences have compromised viability because climatic conditions are not ideal for it to compete. The only way to clarify the status of this species is for qualified botanists to conduct multiple surveys over time.

Threats Urbanization is not currently a threat in the regions of the state where Carex ormostachya has been documented. This may change as areas like State College and southeastern New York continue to expand. Some areas where this species once grew have probably been lost to agriculture, mining, or other human activities, and fracking for gas trapped in deep shales is a significant threat going forward. The climate is warming, and this represents the most significant threat to the continued presence of Carex ormostachya in Pennsylvania, as only in colder areas to our north is this species secure.

Status Justification There is currently only two extant occurrences of Carex ormostachya documented in Pennsylvania. We are never certain that there are no undiscovered occurrences of any species, and that uncertainty is higher for species that are difficult to distinguish from their close relatives. This said, obscure species do get collected, and this species has been collected only 14 times in Pennsylvania, compared with its closest relative, Carex gracilescens, with 166 collections (Morris Arboretum 2017, adjusted for a few specimens the author knows to have been re-identified). It is clear that this is not a common species. When historical collection sites for Carex ormostachya have been searched, it has only been found at one site, despite apparently suitable habitat being encountered at other sites. If the species survives at these sites, it is in low numbers. Despite accounting for some uncertainty in both directions regarding how current knowledge reflects reality, i.e., there might be few more than the single known occurrence, or there might be even more than the total number of collections, the NatureServe Rank Calculator returns S1. Although the current number of extant sites known is consistent with Pennsylvania Endangered, Pennsylvania Threatened is recommended to account for some uncertainty regarding the total number of populations that may exist in the state.

References

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• Brouillet L., F. Coursol, S.J. Meades, M. Favreau, M. Anions, P. Bélisle, and P. Desmet. 2017. VASCAN, the database of vascular plants of Canada. data.canadensys.net/vascan/. Accessed 16 March 2017. • Bryson, C.T. and R.F.C. Naczi. 2002. Carex sect. Laxiflorae. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 23. • Kartesz, J.T. 2013. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press) • Morris Arboretum. 2017. Pennsylvania Flora Project of Morris Arboretum. paflora.org. Accessed 20 March 2017. • NatureServe. 2015. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia. Available http://explorer.natureserve.org. (Accessed: March 17, 2017 ). • Rhoads, A.F. and W.M. Klein, Jr. 1993. The Vascular Flora of Pennsylvania: Annotated Checklist and Atlas. American Philosophical Society, Philadelphia. • Southwest Environmental Information Network, SEINet – Mid-Atlantic Herbaria Consortium Chapter. 2017. http://midatlanticherbaria.org. Accessed in March. • Voss, E.G. and A.A. Reznicek. 2012. Field Manual of Michigan Flora. The University of Michigan Press. Ann Arbor.

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Short’s Sedge – Carex shortiana Dewey Author: Loree Speedy, Carnegie Museum of Natural History. Edited: Jessica McPherson, PNHP.

Introduction Short’s sedge is a tufted perennial grass-like plant that can reach from 8 to 35 inches in height. The flowers are held in densely packed, cylindrical spikes that become dark brown as they mature. It is notable among sedges for having chocolate-colored brown fruits and lateral spikes with female flowers at the top and scattered male flowers below.

Proposed PA Status The PABS-Vascular Plant Technical Committee currently recommends a status of Pennsylvania Rare for Short’s sedge, and we concur with this recommendation because of the limited numbers of populations and individuals known in the state, the relatively few occurrences with good ecological and reproductive viability, relatively limited habitat available in the state, and a moderate level of threat from invasive species, development, energy infrastructure, and changing agricultural practices.

North American Distribution and Global / Regional Conservation Concerns Short’s sedge can be found in the interior of the from Pennsylvania west to the very eastern edge of Kansas, south to northern Arkansas. Interestingly, its range is predominantly Midwestern, essentially ending at the western edge of the Coastal Plain, and it also occupies a fairly narrow north-south range (Kartesz 2003). It appears fairly widely distributed in the Midwestern states, while it is of conservation concern in several states at the eastern edge of the range (Vulnerable S3 in Virginia, West Virginia, Pennsylvania; Imperiled S2 in Maryland) and the northern edge (Critically Imperiled S1 in New York and Ontario). It is also listed as Imperiled in Arkansas.

Pennsylvania Distribution Short’s sedge is known from Allegheny and counties in western Pennsylvania, and from several counties in the southern Ridge and Valley province. Its limited distribution may be partly explained by its requirement for calcareous soils, and partly by our proximity to the northern edge of the global range.

Habitat Habitats range from woods at the base of slopes to wet open ground and bottomlands, often in calcareous soils (Ball and Reznicek 2003). In Pennsylvania, it is reported from calcareous wet meadows and swamps and rich woods (Rhoads and Block 2007).

Extant Locations Short’s sedge is currently known from 31 sites in Pennsylvania. Over half of these sites, however, contain ten or fewer reproducing individuals. Urbanization in the Pittsburgh area has destroyed or greatly curtailed several populations.

Historical Locations and Potential for Relocation Historical specimens have been individually reviewed to determine whether it is likely they might be relocated. Most locations where habitat still exists and location data is reasonably precise have been searched; a few populations have been relocated, while many have not. Most known populations are the result of de novo surveys rather than relocation of historically documented specimens. Since the species seems to have some preference for open, somewhat disturbed habitats (although it is also found

33 in more intact habitats) it is not surprising that many historical locations may no longer exist, while other populations have established in new areas.

Threats Many occurrences are in agricultural areas, perhaps due to this species’ preference for rich, limy soils. Populations in this habitat appear to be persistent rather than ephemeral, and the open character of the habitat favors the species. However, populations in agricultural settings are vulnerable to changes in cultivation practices, such as re-tilling of fields previously left fallow, or expansion of cultivation into marginal wet areas where the plant occurs. Invasive species are also often present in these nutrient-rich habitats, and it is unclear whether these populations may decline in the future due to competition from invasives. The wetland edge habitat utilized by Short’s sedge is also sensitive to fragmentation and hydrological changes from energy infrastructure development.

Maintenance of the currently known populations where Short’s sedge grows will be crucial to its survival. Management of known sites by creating natural vegetation buffers around fragmented habitat and removing invasive species will help to maintain and encourage existing populations. Potential sites for restoration should be evaluated (Pennsylvania Natural Heritage Program, 2015).

Status Justification We recommend a status of Pennsylvania Rare for Short’s sedge because the numbers of populations and individuals known in the state are limited, and consistent with the definition of this status. A status of Pennsylvania Rare is further justified because Short’s sedge has relatively few occurrences with good ecological and reproductive viability, its available habitat is somewhat limited by its requirement for calcareous substrate, and it faces a moderate level of threat from invasive species, development, energy infrastructure, and changing agricultural practices. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S3 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Ball, Peter W. and A. A. Reznicek. 2002. Carex. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 23. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • NatureServe. March 2014. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia. (Accessed 12 January 2015). • Pennsylvania Natural Heritage Program. 2007. • Rhoads, A.F. and T.A. Block. 2000. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia.

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Indian-paintbrush – Castilleja coccinea Mutis Author: Dr. Timothy Block, Morris Arboretum

Introduction Castilleja coccinea is and annual species, and like all Orobachaceae, hemiparasitic, drawing part of its nutrient requirements from the roots of other plants. Annual hemiparasites, like some orchid species, are sometimes irregular in their cyclic appearance at some sites for reasons not well understood.

Parasitic and hemiparasitic plant species are reported to play complex roles in some ecological systems (see Bardgett, et al, 2006; Press and Phoenix, 2005), but the relationships between Castilleja coccinea specifically and its host plants has not been examined.

Castilleja coccinea is easily recognized in flower by its large, bright red bracts which subtend each flower.

Proposed PA Status Castilleja coccinea should be listed as PT (S2) in Pennsylvania. This status is justified by the limited number of extant sites, perceived decline of some populations, and for reasons cited below.

Habitat Castilleja coccinea is a plant of moist to wet meadows, generally on limestone or diabase, or other high pH substrates.

North American Distribution and Global/Regional Conservation Concerns Castilleja coccinea is distributed throughout the eastern U.S. and much of adjacent Canada, although it is considered exotic where it occurs in Nova Scotia. It is ranked S1 in NY, MD, KY, MS, and AL; S2 in CT, PA, NJ, GA, and SC; S3 in IA, IN, KS, VA, and NC; and extirpated, or presumed extirpated in the rest of New England and LA. Its distribution is primarily in the Midwest plus the .

Pennsylvania Distribution Historically, Castilleja coccinea was scattered across much of the southern half of Pennsylvania, extending as far north as Pike County in eastern PA, although restricted to calcareous / diabase habitats. The current distribution is concentrated in Southeastern PA, with most known extant populations in Bucks and Montgomery counties. This is likely due to the intensity of recent botanical field work, and a particular interest in this species, in these counties.

There are between 20 and 25 known extant populations of this species. Population sizes range from very few individuals to several hundred per occurrence. Some extant populations appear to be stable while others seem to be in decline in spite of efforts to properly maintain habitat. Of particular note is the Fulshaw-Craeg diabase meadow site in Montgomery County where Natural Lands Trust has undertaken a study to examine the decline of this and other species of special concern.

Historical Locations and Potential for Relocation Most historical locations in Southeastern Pennsylvania have been checked within the last 10 years or so with mixed results. Some historical locations have been destroyed while others have been placed under protection through purchase or easement.

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Threats Although a species which obtains some of its nutrient and carbon requirements from host species, as with all low-growing, shade-intolerant plant species, the primary threat is competition from taller vegetation.

Status Justification The numbers of extant populations and/or individuals may suggest otherwise, but species with unusual or otherwise poorly understood biological requirements should be given deeper consideration, especially in cases where additional research could yield clarification of conservation concerns.

As with many species for which we lack sufficient biological knowledge, failure to effectively protect extant populations of Castilleja coccinea may well result in a lost opportunity to ultimately understand the importance of the species within a complex ecological system.

References • Bardgett, R.D., R.S. Smith, R.S. Shiel, S. Peacock, J.M. Simkin, H. Quirk, and P.J. Hobbs, 2006. Parasitic plants indirectly regulate below-ground properties in grassland ecosystems. Nature 439: 969-972. • Press, M.C. and G.K. Phoenix, 2005. Impacts of parasitic plants on natural communities. New Phytologist 166: 737-751.

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Short’s Slender Chervil – Chaerophyllum procumbens (L.) Crantz var. shortii Torr. & Gray Author: Steven P. Grund, PNHP.

Introduction Short’s slender chervil is a spring-blooming winter annual in the parsley family (). It can be difficult to distinguish from typical slender chervil (Chaerophyllum procumbens var. procumbens), but it has been treated at the level of species as Chaerophyllum shortii (Small 1933) and will likely be treated as such in the upcoming “New Manual of Vascular Plants of Northeastern United States and Adjacent Canada” (R.F.C. Naczi, pers. comm.). The fruits are usually hairy, and are never hairy in typical slender chervil. The occasional Short’s slender chervil with glabrous (hairless) fruits can be distinguished by having fruits that narrow convexly to the style, compared to typical slender chervil with fruits that narrow more abruptly and have a short concave segment just below the style.

Proposed PA Status We concur with the decision at the PABS-Vascular Plant Technical Committee meeting of 5 October 2013 to recommend a status of PE for Short’s slender chervil. This plant is currently known from only two localities in Pennsylvania.

Habitat Short’s slender chervil grows in rich forests where pH is neutral or higher on floodplains and adjacent lower slopes.

North American Distribution and Global / Regional Conservation Concerns Short’s slender chervil occurs from southern Pennsylvania south to Virginia, and west to Missouri and NW Arkansas. There are apparently disjunct stations in South Carolina and Ontario (Kartesz 2003). It is considered Critically Imperiled (S1) in Ontario and Indiana, and is known only historically in West Virginia. It has not been ranked in other states, likely in part because for many decades its status was obscured by lack of recognition. The global status of G5T3T4Q reflects uncertainty about both the conservation status and the taxonomic status (the “Q”). The “G5” refers to Chaerophyllum as a whole (var. procumbens is Globally Secure). Var. shortii is globally Vulnerable (G3) to Apparently Secure (G4).

Pennsylvania Distribution Short’s slender chervil is currently known from 2 stations, in Greene and Westmoreland counties. There is a historical specimen that was collected in Bedford County.

Extant Locations The two known extant populations are apparently small, probably less than 200 plants between them, but in both cases the available habitat has not been thoroughly surveyed. There is, in our opinion, a strong likelihood that some additional populations will be discovered, but this appears to be more of a habitat specialist than Chaerophyllum procumbens var. procumbens, and we think it is unlikely that this plant will turn out to be too common to be an appropriate conservation target.

Historical Locations and Potential for Relocation The locality data for the single historic collection, combined with geological data showing us where soils with elevated pH are likely to be found provided us with a good guess of the collection site. A visit to that site revealed and area that is partly agricultural field and partly mid-successional forest. No Chaerophyllum was found at the site. It is certainly possible that this species is extant in the vicinity of the site where it was once found in Bedford County.

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Threats Plants with an affinity to elevated pH, like Short’s slender chervil, are particularly vulnerable to displacement by invasive alien plants. Also, this species is found mostly on and immediately adjacent to floodplains, which are prime targets for conversion to agriculture, industry, and other land developments.

Status Justification Despite a number of targeted surveys, Short’s slender chervil is currently known from only two localities in Pennsylvania. A high proportion of the floodplains that provide potential habitat have been converted for various human uses. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1S2 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012). The potential for additional discoveries is what produced the S2 portion of the range rank. The rounded rank is S1, and with only two extant occurrences known and a higher than average threat level, Pennsylvania Endangered is in our opinion the appropriate status.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Small, J.K. 1933. Manual of the southeastern flora. New York. P. 969

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Strawberry Goosefoot – Chenopodium capitatum (L.) Asch. Author: Jessica McPherson, PNHP

Introduction Strawberry goosefoot (Chenopodium capitatum, also recognized as Blitum capitatum by some taxonomists) is a member of the goosefoot family that is historically known from Pennsylvania, but has no known extant populations currently. Although members of the genus Chenopodium can be difficult to distinguish, this species is quite distinct due to its halberd-shaped leaves and the bright red clusters of seeds that develop in the summer. The seeds and leaves are both edible, and this species is sold in some seed catalogs as a vegetable, often under the name “strawberry spinach.”

Proposed PA Status This species is proposed to be listed as Pennsylvania Endangered, as a surrogate for Pennsylvania Historic, because it has been known from the state in the past, no extant locations are currently known, and yet the species cannot be proven to have been extirpated from the state. Pennsylvania Endangered is the current status recommended by the PABS-Vascular Plant Technical Committee.

Habitat In the main part of its range, the habitat is described in the Flora Novae-Angliae as “anthropogenic (manmade, disturbed) habitats, meadows, fields” (Haines, Farnsworth and Morrison 2011) or “sandy or grassy meadows, thickets, open woods, old fields of clearings in forest” in Flora of North America (Clemants and Mosyakin 2003). Michigan Flora describes it as growing in “sandy or gravelly (often very calcareous) sites one to two years after disturbance (e.g., bulldozing) and dying out (if disturbance is not maintained) within another 2 or 3 years.” Michigan Flora also lists some natural disturbance settings where the plant may be found such as burned ridges or limestone ledges (Reznicek, Voss and Walters 2011).

North American Distribution and Global / Regional Conservation Concerns The geographic range of the species is predominantly north of Pennsylvania; occurrences in Pennsylvania were at the southern edge of the range in eastern North America (Kartesz 2003).

Pennsylvania Distribution Historical specimens have been documented from 11 scattered Pennsylvania counties, with little apparent pattern to the distribution. No county has more than one specimen documented except Northampton, which has two specimens.

Extant Locations There are no known extant locations in Pennsylvania at this time.

Historical Locations and Potential for Relocation In Pennsylvania, there are 13 historical records. The most recent record is from 1969, two records are from 1909, and all others were collected before 1900.

The timing of most collections of this species, in the late 19th and very early 20th century, coincides with widespread timbering and fire in Pennsylvania. A possible explanation for why this species was observed at that time, but has not been observed since, is that timbering and fire opened up new habitat for the species, and it moved south during this time, but then failed to persist as early successional habitats matured to forest and fires were suppressed in 20th century.

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This species is a low priority for further fieldwork due to the extreme age of the records, difficulty in targeting fieldwork, and low conservation value of the its typical habitat. Historical records are not useful in targeting searches because the species requires early successional habitat, yet because of the timing and extreme age of the records it is likely the landscape has transformed in the intervening time. It is considered a poor use of resources to survey for this species in generic early successional habitats because this type of habitat is very extensive in the state, yet the species appears to be very rare, so the likelihood of success is very low while the area of available habitat is large enough that it is impossible to comprehensively survey to determine it is absent. Furthermore, these habitats have little conservation value, and highly mobile, early-successional species reap relatively little benefit from land conservation measures typically associated with environmental review.

Status Justification Given the current knowledge of this species, the most appropriate status is Pennsylvania Historic, because it is clearly documented to have occurred in the state in the past, but appears to no longer be present. Since Pennsylvania Historic is not currently available as an official status, proposal of the species may be delayed until such time as this status is officially adopted, or it may be categorized as Pennsylvania Extirpated or Pennsylvania Endangered with no practical distinction since no occurrences are known. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of SH (Historic) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Clemants, Steven E. and Sergei L. Mosyakin. 2003. Chenopodium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 4. • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. October 6, 2016. http://www.michiganflora.net/species.aspx?id=67.

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Hawthornes – Crataegus spp. Author: Jessica McPherson, PNHP Editors: Steven P. Grund, PNHP; Dr. Timothy Block, Morris Arboretum.

The genus Crataegus requires extensive work to reach a sufficient level of understanding to assign conservation statuses to taxa in Pennsylvania. We have not attempted this work within the scope of the Plant Status Updates projects. Below we outline the relevant problems and further work needed (listed in bullet points). Dr. Timothy Block at Morris Arboretum has examined the genus in light of the recent Flora of North America treatment by Phipps; the following represents a synthesis of Block’s comments and information included in the Flora of North America genus treatment.

There are significant differences between the taxonomy used in Plants of Pennsylvania and in the recently published treatment by Phipps in Flora of North America (Block, pers. comm.). According to Phipps (2014), taxonomy is complicated by asexual reproduction, polyploidy, hybridization, and extraordinary morphological variation within taxon concepts (potentially related to the interplay of the previously mentioned factors). Crataegus is also popular in the landscaping trade; the origins of cultivated taxa are in some cases not well documented, and it is unknown whether they are naturalizing or hybridizing with locally native taxa. However, unlike in the case of Amelanchier, there is not much active research on Crateagus; so while the above-mentioned problems create persistent problems in applying the taxonomy, the concepts are not actively being revised (Block, pers. comm). • A greater understanding of Crateagus in Pennsylvania could be gained through the project of going through existing herbarium specimens to attempt identification using the Phipps FNA taxonomy.

It is clear that the traditional species concept of reproductive isolation does not apply cleanly within the genus. As is discussed with Amelanchier and Trillium, actively evolving taxa have different conservation needs than those with more static species concepts. Protection of evolutionary potential must be considered along with protection needs of the organisms identified as rare taxa. If and when taxa of conservation concern are identified within this genus, the same principles and guidelines for conservation of species complexes as were described for those genera should be applied here.

A further problem in Pennsylvania is that the distributions of Crataegus taxa within the state are very poorly understood, to a degree that is unusual among vascular plant taxa today, because the herbarium specimen record cannot be relied upon, and because very few active botanical surveyors are recording the presence of these species. According to Dr. Block, “many specimens in herbaria are either misidentified, or, worse yet, of insufficient material to make a proper determination to species. It really is necessary to obtain, or at least observe, flowers, mature short-shoot leaves, and mature fruits of each individual in order to be 100% certain of the identification of many (if not most) species of Crataegus.” Beyond herbarium collections, there are very few sources of information about the abundance and distribution of Crataegus taxa in Pennsylvania. Most botanical workers do not even attempt to identify the plants beyond genus, and therefore there is almost no data available from general botanical surveys. • After assessment of data gaps through the above mentioned herbarium review, a statewide effort to obtain identifiable collections of Crateagus could improve our understanding of taxa distributions and rarity.

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Despite the data gaps, it appears that at least some taxa within the genus are widespread in the state. Crataegus are encountered fairly frequently. For most Pennsylvania taxa, the habitat appears to be generic, mid-successional, and inclusive of a moderate level of disturbance. These factors suggest that many species in the genus may not be of especially high conservation concern. For a small number of taxa, there is additional information to suggest some conservation concern; Crateagus calpodendron, Crateagus mollis, and Crateagus uniflora were assigned higher Coefficients of Conservatism values (Chamberlain and Ingram 2012). Crataegus pensylvanica is a taxon that is both distinctive and appears to be extremely rare in the state (Grund and Block, pers. comm. 2017). For the remaining taxa, although there is some indication that they are not high conservation priorities, conservation status cannot be reliably assessed or assigned without a better understanding of taxonomy, the nature of blurry boundaries between taxa and their evolutionary implications, and the actual distribution, abundance, and ecology of taxa within the state.

References • Chamberlain, S.J., and H.M. Ingram. 2012. “Developing Coefficients of Conservatism to Advance Floristic Quality Assessment in the mid-Atlantic Region.” Journal of the Torrey Botanical Society 139 (4): 416–427. • Phipps, James B. 2014. Crataegus. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 9.

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Smartweed Dodder – Cuscuta polygonorum Englemann Author: Dr. Timothy Block, Morris Arboretum

Introduction The dodders are annual, parasitic plants having as their hosts a wide range of other vascular plant species. Cuscuta polygonorum is a bit unusual in the genus seeming to specifically, but not exclusively, prefer hosts from the genera Polygonum or Persicaria in the Polygonaceae.

Species of the genus Cuscuta are similar in their vegetative characteristics, leaving only flower and fruit characters to distinguish species. Generally, the merosity of the flowers and the morphology of the and petals are the most important characters to examine.

Cuscuta polygonorum flowers are 4-merous, the corolla lobes are acute, and the persistent styles of the fruits are less than 1 mm long.

Taxonomically, C. polygonorum has been the subject of some confusion and controversy. While Costea, et al (2006) include C. polygonorum at the species level, Beliz (1986) considered this species to be a variety of C. pentagona.

As Costea, et al (2006) point out, little is understood of the reproductive biology of Cuscuta, limiting our understanding of taxonomy and evolution of the genus in general.

Proposed PA Status The conservation status of Cuscuta polygonorum in Pennsylvania should be PE (S1). This assessment is justified by our current knowledge of the limited number of occurrences and individuals of this species.

Habitat Shores, riverbanks, swamps, and marshes. Mostly on Polygonum or Persicaria, but occasionally on Impatiens, Ipomoea, Lycopus, or others.

North American Distribution and Global/Regional Conservation Concerns Generally distributed from southern New England to Virginia, west to North Dakota and Ontario, thence south to Texas, but absent from most of the southeastern U.S.

Cuscuta polygonorum is listed as S1 in Ontario, New York, Wisconsin, Maryland, Washington DC, and West Virginia; S2 in Pennsylvania, New Jersey, Michigan, Virginia, and Kansas; S3 in Massachusetts; and extirpated in North Dakota and Minnesota.

Pennsylvania Distribution Historical and extant collection locations are widely scattered across Pennsylvania from Crawford County in the northwest to Bucks County in the southeast.

One extant location is in Centre County, while a cluster of extant sites is in Bucks County in the Bristol/Croyden area and in Neshaminy State Park, all in tidal areas of the Delaware River. The Bucks County populations, in some cases, cover areas of several hundred square feet, but it’s problematic to determine how many individuals make up these occurrences.

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Historical Locations and Potential for Relocation Given the widely scattered nature of historical occurrences, and the specific attention that needs to be paid to identification to the species level, it would be a monumental undertaking, indeed, to attempt to relocate populations that were collected from the mid-1800s to very recent times.

It would be difficult to specify where to look for additional populations of this species other than wet areas where Polygonaceae species can be found.

Threats Threats to parasitic plant species are linked to threats to their hosts. In this case, since C. polygonorum parasitizes some extremely common host species, it would be difficult to sensibly assess the potential threats.

Status Justification PE is the appropriate status for this species due to its limited occurrences in the state.

References • Beliz, T., 1986. A Revision of Cuscuta sect. Cleistogrammica Using Phenetic and Cladistic Analyses with a Comparison of Reproductive Mechanisms and Host Preferences in Species from California, Mexico, and Central America. Ph.D. Dissertation, University of California, Berkeley. • Costea, M., G.L. Nesom, and S. Stefanović, 2006. Taxonomy of the Cuscuta pentagona complex (Convolvulaceae) in North America. Sida 22(1): 151-175.

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Laurentian Bladder Fern – Cystopteris laurentiana (Weath.) Blasdell Author: Steven P. Grund, PNHP

Introduction The laurentian bladder fern ( var. laurentiana) was described by Weatherby (1926). Wagner (1955) provided evidence based on chromosome morphology of hybrids involving C. bulbifera. Blasdell (1963) found one of Wagner’s hybrids to be morphologically identical to C. fragilis var. laurentiana, and made for it the combination Cystopteris × laurentiana. The “×” indicates a sterile hybrid (incapable of normal meiosis because the chromosomes cannot line up in pairs), but Wagner’s chromosome observations indicated that this taxon has twice the number of chromosomes that would be found in a sterile hybrid of and C. fragilis. When sterile hybrids double their chromosomes, they become capable of normal meiosis and thereby are fertile, so we can abandon the “×”, as Cystopteris laurentiana is full species, a polyploid of hybrid origin that is perfectly fertile, and is treated as such in all modern floras in which it is included (e.g., Gleason and Cronquist 1991, Haufler et al 1993, Rhoads and Block 2007).

Proposed PA Status There are only two known extant occurrences of this species in Pennsylvania. We concur with the 1994 decision by the VPTC to recommend to DCNR a status of PE.

Habitat Like most bladder , this species grows in the crevices of rock outcrops. In this case, specifically outcrops of limestone or other calcareous rock.

North American Distribution and Global / Regional Conservation Concerns Laurentian bladder fern has been documented sporadically from Newfoundland and northern Ontario to eastern Minnesota and , east to central Pennsylvania (Haufler et al. 1993, Kartesz 2003). It has not been found in any state directly bordering Pennsylvania, but across Lake Erie it is considered Vulnerable (S3) in Ontario, where it may be locally common on the Niagara Escarpment on the north shore of Lake Superior (Mike Oldham, pers. comm. as reported by NatureServe 2015). The global conservation rank is Vulnerable; G3 (NatureServe 2015).

Pennsylvania Distribution Laurentian bladder fern is known in Pennsylvania only from two sites in Centre County, both along the same stream.

Extant Locations The two locations, both along Spring Creek in Centre County, are both small. Estimating population size is difficult because of the presence of similar looking species, especially , but the number of rosettes is probably less than 100, perhaps much less, at each site.

Historical Locations and Potential for Relocation The Pike County record was searched for without success in 1989 by Greg Edinger. More potential habitat in the area should be surveyed. A specimen from Butler County formerly referred to as this species has been identified as Cystopteris bulbifera (S.P. Grund, pers. comm.).

Threats

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One of the extant occurrences is near a road and could be impacted by car exhaust or road maintenance activities. Changes to hydrology in the watershed could have a negative impact on this species. Certain invasive species, especially vines like Oriental bittersweet (Celastrus orbiculata) and Japanese honeysuckle (Lonicera japonica), can invade calcareous outcrops and represent significant potential threats.

Status Justification This species is known from only two sites with small populations, and those two sites are along the same stream. Calcareous outcrops have been targeted by botanists more frequently than most habitats, so although the species is difficult to identify, the probability of a large number of extant populations remaining undiscovered is low in our opinion. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012). These facts are consistent with a status of Pennsylvania Endangered.

References • Blasdell, R.F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torrey Bot. Club. 21(4): 1—102. • Gleason, H.A. and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Haufler, C.H., R.C. Moran, and M.D. Windham. 1993. Cystopteris. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. Vol.2. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • NatureServe. 2015. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia. Available http://explorer.natureserve.org. (Accessed: November 1, 2016 ). • Rhoads, A.F. and T.A. Block. 2007. The Plants of Pennsylvania, an Illustrated Manual. 2nd ed. University of Pennsylvania Press, Philadelphia. • Wagner, W.H. 1955. Cytotaxonomic observations on North American ferns. Rhodora 57(680): 219—240. • Weatherby, C.A. 1926. A new North American variety of Cystopteris fragilis. Rhodora 28: 129— 131.

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Heller’s Witchgrass – Dichanthelium oligosanthes (Schult.) Gould Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP

Introduction Heller’s witchgrass is an herbaceous perennial grass characterized, as compared with other Dichanthelium species, by its relatively broad leaves, “stubby” and strongly-veined spikelets, and relatively short and few-branched autumnal growing from the upper nodes of the stem.

Proposed PA Status We recommend Pennsylvania Threatened as the appropriate status for this species, based on the moderate number of populations, generally small population numbers, and the threat that exists from invasive species and other competing vegetation, since Heller’s witchgrass is generally found in early- successional conditions. Since many of the occurrences do not receive active management to maintain the proper successional stage, the risk level for the species is elevated. Pennsylvania Threatened is the status currently recommended by the PABS-Vascular Plant Technical Committee.

