ABSTRACT
A MORPHOLOGICAL INVESTIGATION OF DICHANTHELIUM SECTION LANUGINOSA (POACEAE)
by Justin R. Thomas
Much debate and confusion has persisted regarding the morphological ranges and limits of those taxa found within section Lanuginosa of the genus Dichanthelium. Phenetic analysis, principal components analysis and pair-wise t-tests of character states were conducted with 285 specimens using 32 morphological characters. An additional examination of 458 herbarium specimens including all relevant type material resulted in the recognition of 16 species and two subspecies. Many of the entities subsumed within the D. acuminatum complex are found to represent distinct species in terms of morphology. Their geographic and habitat distinctions are also discussed. A review of all 160 names associated with the section was conducted and an accurate nomenclature was applied to the group. Dichanthelium lanuginosum is reinstated as distinct from D. acuminatum and two new nomenclatural combinations (D. thermale and D. thermale subsp. sericeum) are presented.
A MORPHOLOGICAL INVESTIGATION OF DICHANTHELIUM
SECTION LANUGINOSA (POACEAE)
A Thesis
Submitted to the
Faculty of Miami University
in partial fulfillment of
the requirements for the degree of
Master of Science
Department of Botany
by
Justin Ray Thomas
Miami University
Oxford, Ohio
2008
Advisor ______Dr. Michael A. Vincent
Reader ______Dr. R. James Hickey
Reader ______Dr. W. Hardy Eshbaugh
©
Justin Ray Thomas
2008
TABLE OF CONTENTS
List of Tables…………………………………………………………………...... iv
List of Figures……………………………………………………………………. v
Acknowledgements………………………………………………………………. vi
Introduction………………………………………………………………………. 1
Materials and Methods…………………………………………………………… 4
Results…………………………………………………………………………….. 6
Discussion…………………………………………………………………………. 8
Taxonomic Treatment…………………………………………………………….. 21
Literature Cited…………………………………………………………………… 62
Appendix A – Characters Sampled……………………………………………….. 87
Appendix B – Histograms...... …...... ………...…………….. 91
Appendix C – Raw Data (Long Ligule Group)…...... ………...…………….. 106
Appendix D – Raw Data (Short Ligule Group)………...... …….……….. 112
Appendix E – Specimens Sampled……………………………………………….. 116
iii
LIST OF TABLES
Table
1. Summary of Past Treatments…………………………………………………. 65
2. Characters Sampled for Analysis……………………………………………... 66
3. Results of T-tests for Long Ligule Group………...... …………………… 67
4. Principal Components Loadings for Long Ligule Group…...... ………… 68
5. Results of T-tests for Short Ligule Group………...... ……………….. 69
6. Principal Components Loadings for Short Ligule Group………...... 70
iv
LIST OF FIGURES
Figure
1. Phenogram of all OTUs...... 71
2. Box plot of ligule lengths...... 72
3. Phenogram of Long Ligule Group………...... …………………………. 73
4. Scatter plot of Principal Components for Long Ligule Group…...... …. 74
5. Scatter plot of Principal Components for D. implicatum and D. lanuginosum. 75
6. Phenogram of Short Ligule Group………...... …………………...… 76
7. Scatter plot of Principal Components for Short Ligule Group...... … 77
8. Illustrations of ligule and pubescence types within section Lanuginosa……… 78
9 - 24. Distribution maps.....…………………………...... ………………..... 79-86
v
ACKNOWLEDGMENTS
I would like to thank Dr. Michael Vincent for his patience, sage advice and technical support throughout the life of this project. I am grateful to Dr. R. James Hickey, Dr. Linda Watson and Dr. W. Hardy Eshbaugh for their technical assistance as well as for shedding light on the opportunities and challenges presented in this project. I would also like to thank Doug Ladd of the Nature Conservancy of Missouri for making a kid from Brumley believe he could be a botanist. Ultimately, I cannot express enough thanks to my wife and partner, Dana Thomas, for her tireless support and encouragement. Without her pure smiles and gentle understanding this would not have been possible.