The latest taxonomic treatment in Flora of North America divides Heller’s witchgrass into two varieties, variety oligosanthes and variety scribnerianum, which are separated by spikelet shape and dimensions, leaf blade dimensions, and ligule length. There is some evidence that the differences between the two varieties are not always consistent and there may be some intergradation between the two taxa. Up until the present, all Pennsylvania plants of Heller’s witchgrass were believed to represent variety oligosanthes, but Jim Bissell of the Cleveland Museum of Natural History has indicated that the plants growing at Presque Isle, Erie County, appear to be variety scribnerianum. Based on this information, we recommend that variety scribnerianum should have a rarity status of Pennsylvania Endangered, while variety oligosanthes should continue at a rarity status of Pennsylvania Threatened. The VPTC has not yet considered the varieties individually.

North American Distribution and Global / Regional Conservation Concerns On a global basis, both varieties of the species are considered to be globally secure (G5) and are wide- ranging in North America (Kartesz 2003).

The remaining paragraphs address variety oligosanthes and whether it deserves the current rarity status of Pennsylvania Threatened.

Habitat and Pennsylvania Distribution Heller’s witchgrass generally grows in well drained, sunny conditions, and appears to have preference for substrates such as serpentine, limestone, diabase, schist, shales, and certain metasedimentary rocks that are present in the Wissahickon Formation in southeastern PA (Chester County). It is a characteristic and sometimes locally common component of the flora in serpentine grasslands. Outside the serpentine region, the plant inhabits limestone and diabase glades, grasslands, open woodlands and woodland edges, rock outcroppings, and some human-caused disturbances, such as pastures and abandoned fields (if the proper substrate is present).

Extant Locations The current number of extant locations in Pennsylvania, based primarily on PNHP field forms and other recent observations from the last 30 years, is conservatively estimated at ca 30. About two-thirds of these locations are at serpentine barrens. The larger, more accessible barrens with numerous actively managed grasslands, such as Nottingham and New Texas, probably support hundreds of stems of

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Heller’s witchgrass, and the species seems secure at these locations. The populations at the smaller, less accessible and less managed barrens, most of which have not been visited in recent years, probably only contain a fraction or less, maybe much less, of that number of stems. Even if the smaller barrens were to have active management, the populations of Heller’s witchgrass would probably remain relatively small because these sites are limited in area and because it is the nature of the species to be relatively infrequent. Overall, the serpentine barrens, particularly the larger ones, appear to be the stronghold of this species in the state.

Much less is known about the current status of Heller’s witchgrass at the relatively few extant sites occurring outside of the serpentine region. The species was confirmed at diabase grasslands in Adams and York counties in 2014, and at several metasedimentary sites in Chester County in the last fifteen years. These non-serpentine sites tend to be small in area and in the number of stems present, receive minimal if any active management, have invasive plant problems, and are located on often inaccessible private land. In addition to the documented sites, there is potential for finding new occurrences of the species in the limestone, diabase, and metasedimentary rock areas, but it seems unlikely that there will be any significant number. If attempting to conserve the overall range of the species in the state is important, these non-serpentine locations can’t be marginalized, however.

Historical Locations and Potential for Relocation Based on museum specimens, there appear to be ca 40 historical locations, depending on how an occurrence is defined, of Heller’s witchgrass in Pennsylvania. Most of these herbarium specimens are located along the Delaware River and extreme southeastern PA. The ultimate value in taking the time and expense to attempt to visit each of these historical sites and confirm the presence of Heller’s witchgrass is questionable. Because of their age, subsequent habitat changes, and the great increase in invasive plants in recent decades, it appears likely that very few of these historical sites will be found to still support the species and so add to the number of extant locations. For this reason, most of them have not been entered into Biotics.

Threats One of the most significant threats faced by this species is woody plant encroachment on the early- successional habitats it requires; without active management open, herbaceous communities on serpentine, diabase, or limestone in Pennsylvania typically revert to shrub and tree cover, conditions in which the Heller’s witchgrass declines and may eventually be lost. Another major threat is competition from invasive species. Currently, very few of the sites hosting this species are receiving any management, either to maintain early successional conditions or to control invasive species. Few sites are currently protected, and land ownership and use patterns in the regions inhabited by Heller’s witchgrass create risk that some existing sites may be lost to development or mining.

Status Justification Due to the moderate number of populations known in the state, the small size of most known populations, the vulnerability of all unmanaged populations to decline or loss from succession and invasive species, and the scarcity of appropriate habitat in the state, we recommend a status of Pennsylvania Threatened at this time. The number of known populations is slightly high for Pennsylvania Threatened, but very few sites are receiving active management, and therefore most sites are at risk of decline or loss; if a substantial number of populations are stabilized and protected across the range of the species in PA, it may be appropriate to consider changing the status to Pennsylvania Rare. We do not believe that significant numbers of new populations will be found at this point because most appropriate habitat has already been surveyed, but there is some chance it has been overlooked

48 because Dichanthelium species are difficult to identify and not well known by many botanists; if it does occur that substantial new populations are discovered, the status should also be reconsidered. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the currently available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Log Fern – Dryopteris celsa (W.Palmer) Small Author: Dr. Tim Block, Morris Arboretum; Jessica McPherson, PNHP

Introduction Log fern is a plant of wet woods on generally acidic soils, often on rotting logs or other organic substrates. The characteristics that distinguish this species from other Dryopteris species are subtle. This results in occasional misidentifications of specimens and possible overlooked populations. Specific identification characteristics have to do with the overall shape of the fronds and the relative locations of the sori to the veins.

Log fern is a fertile allotetraploid (4X) with D. goldiana and D. ludoviciana as putative parents. Genetic analysis supports D. goldiana and D. ludoviciana as the most likely parent species, and also indicates a single hybridization event rather than multiple independent origins for the species (Werth 1991, Sessa et al. 2012). In particular, Werth’s analysis includes multiple samples from 11 populations from across the geographic range, and indicates all arose from a single hybrid ancestor.

Proposed PA Status This species is proposed to move from unlisted (N) to Pennsylvania Endangered (PE), due to the extremely low number of populations and individuals known in the state. There are currently six known extant populations, and about ~360 plants known from these sites. Pennsylvania Endangered is the status currently recommended by the PABS-Vascular Plant Technical Committee.

Habitat Flora of North America (Montgomery and Wagner 1993) describes the habitat of log fern as “Seepage slopes, hammocks and logs in swamps, mostly on the Piedmont and Coastal Plain.” In Pennsylvania, extant occurrences are known from swamps and seepage wetlands. Five of the six extant populations have associate species that indicate some degree of calcareous influence, although this characteristic is not mentioned in other sources. Specimen label data describes a variety of minimally to significantly wet habitats such as “springhead in woods along creek,” “swampy woods,” “moist depression among rocks in wooded ravine,” “swamp,” “wet thicket,” “springy place adjacent to streamlet,” “low ground in woods,” and “on bank in sandy leaf mold.”

North American Distribution and Global / Regional Conservation Concerns The global range of log fern includes south-eastern North America and several geographically disjunct populations around the Great Lakes. Although documentation may be hampered by undercollection due to identification difficulties, the current known distribution is very scattered, and the species does not appear to be abundant or widespread anywhere within its range (Kartesz 2003).

Log fern is ranked as Imperiled (S2) or Critically Imperiled (S1) throughout most of its range, Vulnerable (S3) in Maryland and North Carolina, and Apparently Secure (S4) only in Virginia (although it is not ranked in , , or South Carolina). NatureServe lists a global rank of Apparently Secure (G4) in North America. The NatureServe rank of S1 in Pennsylvania is well justified based on current information.

Pennsylvania Distribution In Pennsylvania, log fern has a disjunct distribution, with population clusters in the southeastern portion of the state and also in Erie County.

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Extant Locations Although all historical records are from the SE, only two extant populations are known from that region today, both of which are very small (11-15 plants). Since 1995, four populations have been found in Erie County: one privately held site where 53 plants were documented; a site on state game lands with 27 plants; and the largest known population in the state, in a swamp on an Audubon Society refuge where ~250 plants were documented. Population data was not recorded at the fourth Erie County site.

One Pennsylvania population from Berks County is included in Werth’s genetic analysis, and was found to be genetically similar to other populations across the range and strongly likely to have derived from the same origin. Notably, the Erie County populations had not been discovered at that time and have not been assessed in regards to their similarity to populations in SE PA or the rest of the range.

Historical Locations and Potential for Relocation All historical locations are from SE Pennsylvania. Many have likely been lost due to the high rate of land conversion in this region.

Threats Log fern is threatened by potential habitat loss and/or degradation in Pennsylvania. Although most of the known populations are on protected lands, several have very small populations that are vulnerable to loss due to random disturbances. Most populations occur in small fragments of habitat, which decreases long-term viability and increases vulnerability to invasive species.

Status Justification Pennsylvania Endangered is clearly the appropriate status for the log fern, due to the small number of populations and individuals known in the state, the species’ geographic disjuncture into two even smaller groups of populations, and the threats to long-term viability that these populations face. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Montgomery, James D. and Warren H. Wagner, Jr. 1993. Dryopteris. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 2. • Sessa EB, EA Zimmer, and TJ Givnish. (2012) Unraveling reticulate evolution in North American Dryopteris (). BMC Evolutionary Biology 12(1): 104. • Werth, Charles R. (1991). Isozyme Studies on the Dryopteris "spinulosa" Complex, I: The Origin of the Log Fern Dryopteris celsa. Systematic Botany, Vol. 16, No. 3. (Jul. - Sep., 1991), pp. 446- 461.

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Clinton’s Fern – Dryopteris clintoniana (D.C. Eaton) Dowell Author: Jessica McPherson, PNHP

Introduction This species originated as a fertile allopolyploid (hexaploid) hybrid between Dryopteris goldiana (2x) and (4x). Genetic analysis suggests it originated about 7.3 million years ago, and indicates a single hybridization event as the origin rather than multiple independent hybridization events, although the authors also suggest sampling more individuals will improve confidence in this assessment (Sessa, Zimmer, and Givnish 2012).

Proposed PA Status We recommend the classification of Clinton’s fern (Dryopteris clintoniana) as Pennsylvania Threatened, due to the low number of known locations in the state, and very low number of total individuals in the state. Pennsylvania Threatened is the status currently recommended by the PABS-Vascular Plant Technical Committee.

Habitat This species is found in swamp forests.

North American Distribution and Global / Regional Conservation Concerns The North American distribution of Clinton’s fern appears concentrated in New England, although it extends south to northern Virginia, and west around the Great Lakes through Michigan, where it is somewhat common, to a few isolated occurrences in Illinois and Wisconsin. It is listed of conservation concern by NatureServe in many states and provinces across its range; while it is Secure (S5) in New York, it is Vulnerable (S3) in New Jersey and Quebec, Critically Imperiled (S1) in Maryland, Delaware, Ohio, and New Brunswick, Possibly Extirpated (SH) in Virginia and Wisconsin, and Presumed Extirpated (SX) in Indiana.

Pennsylvania Distribution and Extant Locations Most known extant locations are in the glaciated northwestern corner of the state (17), with a smaller number of occurrences scattered across the NE (5) and SE (3). The habitat has been fairly thoroughly surveyed, so we do not anticipate finding a large number of additional populations. Known populations of this species range from small to extremely small; 23 of 24 have 50 individuals or fewer, and 6 have 10 individuals or fewer. The estimate for the state’s total population is 300-900 individuals. This number may actually be substantially lower due to the inclusion of sterile hybrid individuals in population counts, as detailed below.

Hybridization in Clinton’s fern and its impact on conservation assessment

Clinton’s fern does hybridize with five other species in the genus Dryopteris, but the hybrids are very rarely fertile. Hybrids can most reliably be identified by examining spores to detect whether they are misshapen or aborted; however, hybrids can often be identified in the field with fair reliability based on the appearance of sori, and the presence of traits intermediate between the parent species (Warren H. Wagner and Chen 1965; Whittier and Wagner 1971; W. H. Wagner, Wagner, and Taylor 1986). Based on our review of Carnegie Museum specimens, specimens of fertile fronds from at least mid-July or later are required to reliably determine hybrids based on spores.

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For a substantial minority of known occurrences (6 of 24), specimen documentation is a hybrid individual with D. clintoniana as one parent, rather than a fertile individual of the species.* However, since the hybrids are not fertile, their presence indicates that the parent species are also present, or were present at the time the hybrid individual originated (Montgomery 1976). Montgomery studied Dryopteris hybrids in New Jersey, and found that both parents were nearby in most cases, and the only exceptions were situations in which the missing parent was a very common species such as Dryopteris intermedia. D. clintoniana is an uncommon species, and Montgomery found no hybrids involving D. clintoniana where D. clintoniana was not nearby. Therefore, we are considering hybrid specimens for which D. clintoniana is one parent as sufficient evidence for the creation of an element occurrence record.

Montgomery’s documentation of Dryopteris hybrids in New Jersey found that D. clintoniana frequently forms hybrids, and most frequently with species who co-occur in the same habitat; notably, he found hybrids at 58.8% of 17 sites where both D. clintoniana and D. cristata were present, and at 40% of 15 sites where D. clintoniana and D. intermedia were both present. If hybridization occurs at a similar rate in Pennsylvania, we would expect to find hybrids at more than the ¼ of extant sites for which they are currently documented.

The relative frequency of hybrid individuals vs. D. clintoniana at sites with both taxa present has not been investigated. In most cases, surveyors did not identify hybrids in the field, and recorded population data for D. clintoniana without realizing that hybrids were present. Hybrids were identified later through expert annotation of the specimens. There are several problems that would make determining the relative frequency a difficult if not impossible project: the scattered nature of the individuals, the similar appearance of D. clintoniana, D. cristata, and D. clintoniana hybrids, and the fact that not all individuals produce spores every year and it is frequently not possible to reliably distinguish vegetative individuals of the three above-mentioned taxa. However, we can conclude with the information currently available that the population estimate for D. clintoniana in Pennsylvania may be somewhat inflated, because at least ¼ of the extant occurrences include hybrid individuals.

Historical Locations and Potential for Relocation There are 31 historical locations documented through specimens, 3-15 of which could be potentially relocated. For the remaining historical locations, they have either already been searched without success, the location data recorded is too vague to relocate, or suitable habitat no longer appears to be present at the location described.

Threats In the short term, Clinton’s fern populations appear to be relatively secure. In northwestern Pennsylvania, swamp forest habitat is relatively abundant with many sites protected on state or federal lands. Invasive exotic plant species are not currently abundant in these habitats, although they pose a significant long-term threat. There are many invasive species that are present in the region and would seriously degrade the habitat for Clinton’s fern if they should become established at the sites where it occurs. Deer browse threatens some populations, especially in eastern Pennsylvania; although Dryopteris spp. ferns are not generally a preferred food of deer, in conditions of overpopulation, browse has been observed on Clinton’s fern, especially in eastern Pennsylvania (PNHP Biotics database).

Status Justification Based on the number of extant populations and individuals in Pennsylvania, and the low-to-moderate threat level faced by this species, we have determined that Pennsylvania Threatened is the appropriate

53 conservation rank for this species. Although the number of populations is at the higher end of the usual range for Pennsylvania Threatened, the number of individuals is on the lower end, and this estimate is likely inflated somewhat by the inclusion of some hybrids. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

* All historical records and 23 of 24 extant records for D. clintoniana are based on specimens. All specimens deposited at Carnegie Museum of Natural History and Cleveland Museum of Natural History (19 of 24 extants, 28 of 57 including historical locations) have been examined to determine whether they are hybrids. Four of these were collected too early in the year to have spores, and thus could not be determined with certainty. The remaining records are based on specimens deposited at the Pennsylvania Academy of Natural Sciences or other herbaria, and have not yet been examined.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Montgomery, James D. 1976. “The Distribution and Abundance of Dryopteris in New Jersey.” American Fern Journal 66 (2): 53–59. doi:10.2307/1546519. • Sessa, Emily B., Elizabeth A. Zimmer, and Thomas J. Givnish. 2012. “Unraveling Reticulate Evolution in North American Dryopteris (Dryopteridaceae).” BMC Evolutionary Biology 12 (1): 104. doi:10.1186/1471-2148-12-104. • Wagner, Warren H., and Katherine Lim Chen. 1965. “Abortion of Spores and Sporangia as a Tool in the Detection of Dryopteris Hybrids.” American Fern Journal, 9–29. • Wagner, W. H., F. S. Wagner, and W. Carl Taylor. 1986. “Detecting Abortive Spores in Herbarium Specimens of Sterile Hybrids.” American Fern Journal, 129–40. • Whittier, P., and Warren H. Wagner. 1971. “The Variation in Spore Size and Germination in Dryopteris Taxa.” American Fern Journal, 123–27.

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Schweinitz’s Water-weed – Elodea schweinitzii (J.E. Planchon) Caspary Author: Dr. Timothy Block, Morris Arboretum

Introduction “A taxon of unknown provenance” and “. . . something of a taxonomist’s nightmare” are phrases used by Cook and Urmi-König (1985) in describing Elodea schweinitzii. Globally, all that is known of this taxonomic entity are collections from only two or three poorly defined locations in New York and Pennsylvania, and only between the years 1816 and 1832. In both areas, it would have been sympatric with E. canadensis and E. nuttallii. The specimens in question may be considered within the natural range of variation of E. canadensis or E. nuttallii, in which case E. schweinitzii is not a distinct taxon; some authors also suggest the specimens may be non-persistent hybrids. Even if this limited set of ambiguous historical specimens are recognized as a distinct species, it would now appear to be extinct because it has never been relocated or collected again since 1832, despite repeated searches of the probable locations of the historical collections.

The last major taxonomic work to treat Elodea schweinitzii as anything more than an extinct taxon was the monographic treatment of the genus by St. John (1965). Gleason and Cronquist (1991) note that it is “. . . apparently extinct. It may reflect hybridization between E. canadensis and E. nuttallii.” Additionally, Elodea schweinitzii was not accepted as a good taxon by Haynes in Flora of North America, volume 22 (2000).

Proposed PA Status For reasons noted elsewhere in this report this species should be listed as PX, or, if deemed appropriate based on taxonomic doubt, removed from the list of PA species altogether.

Habitat Probable flowing water of slow-moving streams.

North American Distribution and Global/Regional Conservation Concerns Only known from two or three locations globally. Fishkill Creek, Dutchess County, NY; and vicinity of Bethlehem, PA. Currently listed as PX/SH in PA and as SX in NY.

Pennsylvania Distribution Only one specimen was ever collected in PA, with vague locational data. Possibly from the Lehigh River near Bethlehem (1829-1832), although this time period would not preclude the possibility that the plants were collected from the Lehigh Canal, or a pond of some sort.

Extant Locations None.

Historical Locations and Potential for Relocation See above, essentially no potential for relocation.

Threats Not applicable.

Status Justification

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A consistent character state of the known Elodea schweinitzii specimens is the presence of bisexual flowers. Although bisexual flowers are known from other Elodea species, the vast majority of specimens are dioecious. Cook and Urmi-König (1985) do note, however, “. . . the closest patristic relative (of Elodea) is probably Apalanthe, a genus with exclusively bisexual flowers.” A detailed description of the species is given on pages 151-152 of Cook and Urmi-König, but they conclude by noting that the leaf morphology of E. schweinitzii falls within the range of variability exhibited by E. nuttallii and E. canadensis.

Ironically, Caspary notes in his publication of this species (Caspary, 1858) that his description is based on a specimen having only two bad, half-destroyed flowers (“. . . zwei schlecten, halb zerstörten Blüthen . . .”). He further notes that Elodea schweinitzii is a highly questionable species and probably only a form of E. canadensis (. . . ist jedenfalls eine höchst fragliche Art und wohl nur eine Form von Elodea canadensis . . .).

Whether a truly extinct, but distinct taxon of, perhaps, hybird origin, or an example of a spatially and temporally limited form of a more common taxon, conservation concern is unwarranted in the case of Elodea schweinitzii. Although none of the collection locations were well described, repeated searches in the probable areas where this plant was found have not relocated it nor turned up any new locations.

Additional research into this taxon would not be productive given the age and condition of the few specimens held in collections, and the improbability of this species reoccurring. The PABS – VPTC recommended in 2017 to remove this taxon from the list of those present in the state, with a caveat to remain alert for any perflect-flowered individuals when E. nutallii is encountered.

References • Caspary, R., 1858. Die Hydrilleen (Anacharideen Endl.). Jahrbücher fur Wissenschaftliche Botanik 1: 377-513. • Cook, C.D.K. and K. Urmi-König, 1985. A revision of the genus Elodea (Hydrocharitaceae). Aquatic Botany 21: 111-156. • Gleason, H.A. and A. Cronquist, 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd edition. New York Botanical Garden, New York. • Haynes, R.R., 2000. Hydrocharitaceae. In Flora of North America. Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 22. • St. John, H., 1965. Monograph of the genus Elodea (Hydrocharitaceae). IV. The species of eastern and central North America and summary. Rhodora 67: 155-180.

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Slender Wheat Grass – Elymus trachycaulus (Link) Gould ex Shinners Author: Jessica McPherson, PNHP

Introduction Slender wheat grass can be confused with some other similar grass species, so care should be taken in identification of specimens and in training of botanical surveyors for this species.

Proposed PA Status Slender wheat grass currently has a status of Tentatively Undetermined as recommended by the PABS- Vascular Plant Technical Committee, and there is not sufficient information available at this time to assign a more precise status. We recommend additional training among botanical surveyors to raise awareness of this species, and reassessment in several years when more field data may be available.

North American Distribution and Global / Regional Conservation Concerns The major portion of this species’ geographic range in eastern North America is north of Pennsylvania, with Pennsylvania at the southern edge of the range (Kartesz 2003). NatureServe lists slender wheat grass as Imperiled (S2) in Ohio, West Virginia, and Virginia, while it is listed as Critically Imperiled (S1) in Illinois, New Jersey, and North Carolina.

Habitat In the major part of its range, the habitat is reported to be quite variable in Flora Novae-Angliae: “, cliffs, balds, or ledges, fens, forests, meadows and fields, ridges or ledges, shores of rivers or lakes, talus and rocky slopes, woodlands” (Haines, Farnsworth and Morrison 2011). In Pennsylvania, it is known from dry woodlands and openings, often at the edge of a road or right-of-way, although sometimes in natural settings.

Pennsylvania Distribution In Pennsylvania slender wheat grass is known predominantly from the northeast, consistent with a species that requires a more northern climate.

Extant Locations There are 19 extant populations known. Populations sizes are generally fairly small, ranging between 10 or fewer individuals (5 occurrences ) to 100-300 stems (6 occurrences); most fall in the middle of these extremes. The total number of plants in the state is estimated between 680 and 1200. The species’ ability to use early successional habitat, including some anthropogenic types, suggests it may be more common than current documentation indicates.

Historical Locations There are 11 historical locations documented at this time, based on museum collections. Because this species requires somewhat open, early successional habitat, which typically transitions to less suitable forest habitat without active disturbance, many historical locations may no longer host the species. Most historical locations also have quite vague locational data. It may be possible to relocate a few populations among the historical locations, but not enough to contribute to any significant change in the total number of known populations.

Status Justification The current status proposed by the VPTC is Tentatively Undetermined, and at this time we do not believe enough information is available to further refine this status. While the number of populations

57 and individuals currently known are well within the guidelines for Pennsylvania Threatened, the species’ ability to use anthropogenic early successional habitat suggests that sufficient numbers may eventually be found to justify Pennsylvania Rare or Watch List status. Anecdotal reports from surveyors indicate it may be more common than previously thought. As a counterpoint, however, if it does require cooler climates, it may be at risk from anthropogenic climate change.

Despite the current uncertainty regarding the appropriate status, we do not believe that this species is a high priority for targeted fieldwork, because anthropogenically disturbed early successional habitat is the most likely terrain for finding additional populations, and this type of habitat is difficult to effectively target for successful surveys. While there may be additional populations, there is so much potential habitat that the likelihood of finding the species on any given survey is low. Surveys targeted for this habitat type are also inefficient because it is unlikely to host other elements of conservation concern. It is more efficient to gather knowledge by educating botanical surveyors who work in the appropriate region to look for this species while doing other work. We recommend offering training to encourage this work, reviewing available information for this species again in several years, and revisiting the status of this species if substantial new information has been added.

References • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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White Trout-lily – Erythronium albidum Nutt. Author: Steven P. Grund, PNHP

Introduction White trout-lily is a spring-blooming, clonal, herbaceous annual in the lily family (). The white petals, suffused with a pale blue-violet on the back, as well as more subtle differences in flower and fruit characteristics distinguish this species from Erythronium americanum (yellow trout-lily), the only other member of the genus known from Pennsylvania. It grows on floodplains and lower slopes in rich forests.

Proposed PA Status The Department of Conservation and Natural Resources proposes to classify this species as Pennsylvania Rare (PR) because the species has likely declined significantly due to conversion of habitat to agriculture, industry, and other uses. Although regulations and economic factors have slowed such conversions, floodplains are under continued threat from a variety of other factors, leading us to conclude that the decline will continue.

North American Distribution and Global / Regional Conservation Concerns Erythronium albidum has a distribution centered in the Mississippi Valley. From Minnesota south roughly along the 97th parallel to central Texas. From there northeast to eastern Maryland, north to Massachusetts, New York, southern Michigan and adjacent Ontario, and Wisconsin (Kartesz 2017, Brouillet et al 2017). Globally secure, this species is of conservation concern in a number of states at the periphery of its range: S1 in North Dakota and ; S2 in Louisiana, Mississippi, Virginia, and Maryland.

Pennsylvania Distribution White trout-lily is most abundant in southwestern Pennsylvania and along the lower Susquehanna River, with scattered stations elsewhere, but apparently absent from the north-central portion of the state.

Habitat Floodplains and rich deciduous forests on lower slopes, especially in limestone regions (Rhoads and Block 2007, Pennsylvania Natural Heritage Program 2017).

Threats The most significant threat is invasive species. The floodplain habitat favored by this species is particularly prone to invasion by a number of species, including (but far from limited to) some of the most invasive species in Pennsylvania, Japanese knotweed (Fallopia japonica), and Japanese stiltgrass (Microstegium vimineum). Other threats, individually moderate but additive in effect, include logging, grazing, construction and maintenance of roads, railroads, sewer lines, and pipelines, habitat degradation by all-terrain-vehicles, and others. None of these threats are unique to floodplains, but all are more severe for floodplains.

Status Justification The number of occurrences known to be extant (48) is higher than typical for a species we consider appropriate for the PR category, but the number of individuals (well under 100,000) is low for a PR species. Combining the rarity with trend data and a threat analysis, we concur with the rank of S3 generated by our use of NatureServe’s Conservation Rank Calculator (NatureServe 2015). This is

59 consistent with a status of Pennsylvania Rare under the Wild Resource Conservation Act and associated regulations.

References • NatureServe. 2015. NatureServe Conservation Status Assessments: Rank Calculator Version 3.185. NatureServe, Arlington, VA. Online at http://www.natureserve.org/conservation- tools/conservation-rank-calculator • Pennsylvania Natural Heritage Program. Biotics Database. Accessed March 2017. • Rhoads, A.F. and T.A. Block. 2007. Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press. Philadelphia.

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Godfrey’s Eupatorium – Eupatorium godfreyanum Cronquist Author: Jessica McPherson, PNHP

Introduction According to Flora of North America, “Eupatorium godfreyanum is an apomictic polyploid derivative that includes genomes from E. rotundifolium and E. sessilifolium.” This indicates that the species does not reproduce sexually, and offspring are genetically identical to their parents. It may have originated through hybridization of E. rotundifolium and E. sessilifolium. Taxa derived from hybrid origin can have multiple origins (in which case several genetically distinct clone lines may be included in a concept with a single name); information on whether E. godfreyanum has single- or multiple- hybrid origin does not appear to be available yet. Hybrid-origin taxa can also be ephemeral (in the case of hybrids offspring that are sterile) or persistent (in the case of a hybrid which does have reproductive capacity and competitive ability, and spreads to establish its own range). To comment on E. godfreyanum’s place on this spectrum, FNA continues: “Although it is relatively narrow in distribution, it is known to occur in localities where both progenitor species are absent and it seems to be persistent where it occurs.” (Siripun and Schilling 2006).

Proposed PA Status We concur with the PABS-Vascular Plant Technical Committee’s recommendation that Godfrey’s eupatorium be listed as Pennsylvania Threatened, due to the very low number of populations and individuals known in the state. Although the numbers are most consistent with Pennsylvania Endangered, Pennsylvania Threatened is proposed instead of Endangered because the species’ tolerance of early successional habitat and ability to establish in non-pristine environments leaves some uncertainty around the number of additional populations that may eventually be found.

North American Distribution and Global / Regional Conservation Concerns The documented global range is predominantly south of Pennsylvania, centered around the southern Appalachians; it is most widespread in Virginia, West Virginia, and North Carolina, with outlying populations in Kentucky, Tennessee and Pennsylvania (Kartesz 2003). It is listed of conservation concern by NatureServe in most states where it is known to occur; Imperiled (S2) in West Virginia, Vulnerable (S3) in North Carolina, and Critically Imperiled (S1) in Tennessee and Georgia. It has a global rank of Apparently Secure (G4).