vi INTRODUCTION
The genus Panicum, sensu lato, is one of the largest genera of New World grasses. Because of its diversity and wide distribution, a taxonomic understanding of this notoriously difficult group is vital to floristic works, monitoring projects, restoration efforts and other areas of conservation concern. In 1910 Hitchcock and Chase published Panicum subgenus Dichanthelium to accommodate those taxa that produce terminal and axillary inflorescences (floral dimorphism), possess winter rosettes in addition to cauline leaves (foliar dimorphism), and rounded rather than pointed spikelets. This is in contrast to members of the subgenus Panicum which only produce terminal inflorescences, lack winter rosettes and generally have pointed spikelets. In addition to these differences, more contemporary studies have demonstrated further distinctions between subgenus Dichanthelium and subgenus Panicum. Brown (1948), Smith and Brown (1973) and Brown and Smith (1975) demonstrated that subgenus Dichanthelium utilize C3 photosynthetic pathways as opposed to subgenus Panicum which are primarily C4 grasses. Boyle (1945) reported that subgenus Dichanthelium has an extremely low incidence of polyploidy (only three out of approximately 100 species) whereas 70-80 percent of subgenus Panicum species are polyploid. Brown et al. (1957) noted that subgenus Dichanthelium has two layers of tunica cells in the shoot apices whereas subgenus Panicum has a single layer.
Given the extent of such differences, Gould (1974) elevated subgenus Dichanthelium to the rank of genus. However, some taxonomists have been hesitant to accept this status change stating an abundance of tropical species that appear to be intermediate between the two genera (Lelong 1986; Webster 1988; Crins 1991; Gleason and Cronquist 1991; Yatskievych 1999). The argument against elevation of Dichanthelium to generic rank was further supported by Morrone and Zuloaga (1991) and Zuloaga et al. (1993a, b) who tested the evidence defending Dichanthelium as a distinct genus. Their results demonstrate intermediates and exceptions regarding foliar and floral dimorphism, ploidy level and C3 versus C4 photosynthesis, especially in Central and South American species.
1 More recent evidence has come from research involving the molecular phylogeny of Panicoideae (Giussani et al. 2001; Aliscioni et al. 2003; Morrone et al. 2008). These studies demonstrate that Panicum s.l. is polyphyletic unless Dichanthelium, among others, is treated as a separate genus. Additionally, many of the presumed intermediate taxa (most notably section Cordovensia) that were used to maintain Dichanthelium as a subgenus of Panicum have been shown to represent distinct lineages (Aliscioni et al. 2003; Morrone et al. 2008). Though neither the molecular nor morphological data offer a completely satisfying solution to the debate, their net effect seems to provide reasonable evidence supporting Dichanthelium as a distinct genus. The evolution of an over- wintering rosette in this group of grasses may have led to the diversification of Dichanthelium into temperate North America (Morrone 1991; Zuloaga et al. 1993b). This foliar dimorphism offers a reasonable synapomorphy for the genus.
Dichanthelium is a new world genus of between 72-109 species (Gould and Clark 1978; Freckmann and Lelong 2003; Hitchcock and Chase 1910), of which 34 are native to North America (Freckmann and Lelong 2003). Members of the genus are perennial grasses that have erect to decumbent stems which range from 5-150 cm long, vary greatly in degrees of pubescence, and often have short, wide leaves relative to other grasses. Dichanthelium, as its name implies, experiences two flowering periods per growing season (Freckmann and Lelong 2003). All taxa in the genus produce a terminal paniculate inflorescence in spring followed by mid-summer inflorescences on axillary branches that continue to bloom through autumn. The spikelets of the vernal inflorescences are chasmogamous, while those of the autumnal inflorescences are occasionally cleistogamous (Zuloaga et al. 1993b). The spikelets of Dichanthelium range from 0.8-5.2 mm in length and typically have morphologically dissimilar glumes, one sterile lemma and a single fertile floret (Freckmann and Lelong 2003).
In 1910, Hitchcock and Chase arranged the taxa of subgenus Dichanthelium into 17 informal groups based on morphological characters. Many authors since have incorrectly used these names as subgenera, sections or sub-sections, but only recently have any groups been validly published as sections within Dichanthelium (Freckman and
2 Lelong 2002). In so doing, Freckmann and Lelong (2002) reduced the 17 informal groups of Hitchcock and Chase (1910, 1951) to 13 sections (Angustifolia, Clandestina, Dichanthelium, Ensifolia, Lancearia, Lanuginosa, Linearifolia, Macrocarpa, Nudicaulia, Pedicellata, Oligosantha, Sphaerocarpa and Strigosa). Section Lanuginosa has the widest range of variation in the genus. Sections Dichanthelium, Lancearia, Angustifolia and Strigosa have the most morphological similarity to section Lanuginosa. The informal groups Lanuginosa, Spreta and Columbiana of Hitchcock and Chase (1910, 1951) (Table 1) were subsumed into section Lanuginosa by the treatment of Freckmann and Lelong (2002). Section Lanuginosa differs from other sections by the following combination of characters: vernal leaves distributed along the stem (not basally disposed), ciliate ligules ranging from 0.2 to 4.7 mm long, pubescent spikelets that are elliptical in outline and primary glumes that are less than half the length of the spikelet.