Pennsylvania Distribution, Extant and Historical Occurrences Godfrey’s eupatorium is limited to the south-central region of the state, and these populations are the northern extent of its known global range. Within Pennsylvania, it is clearly more established than a waif, with 5 extant and 4 historical populations known from 5 south-central counties, including multiple populations in Dauphin and Lebanon counties; but it still very scarcely documented and regionally limited. The total number of individuals at known extant sites is estimated to be 1800-2000.

Habitat This species requires early successional habitat (“Dry, open, disturbed sites, edges of deciduous woods” according to Flora of North America) and may not persist at sites where such conditions are not maintained by disturbance.

Threats This species faces threats in several of its known locations from roadside spraying, which is not at this time being appropriately managed by PennDOT to avoid damage to plants of conservation concern.

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Status Justification The current PABS-VPTC proposed status is Pennsylvania Threatened, which is consistent with our knowledge of the species at the present time. Although a small number of populations are currently known, the species is tolerant of disturbance, clearly has the capacity to establish in non-pristine sites, and suitable early successional habitat is abundant in Pennsylvania. Therefore it is not considered to be Pennsylvania Endangered. With present knowledge, Pennsylvania Rare status is also inappropriate due to the global rarity of the species and the extremely limited, range-edge distribution currently documented in Pennsylvania, along with the threats faced by some populations. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

Although there is some uncertainty regarding the number of potential additional populations that may be undocumented, we believe targeted surveys for this species are an inefficient use of resources, due to the abundance of suitable early-successional habitat. Because the species is uncommon, but the habitat is very common, any given survey has a low likelihood of success, and surveys in disturbed early successional habitat typically do not contain other high-value conservation targets. It is more efficient to gather knowledge about this species by educating botanical surveyors who work in the appropriate region to look for this species while doing other work.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Siripun, Kunsiri Chaw and Edward E. Schilling. 2006. Eupatorium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 21.

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Threepetal Bedstraw – Galium trifidum L. Author: Jessica McPherson, PNHP

Introduction Threepetal bedstraw is a perennial in the madder family (Rubiaceae). It is similar in appearance to Galium tinctorium, but can be distinguished when fruit are mature by pedicels longer than 1 cm, which have small barbs pointing backwards away from the fruit (retrorsely scabrous). Galium tinctorium has flower/fruit pedicels shorter than 1 cm, which are usually smooth, although sometimes have small barbs pointing towards the fruit (antrorsely scabrous).

Proposed PA Status We recommend a status of Pennsylvania Rare for threepetal bedstraw due to the moderately low number of populations known in the state, in combination with the threats of invasive species and climate change. Pennsylvania Rare is the status currently recommended by the PABS-Vascular Plant Technical Committee.

Habitat This species occupies open wetlands of a variety of types (marsh, fen, swamp). The pH is usually on the higher end, although it is not as restricted as some calcium-loving species. It can tolerate some disturbance; it is found both in intact fens and in more successional marsh-type habitats. However, only a small minority of occurrences are known from highly disturbed locations such as wet areas of ROWs. It is believed that earlier comments attributing “weediness” to this species refer to its ability to occupy a broader range of habitat than some more conservative fen species, as the plant has not demonstrated aggressive or problematic growth.

North American Distribution and Global / Regional Conservation Concerns Threepetal bedstraw is widespread in western North America, but in eastern North America it is concentrated in more northern regions, fairly widespread from North Dakota east to and more scattered further south in Illinois, Indiana, Ohio, Pennsylvania, and Maryland (Kartesz 2003). It has a NatureServe rank of Vulnerable (S3) in Vermont and Iowa.

Pennsylvania Distribution Threepetal bedstraw is near the southern edge of its range in Pennsylvania. The known distribution in Pennsylvania is similar to the typical pattern of northern fen species, with concentrations in the NW, the eastern Ridge and Valley / Piedmont, and the central Ridge and Valley; however, it has a more expansive presence in the NE than is typical for calcareous species, likely due to its ability to utilize the lower end of higher pH habitats.

Extant Locations As part of Plant Status Collaborative fieldwork in 2014 and 2015, Jim Bissell located five new populations of threepetal bedstraw. • Tioga County: Hills Creek Lake State Park (fen) and Ross Tract on Marsh Creek (emergent marsh). • Erie County: Edinboro Lake Fen. • Warren County: Tamarack Swamp (inundated fen). • Crawford County: SGL 214 (a fairly extensive population occupying a variety of habitats including calcareous emergent marsh, emergent fen, and swamp forest/shrub fen).

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Heritage staff also discovered 2 new locations in Monroe County, 3 new locations in Wayne County, and 1 new location in Luzerne County. There are now 51 extant populations known.

Historical Locations and Potential for Relocation There are 12 historical locations known for threepetal bedstraw; about half of these may be possible to relocate, based on the existence of potential habitat matching reasonably precise location descriptions from specimens. However, the number of populations this could potentially add to our knowledge of the species in Pennsylvania is small enough that it will not greatly impact determination of the appropriate status for threepetal bedstraw.

Threats The majority of known populations for this species are in the glaciated NW portion of Pennsylvania, where invasive species are already prevalent in wetlands. Common reed (Phragmites australis spp. australis), reed canarygrass (Phalaris arundinacea), and glossy buckthorn (Frangula alnus) are well established and have already transformed many fen habitats to the detriment of native species. Over the next 30 years, unless dramatic action is taken, habitat loss to invasive species is likely to continue, significantly reducing the extent of this species. Pennsylvania populations of Galium trifidum may also be vulnerable to climate change, because they are near the southern edge of the global range for the species.

Status Justification Despite the addition of some new locations, the available data on threepetal bedstraw currently support a designation of Pennsylvania Rare. The number of known extant populations (51) is consistent with this status, in combination with the threats it faces from invasive species and climate change. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S3 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Canada Hawkweed – Hieracium umbellatum L. Author: Dr. Timothy Block, Morris Arboretum

Introduction Hieracium umbellatum has a complex taxonomic history which includes many synonyms and its current circumscription remains somewhat unresolved. See Flora of North America, volume 19, pg. 287 (2006) for a thorough list of synonymy.

The Pennsylvania specimens of this species were formerly referred to H. kalmii (Rhoads and Klein, 1993), and Gleason and Cronquist (1991) treated both H. kalmii and H. umbellatum as distinct species, noting that H. umbellatum might best be recognized as a variety of H. scabriusculum. However, Rhoads and Block (2007) chose to follow the treatment by Strother in Flora of North America (2006), placing H. kalmii in direct synonymy with H. umbellatum.

In any case, specimens of Hieracium umbellatum herein treated are distinguished by being generally less pubescent than most other species of the genus, and by possession of short, stout hairs on the leaf margins, as well as some stellate hairs on the leaf surfaces.

Worthy of note also is the fact that Hieracium umbellatum is a circumboreal plant of northern affinities, occurring with regularity in boreal areas of Canada and at higher elevations to the south.

Proposed Status and Justification Hieracium umbellatum should be ranked as PE (S1) in Pennsylvania. It is currently known only from two small, local populations in Erie County.

Habitat Habitat, in the broad sense for this species, consists of fields, roadsides, rocky slopes, forest openings, prairies, and thickets. In Pennsylvania, current habitat is in thickets and thin woodlands.

North American Distribution and Global/Regional Conservation Concerns Hieracium umbellatum is ranked S1 in Pennsylvania, West Virginia, and Nebraska; and S2 in Ohio. Not currently of conservation concern throughout the rest of its range, it is, curiously, considered exotic in Ontario and Iowa.

Pennsylvania Distribution As noted above, two locations are known from Erie County, Erie Bluffs, and Presque Isle state parks. All other locations are historical with the possible exception of a collection made by Reif in 1984 near Beaumont in Wyoming County. This specimen is in the herbarium of Wilkes College (WILK), but has not been examined to confirm identification.

Most historical collections of this species are from northeastern PA with a few from central PA in the Ridge and Valley Province of Centre or Huntingdon counties.

Extant Locations As noted above.

Historical Locations and Potential for Relocation

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Most historical collections include very little specific location information, not an unusual situation. The probability of relocation based on given information would be marginal, at best. The best strategy for location of new populations would be to conduct extensive surveys of rocky ridge tops in northeast Pennsylvania.

Threats The most apparent threat to the extant populations is the instability of the habitat and the relative smallness of the populations present.

Status Justification Hieracium umbellatum was likely never common in Pennsylvania. The earliest collection from PA was from Huntingdon County in 1865. Most other collections were made over a period from the 1890s until the 1940s. Since 1946, the species has been collected just three times as noted above.

This species probably benefited from the vast amount of open habitat provided by mass clear-cutting of forests from the late 19th to the early 20th centuries. Even so, most collections for which habitat information is included were made from ridge tops, open rocky woodlands, and other places not unlike the species’ more northern locations.

Based on the limited Erie County occurrences, and including the unconfirmed Wyoming County record, there are still so few locations and so few plants as to justify a ranking of Pennsylvania Endangered.

References • Gleason, H.A. and A. Cronquist, 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd edition. New York Botanical Garden, New York. • Rhoads, A.F. and W.M. Klein, 1993. The Vascular Flora of Pennsylvania: Annotated Checklist and Atlas. American Philosophical Society, Philadelphia. • Rhoads, A.F. and T.A. Block, 2007. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia. • Strother, J.L., 2006. Hieracium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 19.

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Virginia Sweetspire, Virginia-willow, Tassel-white – Itea virginica L. Author: Dr. Timothy Block, Morris Arboretum

Introduction Itea virginica is the only North American species of a genus that is much more diverse in East . Although I. virginica is widely planted and dozens of cultivars are sold in the horticultural trade, this species appears to seldom naturalize from cultivated sources. The small, smooth seeds probably don’t allow for ready dispersal over a significant distance from the parent plant. Individuals of I. virginica are strongly clonal and it is always difficult to determine the limits of each genetically distinct plant.

Proposed PA Status Itea virginica should be listed as Pennsylvania Endangered (PE) due to the fact that only one natural population exists in the state. Although that one population is in a county park, and presumably afforded some level of protection, the population size is small and probably consists of only one or a few individuals.

Habitat Morin (2009) lists swamps, wet woods, stream banks, stream heads, and beaver ponds as habitat for this species. The only extant site in PA is, indeed, a wet woodland. However, the use of I. virginica as a “foundation” plant in the horticultural trade speaks to this species’ ability to, at least, survive in a broad range of habitat types including much drier conditions that would be expected for natural occurrences.

North American Distribution and Global / Regional Conservation Concerns Itea virginica ranges from Pennsylvania south along the Coastal Plain to Florida, and hence west to Texas. It occurs inland as far west as Oklahoma and north to Illinois and Indiana, especially along the Mississippi River Valley. This species is of conservation concern only in Indiana and Oklahoma where it is listed as S1, as well as in Pennsylvania where it is currently listed as Extirpated (PX), but suggested by the PA Biological Survey as Endangered (PE).

Pennsylvania Distribution The current distribution for Itea virginica consists of one location on the Atlantic Coastal Plain in lower Bucks County. This SE PA location is at the extreme northern edge of the global range.

Extant Locations The specific extant location is in Black Ditch County Park in Bucks County at coordinates 40.131667, - 74.888333 (WGS84). This population covers an area of approximately 80 square meters.

Historical Locations and Potential for Relocation The only other Pennsylvania location for Itea virginica is based on a collection made in 1841 by McMinn. The specimen label states only that it was collected along the Susquehanna River, opposite Harrisburg in Columbia County. Due to the extensive development of this area, it would not be reasonable to expect that population to be extant.

Threats Some evidence of browsing by deer exists on the plants at the Black Ditch County Park site. Although this does not seem extensive enough to be a threat to long-term survival of the population, browsing of stem tips and inflorescences may be limiting to possible dispersal. However, we don’t currently know whether or not this population is producing viable seed.

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The site is a disturbed woodland and home to a number of non-native, invasive plant species such as Berberis thunbergii, Celastrus orbiculatus, Lonicera japonica, Viburnum spp., and many others. These, however, don’t seem to pose a serious threat to the Itea virginica population at this time.

Scion collection for horticultural purposes has been done from time to time over the years, but likely presents little threat unless undertaken more intensely.

Status Justification The lone extant population of Itea virginica in Pennsylvania was discovered by Rhoads and Block while conducting the Natural Areas Inventory of Bucks County in 1998. This population is believed to be a natural occurrence because 1) correct habitat type is present, and 2) low propensity of the species to naturalize from cultivated sources, and no cultivated sources nearby. Being a clonal species, this population may be a remnant of a formerly larger population that managed to survive repeated major disturbance events of the past.

Many other wooded Coastal Plain sites have been checked over the years without finding other natural populations, or indeed, any other populations of this species outside of cultivation.

It might be conceivable that Itea virginica could turn up again somewhere in the lower Susquehanna Valley in Pennsylvania, if sufficient effort was made to survey in detail all of the available habitat.

References • Morin, N.R. 2009. Itea virginica. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 22, pp. 6-7.

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Grass Rush – Juncus biflorus Elliott Author: Dr. Timothy Block, Morris Arboretum; Jessica McPherson, PNHP

Introduction Grass rush (Juncus biflorus) is a rhizomatous rush species; it often grows to one meter or more in height and is distinguished from the more common J. marginatus in Pennsylvania by its larger size and greater number of flowers (up to 100) per . The mature capsules turn chestnut brown. Although Brooks and Clemants in Flora of North America (2000) consider this species as part of J. marginatus, the appearance and habitat of J. biflorus in Pennsylvania is distinctive enough to maintain this taxon at the species level, which is supported by Knapp and Naczi (2008). Michigan Flora, Flora Novae Angliae, and the New York Flora Atlas (Weldy, Werrier and Nelson 2016) also follow this work.

Proposed PA Status We concur with the 2015 recommendation of the PABS-Vascular Plant Technical Committee that the appropriate status for grass rush is Pennsylvania Rare (PR). Although current knowledge of the rarity, threats towards, and viability of this species are either consistent with Pennsylvania Threatened or ambiguous in their indication of Rare or Threatened, Pennsylvania Rare is recommended because the species has a demonstrated ability to occupy disturbed habitats and to establish in new locations.

Habitat The grass rush is found in wet, open woods, wet meadows, boggy fields, swales, ditches, and even gravel pits. In Pennsylvania it is found exclusively on calcareous or diabase substrates, although this characteristic is not mentioned in sources such as Michigan Flora (Reznicek, Voss, and Walters 2011) or Flora Novae Angliae (Haines, Farnsworth and Morrison 2011) that address other portions of the range.

North American Distribution and Global / Regional Conservation Concerns The global range of this species is centered in southeastern North America, from Florida extending west to Texas, and north to Michigan, New York, and Ontario.

Grass rush is ranked by NatureServe as Critically Imperiled (S1) in New York and Ontario; as Imperiled (S2) in West Virginia; and as Vulnerable (S3) in Michigan and Pennsylvania. In all other states where a conservation rank has been assigned it is listed as Secure (S5) or Apparently Secure (S4). Its global ranking is Secure (G5). It is not clear whether or not various state authorities treat J. biflorus as a variety of J. marginatus or as a species in its own right, and what effect on ranking the taxonomic issue might have.

Pennsylvania Distribution and NatureServe State Conservation Rank In Pennsylvania, grass rush is concentrated in the southeastern portion of the state, with scattered populations in south-central PA and two extant records in western PA (Butler County and Erie County). It appears to be exhibiting range-edge characteristics in its Pennsylvania distribution, even though scattered populations do extend further north into New York and Canada. The distribution is also likely affected by its requirement for calcareous or diabase soils, which are largely absent from northern Pennsylvania except in the glaciated northwest.

Extant Locations In Pennsylvania there are 39 extant occurrences (25 in SE PA) and 25 historical occurrences currently documented, with an estimated 6000-7000 plants in total throughout the state. Most populations are extremely small, and without active management to maintain open habitat, populations decline; only 7

69 populations are considered viable based on PNHP rankings. The one very healthy population is at Natural Lands Trust’s Crow’s Nest Preserve in Chester County where NLT actively maintains open meadows through suppression of woody succession. That population consists of approximately 4,000 above-ground stems, the majority of all plants known in the state (all other documented populations combined contribute 2000-3000 additional stems). As with all strongly rhizomatous species, only genetic analysis can tell us how many distinct individual plants are present in a given population.

Historical Locations and Potential for Relocation Historical locations are unlikely to be a particularly fruitful source for locating additional extant populations; most historical collections are described as occurring in early-successional habitats which have likely been altered in the intervening time. Most also occur in SE Pennsylvania, where the extent of landscape change and the level of redundancy in early collections (combined with vague location data) make it very unlikely that individual known historical collections correspond to individual extant populations.

Threats The primary threats faced by this species are loss of habitat due to woody succession in wet meadows and old fields, and out-competition by invasive species. Some populations are also threatened by development or roadside maintenance. Active management of woody vegetation would be necessary to reverse the current downward trend in this species in eastern Pennsylvania.

Although the species is currently quite rare, faces significant threats, and is limited by its habitat specificity, it is also clearly not confined to relict natural habitats. Many of the currently known populations are in old fields, ditches, or roadsides where it is clear from the land use history they cannot have persisted from pre-settlement vegetation. This demonstrates an ability of the species to establish new populations in disturbed habitat types that are fairly abundant, although the requirement for calcareous/diabase substrate does restrict the available habitat somewhat.

Status Justification Although the rarity of the grass rush and the level of threat it faces are consistent with the definition of the Pennsylvania Threatened status, we recommend a status of Pennsylvania Rare based on the number of extant populations in combination with the species’ demonstrated ability to establish new populations and to utilize disturbed, anthropogenic habitat. Habitat conservation tools invoked by the PT status are less useful in the conservation of this species than many others, due to its utilization of anthropogenic habitats and the natural mobility of populations over time. However, active management of some known populations to maintain early-successional conditions will likely be critical to its long- term success in the state. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S3 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Brooks, Ralph E. and Stephen E. Clemants. 2000. Juncus. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 22. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA.

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• Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Knapp, Wesley M., and Robert F. C. Naczi. 2008. Taxonomy, Morphology, and Geographic Distribution of Juncus Longii (Juncaceae). Systematic Botany, vol. 33, no. 4. 685-694. • MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. September 26, 2016. http://michiganflora.net/species.aspx?id=1502. • Weldy, Troy, David Werier, and Andrew Nelson. 2016 New York Flora Atlas. [S. M. Landry and K. N. Campbell (original application development), USF Water Institute. University of South Florida]. New York Flora Association, Albany, New York.

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Fetter-bush — Leucothoe racemosa (L.) Gray Author: Dr. Timothy Block, Morris Arboretum; Jessica McPherson, PNHP

Introduction Fetter-bush (Leucothoe racemosa) is a strongly rhizomatous shrub in the heath family (Ericaceae). It is distinguished from other shrubby members of the Ericaceae by technical characters that include the presence of two distal bracteoles on the pedicels and by apical awns on the anthers. Otherwise, fetter- bush is generally similar in overall appearance to highbush blueberry (Vaccinium corymbosum) and other ericaceous shrubs.

Proposed PA Status Fetter-bush is proposed to move from Tentatively Undetermined (TU) to Pennsylvania Rare (PR), based on the moderate number of populations and individuals known, the apparent viability of many of these populations, and the existence of ongoing threats to many populations. This status change is suggested due to recent data collection efforts which helped clarify the conservation situation with this species. The PABS-Vascular Plant Technical Committee recommended the Pennsylvania Rare status in 2015.

North American Distribution and Global / Regional Conservation Concerns The global range of fetter-bush is along the gulf coast of North America from Texas to Florida, and along the Atlantic coast from Florida to Massachusetts; although it is concentrated in coastal regions, it also extends inland somewhat in many of the coastal states where it is found. Fetter-bush is listed as globally secure (G5) by NatureServe and as Imperiled (S2) only in Tennessee and Vulnerable (S3) in Pennsylvania; Pennsylvania has very little coastal plain habitat, while Tennessee is the only landlocked state where it occurs and hosts scattered inland populations. In all other states where a rank has been assigned it is listed as secure (S5) or apparently secure (S4).

Pennsylvania Distribution and Habitat In Pennsylvania, the species is known primarily from wet, acidic soils in the limited area of Atlantic Coastal Plain habitat in the SE corner of the state, but is also found in swampy woods and vernal pools mainly in the region of the Delaware River and in scattered other Ridge and Valley locations.

Extant and Historical Locations There are 34 extant populations known in the state; 16 of these are from the Coastal Plain, where most historical records were also recorded, and it is estimated there may be substantially more populations in this area. There are several historical locations in Berks, Lancaster, Lehigh, and York counties which have not been relocated; the total number of historical records is 24. There are 18 extant populations known from other portions of the Ridge and Valley; interestingly, historical specimens were not known from this region and these populations have all been documented in recent times. However, habitat is more limited here and it is estimated that fewer additional populations (possibly about 10) may exist.

The largest known population is in the Delhaas Woods section of Silver Lake Nature Center in Bucks County, which is home to several sub-populations of fetter-bush totaling more than 50,000 above- ground stems. However, as with all strongly rhizomatous species, we do not know how many genetically distinct individuals these plants represent. Most other known populations are much smaller, with a few dozen plants present.

Threats

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The setting and threat level faced by populations in the Coastal Plain and in the Ridge and Valley are distinct. In the Coastal Plain, natural habitat has been greatly reduced from pre-settlement times, and remaining areas of natural habitat are generally small and very fragmented, under high threat from invasive species and continued development. However, fetter-bush can be found in even small 1-5 acre fragments, and appears to be fairly persistent despite the challenges to viability that such fragments generally face; it seems to be somewhat minimally affected by invasive species, and is not a preferred deer-browse species.

In the Ridge and Valley, fetter-bush is often indicative of higher quality habitats and found in more intact landscape settings. The level of threat is also lower, because the rate of development in this region is currently lower than in the coastal plain, and because wetlands are afforded some legal protections.

Status Justification Pennsylvania Rare is recommended as the appropriate status for fetter-bush in the state, because the number of populations known and the number of reproductive units (individuals cannot be estimated with a clonal species) known in total are consistent with the definition of this status, supported by the persistence and apparent viability of these populations. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S3 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Red Mulberry – Morus rubra Linneaus Author: Jessica McPherson, PNHP

Introduction The red mulberry is the only native species of mulberry known in Pennsylvania; the white mulberry (Morus alba) is an exotic member of the genus Morus that has been introduced and is now widespread. Today, the exotic white mulberry is much more commonly encountered than the native red mulberry. The two species do hybridize and produce fertile offspring with intermediate physical characteristics. The species are best distinguished by leaf, bark, and bud characteristics; despite the common names, fruit color is not a reliable way to distinguish the two. Red mulberry has dark purple fruit, while white mulberry has fruit ranging in color from dark purple to white. White fruits do indicate white mulberry, but purple fruits may be either species.

Proposed PA Status We concur with the PABS-Vascular Plant Technical Committee’s 2014 recommendation of the Pennsylvania Threatened status for the red mulberry, because its population in Pennsylvania appears to have dramatically declined over the last century, and very few extant populations are currently known in the state. It faces ongoing threats from hybridization with the exotic white mulberry (Morus alba), disease, and environmental stressors.

North American Distribution and Global / Regional Conservation Concerns The red mulberry is known from a wide geographic area, from Florida to Texas and north to Minnesota, Ontario, and Vermont (Kartesz 2003). It is rare across the northern edge of its range, and even in Pennsylvania historical collections are distributed far more abundantly in the southern half of the state than in the northern half (PA Flora Project). Sources across many parts of the range report that the species is presently uncommon or has declined (see below). However, it is only listed by NatureServe in states and provinces in the northernmost extent of its range (Imperiled S2 in Ontario and Michigan, Critically Imperiled S1 in Vermont and Massachusetts, Vulnerable S3 in Delaware). Most states have not assigned a conservation rank, although Illinois, Kentucky, West Virginia, New York, and North Carolina consider it Secure (S5).

Pennsylvania Distribution In Pennsylvania, the species is historically recorded primarily from the southern half of the state. This may reflect an affinity for calcareous habitats, which are much less common in the northern half of the state. It may also reflect approaching the northern edge of the range, such that suitable climatic conditions are more limited on the landscape.

Habitat A fairly wide range of habitats are described for the red mulberry in different sources, possibly indicating that the species behaves differently in different parts of its range, or that it has become ecologically constrained in some way related to its decline in recent years. The following assessment of the species’ habitat in Pennsylvania is based on a synthesis of habitat descriptions in the literature and field experience.

Preference for base-rich soils is mentioned by several sources (see below), and one assessment states the optimal soil pH is 6.0-7.5 (Spurway 1941). The few extant locations known for the species in Pennsylvania are all on soils heavily influenced by calcareous or diabase rock. Further to the northeast,

74 drier, rockier habitats are described than in the central part of the range, where floodplains and moist forests are typically mentioned. • Michigan Flora: “A scarce small tree... Floodplains, river bottoms, and swamps.” • Flora of Virginia: "well-drained floodplain forests, mesic and dry-mesic upland forests, old fields, and fencerows; typically, but not exclusively, in base-rich soils." • The Pennsylvania Flora Project: "rich, moist alluvial soils and wooded slopes" • NY Flora Atlas: "fairly local and populations are often small. Rich dry to dry-mesic forests often on calcareous bedrock or soils. Not weedy like M. alba." (This matches our experience in western PA and the Ridge and Valley province.) • Flora Novae-Angliae: “Rocky forests, ridges, edge of talus slopes, base of ledges.”

The PA Flora Project description is likely drawn from specimen habitat descriptions, as the authors have very little personal field experience with the species (personal communication, 2013 Rare Plant Forum). It seems clear from specimen descriptions that the species was at one time known from moist woods and floodplain areas in Pennsylvania, but no extant populations are currently known from floodplains.

Extant Locations This species was first considered by the VPTC at the 2013 Rare Plant forum. It is unusual in that historical specimens are fairly abundant, but very few extant locations are currently known, suggesting that the species may have declined in recent decades. Review of museum specimens revealed that although there are about 150 locations for historical collections, very few recent collections exist. Although there are a variety of potential explanations for a lack of recent collections, a decline in the species is reinforced by the experience of the state’s expert botanists: at the 2013 Rare Plant Forum, only three people in attendance had ever seen the species, despite the attendance of many experienced botanists who had worked in the state for decades. This is highly unusual for a native woody plant species. Combined observations among the group totaled fewer than 10 individuals.

Dedicated surveys to further pursue this species were conducted through the Plant Status Updates project 2014-2016; PNHP field surveys in this time period have located one population of 7 individuals, on diabase soil in eastern Pennsylvania, and three plants that appear to be M. rubra x M. alba hybrids in Greene County. An additional two individuals, one sapling and one mature tree, were discovered by a consultant along a roadside on the shore of the Cheat River at the boundary of Fayette and Greene counties. Plot work for a study of limestone habitats in 2012 found three different locations with one seedling present at each on central-Pennsylvania limestone slopes. Review of past PNHP surveys found three locations in central PA where the species was included on a species list but no further population data was submitted, as the species was not then tracked. One of these sites was visited in 2012 but not found at that time. Other PNHP survey work documented one individual in Allegheny County; the surrounding natural area has been fairly well surveyed with no additional individuals discovered. All other attempts to relocate historical specimens and locate additional populations through de novo searches have been unsuccessful. A total of 6-9 extant populations (3 have not been recently verified) are now known, with a total of 13 individuals.

Historical Locations and Potential for Relocation Historical collections of the red mulberries are fairly abundant, with about 150 locations in the PA Flora Project Atlas, (39 in the Carnegie Museum of Natural History, the remaining records in the Pennsylvania Academy of Natural Sciences or other herbaria). Records are most abundant in the eastern part of the state; in this region, we do not believe the relocation of extant populations based on historical specimen data is likely, because of the extent of landscape change and the poor location data recorded for most

75 specimens. Furthermore, the combination of landscape fragmentation and the prevalence of white mulberry suggests that hybridization may have already greatly reduced the presence of pure M. rubra.

Threats Although historical collections suggest that red mulberry was at least somewhat common in the state, a variety of sources indicate it has declined dramatically in recent decades. Evidence for the decline comes from literature reports, botanical expert experience, a decline in museum collections, and recent field survey efforts that have been largely unsuccessful in finding the species.

Several studies have been conducted on hybridization of red and white mulberry which show that this process is a clear threat to the red mulberry. Red mulberry is a federally endangered species in Canada, where there are only a few populations known from the southernmost extent of Ontario. Because of the species’ legal status, these populations have been studied extensively. Research conducted in Ontario, Canada has demonstrated asymmetrical introgression where red mulberry and white mulberry co-occur (i.e., more offspring are more similar to white mulberry than red mulberry). White mulberry was more abundant at these sites. This is likely to be a common condition because red mulberry appears to be naturally infrequent (often listed as “uncommon,” “infrequent,” or “scattered” in habitat descriptions, even older ones), while white mulberry reproduces vigorously once established. Genetic studies documented that the hybrids contained a higher proportion of white mulberry DNA than red mulberry DNA (Burgess and Husband 2006). Furthermore, seed production in red mulberry was impaired by an abundance of white mulberry pollen, and white mulberry proved more fit than red mulberry in common garden experiments (Burgess et al. 2005).

Reports of diseases affecting red mulberry have been published for several decades, although no causative agent has been definitively agreed upon to be the cause of the apparently widespread decline. The evidence is summarized below:

The USDA Silvics Manual (1990): “Red mulberry seems to be vanishing from at least a portion of its central range, possibly due to a bacterial disease. The effects and extent of this disease have not been investigated thoroughly, but it is known that red mulberry trees are becoming increasingly scarce(Core 1974). The only noteworthy leaf pathogens of red mulberry reported in the United States are leaf spots caused by a species of Cercospora, Mycosphaerella mori, and Pseudomonas mori (Halls 1973).” The silvics manual cites a species account by Earl Core, author of the Flora of West Virginia, which reads as follows: “there is evidence that red mulberry, once common, is vanishing from at least a portion of this range, possibly because of a bacterial disease. Mature fruiting trees are now scarce in many areas. (Affeltranger 1963[sic 1965], Core 1966: 75) …Red mulberry stands of any size are not mentioned in the literature, and the species is becoming scarce in many areas, perhaps because of disease. Pseudomonas mori is reported as causing a moderately severe disease, manifested chiefly as a leaf spot and blight (Pearce and Spaulding 1942). Mycosphaerella mori, the cause of leaf spot in red mulberry, is widespread and abundant. Mulberry is also attacked by the Texas root rot caused by Phymatotrichum omnivorum (Van Dersal, Mulford, and Thornthwaite 1938). The cerambycid borer Dorcaschema wildii is a destructive insect pest southwards (Solomon 1968).” Affeltranger 1965: “Today Red Mulberry is still found on farm lands, in coves and along stream banks but a scarcity is becoming apparent, and large trees of this species are seldom ever seen. Small seedlings may

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not grow to tree size largely because of a bacterial blight caused by Pseudomonas mori. This disease rarely kills large trees, but is quite virulent in nurseries. When wanting to secure ornamentals for planting, many homeowners, directors of arboretums, and other interested people have discovered that Red Mulberry is a difficult seedling to obtain. Bacteriosis of Red Mulberry is characterized by water soaked lesions on leaves and shoots which later become enlarged, sunken and blackened. A most noticeable symptom is the bacterial ooze that comes out of the lenticels in the form of curled strands known as cirri. Many other diseases have Red mulberry as their host, but only Pseudomonas mori is known to be prominent enough to cause appreciable losses.” The Ontario Ministry of Natural Resources’ Recovery Strategy for Red Mulberry (2013) summarizes a great deal of research done in Ontario and elsewhere, and addresses health declines as follows: “Red Mulberry is known to suffer from twig blight, twig dieback, cankers, and root rot (Ambrose et al. 1998). Health assessments of four populations of Red Mulberry indicate that some populations are in very poor health, suffering population-level declines described as a “gradual, general deterioration” (McLaughlin and Greifenhagen 2002; Spisani et al. 2004). The former study concluded that no single pathogen was responsible for the disease symptoms. Rather, several opportunistic, canker-causing pathogens and two opportunistic root disease pathogens affected the diseased trees. These pathogens are not known to infect healthy tissues, but can successfully cause damage to stressed and weakened hosts. Probable factors causing such stress include drought, low soil fertility and/or poor or suppressed canopy position….” Another source (Little 1995) indicates that the species is highly sensitive to air pollution, and that pollution likely makes the species more susceptible to disease: “In West Virginia, ozone damage to Red Mulberry leaves is believed to increase susceptibility to an opportunistic twig canker disease (Nectria cinnabarina) leading to the death of whole trees (O. Loucks pers. comm. 1996).” The cited source of this information is Orie Loucks, professor emeritus at Oxford University in Miami, Ohio, but he could not be contacted due to ill health.

The breeding system of red mulberry is somewhat unique and potentially relevant to its conservation. It is sub-dioecious: most individuals produce only male flowers or only female flowers, but a small fraction produce both male and female flowers. In the only detailed study available (Nepal et al. 2015), conducted at Konza Prairie Long Term Ecological Research station in Kansas, a small fraction (12%) of hermaphroditic trees were observed in each of three study years, and about 10% of trees also switched their sexual status between years. All populations had a much higher proportion of males than females, which was independent of the size of the individual. This may indicate pollen availability is a limitation for red mulberry’s breeding success, which is consistent with its ecology as a small forest understory tree where individuals are often scattered widely on the landscape. Hermaphrodism was more common in trees with a greater distance to their nearest neighbor. It is not known whether all individuals are capable of hermaphrodism, or if they consistently display this trait upon isolation. If hermaphrodism is limited, pollen limitation may be an increasingly serious problem as the species becomes rarer, with isolated male or female individuals unable to produce fruit.

Status Justification We recommend Pennsylvania Threatened as the appropriate status for the red mulberry, due to the very small number of extant populations and individuals known, in combination with ongoing threats from disease and/or environmental stressors, as well as hybridization with the white mulberry, that have led to a dramatic decline in this species. Although dedicated fieldwork has been conducted for a relatively short period of time compared to some species, the threats to this species and the trend of decline are supported by scientific literature, expert botanical experience, the results of dedicated

77 fieldwork, and trends in herbarium documentation. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Affeltranger, C. 1965. Is Red Mulberry Vanishing? W. Va. University Arboretum Newsletter 15:2 p. • Ambrose, J. D. and the Red Mulberry Recovery Team. 1998. National Recovery Plan for Red Mulberry (Morus rubra L.) (Revised 2001). 28 pp. • Burgess, K.S. and B. C. Husband. 2006. Habitat differentiation and the ecological costs of hybridization: the effects of introduced mulberry (Morus alba) on a native congener (M. rubra). Journal of Ecology. 94: 1061-1069. • Burgess K. S., M. Morgan, L. DeVerno, and B. C. Husband. 2005. Asymmetrical introgression between two Morus species (M. alba, M. rubra) that differ in abundance. Molecular Ecology 14: 3471-3483. • Core, Earl L. 1974. Red mulberry Morus rubra L. In Shrubs and vines for northeastern wildlife. p. 106-107. John D. Gill and William M. Healy, comp. USDA Forest Service, General Technical Report NE-9. Northeastern Forest Experiment Station, Upper Darby, PA. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Lamson, Neil L. 1990. Morus rubra L. In: Burns, Russell M., and Barbara H. Honkala, tech. coords. 1990. Silvics of North America: 1. Conifers; 2. Hardwoods. Agriculture Handbook 654. U.S. Department of Agriculture, Forest Service, Washington, DC. vol.2, 877 p. • Little, Charles E. 1997. The Dying of the Trees. Penguin Books. New York, New York. • McLaughlin, J. and S. Greifenhagen. 2002. A Pathological Survey and Disease Management Analysis for Red Mulberry in Southern Ontario. Ontario Ministry of Natural Resources, Ontario Forest Research Institute. Sault Ste. Marie, Ontario. • MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. October 6, 2016. http://www.michiganflora.net/species.aspx?id=1703. • Nepal, Madhav P.; Ferguson, Carolyn J.; Mayfield, Mark H. 2015. Breeding system and sex ratio variation in mulberries (Morus, Moraceae). Journal of the Botanical Research Institute of Texas . 2015, Vol. 9 Issue 2, p383-395. 13p. • Ontario Ministry of Natural Resources. 2013. Recovery Strategy for the Red Mulberry (Morus rubra) in Ontario. Ontario Recovery Strategy Series. Ontario Ministry of Natural Resources, Peterborough, Ontario. iii + 5 pp + Appendix vi + 47 pp. Adoption of the Recovery Strategy for the Red Mulberry (Morus rubra) in Canada (Parks Canada Agency 2011). • Pennsylvania Flora Project, Morris Arboretum, University of Pennsylvania. www.paflora.org. • Rhoads, Ann F. and Timothy A. Block. 2007. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia; 2nd edition. 1056 pg.

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• Spisani, S. R., J. L. Sylvester, and C. J. Rothfels, 2004. Inventory and health assessment of the nationally endangered Morus rubra (red mulberry) on Royal Botanical Gardens’ properties and adjacent areas. Unpublished report for the Ontario Ministry of Natural Resources, Guelph District. Guelph, Ontario. 62 pp. • Spurway, C.H. 1941. Soil reaction (pH) preferences of plants. Mich. State Coll. Agr. Exp. Sta. Spec. Bull. 306. 36 p. • Weakley, Alan S., J Christopher Ludwig, John F Townsend, Bland Crowder. 2012. Flora of Virginia. Fort Worth, Texas. Botanical Research Institute of Texas Press. • Weldy, Troy, David Werier, and Andrew Nelson. 2016 New York Flora Atlas. [S. M. Landry and K. N. Campbell (original application development), USF Water Institute. University of South Florida]. New York Flora Association, Albany, New York.

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Sourwood – Oxydendrum arboretum (L.) DC. Author: Loree Speedy, Carnegie Museum of Natural History. Edited: Jessica McPherson, PNHP

Introduction Sourwood is a small native tree found within the forest and at its edges; sites that are sunny, open, well drained and with few competitors are best. It appears to require acid soil, and generally grows with oaks and pines on valley slopes and in uplands. Sourwood trees can be easily identified in flower and in fruit by the unique bell-shaped white flowers or fruits held on one side of a long drooping spray. Its leaves turn a bright crimson in the fall, and winter twigs have a unique zig-zag appearance.

Proposed PA Status We concur with the PABS-Vascular Plant Technical Committee’s 2016 recommendation to delist sourwood due to the moderate number of populations known, the large number of individuals documented in some populations, the apparent trend of increase within Pennsylvania, and the observed ability to thrive in disturbed conditions that are regionally abundant.

North American Distribution and Global / Regional Conservation Concerns Sourwood is found from southwest Pennsylvania to southern Indiana, south to southeastern Louisiana and the coastal region of Mississippi, Alabama, and northwest Florida (Kartesz 2003).

Pennsylvania Distribution and Historical Locations Southwestern Pennsylvania is the northern edge of its range. Historically in Pennsylvania, sourwood was known in limited numbers in Greene and Fayette counties, with a known extirpated site in Westmoreland County (Jennings and Avinoff, 1953) that has not since been relocated. More sites, however, in Greene and Fayette counties have been successfully located recently.

Extant Locations Thirty one extant occurrences are known in Pennsylvania; at least 25 have been documented since 2000. Six of these sites have over 1000 individuals, ten have over 100 individuals. The documented sites total an estimated 25,000 to 30,000 individuals. Many documented occurrences, however, have not been fully delineated and appear likely to extend across property lines. The constraints of time, private property access, and defined boundaries of an environmental survey project have limited the ability to fully evaluate the extent of these larger populations. Three large populations covering at least 400 acres on the west-facing flank of Chestnut Ridge may be connected. An additional outlying occurrence has recently been documented in Perry County.

Habitat Documentation of sourwood habitat in Pennsylvania shows that it appears in a variety of acid habitats and in a variety of successional stages, such as the following: • Ridges and slopes with mature forests – typically oak-dominated, dry-mesic • Successional forests that have experienced logging in the last few decades, including clear-cut areas • Young forests regenerating from pasture, or surface mining, or other clearing • Woodland edge habitats created by roads, powerline rights-of-way, mining, etc. • Open, well-drained old fields, abandoned pastures, and nutrient-poor soil of unreclaimed strip mines

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Threats Over the past 50 years, the Pennsylvania portion of the range of sourwood appears to have experienced a notable increase in population size, due to the disturbance regimes in acidic oak forests and fallow land in Fayette and Greene counties. Sourwood can benefit from disturbances that cause nutrient-poor soils and high light availability. Disturbance activities potentially benefiting sourwood include the following: • An increase in strip-mining activity and in strip-mined areas reverting to woodlands, especially those affected prior to the Surface Mining Control and Reclamation Act (SMCRA) of 1977 which mandated topsoil reclamation with fescues and legumes • An increase in edge habitat due to commercial, agricultural and mineral (coal, oil and gas) development • An increase in the regeneration of fallow agricultural lands to forest • Prevalence of working forests managed for timber

Sourwood appears to be thriving in a wide range of natural and disturbed conditions and appears to be expanding its range in the two counties due to these changes in land use over the past 50 years.

Status Justification While the number of documented populations is consistent with the current Pennsylvania Rare status, the large number of individuals documented in some populations, the apparent trend of increase within Pennsylvania, and the observed ability to thrive in disturbed conditions that are regionally abundant all indicate that this species does not require protection to continue to thrive in Pennsylvania. We recommend delisting of sourwood at this time. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S4 (Apparently Secure) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Burns, R. M., & Honkala, B. H. 1990. Silvics of North America (Vol. 2, p. 119). United States Department of Agriculture. • Carnegie Institute, Avinoff, A., & Jennings, O. E. 1953. Wild Flowers of Western Pennsylvania and the Upper Ohio Basin. University of Pittsburgh Press. • Grimm, W. C. (1983). The illustrated book of trees: with keys for summer and winter identification. • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • NatureServe. March 2014. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. (Accessed 12 January 2015). • Pennsylvania Natural Heritage Program. 2007.

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Stiff Cowbane – Oxypolis rigidior (L.) Raf. Author: Loree Speedy, Carnegie Museum of Natural History. Edited: Jessica McPherson, PNHP

Introduction Stiff cowbane is a wetland species in the Carrot Family with the characteristic small white flowers in a flowering cluster, or umbel, whose main branches arise from a single point and radiate outward. The blooming period occurs from late summer to early fall. Leaves are divided into five to nine narrow leaflets with a few scattered teeth.

Proposed PA Status We concur with the PABS-Vascular Plant Technical Committee’s 2016 recommendation to assign the Pennsylvania Rare status to stiff cowbane, because while there are a moderate number of populations known in the state, there are relatively few with good ecological and reproductive viability, and ongoing threats to the species from wetland habitat degradation and invasive species.

North American Distribution and Global / Regional Conservation Concerns Stiff cowbane can be found in North America from New York south to Virginia and west to Minnesota, Iowa, Missouri, Oklahoma and Texas (Kartesz 2003).

Pennsylvania Distribution Pennsylvania is at the northern edge of the range of stiff cowbane; it extends northward in two separate prongs to southeastern and southwestern Pennsylvania.

Habitat The habitat of stiff cowbane is wet to moist swamps, bogs, edge meadows, sandy shores, and abandoned railroad beds. In Pennsylvania it occurs in shaded to partially shaded wetland openings and in western Pennsylvania, it also frequents sandy cobble shores of streams. These habitats lack significant disturbance such as heavy infestation of invasive species, sedimentation, and soil disturbance. There is often a high diversity of native shrubs and wetland plants. In western PA, the plant is not found in degraded wetland habitats. In eastern Pennsylvania, however, most known populations are in fairly degraded wetland habitats such as unmanaged portions of cow pastures, etc. This may reflect remnant populations that have persisted despite habitat degradation.

Extant Locations Although there are currently 66 known extant populations in Pennsylvania, more than half of the populations have less than 25 individuals. Only thirteen of the extant populations have more than 50 individuals.

Historical Locations and Potential for Relocation Some historical populations have been relocated for this species, although most known extant locations are the result of de novo survey work. All known historical locations have been reviewed to determine if there is potential to relocate the population. The eastern historical specimens are considered to have very low potential for relocation because of the degree of landscape change that has occurred in that region. The western historical locations include a mixture of specimens that are too old or vague to relocate, and some that should be surveyed for; most of these surveys are complete, although about six records remain that could still be searched for.

Threats

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The long-term viability of many eastern Pennsylvania populations is questionable, with small populations occurring in fragmented and degraded habitat. Residential and urban development is also ongoing in this region and could result in further habitat loss and degradation. Temporary wetland impacts allowed in pipeline developments may also degrade the current wetland and shoreline habitat in western Pennsylvania. Deer browsing has been observed in scattered sites.

Status Justification Because of the relatively few occurrences with good ecological and reproductive viability, and the species’ questionable ability to persist long-term in degraded wetland habitat, we recommend that stiff cowbane be included on the Chapter 45 list of Rare plant species. This status is supported by the recommendation of the Vascular Plant Technical Committee (2015) and the results of the NatureServe Rank Calculator Version 3.185, which calculated a status of S3 (Vulnerable) in Pennsylvania. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned the S3 status to stiff cowbane in Pennsylvania.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Hilty, John. 2014. Wetland Wildflowers of Illinois: Cowbane (Oxypolis rigidior). Available at: http://www.illinoiswildflowers.info/wetland/plants/cowbane.htm. (Accessed on 12 January 2015). • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • NatureServe. March 2014. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia.. (Accessed 12 January 2015). • Pennsylvania Natural Heritage Program. 2007. • Rhoads, A.F. and T.A. Block. 2000. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia.

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Yellow-fringed Orchid — Platanthera ciliaris (L.) Lindl. Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP.

Introduction Yellow-fringed orchid is a perennial herb that may grow to three feet in height. It is characterized by a hairless stem with several entire and alternate sheathing leaves that decrease in size going upward. The species, which blooms from late July into August, has a terminal cluster of orange flowers with each individual flower having the lowest petal (“lip”) narrowly and delicately fringed on the margin. Without doubt, this species is one of the most spectacular wildflowers in Pennsylvania.

Proposed PA Status We concur with the recommendation of the PABS-Vascular Plant Technical Committee to change the status of yellow-fringed orchid from Tentatively Undetermined to Pennsylvania Threatened (PT). The main reasons for this status include the relatively small population sizes; the threats of excessive deer browse, poaching by gardeners, and non-selective means of controlling vegetation on utility rights-of- way (a frequent habitat for the species); and the generally unpredictable appearance and viability of orchid populations.

North American Distribution and Global / Regional Conservation Concerns Yellow-fringed orchid has a global rank of Secure (G5). This rank is probably in effect because the species is still relatively common in the southern parts of its global range; however, it is generally of conservation concern in the northerly portions.

The global distribution extends from Ontario east into Massachusetts, and south into Texas and Florida.

Yellow-fringed orchid has been documented in all six states that are contiguous to Pennsylvania, and all of the states consider the species to be of conservation concern: Delaware (Historical SH), Maryland (Imperiled S2), New Jersey (Imperiled S2), New York (Critically Imperiled S1), Ohio (Imperiled S2), and West Virginia (Vulnerable S3).

Pennsylvania Distribution Yellow-fringed orchid has a NatureServe state rank of Vulnerable (S2).

The historical distribution of the species occurs throughout the southeastern and southcentral counties, becoming much less frequent or absent in the northern and far western counties. The relatively large number of locations in the southeastern counties is probably due in large part to over-collection by botanists.

Habitat Based on the extant occurrences known to PNHP and the historical records that list habitat data, yellow- fringed orchid grows primarily in wetlands, particularly in places where sphagnum moss is present, but there are also extant locations where the species occurs on well-drained substrates. Similarly, most of the occurrences are in open or partly-open habitats, but there are exceptions where the species grows in a more shaded situation. Perhaps the most interesting item about the habitat of the species involves its ability to colonize and apparently thrive in some human-created/maintained environments, such as at certain utility rights-of-way and road banks and ditches (as long as the substrate is suitable). For example, one of the larger extant occurrences of this orchid is situated on a powerline right-of-way that has a sandy substrate, very early-successional vegetation (thanks to unwittingly-appropriate

84 maintenance activity by the utility company), and several small spring-fed seepages. All of these factors, and perhaps others, provide the right mix for this species to become established. Of course, yellow- fringed orchid also grows at undisturbed or little-disturbed habitats as well.

The habitats provided on Pennsylvania herbarium specimens from the past 150 years include “boggy thicket; boggy meadow; bog; peat bog; boggy ground along railroad; swamp; around spring; marsh; meadow; along mountain road; swampy woods; swampy meadow; moist grassland; dry serpentine; wet meadow; sandy slope; damp oak barren; moist woods; moist woods road; along streamlet; field; open swamp.”

Extant locations Numbers At present, yellow-fringed orchid has been documented by PNHP at 32 extant locations in 14 counties. Approximately one-third of these occurrences are clustered in Adams and Cumberland counties in the South Mountain region. Nearly two-thirds of the 32 locations are situated on public land (state forest 13; state parks 2; state game lands 5) or municipal, usually watershed, properties (4).

For this species more than most, the number of extant occurrences is subject to considerable revision. It should be noted that approximately 40% of the 32 occurrences have not been revisited in more than ten years, so the documented number may be lower than reported; on the other hand, a network of orchid enthusiasts exists in the state, and this group sometimes is hesitant to share locations with PNHP, so this could be a reason why the number of populations might be larger than that known to PNHP.

Population sizes The population sizes at the 32 extant locations vary from one plant to ca 130 plants, although nearly one-half of the occurrences have ten plants or fewer and four occurrences have a single plant. Even at sites where the population is considered to be established, the number of plants that make an appearance in flower from year to year can vary, and plants may only be vegetative in a given year or more.

Potential for additional EOs There is potential for additional de novo populations, given the unpredictability of orchids, the abundance of public land, and the ability of the species to colonize certain types of human-related disturbance. An estimate of the total number of these potential populations is impossible.

Historical locations Numbers An estimated 84 historical occurrences in 28 counties, based on herbarium specimens, are known from Pennsylvania, particularly in the eastern counties. This number is difficult to estimate because only a few Academy of Natural Sciences records have been processed into Biotics and the procedure of delineating occurrences when mapping. Approximately 30% of these records were collected prior to 1900, approximately 80% were collected prior to 1925, and the most recent collection was from 1952. As is usually the case with herbarium specimens, the size of the populations at the time of collection is unknowable and many of the locations are rather vague.

Potential for extant EOs Given the age of these collections and the part of the state involved, it seems likely that there has been habitat degradation, or even destruction, at many of these locations. This negative factor is mitigated

85 by the ability of the species to colonize certain types of disturbances. Although there do appear to be a few instances of a historical location, sometimes from as long as 70 years ago, still having a population of the species today, these cases are probably rare exceptions. Most of the historical occurrences of yellow-fringed orchids will probably not be found to have current populations, given the age of the collections, the vague locational data, and the portion of the state where most of them are located.

Threats Threats to extant occurrences include deer browse, poaching by gardeners, exotic species, and improper vegetation control methods (non-selective mowing or herbicide spraying) in cases where the species is occupying a human-created habitat. It is unclear how persistent these later occurrences are, even if the management is generally benign.

Status justification We recommend a status of Pennsylvania Threatened for yellow-fringed orchid. Although the number of extant populations is somewhat higher than typical for this status, many of the populations are extremely small, with uncertain viability. Furthermore, the species has numerous threats, such as excessive deer browse, poaching, improper management when growing in human-maintained habitats, and invasive species, although balanced to some degree against those concerns is the capacity of the species to utilize certain types of human-created-and-maintained disturbed areas, which increases the possibility (although not necessarily the viability) of new populations. In view of the numbers of populations and individuals known and the level of threat faced by the species, a status of Pennsylvania Threatened is appropriate at this time. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • NatureServe. 2015. NatureServe Conservation Status Assessments: Rank Calculator Version 3.185. NatureServe, Arlington, VA. Online at http://www.natureserve.org/conservation- tools/conservation-rank-calculator • Pennsylvania Natural Heritage Program. Biotics Database. Accessed March 2017. • Rhoads, A.F. and T.A. Block. 2007. Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press. Philadelphia.

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Bog Bluegrass – Poa paludigena Fern & Wieg. Author: Jim Bissell, Cleveland Museum of Natural History; Jessica McPherson, PNHP; John Kunsman, PNHP

Introduction Bog bluegrass is a perennial herbaceous member of the grass family (). The grass is easily overlooked due its short stature, narrow leaves, and tendency to lean over when in flower and fruit. Peak bloom is in mid-June in western PA, although substantially earlier in eastern PA, where it may be sufficiently past bloom as to be difficult to identify at that time. Bog bluegrass can flower as early as mid-May. A few plants in most populations have intact fruiting spikelets as late as mid-August.

Proposed PA Status The status currently recommended for this species by the PABS-Vascular Plant Technical Committee is Pennsylvania Rare, although it was last considered in 1999. The information currently available about bog bluegrass is roughly in line with Pennsylvania Rare, with some indications towards Pennsylvania Threatened as well; however, significant information gaps still exist that could affect the appropriate status designation.

North American Distribution and Global / Regional Conservation Concerns Bog bluegrass ranges from Minnesota and Iowa, east to New York, Pennsylvania, West Virginia, and North Carolina. No populations are currently known within Canada. It is listed by NatureServe as Globally Vulnerable (G3). It is listed of conservation concern in every state where it occurs: Critically Imperiled (S1) in New York, West Virginia, and North Carolina; Imperiled (S2) in Virginia, Michigan, Minnesota, and Iowa; Vulnerable in Wisconsin, Indiana, Ohio, and Pennsylvania; and Extirpated (SX) in Illinois. In some states, only small populations (10-50 individuals) have been documented. This species is likely underdocumented to some degree because it is part of an obscure and difficult genus, the plants are small and inconspicuous, and the survey window is narrow. Wisconsin, for example, removed bog bluegrass from its list of threatened and endangered species in 2014 (switching to the current status of “Vulnerable”) because inventory work had revealed new populations.

In the late 1980s, bog bluegrass was on track to be listed under the federal Endangered Species Act by the U.S. Fish and Wildlife Service. The discoveries of new populations within northwestern Pennsylvania, northeastern Ohio, Indiana, and Minnesota between 1987 and 1995 convinced the U.S. Fish and Wildlife Service to withdraw the proposal for federal listing. However, since the end of USFWS’s consideration of the species, little current information has been published on population sizes or survey effort across the range, making it difficult to further assess the degree to which the species is truly globally rare or whether its apparent rarity is an artifact of underdocumentation.

Pennsylvania Distribution Bog bluegrass has a somewhat bi-modal distribution in Pennsylvania, with populations concentrated in the glaciated northwest and in eastern Pennsylvania. A few scattered populations are also known from south-central Pennsylvania. The distribution somewhat resembles a northern species, as it occupies the cooler parts of Pennsylvania and further south is only known from the mountains, but this characterization is at odds with the lack of populations in Canada or further north than New York or Michigan.

Habitat

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Bog bluegrass appears to occupy different kinds of habitat in eastern versus western Pennsylvania, although this may also reflect the occurrence of appropriate microhabitats in different landforms regionally. In northwestern PA, habitat for bog bluegrass is on the perimeters of high pH fens. In 1990, pH readings were taken for several bog bluegrass populations in northwestern Pennsylvania and northeastern Ohio. All of them measured 6.4 pH. Bog bluegrass is not clonal. Sometimes a single three- foot-diameter hummock can support as many as 50 to 100 individual plants. Bog bluegrass is often found growing in wetland hummocks covered with fern moss (Thuidium delicatulum).

In eastern Pennsylvania, populations of bog bluegrass do occur in swamps and wet woods, but most eastern populations have a much different habitat, which is in low-volume, spring-fed streamlets and seepages in wooded situations, frequently on diabase. The overall habitat could hardly be called a swamp, and the adjacent woods may actually be quite well drained just a few feet from the streamlet or seepage. The bluegrass plants tend to be rooted, individually or in small clumps, on bryophyte-covered rocks or in crevices of rocks located in the streamlet/seepage bed itself (just above the normal waterline so that the bryophytes and the roots of the bluegrass remain damp) or along the margins of the streamlet or seepage. Occasional bluegrass plants may be found farther downstream where the volume of water is much higher, but the species appears to do best in the low-volume headwaters situations, so protecting the spring source and its water quality and chemistry are particularly important. Another ecological difference between the eastern and western populations is that shading doesn’t appear to be a great threat for the eastern ones.

The Laurel Mountain population has a similar habitat to that described for eastern Pennsylvania.

Extant Locations There are 32 extant populations of bog bluegrass in Pennsylvania; twelve (12) populations are in northwestern Pennsylvania, 17 populations are in eastern Pennsylvania, and 3 populations are in south- central Pennsylvania. Population sizes are fairly well documented for most extant locations, especially with recent fieldwork. Although there are more bog bluegrass populations in eastern Pennsylvania than in the west, the largest populations known in the state are in western Pennsylvania. During the 2014 field season, Cleveland Museum of Natural History staff members carefully mapped and counted bog bluegrass populations within PA State Game Land 214 and PA State Game Land 122 and private lands adjacent to PA State Game Land 122 in Steuben Township, Crawford County. There are more than 1,500 plants within PA State Game Land 214 and more than 3,000 plants within Steuben Township, Crawford County, PA. The population in Westmoreland County has 3,000-6,000 plants. To our knowledge (not all states publish information on population sizes) these are some of the largest populations of bog bluegrass known in the world.

The total number of individuals estimated to be at the known extant occurrences is 4,800-14,500. This is a very wide ranging estimate, in part because many have been visited multiple times with substantially different counts at different visits. In the east, most revisits have resulted in a lower count than the previous visit and the populations are often small and localized enough to provide a fair degree of confidence in the count per visit. This may reflect decline for these populations, in part due to the effects of out-competition by invasives at some sites. There are 8 populations with 30 plants or fewer, 4 populations with 75-150 plants, and 4 with 200-350 plants. In the west (including the south-central populations), the numbers documented at the several large populations mentioned previously are greater than those recorded at initial surveys, although this most likely reflects more careful counting on the later visits. In addition to the four large populations documented in the west, there are 5 populations with 50 plants or fewer, 4 populations with 50-500 plants, and 2 populations where counts

88 were not recorded. The fairly large population documented in a headwaters rocky seep in Westmoreland County in 2017 (this location previously was documented only from a 1995 specimen and had no population data recorded) suggests that similar habitats should be further explored in this region of the state.