According to Freckmann and Lelong (2003) section Lanuginosa contains three North American species and 14 subspecies (Table 1). This is a significant reduction in the number of taxa from the treatment of Hitchcock and Chase (1910, 1951), who recognized 33 species and four varieties (Table 1). While no discussion was provided regarding sections, Gould and Clark (1978) recognized three species and twelve varieties within what is now section Lanuginosa (Table 1). Other regional treatments (Braun 1967; Freckmann 1981; Gleason and Cronquist 1991; Hansen and Wunderlin 1988; Lelong 1984; Mohlenbrock 1986; Radford et al. 1968; Steyermark 1963; Strausbaugh and Core 1978; Swink and Wilhelm 1994; Voss 1972; Weishaupt 1968) are based largely on Fernald’s (1950) treatment of those taxa occurring in northeastern North America. Though each of these treatments provides insight into the group, confusion still exists regarding these taxa. Much of the confusion comes from conflicting interpretations and degrees of emphasis on ligule, pubescence and spikelet characters (Fernald 1921). It has been suggested that detailed studies of the morphological boundaries and population- level variation are necessary in order to better understand the species complexes within Dichanthelium (Freckmann 1981; Gould and Clark 1978; Hansen and Wunderlin 1988; Swink and Wilhelm 1994; Yatskievych 1999).
3 Though Shinners (1944) conducted an informal investigation into the “empirically existing discontinuities” of several taxa, a formal analysis of the morphological diversity within section Lanuginosa has never been published. Various splitting and lumping at the levels of species, subspecies and variety have occurred with little or no justification. As stated above, the taxonomic ambiguity of the section primarily rests on the widely varying interpretations of ligule, pubescence and spikelet characters. The present study examines the morphological variation within section Lanuginosa in order to elucidate the natural boundaries of each taxon and to apply an accurate and useful nomenclature. It is important to note that this analysis is not an attempt to explain or demonstrate the phylogenetic relationships of these taxa. Rather, this analysis is based on a morphological species concept.
MATERIALS AND METHODS
A total of 2780 specimens, including type specimens, from throughout the geographical range of section Lanuginosa was examined from several herbaria (F, GH, MIN, MO, MU, NCU, NY, URV, US, WIS, WS). A subset of specimens representing the morphological and geographical range of the section Lanuginosa was chosen. Specimens that lacked ample fertile or vegetative material for sampling were not included in the data set. Specimens were sampled for 104 vegetative characters and 12 floral characters (Appendix A). Each unique specimen served as a separate operational taxonomic unit (OTU) within the data set. These data were then analyzed via histograms (Appendix B) and mean analysis in order to determine which characters might demonstrate significant morphological differences and to eliminate any invariant or unimodal characters. This was also done to reduce the number of measured characters and thereby increase the number of OTUs that could be sampled during the time frame of the study. Using the remaining 32 characters (Table 2), 285 OTUs were sampled (Appendix C). All type specimens that possessed sufficient characters were included in the sampling. In order to maximize efficiency, more specimens were sampled from the notoriously problematic species complexes within section Lanuginosa than from the better understood groups.
4
The data were entered into Microsoft Excel and imported into NT-SYS-pc 2.1 (Exeter Software, Setauket, NY) for phenetic analysis. In NT-SYS, the data were standardized, a similarity matrix was created, distance coefficients were computed and a phenogram was constructed by the sequential, agglomerative, hierarchical, and nested clustering method (SAHN) using the unweighted pair group method with arithmetic mean (UPGMA). This analysis resulted in a phenogram with two very distinct clusters. These clusters are here referred to as the Long Ligule Group and the Short Ligule Group. In order to more thoroughly and adequately investigate the significance of the morphological variation in the analysis as well as elucidate the morphological characters of taxonomic importance, the data were divided into these two groups and analyzed independently for the remainder of the analysis.
Invariant characters were eliminated from the two data sets, leaving 24 characters (Table 2) for the 142 OTUs within the Long Ligule group and 12 characters (Table 2) for the 143 OTUs within the Short Ligule group. Each of the two data sets was then analyzed by the statistical methods used in the phenetic analysis outlined above as well as by principal components analysis (PCA). All data in the PCA were standardized so that each character had an equal contribution to the variance in the analysis. The clusters of OTUs generated by the phenetic analysis and PCA were evaluated for similarity and consistency. These clusters were then analyzed by pair-wise comparisons, using two sample T-tests, in order to investigate significant differences between clusters for each character. A PCA was conducted in order to further investigate the morphological variance of slightly overlapping taxa in the Long Ligule Group.