Historical Locations and Potential for Relocation All but a few historical locations have been investigated; a few have been relocated, several were determined to have failed or been extirpated.

Threats Invasive species threaten to outcompete the diminutive bog bluegrass throughout Pennsylvania, although the most problematic species differ from east to west. In the west, large populations of narrow-leaf cattail, non-native Phragmites and reed canarygrass threaten bog bluegrass. The spread of these invasive species within the partially shaded hummocks that support bog bluegrass is a major threat to the populations. In the east the main threat is Poa trivialis, which can aggressively dominate and clog up the streamlets, seepages, and springheads. Japanese stiltgrass (Microstegium vimineum) can also dominate this type of habitat; although it has not yet been observed at many of the known sites, it is a fast-moving species that is a potential future threat.

Shading through succession has been observed to cause decline within bog bluegrass populations in the western part of the state. One population found within PA State Game Land 214 in 1987 has declined from more than 200 plants to less than 50 plants in 2014. The site was partially shaded in 1989. In 2014, the canopy was fully shaded. Observers of eastern populations report that shading does not appear to be a major threat at these sites.

The impending die-back of black ash from northwestern Pennsylvania may also have a negative effect on bog bluegrass populations within the commonwealth. Many bog bluegass populations within northwestern Pennsylvania seepages and swamps are confined to prostrate, rotting black ash logs. Several populations of bog bluegrass in Ohio are also restricted to black ash logs.

Status Justification The information currently available about bog bluegrass is roughly in line with Pennsylvania Rare, with some indications towards Pennsylvania Threatened as well; however, significant information gaps still exist that could affect the appropriate status designation. Pennsylvania Rare is indicated by the number of populations that are known (32) and the fact that this species is almost certainly underdocumented to some degree. However, the number of populations is towards the low end of the range usually considered for PR, and the combination of small population sizes at about half the sites and pervasive threats from invasive species and habitat alteration casts doubt on the long term viability of many populations, suggesting that perhaps Pennsylvania Threatened might be more appropriate. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a range rank of S2S4 (Imperiled to Apparently Secure) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

Because the species may be globally rare, it is also important to consider the degree to which Pennsylvania populations affect its global conservation status. Because large populations are known here and have not been reported from other states in the range, it may be a species for which Pennsylvania bears a large share of the responsibility for its overall success. However, there remains

89 uncertainty about whether the apparent rarity in other states is real or an artifact of underdocumentation, and little information is available on its status in the rest of the global range.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • NatureServe. March 2016. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia.. (Accessed 31 October 2016).

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Racemed Milkwort – Polygala polygama Walter Author: Jim Bissell, Cleveland Museum of Natural History. Edited: Jessica McPherson, PNHP

Introduction Racemed milkwort is an herbaceous biennial in the milkwort family (Polygalaceae). Racemed milkwort plants are several-stemmed and decumbent, with alternate, entire leaves which are narrowly oblanceolate. Elongate, arching racemes with bright, pink flowers bloom during the middle of June.

Proposed PA Status We recommend the assignment of Pennsylvania Endangered status to racemed milkwort, due to the low number of extant populations known in the state, the low number of plants in most known populations, and the apparent decline of the species due to loss of appropriate early successional habitat.

North American Distribution and Global / Regional Conservation Concerns Racemed milkwort has a somewhat unusual global distribution. Although it ranges from Ontario and Quebec south through the eastern half of North America to Texas and Florida, it is concentrated in northern and southern portions of the range, and scarce in the middle portion, mainly following the Appalachian Mountains with scattered outlier populations across Ohio, Indiana, and Illinois (Kartesz 2003).

Habitat This species requires open, sandy habitats, and its distribution likely in part reflects the regional abundance of this type of habitat. In Pennsylvania, it is largely confined to open sand barrens, rock barrens, and shale barrens.

Pennsylvania Distribution In Pennsylvania, most records known for the species are from the south central section of the state, concentrated in Bedford and Fulton counties and scattered northwards in the Ridge and Valley. A smaller cluster of records is known from the northwestern portion of the state, and there is one record known from Berks and Monroe counties.

Extant Locations, Historical Locations and Potential for Relocation There are 10 extant records and 17 historical records. Most extant records have very small numbers of plants recorded. Of the historical records, about 1/3 have been searched for without success, while 3 extant populations were located by searching for historical specimens. Because this species requires early successional habitat, historical records are unlikely to be relocated after substantial time has passed, although they may indicate general areas to search for present-day suitable habitat.

The Cleveland Museum of Natural History has found only two populations of racemed milkwort in northwestern Pennsylvania. In 2009, museum staff and volunteers spent several days searching for a historical record collected in eastern Erie County near Cory, PA and no open sand barren habitat suitable for the rare plant could be found. As a few new populations have been discovered in Bedford County in recent years, and habitat that appears suitable is scattered in that region, some additional locations may continue to be found through de novo survey work.

Threats Survey work and repeated observations of this species’ populations in northeastern PA and Ohio by the Cleveland Museum of Natural History have documented the vulnerability of this species to successional

91 habitat change, but also positive response to active management. The Cleveland Museum of Natural History owns the only extant population of racemed milkwort in Northeast Ohio. The only other extant populations in Ohio are within the Oak Openings region west of Toledo, Ohio. The population at the Museum’s North Kingsville Sand Barrens Preserve, nine miles west of the Springfield Barrens, has crashed several times during the last 25 years, from hundreds to a few dozen plants when succession was permitted to choke out the rare plant. Mowing or mechanical disturbance of the sands at the North Kingsville Sand Barrens has increased populations from a few dozen to several hundred during the year following the disturbance.

None of the known populations in central Pennsylvania are currently receiving active management, and thus all may be vulnerable to decline due to succession. The extremely small size of the populations raises questions about whether it can successfully disperse and colonize other early successional habitats. Additional threats are invasive species and, for roadside occurrences, improper road maintenance procedures.

Status Justification The small total number of individuals known in the state, the trend of decline in this species’ populations without active management, and the existence of additional threats (invasive species and roadside maintenance) indicate a status of Pennsylvania Endangered. If, in the future, additional populations are discovered through more survey work, and/or if some populations are increased through protection and active management, it may be appropriate to consider changing the status to Pennsylvania Threatened. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Lion’s Foot – Prenanthes serpentaria Pursh Author: Jessica McPherson, PNHP

Introduction Lion’s foot is a member of the genus Prenanthes; several members of this genus occur in Pennsylvania, including some that are relatively common. They typically have a monocarpic life history, in which many juvenile individuals with only basal leaves develop each season, but only a few mature to produce flower stalks and seeds, after which the plant dies. Several species, including lion’s foot, cannot be reliably identified by their juvenile leaves, which can be quite variable in shape and also somewhat similar between species.

Proposed PA Status Based on currently available data, we recommend a status for this species of Pennsylvania Threatened, in concurrence with the PABS-Vascular Plant Technical Committee recommendation. However, recent surveys have increased the number of known populations, so we recommend delay in formalizing the status of this species in combination with ongoing survey efforts, as enough new populations may eventually be found to redesignate it as Pennsylvania Rare.

North American Distribution and Global / Regional Conservation Concerns This species is known from a fairly broad geographic range in eastern North America. Although most of the range is south of Pennsylvania, the northern extent is in New England, so Pennsylvania populations are not at the edge of the range. The range does appear to constrict towards the coast in its northern portion, starting in Pennsylvania, where it is not known from the western part of the state (Kartesz 2003).

Pennsylvania Distribution Lion’s foot is found mainly in southeastern Pennsylvania; most extant occurrences are known from Chester County, with a scattering of others in Adams, York, Lancaster, and Berks counties. Scattered historical specimens are known from as far north as Monroe County and as far west as Bedford County.

Habitat Flora of North America describes the habitat as “Oak-hickory woodlands, borders, oak flats, pine woods, sandy areas.” In Pennsylvania, the habitat of documented sites tends towards wooded roadbanks, successional but not weedy, often on serpentine soils.

Extant and Historical Locations There are 17 extant occurrences currently documented in Pennsylvania, and 35 historical occurrences. Historical locations are concentrated in Bucks, Montgomery, Chester, and Delaware counties; due to imprecise location information, a high degree of landscape change in this region, and some preference for early successional habitat which often makes populations transient on the landscape, most of these are unlikely to be relocated.

Status Justification The current status proposed by the VPTC is Pennsylvania Threatened, which is consistent with the number of populations and individuals known in the state. However, recent survey work, especially in Chester County, has expanded the number of known occurrences, and enough may eventually be found to merit redesignation as Pennsylvania Rare.

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Despite the potential for more locations to be found, we do not believe that this species is a high priority for targeted fieldwork, because a variety of factors combine to make targeted survey work inefficient. Early successional habitat is the most likely terrain for finding additional populations, and this type of habitat is difficult to effectively target for successful surveys. While there may be additional populations, there is so much potential habitat that the likelihood of finding the species on any given survey is low. The likelihood of survey success is further limited by the life cycle of the plant; the survey window is narrow because the plant cannot be reliably distinguished by its basal leaves, and most individuals do not flower in a given year. Surveys targeted for this habitat type are also inefficient because it is unlikely to host other elements of conservation concern. It is more efficient to gather knowledge by educating botanical surveyors who work in the appropriate region to look for this species while doing other work. We recommend reviewing available information for this species again in several years, and revisiting the status of this species if substantial new information has been added.

References • Bogler, David J. 2006. Prenanthes. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 19, 20, 21. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Swamp chestnut oak – Quercus michauxii Nuttall Author: Dr. Timothy Block, Morris Arboretum; Jessica McPherson, PNHP

Introduction Swamp chestnut oak is a large tree of the white oak group, very similar in overall appearance to swamp white oak (Q. bicolor). The only completely reliable characters for distinguishing the two species are those of the winter buds. In swamp chestnut oak, the buds are reddish-brown, ovoid, rounded to acute at the apex, and generally somewhat pubescent. In swamp white oak, the buds are light to dark brown (not reddish), globose to ovoid, rounded at the apex, and completely glabrous. The abaxial surfaces of mature leaves of swamp chestnut oak possess both stellate and long unbranched hairs while those of swamp white oak have only stellate pubescence. This character is reliable only if leaves from the upper portions of the trees can be reached as shade leaves of both species have only stellate pubescence. The habitat of these two species is similar, and in areas where the ranges overlap, they are occasionally mistaken for one another.

Proposed PA Status We concur with the PABS-Vascular Plant Technical Committee’s recommendation of a status of Pennsylvania Endangered for swamp chestnut oak, due to the very low number of populations and individuals known to the state, and the extremely limited habitat remaining in the state.

Habitat The habitat for swamp chestnut oak is described by Flora of North America as “bottomlands, rich sandy woods and swamps, on variety of soils.”

North American Distribution and Global / Regional Conservation Concerns The global range of swamp chestnut oak is concentrated on the Atlantic Coastal Plain in south-eastern North America, from eastern Texas to Florida and north to south-eastern Pennsylvania and New Jersey. It also extends up the Mississippi River Valley into southern Illinois and Indiana (Kartesz 2003).

Pennsylvania, which is both at the northern extent of the species’ range and contains very little coastal plain habitat, is the only state in which NatureServe lists swamp chestnut oak as Critically Imperiled (S1). It is listed as vulnerable (S3) in New Jersey (also at the northern edge of the range but with more coastal plain habitat) and Illinois; as Apparently Secure (S4) in Oklahoma; and as Secure (S5) in all other states where a conservation status has been assigned. The general distribution of this species is the southeastern U.S. with historical, but not extant, reports from New York, Connecticut, Massachusetts, and Oklahoma.

Pennsylvania Distribution In Pennsylvania, the species is known only from the Coastal Plain.

Extant Locations The presence of swamp chestnut oak in Pennsylvania was first detected by John Kunsman while working at Delhaas Woods at the Silver Lake Nature Center in Bucks County in 2008. The following year several mature and immature trees were noted along the northeast side of the power line right-of-way at Delhaas Woods. In 2010 another small population was found on Money Island in the Delaware River. One specimen collected in 2007 by Rhoads and Block at Boothwyn in Delaware County, housed at the Morris Arboretum herbarium (MOAR) and initially identified as Q. bicolor, has been redetermined to Q.

95 michauxii by this author. This brings the total known extant sites in Pennsylvania for Q. michauxii to three.

Historical Locations and Potential for Relocation In addition, several herbarium specimens at the Academy of Natural Sciences of Drexel University (PH) collected as Q. bicolor have been examined and are probably Q. michauxii. These specimens need further examination, and if redetermined, will require further field exploration to determine if these occurrences still exist.

Status Justification Swamp chestnut oak is proposed to move from no status (N) to Pennsylvania Endangered (PE) based on current best knowledge of its distribution and population status in the state. With only three known populations the species is extremely rare in the state, and even if several of the specimens at PH prove to be this species, the number of individuals is likely to remain consistent with Pennsylvania Endangered status. The major threat to swamp chestnut oak in Pennsylvania is potential habitat loss. Of the known occurrences, only the Delhaas Woods site is on protected land. Additional field work may turn up other populations in southeast Pennsylvania, but these will always be small occurrences, because the patches of remaining coastal plain habitat are small. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. • Nixon, Kevin C. 1997. Quercus. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 3.

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Water-plantain Spearwort – Ranunculus ambigens S. Wats. Author: Dr. Scott Schuette, John Kunsman PNHP. Edited: Jessica McPherson, PNHP.

Introduction Water-plantain spearwort is an herbaceous perennial characterized by its lance-shaped, marginally- toothed leaves with elongate leaf stalks, solitary flowers with five yellow petals, tiny seed-like fruits, and wetland habitat. The shape, and often the length, of its leaves are distinctive and look quite unlike most of its buttercup relatives in which the leaves are divided into leaflets or lobed/incised segments.

Proposed PA Status In 2017 the PABS-Vascular Plant Technical Committee changed the recommended status from Tentatively Undetermined to Pennsylvania Threatened. Currently available information indicates a status of Pennsylvania Endangered, based on a very small number of known populations and individuals in Pennsylvania combined with an apparent range-wide decline, and ongoing threat from deer browse and invasive species. The VPTC-recommended status of Pennsylvania Threatened is also acceptable to accommodate some uncertainty in the number of undocumented populations that may exist.

North American Distribution and Global / Regional Conservation Concerns On a global basis, the species is listed as Apparently Secure (G4) by NatureServe; however, it appears to be uncommon throughout most of its range. It is distributed in the eastern third of the United States, from Maine south to Tennessee (Kartesz 2003). In the Mid-Atlantic portion of its range, which includes Pennsylvania and adjacent states, it is considered to be Threatened (New Jersey), Endangered (Delaware, Ohio), Rare (Pennsylvania), Historical (Maryland) or Secure (New York, West Virginia). Beyond the Mid-Atlantic portion of its range it is considered Extirpated (Michigan, Maine), Endangered (), or Historic (Massachusetts, North Carolina).

Pennsylvania Distribution The species is historically known from 36 counties, with no particular distributional pattern discernable. Pennsylvania is in the middle of the known geographic range, and within the state, water-plaintain spearwort does not appear to be limited to specific regions, climates, or geology types.

Habitat Water-plantain spearwort grows in various types of wetlands, which have been described on herbarium specimen labels as “streambanks and margins, swamps, wet places, ditches, bogs, marshes, wet thickets, intermittent streamlets, wet woods, pools and ponds, woodland ponds, muddy shores and depressions, swales, low fields, wet pastures, sloughs, ditches, and bottomlands.” Based on these descriptions, the species doesn’t appear to be limited to only certain types of wetlands. However observations of the few extant populations suggest that this plant prefers to be in mucky soils in or near slow-moving water where there is some periodic fluctuation in water levels.

Extant Locations Currently there are six extant locations in Pennsylvania, based on PNHP data, and all but one population have relatively small numbers (25 or fewer plants) of individuals. One population of between 250 and 500 individuals represents the majority of plants known in the state. Three of these locations haven’t been confirmed for ten years or more, so it is likely that some of them are no longer extant. The wide range of the species seems to suggest that it is not restricted to selected watersheds or physiographic provinces, and this may increase the possibility of finding more populations. However this is a large and

97 recognizable species and the lack of documentation suggests that it’s very uncommon throughout the state.

Historical Locations and Potential for Relocation There appear to be ca 60-66 historical locations in ca 30-35 counties, which are well distributed throughout the state. As a group, the historical locations are old - ca 40 per cent from 1900 or previous and ca 84% from 1950 or previous. A re-survey of suitable wetland habitats at some of the historical locations failed to find the species, suggesting that water-plantain spearwort is either extirpated from those sites or is seed-banking and may show up another year. Most of the historical locations have not been surveyed, but their age and accessibility argue against the likelihood of confirming very many of them. In addition, throughout its range this plant is declining and is threatened with extirpation from many locations and in some cases from the entire neighboring states.

Threats Some additional factors that are impacting this species include deer grazing and the encroachment of native and exotic invasive species. At all sites where the plant was confirmed extant, there was clear evidence of severe deer grazing, especially of the flowering stems that are elevated above the ground. The effects of this grazing were compounded by the fact that other, un-grazed, native invasive species were able to out-compete the spearwort for available habitat.

Status Justification With only six known extant populations, most of which haven’t been confirmed recently and some of which may be gone, and low numbers of plants per population in conjunction with the apparent range- wide decline, Pennsylvania Endangered is the most appropriate status for water-plantain spearwort (Ranunculus ambigens). The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012). However, Pennsylvania Threatened is also an acceptable status to account for some uncertainty in the number of populations that may exist but not yet have been documented.

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Pennsylvania Natural Heritage Program http://www.naturalheritage.state.pa.us/Species.aspx • North Carolina Heritage http://www.ncnhp.org/data/species-community-search • Virginia Heritage http://www.dcr.virginia.gov/natural-heritage/document/plantlist16.pdf • Maine Heritage http://www.maine.gov/dacf/mnap/features/ranamb.htm • Massachusetts Heritage http://www.mass.gov/eea/agencies/dfg/dfw/natural- heritage/species-information-and-conservation/rare-plants/plants-historic-list.html • New Hampshire Heritage http://www.nhdfl.org/library/pdf/Natural%20Heritage/TrackingList- PlantGeneral.pdf

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Long-stalked Crowfoot – Ranunculus hederaceus L. Author: Steven P. Grund, PNHP

Introduction Ranunculus hederaceus is one of the aquatic, white-flowered buttercups sometimes segregated from Ranunculus in the genus Batrachium, thus it is sometimes called Batrachium hederacea. It is easily distinguished from the only other white-flowered buttercup in the eastern United States (R. aquatilis var. diffusus) because the leaves are not dissected into linear segments. The other aquatic buttercups in Pennsylvania have yellow flowers, and have leaves very different from the small, three-lobed leaves of R. hederaceus. The lax stems, creeping along the saturated soil or near the surface of the water, rooting at the nodes, are also distinctive.

Proposed PA Status Ranunculus hederaceus has a current status of Pennsylvania Extirpated. If examination of the evidence shows the species to be native in Pennsylvania, a status of Pennsylvania Endangered will be proposed.

Habitat R. hederaceus typically occurs in disturbed, often eutrophic conditions in flushes in heathland, gateways and tracks, as well as on the margins of rivers, streams, and ponds (Lansdown 2011). In Pennsylvania it is known from an abandoned farm pond (Brickhouse Environmental Consultants and Engineers 2007) and from a springy brook (Pennell 1929).

North American Distribution and Global / Regional Conservation Concerns Ranunculus hederaceus occurs in western , North Africa and the northeast coast of North America. In Europe, it is restricted to Sweden, Denmark, , the United Kingdom, Ireland, the Netherlands, Belgium, France, , Portugal, Norway, and Croatia; it is protected as Endangered in Germany and Sweden, and considered introduced in Norway (Lansdown 2011). In North America, opinions have been mixed, with agreement only that it was not likely native in South Carolina, where it was discovered in 1821 and declared “if not extinct it has become rare” (Elliot 1823). It has not been documented in South Carolina since its original discovery there. Cook (1966) and Fernald (1929, 1950) considered Ranunculus hederaceus to be native in Newfoundland, but recent authors have considered the species to be introduced, primarily because the known localities are in the regions of the island with ports, and not in the southwest region where disjuncts from further south are typically encountered (Meads et al 2000, Rouleau and Lamoureux 1992). Recent rediscoveries of the species in both Pennsylvania and Maryland have revived the debate regarding the native status of Ranunculus hederaceus in North America.

Pennsylvania Distribution Ranunculus hederaceus has been documented in Pennsylvania only in Chester County, in 1929 and 2007.

Extant Locations The single extant location for Ranunculus hederaceus is on the edge of an abandoned farm pond in Chester County (Brickhouse Environmental Consultants and Engineers 2007).

Historical Locations and Potential for Relocation The 1929 collection from a tributary of the lower Susquehanna River could warrant more investigation. It is near some well-known serpentine barrens, and the general area has been extensively botanized, but

99 the possibility of that population persisting in the area cannot be ruled out. This species is tolerant of disturbance, so ephemeral populations may be present, but given the rarity of the species, seeds are likely not commonly present waiting for the right conditions for germination and growth.

Status Justification A specific recommendation must await completion of an analysis of the nativity of the species in our region (Grund et al, study in progress). If it is determined that the species is native here, a status of Pennsylvania Endangered would be appropriate, as we have only one known extant population, and there is only one other known extant population in the United States (Maryland).

References • Brickhouse Environmental Consultants and Engineers. 2007. Report detailing field surveys conducted by Janet Ebert and Jack Holt at Fox Hunt Farm, Chester County of April 7 and May 27, 2007. Unpublished report to the Pennsylvania Department of Natural Resources. • Cook, C. D. K. 1966. Studies in Ranunculus subgenus Batrachium (DC.) A. Gray. III. Ranunculus hederaceus L. and Omiophyllus Ten. Watsonia 6(4): 245-259. • Fernald, M. L. 1929. Some relationships of the floras of the Northern Hemisphere. Proc. Int. Congr. Pl. Sci. Ithaca 2: 1494-1500. • Fernald, M. L. 1950. Gray’s manual of botany, 8th ed. American Book Company, New York. • Grund, S.P., W.M. Knapp, and C.S. Keener. Unpublished study in progress. Rediscovery of Ranunculus hederaceus L. () in North America: Conservation Implications for Native and Naturalized Species. • Lansdown, R.V. 2011. Ranunculus hederaceus. The IUCN Red List of Threatened Species 2011: e.T164528A5904073. Downloaded on 21 October 2016. • Pennell, F.W. 1929. Herbarium specimen label, Pennell collection number 14749, duplicate specimens at Carnegie Museum of Natural History (CM) and the Academy of Natural Sciences of Philadelphia (PH)).

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Swamp Currant — Ribes lacustre (Pers.) Poir. Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP

Introduction Swamp currant is a low spreading shrub to about three feet in height and has conspicuously prickly and spiny branchlets. The leaves of this species have the maple-like shape typical of the genus Ribes, but can usually be distinguished from the other Pennsylvania species of the genus by their deeper and narrower sinuses and prominently doubly-toothed lobes. The small flowers, which appear in May, are reddish and are arranged in a raceme-like inflorescence. The not-very-tasty fruits are dark purple to black with bristly/sticky hairs.

Proposed PA Status Although the swamp currant has a PA legal status of Tentatively Undetermined, it has been assigned a PABS-Vascular Plant Technical Committee suggested status of Pennsylvania Endangered because only a single extant population has been documented by PNHP during its 35 years of activity. The Endangered status recommendation for this species has been in place for at least twenty years, and current information continues to support this as the appropriate status.

North American Distribution and Global / Regional Conservation Concerns Swamp currant has a global rank of Secure (G5).

Its global range is transcontinental across northern North America, with southern extensions in the Rocky Mountains and in the Appalachian Mountains to West Virginia (Kartesz 2003).

Of the six contiguous states to Pennsylvania, the swamp currant has been found in New Jersey (not ranked), New York (Apparently Secure S4) and West Virginia (Imperiled S2).

Pennsylvania Distribution In Pennsylvania, the species has a PNHP rank of S1 (Critically Imperiled) because there is only a single known population.

The historical state distribution of swamp currant includes seven counties in the glaciated northeast (Luzerne, Monroe, Sullivan, Susquehanna and Wayne) and glaciated northwest (Crawford and Warren), as well as two southern Allegheny mountain counties, Cambria (this specimen may be misidentified) and Westmoreland. From a Pennsylvania perspective, swamp currant is definitely a northern species. Except for a few locations in West Virginia, the swamp currant in Pennsylvania is near the southern end of its eastern North America global range.

Habitat In its global range, swamp currant grows in cool, damp to wet woods, thickets and swamps, but may also be associated with rock outcrops and talus slopes. The latter is certainly true for Pennsylvania’s only known population in Ricketts Glen State Park in Luzerne County, where the species grows in a hemlock (Tsuga canadensis) and northern hardwood forest within a cool, shaded ravine featuring a mountain stream with numerous waterfalls and humid growing conditions. The species grows on damp to seepy rocky outcrops and talus slopes along a relatively short stretch of this ravine.

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Only two of the approximate fourteen herbarium specimens of this species from Pennsylvania include habitat data, which is described as “dense woodland” (Warren County 1943) and “deep woods on stream” (Wayne County 1939).

At least superficially, there would appear to be considerable potential habitat for swamp currant in northern Pennsylvania, but the species may have subtle ecological requirements that aren’t yet apparent to botanists.

Extant Locations Numbers The population of swamp currant at Ricketts Glen State Park currently represents the only known extant location in Pennsylvania. This occurrence was documented by PNHP in 1991, but there are herbarium records from this location dating back to 1929 and possibly back to the 1890s (depending how locations on herbarium collections are interpreted).

Population size Based on periodic observations by PNHP over the last 25 years, swamp currant appears to be thriving in Ricketts Glen, although the area occupied by the plants is relatively limited and the number of plants relatively small. Definite population numbers are difficult to quantify because the growth habit of the species can make distinguishing individual plants very challenging, and some plants are situated in inaccessible places on cliffs, but the current population estimate is approximately 100 individuals.

Potential for additional EOs In the past 35 years, PNHP has conducted hundreds of de novo surveys in northern and/or mountainous Pennsylvania, which appear, at least superficially, to have potential for the presence of swamp currant, but the species has not been found. The species is also relatively distinctive looking and unlikely to be overlooked or misidentified as another species of Ribes.

An especially favorable de novo site would appear to be the Fishing Creek watershed located just a few miles west of Ricketts Glen. A stretch of this stream contains terrain, geology, habitats, and species, including rare species like twisted-stalk ( amplexifolius) and Braun’s holly fern (Polystichum braunii), which are very comparable to that where the swamp currant grows in Ricketts Glen. Despite these similarities, several PNHP surveys in this section of Fishing Creek have failed to detect swamp currant.

Nevertheless, there remains at least some potential of documenting new occurrences of swamp currant in Pennsylvania.

Historical Locations Numbers Historical locations of swamp currant in the form of herbarium specimens are known to PNHP from Cambria County in 1920 (although this specimen may be misidentified), Luzerne County in 1929 and 1939, Monroe County in 1840, Sullivan County in 1896, 1897, 1898, and 1903, Warren County in 1943, Wayne County in 1920 and 1939, and Westmoreland County in 1949. The locational data provided on these particular specimens is even vaguer than is usual on old herbarium specimens, and no information is available concerning how large these populations were at the time of collection.

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The three Luzerne County and possibly the four Sullivan County specimens may refer to the Ricketts Glen extant population, which leaves only five potential historical occurrences to be searched. None of these five have been vigorously searched by PNHP, mainly due to the relatively vague locations given.

Flora of Pennsylvania (Porter 1903) listed five counties (Dauphin, Huntingdon, Schuylkill, Tioga, and Somerset) as having records for swamp currant, but none of these counties are included in The Vascular Flora of Pennsylvania (Rhoads & Klein 1993), and must have been based on misidentifications.

Potential for extant EOs Confirming an extant population at any of the historical locations would appear to be a long shot.

Because the historical locations aren’t known with any specificity, the degree of past disturbance or current threat is unknown. It is unclear if the Ribes eradication programs in the 1900s for control of the white pine blister rust (Cronartium ribicola) had any effect on the frequency of swamp currant.

Threats Despite the apparent health of this population and its location in a portion of a state park where maintaining natural conditions is a priority, this occurrence of swamp currant does have some threats. The main one is the potential negative effects of the hemlock woolly adelgid (Adelges tsugae) in the ravine, as the loss of hemlocks could markedly change the cool, moist, and shaded growing conditions that currently are present, and be detrimental to swamp currant. Another concern for the species may be the relatively recent increase in the abundance of wood-nettle (Laportea canadensis) in the ravine. This native plant can become relatively tall and grow in dense concentrations, which could affect seedlings or immature individuals of swamp currant. Because of densely wooded and steep terrain at this location, and the considerable hiking traffic, the swamp currant may be protected to some degree from deer browsing, which may be significant on other species of Ribes at some locations in the state.

Status justification The state status of Endangered appears to be very appropriate for swamp currant, based on the single documented extant occurrence with its relatively small population, and the lack of success of discovering any additional populations despite decades of surveying in potential habitats in the state. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Porter, Thomas Conrad. 1903. Flora of Pennsylvania. Editor: John Kunkel Small. Boston, Ginn & Company. • Rhoads, Ann Fowler and William McKinley Klein, Jr. 1993. The Vascular Flora of Pennsylvania: annotated checklist and atlas. American Philosophical Society, Philadelphia, PA.

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Meadow Rose – Rosa blanda Aiton Author: Jessica McPherson. Reviewed: Steven P. Grund, John Kunsman, PNHP.

Introduction This species is native to eastern North America, but also widely cultivated.

Proposed PA Status Meadow rose is currently classified by the PABS-Vascular Plant Technical Committee as Tentatively Undetermined, and at this time there is not sufficient information available to justify a more precise status classification, because it cannot be determined if known populations are naturally occurring or spread from cultivation. Additional fieldwork is unlikely to provide the needed information; genetic work may be able to resolve the question.

Habitat The habitat in the major part of the range is described as “Anthropogenic (man-made or disturbed habitats), meadows and fields, shores of rivers or lakes, talus and rocky slopes“ in the Flora Novae- Angliae (Haines, Farnsworth and Morrison 2011).

North American Distribution and Global / Regional Conservation Concerns The North American range is predominantly north of Pennsylvania, with Pennsylvania populations near the southern edge of the range. The distribution in New York is more widespread, but concentrated in the northern part of the state (Kartesz 2003). It is listed as Imperiled (S2) in Ohio and West Virginia, and Critically Imperiled (S1) in Maryland. It is not yet ranked in Virginia because there is only one historical specimen known from the state and it cannot be determined whether it occurred naturally or not (Virginia Botanical Associates 2016).

Extant Locations In Pennsylvania, there are three extant occurrences, all situated in disturbed habitats. Because this species can and often does naturally colonize and utilize disturbed habitats, there is no reliable way at this time to distinguish whether the plants at these locations have established naturally or have persisted or spread from cultivated plantings. Genetic work drawing upon a region-wide sample of material may be able to provide further insights into this question; we recommend consultation with an expert in genetic taxonomic assessments to determine whether this avenue of research is likely to prove fruitful, given the limited availability of fresh material, the condition of dried specimens, and the confounding problem of cultivars.

Historical Locations and Potential for Relocation There are 11 historical occurrences documented through museum specimens. The specimens upon which the historical occurrences are based did not include enough habitat data to provide any insight into the species’ requirements in Pennsylvania, aside from further suggesting an affinity for disturbed habitat. In fact, while its presence in adjacent New York and the high elevations of western Maryland and West Virginia suggests it is likely for the species to occur natively in Pennsylvania, it cannot be conclusively determined based on the scanty information provided with the specimens that any of these occurrences are naturally occurring rather than spread from cultivation.

Status Justification

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Because it is not possible with currently available information to determine whether any of the extant occurrences of Rosa blanda in Pennsylvania are naturally occurring, we cannot recommend any change to the current status of “Tentatively Undetermined.”

References • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Virginia Botanical Associates. 2016. Digital Atlas of the Virginia Flora (http://www.vaplantatlas.org). c/o Virginia Botanical Associates, Blacksburg. [Accessed: (10/6/2016)]

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Bristly Blackberry – Rubus setosus Bigelow Author: Jessica McPherson. Reviewed: John Kunsman, William Gleason PNHP.

Introduction Bristly blackberry is a member of the taxonomically difficult genus Rubus. Taxonomists have long struggled to satisfactorily define species in this genus, because asexual seed production (apomixis), spontaneous generation of new taxa by genome multiplication (polyploidy), and hybridization all occur frequently in the genus, creating a great range of morphological types that cannot easily be categorized using the conventional definition of species as a group of individuals that can interbreed with each other but are reproductively isolated from individuals outside the species. Over the course of the last century, estimates of the number of Rubus species in eastern North America by different taxonomists have ranged from 25 to 250. We follow the somewhat conservative treatment of Flora of North America. Although the bristly blackberry is uncommon in Pennsylvania, it appears to be reasonably common northwards in its range, and it is generally recognized as a valid species concept. Flora of North America, the Pennsylvania Flora Project, and other major floristic treatments that address regions within the range of bristly blackberry all recognize it as a valid species concept (Alice et al. 2014; Rhoads and Block 2007; Reznicek and Voss 2011; Haines, Farnsworth and Morrison 2011; Weakley 2015).

Kartesz’s BONAP project recognizes a more limited species concept than other modern authors (Kartesz 2003), choosing to segregate out several other concepts (R. semisetosus, R. elegantulus, R. racemiger) that are included under the name R. setosus by FNA, Pennsylvania Flora Project, and most modern authors. This document recognizes the broader concept, following the Pennsylvania Flora Project and the Pennsylvania Natural Heritage Program.

Proposed PA Status A status of Pennsylvania Threatened is recommended for the bristly blackberry at this time, based on the small number of known extant populations (4), mitigated by its ability to utilize disturbed habitat and uncertainty about the level of survey effort to date; as increased awareness of this species and survey effort in its habitat have recently yielded newly discovered populations, this trend may continue. This species currently has no assigned legal status, and a recommended status of Pennsylvania Endangered from the PABS-Vascular Plant Technical Committee, which has not yet considered the status in light of recent information.

North American Distribution and Global / Regional Conservation Concerns This species has a global range concentrated in northeastern North America. It is listed by NatureServe as Possibly Extirpated (SH) in West Virginia and New Jersey, and as Apparently Secure (S4) in New York. The global rank assigned by NatureServe is G4G5, following the Weakley concept of the species. NatureServe lists two concepts for this taxon, an older less-inclusive one by Kartesz and a newer, broader one by Weakley that includes some older names that were previously recognized as distinct. Kartesz maps the more limited concept south to West Virginia, while Weakley has locations for the broader concept in North Carolina. Both concepts, however, indicate the species as quite uncommon south of New York.

Pennsylvania Distribution Pennsylvania is near the southern extent of the range. Historical specimens document the species in Pennsylvania from the northeastern region, and from the higher elevations of the Alleghenies in southwestern PA. This distribution, which includes the areas of the state with the coolest climate, is typical for species that are at the southern edge of their range in Pennsylvania.

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Habitat In the main part of its range, the habitat is described in Flora Novae-Angliae as “fields, roadsides, forest borders, open rights-of-way, marshes edges, ascending to boreal habitats” (Haines, Farnsworth and Morrison 2011). In Pennsylvania, specimen habitat data and observations at extant sites confirm similar generic early successional edge or field habitats, sometimes wet.

Extant and Historical Locations At the start of the Plant Status Updates projects, no extant populations had been observed in Pennsylvania since 1963. In the course of the project, all museum specimens were reviewed to confirm identification, and all historical locations with reasonably precise locational data and permission to access were surveyed. Surveys at historical locations did not relocate any plants that could be confirmed as Rubus setosus, although several individuals with traits suggesting possible R. setosus parentage were observed. However, four new locations were documented through de novo survey work. All new locations of Rubus setosus were documented in northeastern Pennsylvania. Specimen identification has been confirmed for the first two new locations by Tyler Smith, PhD, research scientist with the Ottawa Research and Development Centre in Canada and taxonomy editor for the journal Castanea. Dr. Smith was conducting research on Rubus including the Setosus group, but this work is not yet complete and is currently on hold. One specimen was also identified as Rubus groutianus by Mark Widrlechner, PhD, of the USDA-ARS North Central Regional Plant Introduction Station in Iowa. Dr. Widrlechner’s expertise includes biosystematics of members of the genus Rubus native to the North Central United States. Under the treatment provided in Flora of North America, R. groutianus is synonymized under R. setosus. Informal commentary by several authors including Tyler Smith suggests that the concept currently recognized as R. setosus may contain more than one taxonomic entity, but work has not yet been completed to resolve this question. Until such work is completed, we recommend following the Flora of North America treatment to consider R. groutianus as a of R. setosus. Two additional fairly extensive populations were documented in disturbed floodplain habitat along the Delaware River in Wayne County. One extensive population was along trails/hedgerows between old fields and forest in formerly farmed lands now owned by PGC. The other was in the river floodplain at a boat launch area.

It is uncertain how far west in Pennsylvania the setose blackberry may extend today. No populations have been relocated in the Alleghenies. Three of the four known extant populations are in Monroe and Wayne counties, which are in close geographic proximity to the known range in New York and New Jersey, where the species appears to be more abundant. However, one of the extant populations was found in Tioga County, which is not near any counties in New York or Pennsylvania where the species has been previously documented.

Threats The major threat faced by this species is potential decline of populations due to loss of suitable habitat in Pennsylvania from climate change. As a northern species at the southern edge of its range in Pennsylvania, historically known only from the parts of the state with the coolest climate, it may not be able to persist if climate conditions are altered in these regions.

Status Justification With currently available data, a status of Pennsylvania Threatened is recommended for this species. Factors indicating a conservative status of Pennsylvania Endangered include its rarity, with only four extant populations currently documented; and its potential vulnerability to anthropogenic climate

107 change, as a northern species at the southern edge of its range in Pennsylvania which is only found in the coldest portion of the state. Factors indicating a less conservative status are its use of disturbed early successional habitat and the abundance of such habitat in the NEPA region; and uncertainty about the level of survey effort for this species. While the setose stem is fairly distinctive, and collections are almost completely absent for several decades, the species can also be easily overlooked if other Rubus species with 5 palmate leaflets are present (R. allegheniensis, R. canadensis, R. pubescens), and these species are very common in potential R. setosus habitats. It seems likely that some new populations will be documented in the northeastern region of PA, but it is uncertain how many, and what kind of long- term trend the species is experiencing.

Given this combination of factors, a status of PT is recommended for this species at this time. Extrapolation of a species with only 4 known extant sites to Pennsylvania Rare is not warranted, while PE is probably too cautionary given its use of disturbed habitat and the likelihood it is somewhat overlooked. It may also be advisable to reassess the status in several years, if survey work continues to document new populations. The Pennsylvania Natural Heritage Program, a member program of NatureServe, recommends a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0. • Lawrence A. Alice, Douglas H. Goldman, James A. Macklin, Gerry Moore . 2014. Rubus. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 9. • MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. October 6, 2016. http://www.michiganflora.net/species.aspx?id=2564. • Pennsylvania Flora Project, Morris Arboretum, University of Pennsylvania. www.paflora.org. • Rhoads, Ann F. and Timothy A. Block. 2007. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia; 2nd edition. 1056 pg. • Weakley, Alan S. 2015. Flora of the southern and mid-Atlantic states. University of North Carolina Herbarium, North Carolina Botanical Garden, University of North Carolina, Chapel Hill.

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Blue Willow – Salix myricoides Muhl. Author: Jim Bissell, Cleveland Museum of Natural History. Edited: Jessica McPherson, PNHP

Introduction Blue willow (Salix myricoides) is a shrub within the willow family () which has a strong resemblance to heat-leaf willow (Salix eriocephala) and pussy willow (Salix discolor).

Proposed PA Status We recommend a status of Pennsylvania Endangered for the blue willow because there are only eight extant populations in the state, the populations contain very small numbers of plants, and the species appears to have declined in recent decades. Two of the populations are within the Presque Isle Peninsula. Some large populations of blue willow, mapped at Presque Isle in 1997, have disappeared due to erosion of sand dunes. All inland populations observed in 2014 were severely browsed by deer. The PABS-Vascular Plant Technical Committee currently recommends a status of PE.

North American Distribution and Global / Regional Conservation Concerns Blue willow has a very limited geographic range within North America: southern Ontario, Michigan, Wisconsin, Illinois, Indiana, Ohio, and Pennsylvania. It is mostly found near the Great Lakes, although a few inland locations are known in Wisconsin, Ohio, and Pennsylvania (Kartesz 2003). NatureServe has assigned a global rank of G4, Apparently Secure. A NatureServe status of Imperiled (S2) is assigned in Ohio and Maine, while it is considered Vulnerable (S3) in Quebec, Newfoundland, and New Brunswick, and Apparently Secure (S4) in Ontario. Blue willow is not listed in Michigan. Blue willow has legal status of Potentially Threatened (rare) within Ohio and Threatened (rare) within Ontario.

Habitat With the exception of sand dunes at Presque Isle State Park in Erie County, all inland populations of blue willow are confined to wetlands. The only upland occurrences known in Pennsylvania are confined to sand dunes within Presque Isle State Park. Michigan Flora describes its habitat as “Sandy shores, dunes, and moist interdunal hollows along the Great Lakes; rarely inland on calcareous shores and fens.” This description appears to be accurate for Pennsylvania based on known locations; blue willow populations in Pennsylvania have been documented at Presque Isle and in glacial fens, river edges, and alkaline swamps. One population is present within an open semi-wet meadow at Jennings Environmental Center.

Extant Locations All extant populations of blue willow within the commonwealth, with the exception of Presque Isle, are quite small. Prior to 1990, a large population of blue willow occurred on sand dunes at Presque Isle. However, this population has declined, and all known within Pennsylvania have 50 or fewer individual shrubs. The largest population is at Jennings Prairie, with about 50 individuals, and the remaining populations range from 1 to 19 shrubs, for a combined total of 90 individuals known in the state. Extant populations are known within Erie County, Fayette County, Franklin County, and Tioga County. While a number of historical locations are known from river edges, today most known locations are in fens, while only one extant location is along a river edge.

Historical Locations and Potential for Relocation There are 11 historical locations of blue willow known from museum collections; several are from floodplains of rivers or large creeks, a few are from fen-like habitats in the northwest, and a few from other wetlands in various parts of the state. Five of these sites are estimated to have any chance of

109 relocation; the remaining sites have either been surveyed unsuccessfully already, had habitat alterations sufficient to eliminate the population, or have location data listed that is too vague to relocate.

Threats Cleveland Museum surveys of blue willow populations in northwestern Pennsylvania, in 2014, found all fen populations severely grazed by deer. Since construction of the off-shore jetties at Presque Isle in 1990, large populations of blue willow on sand dunes, east of the off-shore jetties, were extirpated from Presque isle because the dunes on which they occurred in 1997 were washed away during winter storm surges.

Status Justification We recommend that blue willow should be listed as Pennsylvania Endangered based upon the limited number of known populations, the small population sizes, the very low number of total individuals known in the state, and the ongoing threat from deer browsing. Presque Isle still has the largest population known within Pennsylvania, but the population at Presque Isle has drastically declined since 1997. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Bog Willow — Salix pedicellaris Pursh Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP.

Introduction Bog willow is a low spreading shrub to about 3 feet in height with consistently hairless branchlets. The thickish leaves have prominent veins, entire margins, are hairless on both sides, and often whitened below. The male and female flowers occur on separate plants. The fruit is a many-seeded capsule with the individual seeds each having a tuft of hairs.

Although species in the genus Salix are difficult to identify vegetatively because of variability in hairiness and leaf margin type, bog willow is a notable exception. The lack of hairiness on the vegetative parts, the consistently entire leaves that are often whitened below, and its bog habitat, make it one of the more easily identified willows in Pennsylvania.

Proposed PA Status There is no current legal status for bog willow in PA, since it had not been documented in Pennsylvania at the time of the publishing of the initial state plant regulations. It has been now been assigned a PABS-Vascular Plant Technical Committee suggested status of Pennsylvania Endangered, and has had this status for at least ten years, since the first occurrence was discovered in 1994 in Erie County. Only two additional extant occurrences have been found, in the northeastern counties of Pike and Wayne, since the Erie County discovery. We recommend Pennsylvania Endangered as the appropriate status based on small number of known populations and the relatively thorough inventory of available habitat that has been completed.

North American Distribution and Global / Regional Conservation Concerns Bog willow has a global rank of Secure (G5) and the global range is transcontinental across northern North America (Kartesz 2003).

Of the six contiguous states to Pennsylvania, bog willow has been found in New Jersey (Critically Imperiled S1), New York (Secure S5), and Ohio (Critically Imperiled S1).

Pennsylvania Distribution In Pennsylvania, bog willow has a PNHP rank of S1 (Critically Imperiled) because only three relatively small populations are known in the northern counties of Erie, Pike, and Wayne. This species is distinctively northern from a Pennsylvania perspective, and reaches a southern border of its global range here.

Habitat As its name implies, bog willow typically grows in boggy and peaty wetlands. All three of the extant sites in Pennsylvania would be characterized as fitting this description, but the Erie and Wayne occurrences have also been described as having calcareous qualities based on the other species growing there. This might suggest that bog willow has a preference for boggy and peaty habitats that are less acidic (at least in Pennsylvania), and might explain the apparent rarity of the species in the state since calcareous habitats of that type are believed to be much less frequent than the more acidic types.

Extant Locations Numbers

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Bog willow has been documented at three locations: Edinboro Lake Fen in Erie County, Smiths Swamp in Pike County, and Farrell Corners Fen in Wayne County.

The Edinboro Lake Fen occurrence was first documented in 1994 and revisited in 2002, which is significant because during both visits the population was reported to be having severe competition and deer browse problems. An update is needed to confirm that this population still exists, and if so, an estimate of the size of the colony. This location is privately owned.

The Smiths Swamp occurrence was discovered in 1995 and reconfirmed by PNHP in 1996, 2005, and 2009. This bog willow colony appears to be thriving, at least vegetatively, although flowers or fruits have yet to be seen at this location. The surrounding swamp, which is situated in State Game Lands, has plenty of available habitat for expansion of the population, and possibly supports additional individuals.

The Farrell Corners Fen occurrence was first documented in 2006 and last visited in 2013. This population is the largest of the three populations. The situation at this site appears similar to Smiths Swamp in that the plants are situated in State Game Lands, look vegetatively healthy but flowers or fruits have not been observed, and there is a lot of potential habitat for the species to utilize.

Population sizes Both the Edinboro Lake Fen and Smiths Swamp occurrences consist, at least at this juncture, of vegetative clones of a single individual – the size being described as “one or two small clones” for the former and a single 30 feet by 15 feet clone for the latter.

The Farrell Corners Fen occurrence consists of a few dozen plants that are separated from each other and found scattered over about an acre, which suggests that these plants are separate individuals and that seeds may have been produced at least sporadically in the past. On the other hand, these plants may have been part of a clone in which the individual ramets became separated over time and now resemble individual plants.

Potential for additional EOs In the past 35 years, PNHP has conducted hundreds of de novo surveys in boggy/peaty wetlands in northern Pennsylvania, which appear, at least superficially, to have potential habitat for bog willow, but only three occurrences have been found. The species also has relatively distinctive features and seems unlikely to be misidentified as another species of willow. There is also some suggestion that the species may prefer a more calcareous type of boggy/peaty wetland than is typically found in the state. All of these factors could lead to the conclusion that documenting additional populations of bog willow in Pennsylvania appears unlikely.

On the other hand, many of these surveyed wetlands, as well as the large number of wetlands that have yet to be surveyed, often cover a large area and may be a challenge to survey and maneuver in because of the substrate and dense vegetation. It may be that bog willow occurs in small populations in the Pennsylvania part of its range, which would make new locations even more difficult to find. These factors suggest that there remains at least some potential for finding additional extant occurrences of the species.

Historical Locations No historical occurrences are known. The species was first found in Pennsylvania in 1994.

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Threats The Edinboro Lake Fen population has been described as having problems of deer herbivory and competition from other plant species, as well as the handicaps of very small population size and possibly a reliance on vegetative reproduction only.

The Smiths Swamp and Farrell Corners Fen populations have potential negative impacts from hydrology changes – especially flooding from beaver activity – as well as the handicaps of small population sizes and possibly a reliance on vegetative reproduction only.

Status Justification We recommend the status of Pennsylvania Endangered for bog willow. Pennsylvania Endangered is very appropriate, since only three extant occurrences have been documented despite 35 years of surveys in its potential habitat, the populations at each of the occurrences are very small and possibly reliant exclusively on vegetative reproduction, and the populations are threatened by deer, competition, and potential hydrological changes to the habitats. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S1 (Critically Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T. 2003. A Synonymized Checklist and Atlas with Biological Attributes for the Vascular Flora of the United States, Canada, and Greenland. Second Edition. In: Kartesz, J.T. Synthesis of the North American Flora, Version 2.0.

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Blue-eyed Grass – Sisyrinchium montanum Greene Author: Jessica McPherson, PNHP. Reviewed: Steven P. Grund, PNHP.

Introduction There are two varieties of this species, Sisyrinchium montanum var. montanum and Sisyrinchium montanum var. crebum. S. montanum var. crebum is believed to have more limited distribution in the northeast coastal region, while var. montanum is widespread in the northern regions of North America (Kartesz 2015; Cholewa and Henderson 2002; Haines, Farnsworth and Morrison 2011). It is unclear whether one or both varieties are present in Pennsylvania, and thus the taxa are treated at the species level in this account.

Proposed PA Status The current status recommended for this species by the PABS-Vascular Plant Technical Committee in Pennsylvania is Tentatively Undetermined, and sufficient information is not yet available to assign a more precise status. We recommend further training of botanical surveyors to increase detection of this species in appropriate habitat (generic early successional areas), and reassessment of available information in several years.

North American Distribution and Global / Regional Conservation Concerns The North American range is concentrated north of Pennsylvania (Kartesz 2015). It is listed by NatureServe as Critically Imperiled (S1) in West Virginia, Ohio, Indiana, and Illinois.

Pennsylvania Distribution Pennsylvania is at the very southern edge of the North American range. Within the state, the distribution appears to be concentrated in the northeast, with a few occurrences documented in the northcentral and central regions. Typically, species that exhibit this pattern are limited to northern climates.

Habitat The habitat is early successional openings, described as “fields and pastures” in the New York Flora Atlas (Weldy, Werier and Nelson 2016), and “forest edges, meadows and fields, shores of rivers or lakes” in Flora Novae-Angliae (Haines, Farnsworth and Morrison 2011).

Extant and Historical Locations Although this species was listed as Tentatively Undetermined in 2011, no element occurrences are currently recorded in Biotics. No new locations for the species have been documented, and historical records from museum sources have not yet been added to the database. The Pennsylvania Flora Project is the best available source of distribution data; 38 locations are mapped. (These are all assigned to S. montanum var. crebum, but new work has been done to distinguish the varieties since the map was made, and the specimens have not been reviewed in light of the new work). Specimen data from Carnegie Museum was reviewed as part of this project, including 15 records from a wide scattering of counties. The most recent collections are 1979 and 1982 in Blair County, 1982 in Westmoreland County, 1966 in Clearfield County, and 1960 in McKean County.

Threats It is difficult to assess potential threat level to this species in Pennsylvania when no extant locations are currently known. Its ability to utilize early-successional habitats and establish in such settings, documented elsewhere in the range, make it likely that it faces a fairly low level of threat. However, as a

115 northern species near the southern edge of its range in Pennsylvania, it may face threats from climate change.

Status Justification Because there are no known extant sites currently in Pennsylvania, but museum specimens indicate at least 38 sites with a range of dates from quite old to moderately recent, and the species utilizes early successional habitat which is abundant in the state but makes relocation of historical locations questionable, it is highly uncertain how much of this species might be present in Pennsylvania, and we cannot at this time recommend an update to the proposed status of Tentatively Undetermined.

The status of this species remains unresolved, but it is a low priority for targeted fieldwork. It remains uncertain how much of this species is found in PA, but it appears to occupy disturbed habitats such as grassy road verges or ROWs. Because it prefers generic disturbed habitats that are both extensive and generally very low priority for conservation-oriented botany surveys, it is not a good use of resources to conduct targeted field surveys for this species. It is more efficient to educate botanists who are already visiting these habitats to watch for this species in the course of other work. We recommend reviewing available information for this species again in several years, and revisiting the status of this species if substantial new information has been added.

References • Cholewa, Anita F. & Douglass M. Henderson. 2002. Sisyrinchium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 26. • Haines, Arthur, Elizabeth Farnsworth, and Gordon Morrison. 2011. New England Wildflower Society's flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. [Framingham, Mass.]: New England Wild Flower Society. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. • Pennsylvania Flora Project, Morris Arboretum, University of Pennsylvania. www.paflora.org. • Weldy, Troy, David Werier, and Andrew Nelson. 2016 New York Flora Atlas. [S. M. Landry and K. N. Campbell (original application development), USF Water Institute. University of South Florida]. New York Flora Association, Albany, New York.

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Purple Meadow-rue – Thalictrum dasycarpum Fisch. & Avé-Lall Author: Steven P. Grund, PNHP

Introduction Thalictrum dasycarpum is a Midwestern species. In the east, it is replaced by the closely related T. pubescens, which is very common in Pennsylvania. In the narrow zone of overlap, which includes western Pennsylvania, there is hybridization and/or introgression between the species, and identification can be difficult. In the “first Atlas” (Wherry et al 1979), there is no map for T. dasycarpum, but there is a map for “Thalictrum – intermediate of dasycarpum and pubescens.” Keener (1986, attached) addressed the situation in an unpublished manuscript after conducting research funded by DER. Keener examined specimens from CM, PENN (now part of PH), and PAC, including all of the specimens mapped as intermediates in Wherry et al (1979), and conducted field studies. He recommended at the time that Thalictrum dasycarpum not (emphasis his) be included as a member of the flora of Pennsylvania; however, in 1989, Marilyn Park later discovered a specimen at PH from Warren County collected by H.A. Wahl, annotated it as Thalictrum dasycarpum; and Keener concurred with the identification (it is not clear whether Keener independently identified the specimen or accepted the expertise of Park). Keener recommended that the site be surveyed to determine whether the population is still extant (Ed Dix, pers. comm., quoted from the minutes from the 1997 Pennsylvania Rare Plant Forum).

Proposed PA Status The PABS-Vascular Plant Technical Committee has changed its recommendation for purple meadow-rue from Tentatively Undetermined (adopted spring 1997) to Pennsylvania Extirpated (adopted Spring 2017), because herbarium specimen review has determined there is only one verified specimen of this species collected in Pennsylvania in 1951, and fieldwork failed to relocate the plant at that site.

Habitat Deciduous riparian woods, damp thickets, swamps, wet meadows, and prairies (Park & Festerling 1997). In Pennsylvania, the single specimen known was collected from a brushy bottom (Wahl, H.A. et al. 1951 June 22, specimen at the Academy of Natural Sciences, Philadelphia).

North American Distribution and Global / Regional Conservation Concerns Purple meadow rue ranges from the Yukon south to northern Arizona, west to Quebec and Alabama. It is most common in the Great Plains and Midwest (Kartesz 2015). It is considered to be rare in several states at the periphery of its range, but none of those states are near Pennsylvania.

Pennsylvania Distribution Purple meadow rue has been reported in several locations in northwestern Pennsylvania, but most of these records have been determined to represent intermediates between this species and the closely related Thalictrum pubescens, which is widespread and common in Pennsylvania. The single specimen from Pennsylvania now considered to represent T. dasycarpum was collected in Warren County.

Extant Locations This species has not been documented in Pennsylvania since 1951. Surveys have been conducted in the vicinity of the 1951 collection locality, but there are many wetlands in that area for which landowners denied permission for surveys to be conducted.

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Surveys have been conducted in the vicinity of the 1951 collection locality, but there are many wetlands in that area with suitable habitat for which landowners denied permission for surveys to be conducted.

Threats Threats are not relevant to the appropriate conservation status of this species in Pennsylvania, since we have no evidence of any extant occurrences. If the species is extant, but undetected, somewhere in the commonwealth, it is subject to at least the many threats common to all wetland species.

Status Justification The 1951 Wahl specimen was collected from 0.5 miles south of Clarendon in Warren County. Grund (2015) surveyed areas of the Allegheny National Forest that fit that locality description, but was refused permission to access adjacent private lands with extensive wetlands. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of SH (State Historic) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber- Langendoen et al. 2012). We do not know whether the species is extant at the Wahl station, but it has not been documented there since 1951, and has never been convincingly documented anywhere else in the commonwealth. Because there is no status in the Wild Resource Conservation Act that corresponds directly to State Historic, we recommend a status of Pennsylvania Extirpated.

References • Grund, S.P. 2015. Survey for Thalictrum dasycarpum at West Branch Tionesta Creek. F15GRU25. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. • Keener, C.S. 1986. Possible distribution of Thalictrum dasycarpum in Pennsylvania. Report to Commonwealth of Pennsylvania, Department of Environmental Resources. • Park, M.M. and D. Festerling Jr. 1997. Thalictrum. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 3. • Wherry, E.T., J.M. Fogg, and H.A. Wahl. 1979. Atlas of the Flora of Pennsylvania. The Morris Arboretum of the University of Pennsylvania.

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Nodding Trillium — Trillium cernuum L. Author: John Kunsman, PNHP. Edited: Jessica McPherson, PNHP.

Introduction Among the Pennsylvania species of Trillium that have white flowers, nodding trillium can generally be distinguished by its strongly-declined flower stalk that is often oriented more-or-less parallel to the stem and holds the flower “hidden” beneath the leaves. The leaves alone can sometimes be distinguished from those of the other white species by the presence of a more -like leaf base and the “oily” appearance of the fresh leaf surface.

This species has a notable variation in the size of its flowers, with plants having larger flowers being named as variety macranthum. Whether this variety and typical variety cernuum, which has smaller and less attractive flowers, have distinct habitats and distributions in Pennsylvania, and possibly different conservation concerns, remain to be investigated.

Proposed PA Status Although nodding trillium has a PA legal status of Tentatively Undetermined, PABS recommends a status of Threatened. The primary reasons for this status are the relatively small population sizes at most extant occurrences, and because the mesic to wet woodland habitat that is typical for the species has major threats from invasive species and fragmentation, especially in southeastern Pennsylvania where most of the populations occur.