Based on the results of these data, preliminary groups and their defining morphological characters were determined. The range of character variability found in the data was analyzed by testing the results against an additional 458 additional herbarium specimens from 33 states, 7 Canadian provinces, 4 Central American countries and 3 Caribbean islands.
5 A thorough review of the literature involving Panicum and Dichanthelium was conducted in order to determine the appropriate application of nomenclature within section Lanuginosa. Once an accurate nomenclature was determined, a key was constructed and all 2780 specimens from the original compilation were sorted and annotated.
RESULTS
Of the 116 morphological characters sampled (Appendix A) using histogram and mean analysis, 32 proved informative. The phenetic analysis of all 285 OTUs for all 32 characters (Figure 1) resulted in an obvious split of the data set into two groups here referred to as the Long Ligule Group and the Short Ligule Group. These two groups demonstrated subordinate grouping which matched various past treatments and well represented the type material and associated OTUs. A review of all the characters in the analysis revealed that the length and distribution of ligule hairs were largely responsible for the divison of the OTUs into these two groups (Figure 2). The Long Ligule Group consistently possesses a uniformly long ligule pubescence that is indistinguishable from the pseudoligule and is always 1.7 mm long or longer. The Short Ligule Group primarily possesses a short ligule that is often, but not always, subtended by a longer pseudoligule. When the pseudoligule is short or intermeshed with the ligule, the ligule is often difficult to discern. Regardless of the distribution of ligule pubescence, the first row of ligule hairs is always 1.5 mm long or shorter. Given the strength of the split and its alignment with past treatments within what is now section Lanuginosa and in order to best demonstrate the statistical variation in morphology within the Long Ligule and Short Ligule Groups, each of these two groups were analyzed independently for the remainder of the study. Twenty of the 32 sampled characters were unique to the Long Ligule Group, eight were unique to the Short Ligule Group and four were common to both groups (Table 2).
The phenetic analysis of the Long Ligule Group resulted in nine clusters (Figure 3). Pair-wise comparisons of each cluster using two sample t-tests showed a significant
6 difference of P ≤ 0.05 for no less than nine and as many as 22 of the 24 characters (Table 3). A significant difference of P ≤ 0.005 was found for no less than four and as many as 20 of the characters (Table 3).
Within the Long Ligule Group, the first three principal component axes of the PCA accounted for 72 percent of the variation (Table 4). Nine clusters were separated by the first two principal component axes (Figure 4). Characters involving the length and density of pubescence along leaves and sheaths, as well as spikelet characters, contributed the most variation along the first principal component (PC1). Leaf length and characters involving the pubescence of the inflorescence contribute most to the variation along the second principle component (PC2). Dichanthelium lanuginosum and D. implicatum demonstrated some overlap along PC1 and PC2.
T-tests demonstrated that Dichanthelium lanuginosum differs significantly (P ≤ 0.05) from D. implicatum by 10 of the 24 characters sampled (Table 3). According to the t-tests, maximum leaf length, maximum leaf width and the length of pubescence on the abaxial surface of the terminal vernal leaf are the strongest characters separating D. lanuginosum and D. implicatum. The scatter plot of the first and second principal components (Figure 5) demonstrate little overlap between these two taxa.
The phenetic analysis of the Short Ligule Group resulted in seven clusters (Figure 6). Pair-wise analysis of clusters for each character using two sample t-tests showed a significant difference of P ≤ 0.05 for no less than four and as many as 11 of the 12 characters (Table 5). A significant difference of P ≤ 0.005 was found for no less than three and as many as 11 characters (Table 5).
The first three axes of the PCA account for 78 percent of the variation within the Short Ligule Group (Table 6). The first two principal component axes demonstrate seven clusters (Figure 8). Distribution, orientation and density of pubescence, as well as glume length and characteristics of the ligule contributed the most variation along the first
7 principal component. Ligule length and spikelet length contributed to the most variation along the second principal component.