North American Distribution and Global / Regional Conservation Concerns Nodding trillium has global rank of Secure (G5).

The distribution of nodding trillium extends from Newfoundland west into Saskatchewan, and south into Virginia, West Virginia, Ohio, Indiana, Illinois, Iowa, and South Dakota (Kartesz 2015).

Nodding trillium has been documented in all six states contiguous to Pennsylvania. The species has a conservation status in Delaware (Imperiled S2), Ohio (Extirpated SX), Maryland (Vulnerable S3), and West Virginia (Critically Imperiled S1). The species does not have a rank in New Jersey (NR) and is considered Apparently Secure (S4) in New York.

Pennsylvania Distribution Nodding trillium has a NatureServe state rank of Imperiled (S2), for reasons listed under the status justification.

The current and historical distribution of nodding trillium is concentrated in the southeastern counties, particularly in the Delaware River watershed. To the west, the species also occurs in some counties, notably Lancaster, Lebanon, and York, in the lower Susquehanna River watershed. There are relatively few locations currently documented from the Ohio River watershed in western Pennsylvania.

Habitat Nodding trillium often gets labeled as a “rich moist woods” species, particularly where the forest has a calcareous substrate, or has a geology such as diabase that tends to be beneficial for calcareous species. While many populations in PA fit well into this generalized habitat, some occurrences have been found on sandstone and related rocks. Even more different from the conventional wisdom is the fact that

119 nodding trillium may also be found in forests that are markedly poorly-drained, such as swamps and seepage woodlands.

Based on PA herbarium specimens from the past 150 years, some examples of the habitats utilized by nodding trillium include “rocky woods, moist woods, deep rich woods, rich woods, cool woods, wooded slope, moist woods on sandstone, swamp, swampy woods, spring swamp, alluvial woods, wet woods, swale, bog, streambank, low woods, swampy bottomland, damp hemlock grove.”

Extant Locations Numbers At present, 68 populations of nodding trillium have been documented by PNHP in 15 counties. This extant range strongly resembles the historical range, with nearly three-fourths of the known populations located in the southeastern counties of Chester (23), Berks (9), Northampton (8), Bucks (4), and Lehigh (4). Most of the occurrences are situated on private land, with approximately 10 on federal property or state property of state forests, state parks, or game lands. Approximately 19 of these populations have not been revisited within the past 10 years.

Population sizes The population sizes at the 68 extant locations vary from one plant to over 1000 plants (or stems), although about one-half the populations have 25 plants or fewer. The estimated number of plants in the state is approximately 6000.

Although Chester County has 35% of the extant locations, it has less than 10% of the total number of known plants in the state.

Potential for additional EOs There is potential for additional populations, but exactly how many is unknowable.

Historical Locations Numbers Based on herbarium specimens from Pennsylvania, there have been collections of nodding trillium from at least 80 locations in 17 counties, primarily in southeastern and to a lesser degree southcentral Pennsylvania. The actual number will vary depending on the manner in which individual occurrences are distinguished during the PNHP mapping process, which is made more difficult because many of the collections have vague directions. The dates of the collections vary from the mid-1800s to the 1970s, with well over one-half of the specimens collected prior to 1920. The size of the populations that existed at these collection sites remains completely unknown due to the lack of data on the herbarium label. Many of these historical occurrences have not been processed into Biotics.

Potential for extant EOs Because the locations of these historical records are situated primarily in the southeastern counties, it is likely that habitat loss and degradation (particularly from invasive species) have been substantial. Given their location in the state and the changes that have occurred there over time, the age of the collections, and the effort required to contact landowners for access, it appears unlikely that many of these historical locations would be worth the effort to survey. In fact, de novo surveys in southeastern Pennsylvania, particularly on public lands, watershed properties, and nature preserves, might offer a greater potential for discovering extant populations than searching historical locations.

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Threats Because most of the known populations are located in southeastern Pennsylvania, there are major threats from invasive species, habitat fragmentation, and excessive deer browse. Poaching by gardeners may be a problem at some locations.

Status Justification Pennsylvania Threatened is recommended for nodding trillium, because while the number of known populations is higher than usual for this status, most populations are small (one-half have 25 plants or fewer) and the total state population (6000) is consistent with Pennsylvania Threatened status. Furthermore, the species faces a high level of threat because most of the known populations occur in southeastern Pennsylvania where threats from habitat fragmentation and invasive species are particularly serious. A considerable number of populations have not been revisited recently, and therefore do not have current information available regarding their health; given regional threat trends, it is likely these may have declined. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S2 (Imperiled) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)].

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Declined Trillium – Trillium flexipes Raf. Author: Jessica McPherson, PNHP

Introduction Declined trillium is a member of the Erectum group of the genus Trillium, which includes six species. Three occur in Pennsylvania: declined trillium (Trillium flexipes), red trillium (), and nodding trillium (Trillium cernuum). These species can hybridize, and individuals with hybrid traits are frequently observed in PA populations. The term “syngameon” has been applied to this species complex, to describe a group of taxa that have partial reproductive isolation but can also hybridize.

Proposed PA Status Pennsylvania Threatened is determined to be the appropriate conservation status rank for Trillium flexipes, based on the limited number of locations and individuals known in the commonwealth and the level of threat the species currently faces. This is the status currently recommended by the PABS- Vascular Plant Technical Committee.

Habitat Flora of North America describes the habitat for declined trillium as “rich wooded slopes, floodplains in deciduous forests, especially over limestone” (Case 2003). This is consistent with Pennsylvania populations.

North American Distribution and Global / Regional Conservation Concerns The global range of declined trillium is predominantly midwestern, from eastern Minnesota, Iowa, and Missouri to Western Pennsylvania and New York; there are also semi-disjunct populations in the southern Appalachians and eastern Pennsylvania. It is considered to be of conservation concern in many states at the edges of its range. It is listed by NatureServe as Imperiled (S2) in West Virginia and Pennsylvania, and as Critically Imperiled (S1) in New York, Maryland, and Virginia.

Pennsylvania Distribution Trillium flexipes is known only from localized areas in Pennsylvania. There are several large populations along the lower Susquehanna River in eastern Pennsylvania. In western Pennsylvania, most records for the species are from Allegheny and Beaver counties, with a few scattered locations in surrounding counties, and a scattering of records further in the northwest.

Extant Locations Forty-one (41) extant locations are known for declined trillium.* Twenty-three of these are in eastern Pennsylvania, mainly along the Susquehanna River floodplain. Seventeen (17) are in western Pennsylvania, and 1 record is known from Centre County.** The total population for the state is estimated to be ~14,000 individuals.

*This includes Biotics records for Trillium flexipes and for “Trillium x 1,” as well as 11 eastern Pennsylvania locations that have not been entered into Biotics due to landowner permission issues.

**The Centre County population has a different appearance than other populations and is also situated very near to residential development; in combination with its disjuncture from other population clusters in the state, these facts lead to some suspicion it may have been introduced, although the habitat and surrounding vegetation appear very consistent with the species elsewhere in the state.

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Historical Locations and Potential for Relocation Twenty-three historical populations are known from museum specimens. About half of these have been searched for, while another 6 locations either have no good habitat remaining or have locational data too vague to relocate. Many historical locations are also in fairly close proximity to each other or to extant locations; at this point, relatively few additional sites are likely to be discovered through searching for historical records.

Threats Declined trillium faces several serious threats to its long-term viability in the state. The species is very vulnerable to competition from exotic invasive species, because most locations are in fairly fragmented habitats in regions where invasive species are highly problematic. Furthermore, the mesic, high-pH habitats occupied by declined trillium, especially floodplains, are a type of ecosystem that is particularly vulnerable to invasion. Deer browse is also an ongoing threat, as Trillium spp. are a preferred food for deer. In western Pennsylvania, many extant locations and areas of available habitat are also situated in regions with a fairly high rate of residential development, and vulnerable to either direct or indirect impacts of continued development.

Status Justification A status of Pennsylvania Threatened is recommended for declined trillium, due to the high level of threat the species faces in combination with low numbers of populations and individuals present in the state. Although the numbers of populations (41) and individuals (14,000) are slightly higher than typical for a status of Pennsylvania Threatened, the high threat level faced by the species justifies this status. Furthermore, we recommend that for conservation purposes, declined trillium be considered together with the hybrid Trillium erectum x flexipes, because most populations contain a mixture of both taxa that cannot reliably be distinguished, and because in the case of this species complex, the conservation target is the unique, much less commonly distributed Trillium flexipes genetic material.

Trillium Erectum group Taxonomy, Hybridization and Conservation Implications The following summary reviews current research assessing the degree to which declined trillium is taxonomically distinct, and also addresses the implications of hybridization in this group for conservation policies.

Taxonomic evidence from Biogeography and Morphology The three Erectum group Trillium species that occur in Pennsylvania have traditionally been classified as distinct species because of morphological differences and because they have fairly distinct global ranges with little overlap. However, Pennsylvania is one of the zones of overlap, and the morphological distinctions set forth by Case (2003, 2009) are not entirely consistently observed here.

The North American range of Trillium erectum is predominantly eastern (Kartesz 2015, Error! Reference source not found.), while the North American range of Trillium flexipes is predominantly midwestern (Kartesz 2015); western Pennsylvania is one of the zones of overlap near the edges of both species’ ranges. Two varieties of Trillium erectum are sometimes recognized, one with maroon flowers (Trillium erectum var. erectum), and one with white flowers (Trillium erectum var. album). In Pennsylvania the white-flowered variety is much less extensively distributed than the maroon-flowered variety, and its range is generally congruent with the range of T. flexipes (Pennsylvania Flora Project 2016). However, it is quite common in southwestern Pennsylvania. The maroon-flowered variety is relatively common throughout most of Pennsylvania (Pennsylvania Flora Project 2016).

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Trillium cernuum’s North American range is primarily in the northeast and upper Midwest; these regional populations appear almost disjunct (Kartesz 2015). T. cernuum’s range is also mostly complimentary to the range of T. flexipes, with zones of overlap at the edges. In Pennsylvania, T. cernuum is concentrated in the southeastern portion of the state, which is also the one portion of the state in which T. erectum is uncommon (Pennsylvania Flora Project 2016). Many T. cernuum populations today are quite small, and the threats of deer browse and invasive species are particularly intense in southeastern Pennsylvania. T. cernuum is documented from a few scattered populations in western Pennsylvania, but there are no known extant populations west of Huntingdon County.

Most known populations of T. flexipes show signs of hybridization with T. erectum. The western populations frequently co-occur with T. erectum, while the eastern populations along the Susquehanna do not typically have T. erectum present. Typically, populations contain some individuals that exhibit enough T. flexipes characters to key fairly unambiguously to the species, and other individuals with characters that fall somewhere on a spectrum between what is typical for T. erectum or T. flexipes. In the west, there are also typically individuals that would key to T. erectum, either white flowered or maroon flowered. The populations are usually predominantly white-flowered, with scattered red individuals, or a mixture of patches of both may be present at the site. There is variation in the coloration from pure white or speckled with light pink (both of which are accepted for T. flexipes), to dark maroon (typical of T. erectum); a range of intermediates are usually present with pink, maroon- speckled, or maroon–striped ovary coloration. Anther:filament ratio also varies (4:1 is typical for T. flexipes, while 1:1 is typical for T. erectum), as does ovary shape (pyramidical is typical for T. flexipes while round-oval is typical for T. erectum). In the eastern populations, Trillium erectum is typically red- flowered with possibly a few scattered white individuals, and there is also a distinct “wet dog” odor to the flowers, which is not present in the Trillium flexipes populations.

The “Syngameon” Concept Because the species of the T. erectum group are known to hybridize, there is some debate over whether “species” is the appropriate taxonomic level to recognize them. The term “syngameon,” a group of taxa that hybridize but also have partial reproductive isolation, has been applied to the group (Millam 2006). Although the most conservative definitions of the species concept argue that complete reproductive isolation is necessary to apply species status (Grant), incomplete reproductive isolation is common in the plant world among taxa generally recognized as species (Ereshefsky 1992). The oaks (Quercus) are a notorious example. Ereshefsky states: “The species within a syngameon are often real units in terms of morphology, ecology, genetics, and evolution. For example, the fossil record indicates that balsam poplars and cottonwoods (both from the genus Populus) have been distinct for at least 12 million years and have generated hybrids throughout this period (Eckenwalder 1984). Even though the hybrids are widespread, fertile, and ancient, these tree species have and are maintaining genetic, phenotypic, and ecological cohesion within and distinction between and have maintained themselves as distinct evolutionary lineages for at least 12 million years. Hence, cottonwoods and poplars are real biological units and should not be ignored.”

Taxonomic Evidence from Genetics and Ecology There are several studies that have addressed the taxonomy and evolutionary relationships of the T. erectum group through genetic analysis. A masters’ thesis (Millam 2006) conducted mainly in Tennessee explored the genetics, ecology, and morphology of six of the seven species in the group (T. cernuum’s range is north of the geographic area of the study). This study examined multiple samples of each taxa, but did not find genetic markers that were distinct to individual taxa. However, the study did not use

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Pennsylvania material, and included a very limited sample size of T. flexipes (4 individuals). This author supported the application of the term “syngameon,” based on the lack of physiological barriers to hybridization and the lack of genetic distinctions, but the presence of observed differences in ecology and morphology that suggest some limitations to gene flow between taxa are in place.

Field observations in Pennsylvania anecdotally support this concept. Although the T. flexipes populations almost invariably display some individuals which would key cleanly to T. flexipes, some individuals which have intermediate traits, and some individuals which would key to T. erectum (in the west), they appear ecologically similar to the habitat described for T. flexipes in other literature: rich floodplains and calcareous lower slopes (Fred Case book). T. erectum displays a broader ecological range than the T. flexipes populations, including more acidic soils and a variety of topographic positions. However, some populations of T. erectum where no individuals were observed that would key to T. flexipes do include individuals with traits that deviate from the descriptions provided for T. erectum; an ovary that varies from pyramidical to oval in shape, variation in anther:filament length ratio, and presence in strongly calcareous soils. These traits could represent introgression of T. flexipes genes in the populations in the area of range overlap.

Millam’s thesis documented significant but not complete isolation between plants of red and white flower color based on pollinator behavior. It investigated whether the various taxa had significant association with ecological habitat characteristics, and found some associations for geology type, temperature, and precipitation. There were some distinctions between taxa, especially in regards to T. erectum and T. vaseyi (a southern taxon). It did not find any distinct habitat characteristics associated with T. flexipes. However, the study was limited because it only examined the species in portion of their range, and it only looked at habitat ecology using fairly broad-scale GIS data, which can sometimes oversimplify and/or misrepresent on-the-ground conditions in regards to habitat characteristics.

It is also important to recognize that while the genetic work in Millam’s thesis contributes a new view on the Erectum group, it also has limitations that may affect the universality of its conclusions. It included plant material from only 11 sites, and appears to include DNA from only 4 T. flexipes individuals. The Erectum group is estimated to have diverged 900,000 years ago during the Middle Pleistocene, when the last glacial maximum reached south to the Appalachian mountain range (Millam 2006). However, all study sites except one were within a single Southern Appalachian glacial refugium identified through a study of sessile-leaved (Gonzales et al. 2007). Most species in the group have large portions of their range outside of the sample area; if other glacial refugia existed, they may have contributed genetic diversity that is undocumented by this study.

Conservation for Syngameon Taxa From a conservation perspective, hybridization is often viewed as negative, due to its potential to merge distinct taxa to a point where the individual identity of either may be lost. However, in the context of a syngameon, where partial reproductive isolation exists, natural hybridization may be a source of genetic flexibility without loss of identity. A study of the above-described syngameon within Populus, where distinct taxa have co-existed for thousands of years despite hybridization, argues that introgression (the entry of material from one taxon’s genome into the genome of another taxon) can be shaped by natural selection (Martinsen et al. 2001), and thereby be adaptive over the long-term.

Under this outlook, the taxa within the Erectum group represent genetic diversity that may provide important ecological, morphological, or physiological flexibility to individual populations. Individual taxa appear to exhibit some degree of ecological specialization, such as T. flexipes’ association with higher pH

125 substrate, or T. vaseyi’s association with higher temperatures than T. erectum. There appears to be some degree of pollinator fidelity to either white- or red-flowered individuals. These ecological distinctions may confer partial reproductive isolation among the taxa. At the same time, the possibility of hybridization between the taxa allows these traits to be transferred in the event that ecological or environmental shifts require new adaptations. Millam’s analysis of chloroplast DNA found less genetic diversity within the entire Erectum group than is typically expected to be found within a species. If genetic differences exist between the taxa that are involved in their distinct ecological niches, these may be important to the overall adaptive ability of a group that otherwise has little genetic diversity. In this context, a conservation goal that can preserve this flexibility is to retain existing spatial distribution of genetic variation. In Pennsylvania, since T. erectum and T. flexipes often occur together, yet T. erectum is fairly common while T. flexipes is much less common, populations of T. flexipes or that include morphologically distinct T. flexipes genetic material are the conservation target to reach this goal.

References • Case, Frederick W. Jr. and Roberta B. Case. 2009. Trilliums. Timber Press. • Case, Frederick W. Jr.. 2003. Trillium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 20+ vols. New York and Oxford. Vol. 26. • Ereshefsky, Marc. 1992. The Units of Evolution: Essays on the Nature of Species. MIT Press. • Gonzales, Eva, James L. Hamrick and Shu-Mei Chang. 2008. Identification of glacial refugia in south-eastern North America by phylogeographical analyses of a forest understorey plant, Trillium cuneatum. Journal of Biogeography. Vol 35, number 5. Pg. 844-852. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)]. • Martinsen, Gregory D., Thomas G. Whitham, Richard J. Turek, Paul Keim. 2001. Hybrid Populations Selectively Filter Gene Introgression Between Species. Evolution. Vol 55, number 7. Pg. 1325-1335. • Millam, Kendra C. 2006. The Trillium Erectum Species-complex (): Insights from Molecular Systematics and Biogeography. Master’s thesis, University of Wisconsin. Pennsylvania Flora Project, Morris Arboretum, University of Pennsylvania. www.paflora.org.

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Appalachian Blue Violet – Viola appalachiensis Henry Author: Loree Speedy, Carnegie Museum of Natural History. Edited: Jessica McPherson, PNHP

Introduction The Appalachian blue violet is a low-growing herb with perennial stems that lay flat and allow the plant to spread and form a distinct mat-forming habit. Flowers are pale violet and the leaves are wide and heart-shaped with shallow, rounded teeth (Pennsylvania Natural Heritage Program, 2015). At one time Appalachian blue violet was thought to be a variety of Walter’s violet (, known from the southeastern U.S.) or a hybrid, but studies by Ballard and Wujek in 1994 demonstrated that it is in fact a distinct species.

Proposed PA Status The current recommendation of the PABS-Vascular Plant Technical Committee is for a status of Pennsylvania Rare for the Appalachian blue violet. The number of known extant populations and individuals is consistent with Pennsylvania Rare, but there are almost certainly more populations not yet documented, and the species also appears to be under little threat because it can thrive in disturbed and anthropogenic early successional habitats. These factors indicate consideration of delisting Appalachian blue violet; however, the VPTC recommendation is made out of concern about its limited global range and potential that it may be a species for which Pennsylvania contains a large fraction of all existing populations globally.

North American Distribution and Global / Regional Conservation Concerns The species is currently known from Pennsylvania, West Virginia, Maryland, and North Carolina. It appears that the species is a central-southern Appalachian endemic with a limited global distribution. Although the species is somewhat under-documented, and better documentation may fill some holes in the global range, it is unlikely to change dramatically. The specimen record provides fairly good evidence for the global distribution; Viola is well-represented within herbarium collections, and experts have reviewed regional collections.

Pennsylvania Distribution The northern edge of this species’ known range falls in Pennsylvania. Within the state, the distribution is limited, in two disjunct clusters. Most populations are found in southwestern Pennsylvania, in Somerset, Fayette, Westmoreland, and Cambria counties. A few populations are known from the northwest in Erie and Warren counties; there is one historical record from Tioga County.

Habitat Habitat records throughout this global range show that it has been found in rich mesic forests, serpentine barrens, and riverbank levees, but additional documentation shows it may favor moderately disturbed areas such as old road beds through coves (Weakley, 2012), old fields, and ground that is regularly mowed. Documentation of habitat in Pennsylvania shows that it is a generalist plant species able to thrive in a variety of forested edge and somewhat open habitats that are common in the Allegheny Mountain counties of Somerset and Cambria. The species can appear on forested floodplains, terraces, and moderate slopes, in a variety of successional stages, from recently logged to mature. It appears to occur in smaller numbers in less-disturbed mid-successional to mature habitats, indicating that it may not tolerate shaded conditions of these areas.

The species does appears to favor moderately disturbed open areas; it has been found in high numbers in grassy areas in parks and picnic areas, dirt road edges, sewer line placement areas, powerline rights-

127 of-way, trailsides, and lawns not subject to high-maintenance practices. It is also known from abandoned pastures. Because it effectively colonizes these human-disturbed areas, Appalachian blue violet may be more common today than it was before human settlement, increasing in population numbers due to the available habitat both generated and maintained by anthropogenic processes (Grund and Isaac 2007, NatureServe, 2014).

Appalachian blue violet is also thought to be overlooked (NatureServe, 2015). The species is difficult to identify for those not familiar with the species and may often be mistaken for the more common dog violet (V. labradorica), especially when growing in parkland or ruderal habitats.

Extant Locations There are 39 extant locations in Pennsylvania. Over thirty have been documented since 2000, increasing both the known range and the proportion of the range that is occupied. Recently documented occurrences are in areas that have not been adequately botanized, so are likely long existing occurrences.

Fieldwork on Appalachian blue violet in 2014 focused on moderately disturbed habitats such as fishing access areas, local community parks geared toward picnicking, and state forest roadside and picnic areas. The species was documented in eleven new sites in Somerset and Cambria counties. The fairly high success rate in surveying this type of habitat, which is fairly abundant and by no means comprehensively surveyed, suggests that it is likely additional populations exist in these counties.

Fieldwork in 2016 focused on the area between the two known population clusters, attempting to investigate whether the species’ range in Pennsylvania may in fact be contiguous between them. Similar types of habitats to the 2014 surveys were considered. However, no new additional populations were located, in 36 site surveys across Clearfield, Cameron, Potter, McKean, Warren, and Crawford counties. Because the area is so extensive and the potential habitat generic and abundant, the lack of positive results cannot be taken to signify conclusively that the species is not present.

Historical Locations and Potential for Relocation At this point, few additional populations are likely to be added by attempting to relocate historical records, because most have been searched and a fairly small number were documented to begin with. Of the 11 historical locations known in Pennsylvania, 3 were deemed low priority for survey because extant populations are already known within 1 mile of their mapped location. Six (6) historical records were determined to have sufficiently precise locational data with current available habitat to make attempts to relocate. Two (2) of these locations were surveyed in 2014 and plants were relocated successfully; 1 location (the only known northcentral PA record, collected from Potter County in 1926) was unsuccessfully searched in 2016. Three locations have yet to be surveyed. The 2 remaining historical records had locational data too vague to search effectively.

Threats Surveys conducted in 2014 located fairly large and apparently healthy populations of the Appalachian blue violet in semi-maintained and anthropogenic early successional habitats. These results indicate Appalachian blue violet may adapt to, and respond positively to, a wide range of natural and disturbed conditions, and is not in danger of population decline in Pennsylvania (NatureServe, 2014). It may even be expanding its range in Pennsylvania because changes in land use since European colonization (residential development, agricultural succession, and forestry) have made more habitat available. Therefore, this species may not require protection to continue to thrive in Pennsylvania.

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However, questions remain about the species’ distribution and abundance in Pennsylvania and in the rest of its limited global range. Most importantly, without updated information and more survey data from adjacent regions, it is impossible to assess the degree of responsibility Pennsylvania holds for the species’ global conservation, i.e., how large a fraction the Pennsylvania populations represent of all known populations. The question of how extensive or limited the range is within the state is also not yet conclusively answered, and will be difficult to answer due to the large amount of potential habitat.

Status Justification The lead investigator of this species in the plant status project, Loree Speedy, supports the position that in spite of its restricted global range, there is evidence to support that the species is not in danger of range or population reduction, and it may be feasible to remove it from the Chapter 45 list of classified plants; and alternatively, it may be retained on the Chapter 45 list of Tentatively Undetermined plants until its distribution and abundance in Maryland, West Virginia, and North Carolina is better understood.

In its most recent consideration of the species in 2015, the PABS-Vascular Plant Technical Committee voted to recommend a status of Pennsylvania Rare, acknowledging new information about the abundance of the species in PA but also concern about removing all legal protections before the degree to which it may be a responsibility species for the state is understood.

When the updated information about Appalachian blue violet was entered in the NatureServe Rank Calculator Version 3.185, it received a ranking of S4, Apparently Secure in the Commonwealth of Pennsylvania. However, this assessment does not include concerns about global rank and state responsibility.

References • Ballard, Harvey E and Daniel E. Wujek. 1994. Evidence for the Recognition of Viola appalachiensis. Systematic Botany. 19(4): 523-38. • NatureServe. March 2014. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1. NatureServe, Arlington, Virginia. (Accessed 12 January 2015). • NatureServe. 2015. NatureServe Conservation Status Assessments: Rank Calculator Version 3.185. NatureServe, Arlington, VA. Online at http://connect.natureserve.org/publications/StatusAssess_RankCalculator • Pennsylvania Natural Heritage Program. 2015. (Accessed 11 February 2015) • Rhoads, A.F. and T.A. Block. 2000. The Plants of Pennsylvania: An Illustrated Manual. University of Pennsylvania Press, Philadelphia. • Weakley, A.S. 2012. Flora of the Carolinas, Virginia, and Georgia [working draft of November 2012]. University of North Carolina Herbarium, Chapel Hill, North Carolina. Available online at http://www.herbarium.unc.edu/flora.htm. Accessed 13 January 2015. • United States Department of Agriculture - Natural Resources Conservation Service. 2005. The PLANTS Database [web application]. National Plant Data Center, Baton Rouge, Louisiana. Available at http://plants.usda.gov. Accessed 13 January 2005.

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Netted Chain Fern – Woodwardia areolata (L.) T.Moore Author: Dr. Timothy Block, Morris Arboretum; edited Jessica McPherson, PNHP

Introduction Netted chain fern is strongly dimorphic, producing markedly different vegetative and fertile fronds. The plant is also strongly rhizomatous, often forming large colonies that are genetically a single individual. Sterile specimens of netted chain fern are sometimes confused with the more common sensitive fern ().

Proposed PA Status Netted chain fern is proposed to move from unlisted (N) to Pennsylvania Rare (PR) due to its limited distribution in the state and the moderate number of populations known to exist in the state.

North American Distribution and Global / Regional Conservation Concerns The North American distribution of netted chain fern (Woodwardia areolata) is primarily on the Atlantic Coastal Plain, from East Texas to Florida and north to Maine, but extending inland particularly in states south of Pennsylvania (Kartesz 2015). Netted chain fern is listed with a NatureServe rank of secure (S5) in New Jersey, Delaware, Virginia, and North Carolina, and is probably secure in Coastal Plain habitat throughout the southeastern U.S. It is of conservation concern in the states along the northern edge of its range (Imperiled-S2 in Missouri, Illinois, Indiana, and West Virginia; Vulnerable-S3 in Ohio and New York). It is assigned a global rank of Secure (G5).

Pennsylvania Distribution While a netted chain fern is a common species on much of the Atlantic Coastal Plain, its distribution in Pennsylvania has historically been limited by the rarity of Coastal Plain habitat in our state. However, in recent years an increasing number of populations have been found outside of the Coastal Plain, in the southcentral Ridge and Valley and in Greene and Fayette counties.

Habitat Netted chain fern occurs in wetlands and is restricted to fairly acidic soils, never occurring on limestone or other high pH substrates.

Extant Locations In total, there are 37 known extant populations of netted chain fern; slightly more than half are outside of the Coastal Plain.

The largest known populations of this species are in Bucks County at the Delhaas Woods section of the Silver Lake Nature Center, a Bucks County park. Several smaller populations exist in Coastal Plain remnant forest sites of about five acres or less in size. Many of the newly discovered populations outside of the Coastal Plain are small, suggesting that they may have established recently; they may consist of a single genetic clone.

Historical Locations and Potential for Relocation Many historical populations in Bucks, Delaware, Montgomery, and Philadelphia counties no longer exist due to development of Coastal Plain habitat. Most historical specimens known in the state are from this region, and further searching for these is not likely to result in discovery of many new populations. There are very few historical locations in central or western PA, where new populations have recently been documented.

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Threats The rate of development on the Coastal Plain remains high, potentially threatening populations located in that region of the state. The remaining areas of natural habitat in the Coastal Plain are also small and fragmented, with questionable long-term viability due to threats from edge effects, invasive species, and disturbance from human traffic and recreational activities. Outside of the Coastal Plain, populations face a lower level of threat, mainly from invasive species and various sources of development. The genetic structure of populations in Pennsylvania may also create long-term vulnerability for the species; although a few large populations exist, it is likely that these occurrences represent only a few genetically distinct individuals, the upright leaves arising from a branching, below-ground stem system. Many small populations may consist of a single genetic individual. This lack of genetic diversity and the physical connectivity of clones can leave the population vulnerable to rapid decline or extirpation when faced with disease or environmental changes, in spite of its robust above-ground appearance.

Status Justification Given the moderate number of populations now known in the state, and the apparent expansion of the species in recent years outside of its historically known range in the Coastal Plain, Pennsylvania Rare is recommended as the appropriate status for netted chain fern at this time. The ongoing threats and limited number of known populations, combined with apparent expansion into habitat in moderately rather than highly threatened regions of the state are consistent with the level of conservation concern in the definition of Pennsylvania Rare. The Pennsylvania Natural Heritage Program, a member program of NatureServe, has assigned a rank of S3 (Vulnerable) in the state, based on analysis of the available data on the species with the NatureServe Rank Calculator tool (Faber-Langendoen et al. 2012).

References • Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe Conservation Status Assessments: Methodology for Assigning Ranks. NatureServe, Arlington, VA. • Kartesz, J.T., The Biota of North America Program (BONAP). 2015. North American Plant Atlas. (http://bonap.net/napa). Chapel Hill, N.C. [maps generated from Kartesz, J.T. 2015. Floristic Synthesis of North America, Version 1.0. Biota of North America Program (BONAP). (in press)].

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Appendix A: Annotated Species List Scientific Name Notes Lead Partner Ageratina Dedicated fieldwork not needed, sufficient information available to provide Morris aromatica justification of current status. Geographically restricted to SE PA. Not historically Arboretum restricted to serpentine; but today all known occurrences are on serpentine geology. Revisiting historicals is unlikelty to yield much because it's successional, also historicals in SE PA are highly problematic to relocate due to vagueness, redundancy in collection, and high rate of habitat conversion. Amelanchier This species is listed as Amelanchier canadensis var. canadensis in the recent FNA Morris canadensis revision. Arboretum Amelanchier No taxonomic changes in the recent FNA treatment. This species needs additional Morris humilis herbarium work; 19 extants is too many for PE status, but since many A. humilis Arboretum records were reidentified to A. sanguinea, it is possible records are still in there as extant A. humilis that shouldn't be. Amelanchier This species is listed as Amelanchier canadensis var. obovalis in the recent FNA Morris obovalis treatment. Arboretum Amelanchier No taxonomic changes in recent FNA treatment. Morris sanguinea Arboretum Ammannia This species’ occurrence and distribution in Pennsylvania is likely the result of Morris coccinea human mediated events rather than more natural causes. As Ammannia coccinea is Arboretum an abundant, often weedy, plant of highly disturbed habitats throughout its range, and was not present as a naturalized part of the Pennsylvania flora until some time in the 1960s (perhaps later), it seems unreasonable to consider this species as strictly native to Pennsylvania, let alone to consider it as a species of special concern. Asclepias Fieldwork completed 2014. Current status of PT considered appropriate. PNHP verticillata Astragalus Considered at 2014 RPF; status changed from TU to PE. Although this is a PNHP canadensis disturbance-adapted species, it appears to have always been somewhat limited in the state and to have become more so. Lots of surveys have been conducted in appropriate habitat yet very few extants are known. Many appear to have been lost to succession, and not replaced by new populations. Bartonia Will be proposed at 2017 RPF/VPTC meeting PNHP paniculata Carex ormostachya This species was changed to TU at the 2015 Rare Plant Forum. Although only one PNHP extant population is known in the state despite fairly extensive survey effort to relocate historical populations, and it is at the southern edge of its range in Pennsylvania, the habitat is generic upland forest that is very extensive in the state. While available evidence suggests this species is likely to be very uncommon in the state, survey effort cannot be considered comprehensive because potential habitat is so vast. Carex shortiana Fieldwork completed in 2014. Results support the current status, PR. Habitat: Carnegie marshy old fields, floodplain woods, in a high quality swamp; a calciphile. John Museum Kunsman has seen about 10 occurences. Tend to be small populations though. Castilleja coccinea Most historicals have been visited. Successional; seems to get outcompeted very Morris quickly, especially with taller stuff. Small populations. Diabase / limestone. Also Arboretum likes seasonally wet. Chaerophyllum Further fieldwork supports current VPTC recommendation of PE, despite more PNHP procumbens var. survey work there are still very few populations known. shortii

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Scientific Name Notes Lead Partner Chenopodium During the time of widespread fire after timbering, this species gained a foothold in PNHP capitatum northern PA. Now it has disappeared due to fire succession. Going back to historicals is very unlikely to suceed, due to the early-successional nature of the species and the dramatic transformation of the landscape in the intervening years. Crataegus dilatata There is now a revision in FNA - account included here addressing Crataegus as a PNHP, Morris genus explains what would need to be done to sort these out, if even possible.

Crataegus There is now a revision in FNA – account included here addressing Crataegus as a PNHP, Morris pennsylvanica genus explains what would need to be done to sort these out, if even possible. Cuscuta corylii This species has a drier and more distinctive habitat than many Cuscuta spp., and PNHP also has not been seen in many years. Cuscuta This has a more typical riverbank habitat. Anne Rhoads and Timothy Block looked PNHP polygonorum for it in their study of Cuscuta, and did not find a lot. Can defend PT at this point reasonably well with the argument there might be more, but not a huge amount more. Historicals are useless. Cystopteris We think the taxonomy is done and the proposal is solid as it is. PNHP laurentiana

Dichanthelium Fieldwork completed 2014. Current status of PT considered appropriate. PNHP oligosanthes Dryopteris celsa This species is historically known from SE PA, but now only 2 small extant Morris populations in that region, while 4 extant populations have been found since 1995 in Arboretum the NW. Not sure if it is restricted to acidic habitats in PA, does not appear to be in NW. Dryopteris Most historicals have been searched for, populations are not large; can defend PT PNHP clintoniana for this. Elodea schweinitzii This is a very poorly documented taxon that is only represented by 2 historical Morris specimens in NY and 1 in PA; it is either not a good species, or if it is, it appears to be Arboretum extinct.

Elymus John Kunsman wants to move to Watch List; he feels that because since it can take PNHP trachycaulus advantage of successional habitats (pipelines, edges of fields, wooded roadsides), it seems likely sufficient numbers will eventually be found and it doesn't need help through land conservation efforts. However, current numbers are low for delisting, and it may be vulnerable to climate change because it is a northern species and mainly occurs in NEPA. Wait a few years and keep an eye out for it, see if there are more discoveries in the course of other work. Erythronium Very short window in early spring when it can be distinguished from E. americanum PNHP albidum may have resulted in underdocumentation in the past. Fieldwork completed in 2014; considered at 2015 Rare Plant Forum, status changed to PR.

Eupatorium John Kunsman updated our biggest site this year; decimated by roadside spraying. PNHP godfreyanum Also knew of Fulton site not in database (private land), that was also sprayed. It has a very restricted global range. Selectively weedy; likes bit of opening in woodlands, interface with road and woods. Low numbers plus early successional habitat combine for PT recommendation. Galium trifidum PR is an appropriate status for this species with the number of known extants and PNHP the threats from invasive species and climate change. Likes midrange pH; a bit richer but not a strict fen species. Past comments about "weediness" likely refer to an 133

Scientific Name Notes Lead Partner ability to tolerate some disturbance, and occupy a broader range of habitats than many fen species, including early successional marsh or beaver-impacted wetlands. However, it does not demonstrate an aggressive growth habit, and it is not extremely disturbance tolerant. Fieldwork may continue to add some new sites, but it is not believed it will greatly change its status. Some fieldwork was conducted in the course of Plant Status work 2014-2015, but it was not considered a major priority because it is unlikely to greatly affect the status. Hieracium Only two extants are Presque Isle and Erie bluffs. Some searching for historicals has Morris umbellatum already been completed, unsuccessfully: John Kunsman looked for it in Poconos but Arboretum has never seen it, and John looked for the Centre County records too. Due to the limited number of extants, the limited number of historicals, and the effort already expended, PE is justifiable without further fieldwork. Note that there has been a taxonomic update affecting this taxon's nomenclature: we once listed this as Hieracium kalmii, but now H. kalmii, H. umbellatum, and H. canadense are all combined under the name "H. umbellatum." However, before the name change we had no records of H. umbellatum or H. canadense, so in Pennsylvania, the occurrences previously listed as H. kalmii comprise the total occurrences for the expanded H. umbellatum concept. Itea virginica Nativity not in question, very distinctive species, Ann Rhoads and Tim Block have Morris extensively searched coastal plain. Deer love to eat it. Status is correct. Tim: agrees; Arboretum the one site is native, they have searched for 20 years and found no others. Very defensible. Juncus biflorus Fieldwork completed in 2014. This species was considered at the 2015 Rare Plant Morris forum, status changed from PT to PR. Restricted geology; tends to be on diabase and Arboretum limestone. Early successional; wet and open, but not weedy as in vacant lot. Remains PR despite early successional/disturbed habitat because of habitat restriction, small populations, and some concern about succession-related declines not being matched by new population establishment. Leucothoe Fieldwork completed 2014; considered at 2015 RPF, status changed from PT to PR. Morris racemosa This species is limited by habitat; while present in many coastal plain woodlands, Arboretum this habitat is under heavy threat and mostly in very poor shape. Outside of coastal plain, it is quite uncommon, and often in good habitats. Morus rubra Despite additional fieldwork, only one new location has been found for this species. PNHP Since recent field efforts were unproductive, it is believed at this point that additional fieldwork may reveal a few more locations, but is unlikely to change the status, as known populations are few and generally very small. There is extensive documentation of the current PT status from the proposal adopted in 2013. Review of existing research on this species indicates significant declines in many parts of the range from disease and hybrid introgression. Oxydendrum Fieldwork completed in 2014. This species was considered at the 2015 Rare Plant Carnegie arboreum forum, status changed from PR to Watch List. Museum

Oxypolis rigidior Fieldwork was completed in 2014. This species was considered at the 2015 Rare Carnegie Plant Forum, status changed from PT to PR. Museum

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Scientific Name Notes Lead Partner Parthenium This species not addressed due to time limitations. Typical shale barren species PNHP integrifolium further south; also found on shale road banks though. Easy to see, and not found on any other shale barrens. Look at historicals if they haven't been checked. The northern populations (our only extants) are considered introduced (ask NY about this!); if any southern locations are discovered, they would be considered native. Must be resolved how that should be handled. While sufficient information is available to classify this is PE or PH, which are functionally the same, this species should not be taken through the regulatory process until the question of whether non-native occurences can be handled distinctly from native occurences is resolved. Platanthera ciliaris This species inhabits successional areas; powerline ROWs. Not weedy though. PNHP Collection pressure, deer browse (they love the flowers). Also small populations. So showy that everyone reports it, underdocumentation not a problem. Poa paludigena Despite additional fieldwork that has clarified the status of western PA populations, Cleveland there remains a difference of opinion among eastern and western experts about this Museum of one (Jim Bissell supports PR or delisting, John Kunsman supports PT); also the global Natural status is uncertain, which means the question of PA responsibility cannot be settled. History More information from other states about whether it is truly rare or they just haven’t looked much would be useful; several places have found it once they started looking, resulting in a local rank downgrade. Polygala polygama Fieldwork conducted in 2014; current PABS status is justified. This species appears Cleveland to be disturbance-dependent, and populations have declined due to succession and Museum of competition where disturbance is not maintained. Some historicals were relocated Natural but many were not; small number of extants plus ongoing threat justifies PE. History

Prenanthes Wooded roadbanks; successional but not weedy. Often on serpentine. John PNHP serpentaria Kunsman thinks should be PR. Only recently started tracking this. If we wait a couple of years we might have enough data for PR.

Quercus michauxii Fieldwork completed 2014; no additional sites located, PE status considered Morris appropriate. John Kunsman and Tim Block have no doubt the species is native in PA, Arboretum and do not believe the known sites have any hybridization issues. Ranunculus The ecology and distribution of this species are not fully understood. It occurs over a PNHP ambigens fairly widespread geographic area, but appears to be fairly rare in most of the places it is known from. If it is limited by specialized habitat requirements, it is not clear what these limitations may be; its habitat appears to somewhat to fully open fairly wet areas, with no particular pH constraints. It may be limited by deer browse, as it is a very succulent robust plant. It may also be limited by some combination of succession shading out existing populations and failure of dispersal/establishment for new populations (reasons unknown; maybe deer browse limits fruiting, etc). Due to its requirement of somewhat open areas, historical populations may not persist or may move around as available habitat shifts to due succession or disturbance. Field experience appears to support this idea, as a few historical locations were relocated but more were not. Focused fieldwork to relocate historicals and refine our understanding of habitat did not result in many additional locations. Ranunculus Jack Holt re-discovered this. Steve Grund and others are working on a paper; PNHP hederaceus however, cannot be certain with available information whether it is native. Still gathering information. Ribes lacustre Current status of PE is appropriate; John Kunsman looked for almost all eastern PNHP historicals and couldn't find it; very distinctive for a Ribes; habitat specialist.

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Scientific Name Notes Lead Partner Rosa blanda Cannot assign status because available information suggests it is both native and PNHP cultivated, and currently there is no way to distinguish populations derived from cultivation vs. native ones. Based on global range it would be logical that this species could occur natively here; however, there are very few known extant populations, and all are in very disturbed habitats. Rubus setosus This was a good candidate for fieldwork because historicals had not all been PNHP revisited, and no extant sites had been observed in many years. Fieldwork documented two extant locations; it appears to tolerate edges and disturbed habitats, and more populations may be found with further work. PT is appropriate status with currently available information. Salix myricoides Can defend status for this with available information. Under serious pressure from Cleveland deer browse. Museum of Natural History Salix pedicellaris Can defend PE with available information; habitat has been searched fairly PNHP thoroughly yet only 3 extant sites are known. Sisyrinchium The status of this species remains unresolved, but it is a low priority for fieldwork PNHP montanum under this project. It remains uncertain how much of this species is found in PA, but it appears to occupy disturbed habitats such as grassy road verges or ROWs. Because it prefers generic disturbed habitats that are both extensive and generally very low priority for conservation-oriented botany surveys, it is not a good use of resources to conduct targeted field surveys for this species; better to educate people who are already going to these habitats to keep their eyes open, and see what comes in over time. Stellaria borealis Changed to PT in 2014 because we have visited most available habitat and it remains PNHP uncommon. Thalictrum We are in the zone of geographic overlap between the geographic range of T. PNHP dasycarpum pubescens (east) and T. dasycarpum (Midwest), and some of our plants may be intermediate. There is now only one specimen that is probably (but not certainly) T. dasycarpum from PA, and the site has been checked, unable to relocate the population. PH is appropriate. Trillium cernuum We do not think there is a taxonomic problem, it is usually distinct. John Kunsman PNHP has seen no hybrid populations. Trillium erectum This is addressed under Trillium flexipes; T. flexipes and the hybrid should be PNHP L.× flexipes considered together in terms of status assessment, because most sites include both. Trillium flexipes Fieldwork completed in 2014. Results support the current status, PT. This taxon is PNHP part of the Trillium erectum complex, which includes 6 species that appear to have unusually low genetic difference between them. Most PA populations show evidence of hybridization, but also appear to have some ecological and morphological distinctions from the common T. erectum, by which they can be positively identified as T. flexipes or T. flexipes hybrids. These distinct traits may be of value to the genetic diversity and adaptive capability of the complex and should be protected. The western and eastern populations of this taxon in PA are geographically disjunct, show some morphological distinctions, and may possibly have independent evolutionary origin.

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Scientific Name Notes Lead Partner Viola Fairly extensive fieldwork was conducted in 2014 and 2016 for this species; this Carnegie appalachiensis work has improved our understanding of the species, but not yet fully resolved its Museum status. This species can occupy somewhat disturbed habitat, open semi-maintained lawn; it is also currently documented from disjunct areas in the state (mainly SW, isolated records in NW and NC). It seems likely that it is underdocumented, but field efforts so far have not greatly increased the range within the state. According to currently known records its global distribution appears to be very limited with most populations in PA, but this may also be due to underdocumentation. Efforts are underway to raise the level of awareness and encourage search effort for the species in surrounding states.

Viola renifolia Fieldwork completed 2015. Current status of PE justified. Only one extant location is PNHP known, and available habitat limited because it is a northern species at the edge of its range and a calciphile.

Woodwardia Fieldwork completed 2014. Considered at 2015 RPF, status changed from PT to PR. Morris areolata The known range in PA has been extended from exclusively coastal plain to include a Arboretum number of Ridge and Valley sites, and the expanded area justifies reduced conservation concern. However, PR is still appropriate because of the total number of occurences plus the high threat faced by populations in coastal plain habitat.

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Appendix B: PNHP Field Surveys 2014-2016 for All Plant Status Update Projects Taxa Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Amelanchier humilis F15GLEB3 needs ID review Amelanchier sanguinea F14KUN77 no Amelanchier sanguinea F15GRU50 no Amelanchier sanguinea F14KUN77 update (679) yes Asclepias verticillata F14KUN70 no Asclepias verticillata F14KUN73 new record yes Asclepias verticillata F14KUN74 new record yes Asclepias verticillata F14KUN75 no Asclepias verticillata F14KUN76 new record yes Asclepias verticillata F14MCP45 search for historical record no Asclepias verticillata F14GLE78 new record yes Astragalus canadensis F15GRU31 revisit no Astragalus canadensis F15SCH32 revisit (10700) no Bartonia paniculata F14GLE46 de novo yes Bartonia paniculata F14GLE60 yes Bartonia paniculata F15SCH24 new record yes Bartonia paniculata F15SCH26 no Bartonia paniculata F15SCH27 update yes Bartonia paniculata F15SCH34 no Bartonia paniculata F15SCH35 new record yes Bartonia paniculata F15SCH36 new record yes Bartonia paniculata F15SCH37 new record yes Bartonia paniculata F15SCH38 no Bartonia paniculata F14GLE46 new record yes Bartonia paniculata F14GLE60 new record yes Bartonia paniculata F15GLE57 new record yes Bartonia paniculata F15GLE60 new record yes Bartonia paniculata F15GLE62 new record yes Bartonia paniculata F15SCH27 update yes Bartonia paniculata F16GLE61 new record yes Bartonia paniculata F16GLE59 new record yes Bartonia paniculata F16GLE01 new record yes Bartonia paniculata F15SCH38 update no Carex ormostachya F14GRU24 no Carex ormostachya F14GRU26 no Carex ormostachya F14KUN35 no Carex ormostachya F14KUN36 no Carex ormostachya F14KUN40 no Carex ormostachya F14KUN42 no Carex ormostachya F14KUN43 no Carex ormostachya F14KUN47 no Carex ormostachya F14KUNG1 no Carex ormostachya F14MCP26 search for historical record (8808) no 138

Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Carex ormostachya F14MCP27 search for historical record (17453) no Carex ormostachya F14MCP28 search for historical record (7633) no Carex ormostachya F14MCP29 search for historical record (17452) no Carex shortiana F14GLE10 update (4038) yes Carex shortiana F15MCP04 de novo yes Chaerophyllum F14GRU06 de novo no procumbens var. shortii Chaerophyllum F14GRU07 de novo no procumbens var. shortii Chaerophyllum F14GRU08 de novo no procumbens var. shortii Chaerophyllum F14GRU12 de novo no procumbens var. shortii Chaerophyllum F14GRU14 de novo no procumbens var. shortii Chaerophyllum F14GRU15 de novo no procumbens var. shortii Chaerophyllum F14GRU17 de novo no procumbens var. shortii Chaerophyllum F14GRU18 de novo no procumbens var. shortii Chaerophyllum F14MCP05 de novo no procumbens var. shortii Chaerophyllum F14MCP07 de novo no procumbens var. shortii Chaerophyllum F14MCP08 de novo no procumbens var. shortii Chaerophyllum F14MCP09 de novo no procumbens var. shortii Chaerophyllum F14MCP10 de novo no procumbens var. shortii Chaerophyllum F14MCP12 de novo no procumbens var. shortii Chaerophyllum F14MCP13 de novo no procumbens var. shortii Chaerophyllum F14MCP15 de novo no procumbens var. shortii Chaerophyllum F14MCP16 de novo no procumbens var. shortii Chaerophyllum F14MCP17 de novo no procumbens var. shortii Chaerophyllum F14MCP18 de novo no procumbens var. shortii Chaerophyllum F14MCP19 de novo no procumbens var. shortii Chaerophyllum F14MCP20 de novo no procumbens var. shortii Chaerophyllum F14MCP21 de novo no

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Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) procumbens var. shortii Chaerophyllum F14MCP22 de novo no procumbens var. shortii Chaerophyllum F14MCP23 de novo no procumbens var. shortii Chaerophyllum F14MCP24 de novo no procumbens var. shortii Chaerophyllum F14MCP48 de novo no procumbens var. shortii Chaerophyllum F14MCP49 de novo no procumbens var. shortii Cystopteris laurentiana F14GRU47 update yes Dichanthelium F14KUN70 no oligosanthes Dichanthelium F14KUN73 new record yes oligosanthes Dichanthelium F14KUN74 new record yes oligosanthes Dichanthelium F14KUN75 update (4534) yes oligosanthes Dichanthelium F14KUN76 no oligosanthes Dryopteris celsa F15WOO25 update (680) yes Dryopteris clintoniana F15GRU26 new record yes Dryopteris clintoniana F15WOO60 yes Dryopteris clintoniana F16WOO67 new record yes Elymus trachycaulus F14GLE23 update (18042) yes Elymus trachycaulus F14GRU48 yes Elymus trachycaulus F15GLE50 update (2926) yes Elymus trachycaulus F15GLE81 new record yes Elymus trachycaulus F15GLE84 update (21442) permission denied no Erythronium albidum F14GRU02 update (15761) yes Erythronium albidum F14GRU07 new record yes Erythronium albidum F14GRU06 no Erythronium albidum F14GRU08 no Erythronium albidum F14GRU10 no Erythronium albidum F14GRU12 no Erythronium albidum F14GRU14 no Erythronium albidum F14GRU15 new record yes Erythronium albidum F14GRU16 no Erythronium albidum F14GRU17 no Erythronium albidum F14KUN02 new record yes Erythronium albidum F14KUN03 no Erythronium albidum F14KUN04 no Erythronium albidum F14KUN05 new record yes Erythronium albidum F14KUN06 no 140

Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Erythronium albidum F14KUN15 no Erythronium albidum F14MCP03 no Erythronium albidum F14MCP05 no Erythronium albidum F14MCP07 no Erythronium albidum F14MCP08 no Erythronium albidum F14MCP09 update (25237) yes Erythronium albidum F14MCP10 no Erythronium albidum F14MCP11 no Erythronium albidum F14MCP12 no Erythronium albidum F14MCP13 no Erythronium albidum F14MCP15 no Erythronium albidum F14MCP16 no Erythronium albidum F14MCP17 no Erythronium albidum F14MCP18 no Erythronium albidum F14MCP19 no Erythronium albidum F14MCP20 no Erythronium albidum F14MCP21 no Erythronium albidum F14MCP22 no Erythronium albidum F14MCP23 no Erythronium albidum F14MCP24 no Erythronium albidum F14MCP46 no Erythronium albidum F14MCP47 no Erythronium albidum F14MCP48 no Erythronium albidum F14MCP49 update yes Erythronium albidum F14TRA07 new record yes Erythronium albidum F14WOO09 no Erythronium albidum F14WOO10 update (24613) yes Erythronium albidum F15WOO11 new record yes Erythronium albidum F14WOO12 update (15359) yes Erythronium albidum F14WOO14 no Erythronium albidum F14SCH05 no Eupatorium F13KUN84 yes godfreyanum Galium trifidum F14GRU31 no Galium trifidum F15GLE54 yes Galium trifidum F15GLE53 yes Galium trifidum F16GLE71 update (18263) yes Galium trifidum F16GLE69 yes Galium trifidum F16GLE72 update (15797) yes Galium trifidum F16GLE45 yes Galium trifidum F16GLE46 update (18263) yes Juncus biflorus F16KUN50 new record yes Leucothoe racemosa F14MCP51 de novo yes Leucothoe racemosa F15GLE14 update (5517) yes

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Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Morus rubra F15KUN73 new record yes Morus rubra F15GRU42 search for historical record no Morus rubra F15MCP03 new record yes Morus rubra F15MCP04 searched near historical record no Morus rubra F15MCP05 searched near historical record no Morus rubra F15MCP07 searched near historical record no Morus rubra F15MCP11- floodplain plots, Raccoon Creek SP no 15 Morus rubra F15MCP16- floodplain plots, Raccoon Creek SP no 21 Morus rubra F15MCP26 no Morus rubra F15MCP27 no Morus rubra F15MCP31 no Morus rubra F15MCP34 no Morus rubra F15MCP35 no Morus rubra F15MCP36 no Platanthera ciliaris F14KUN59 update (2308) yes Poa paludigena F14GRU33 update (9693) no Poa paludigena F15GRU09 update (9693) no Poa paludigena F16GLE16 no Poa paludigena F14GRU29 yes Poa paludigena F15KUN14 update (4775) yes Polygala polygama F16GRU57 de novo yes Polygala polygama F15GLEE8 new record yes Polygala polygama F16GLE19 update (23805) yes Polygala polygama F16GRU57 new record yes Prenanthes serpentaria F14KUN52 de novo yes Ranunculus ambigens F15SCH09 update (20217) yes Ranunculus ambigens F15SCH19 update (4804) no Ranunculus ambigens F15SCH22 update (22993) no Ranunculus ambigens F15SCH18 search for historical record (22996) no Ranunculus ambigens F15SCH13 search for historical record (23001) no Ranunculus ambigens F15SCH12 search for historical record (23003) no Ranunculus ambigens F15SCH11 search for historical record (23004) no Ranunculus ambigens F15SCH17 new record yes Ranunculus ambigens F14KUN56 searching for historical record no Ranunculus ambigens F14KUN57 de novo no Ranunculus ambigens F14KUN63 de novo no Ranunculus ambigens F14KUN90 de novo no Rubus setosus F16GLE33 de novo yes Rubus setosus F14KUN41 Search near historical record no Rubus setosus F14KUN48 search near historical record no Rubus setosus F14KUN51 Search near historical record no Rubus setosus F14KUN53 search near historical record no

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Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Rubus setosus F14KUN54 search near of historical record no Rubus setosus F14KUN69 search in near historical record no Rubus setosus F16GLE56 new record yes Salix myricoides F16GRU38 new record yes Salix pedicellaris F16GLE29 update (18256) yes Sisyrinchium montanum F14SCH31 de novo yes Thalictrum dasycarpum F15GRU25 search for historical record (14662) no Trillium flexipes F14MCP24 new record yes Trillium flexipes F14GRU02 no Trillium flexipes F14GRU06 no Trillium flexipes F14GRU07 no Trillium flexipes F14GRU08 no Trillium flexipes F14GRU11 no Trillium flexipes F14GRU12 no Trillium flexipes F14GRU14 no Trillium flexipes F14GRU15 no Trillium flexipes F14GRU17 no Trillium flexipes F14GRU18 no Trillium flexipes F14MCP03 no Trillium flexipes F14MCP05 no Trillium flexipes F14MCP07 no Trillium flexipes F14MCP08 no Trillium flexipes F14MCP09 no Trillium flexipes F14MCP10 no Trillium flexipes F14MCP11 no Trillium flexipes F14MCP12 no Trillium flexipes F14MCP13 no Trillium flexipes F14MCP15 no Trillium flexipes F14MCP16 no Trillium flexipes F14MCP17 no Trillium flexipes F14MCP18 no Trillium flexipes F14MCP19 no Trillium flexipes F14MCP20 no Trillium flexipes F14MCP21 no Trillium flexipes F14MCP22 no Trillium flexipes F14MCP23 no Trillium flexipes F14MCP24 no Trillium flexipes F14MCP46 no Trillium flexipes F14MCP47 no Trillium flexipes F14MCP48 no Trillium flexipes F14MCP49 no Trillium flexipes F14TRA01 no Trillium flexipes F14WOO09 no Trillium flexipes F14WOO10 no 143

Taxon name Reference Survey type (EOID if revisit or Plants found? Code search for historical record) Trillium flexipes F14WOO11 no Trillium flexipes F14WOO12 no Trillium flexipes F14WOO14 no Trillium flexipes F14YEA05 no Trillium x 1 F14MCP04 new record yes Trillium x 1 F14KUN14 new record yes Trillium x 1 F14GRU18 yes Trillium x 1 F14KUN11 new record yes Trillium x 1 F14MCP12 new record yes Trillium x 1 F14MCP13 new record yes Viola appalachiensis F16GRU07 de novo no Viola appalachiensis F16GRU10 de novo no Viola renifolia F14KUN36 no Viola renifolia F14KUN40 no Viola renifolia F14KUN42 no Viola renifolia F14KUN42B no Viola renifolia F14KUN46 no Viola renifolia F14KUN47 no Viola renifolia F14KUN48 no Viola renifolia F14KUN51 no Woodwardia areolata F14KUN57 new record yes Woodwardia areolata F15GLE57 new record yes Woodwardia areolata F16KUN30 new record yes Woodwardia areolata F14KUN56 de novo no Woodwardia areolata F14KUN58 de novo no Woodwardia areolata F14KUN59 de novo no Woodwardia areolata F14KUN63 de novo no Woodwardia areolata F14KUN87 de novo no

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