Each cluster in the phenetic analyses of the Long Ligule and Short Ligule groups contained one or more type specimens. The nomenclature outlined in Figures 3 and 6 was applied to the Long Ligule and Short Ligule groups based on the type specimens contained within each cluster, their published descriptions and a comparison of those types not complete enough to be included in the general sampling. This followed a thorough review of all 160 published names directly associated with these taxa. From the above results, 16 species and two subspecies were found to best represent the morphological diversity found within section Lanuginosa. Lastly, a key utilizing the strongest characters in the analysis was found to well differentiate all taxa when tested against 458 herbarium specimens from throughout the range.
DISCUSSION
An in-depth analysis of the morphological boundaries found within section Lanuginosa resulted in the delineation of 16 species and two subspecies. Based on the phenogram (Figure 3), PCA (Figure 4 and Table 4), a review of specimens and the significance of each character in pairwise comparisons (Table 3), it is determined that the Long Ligule Group consists of nine species and one subspecies. Based on the phenogram (Figure 6), PCA (Figure 7 and Table 6), a review of specimens and the significance of each character in pairwise comparisons (Table 5) it is determined that the Short Ligule Group consists of seven species and one subspecies.
The preliminary analysis of all 285 OTUs for all 32 morphological characters resulted in a phenogram with two very distinct groups (Figure 1). A review of the data showed that this split resulted from a distinct discontinuity in ligule length (Figure 2). These two groups are referred to as the Long Ligule Group and the Short Ligule Group throughout this study. The split also strongly correlates with the presence or absence of a distinct pseudoligule. Those members of the Long Ligule Group do not possess a distinct
8 pseudoligule. All members of the Short Ligule Group possess a pseudoligule that is longer than the ligule. The only exeption to this occurs in glabrate forms of D. columbianum and D. ovale (Short Ligule Group) in which the lack of pubescence on the the adaxial leaf surfaces correlates to the lack of a pseudoligule; with the pseudoligule being associated with the pubescence of the adaxial leaf surface rather than the ligule. This is largely in line with the use of ligule characters in other treatments (Gould and Clark 1978; Hansen and Wunderlin 1988; Hitchcock and Chase 1910, 1951; Pohl 1947; Stone 1911) as a major division between taxa in what is now considered section Lanuginosa. The Long Ligule Group well represents most of the taxa included in the informal groups Lanuginosa and Spreta of Hitchcock and Chase (1910, 1951), while the Short Ligule Group represents all members of Columbiana as well as some members of Lanuginosa. Furthermore, the preliminary phenogram (Figure 1) demonstrates the segregation of OTUs within these two major groups into smaller groups that are well aligned with previous treatments.
Most taxa in the Long Ligule Group were found to be morphologically distinct. While the phenetic analysis (Figure 3) demonstrated considerable separation between D. lanuginosum and D. implicatum, there was some degree of overlap between the two in the scatter plots of the PCA (Figure 4) for the Long Ligule Group. A second PCA involving only D. lanuginosum and D. implicatum (Figure 5) further confirmed that, while these two species are morphological similar, they are distinct. Recent studies (Allen 2001; Naczi 1998; Peterson et al. 2002; Rothrock and Reznicek 2001; Saarela et al. 2003) have used similar levels of distinction for separating closely related species. Furthermore, D. lanuginosum differs from D. implicatum (P ≤ 0.05) by 10 of the 24 characters sampled (Table 3). Maximum leaf length, maximum leaf width and the length of pubescence on the abaxial surface of the terminal vernal leaf are the strongest characters separating these two taxa. Given the distinctness of the majority of specimens and the narrower ecological niche of D. implicatum (mostly restricted to sandy soils) these taxa are here treated as separate species. Past treatments have attempted to separate or lump these taxa based primarily on spikelet length. While the spikelets of D. implicatum average slightly smaller than those of D. lanuginosum (1.52 mm versus 1.59
9 mm), the range of measurements for this character exhibit too much overlap (1.3 – 1.7 mm long for D. implicatum versus 1.3 – 1.9 mm long for D. lanuginosum) to render this a valid taxonomic character.
This analysis failed to capture the full complement of morphological variation found within the D. thermale complex. Specimens that have always been associated with the complex were found in both the Long Ligule and Short Ligule groups. Essentially, the data demonstrated that what is here called D. thermale subsp. thermale clearly possesses a short ligule backed by a longer pseudoligule and is only found along hot springs in northern California. What is here called D. thermale subsp. sericeum possesses a long ligule and only occurs near hot springs and geysers in the Rocky Mountains. However, a few specimens (such as the type for P. ferventicola) from Yellowstone National Park have intermediate ligule lengths and specimens from Banff, Alberta (such as the type for P. ferventicola var. papillosum) have very short and wide leaves. As noted by Hitchcock and Chase (1910), pubescence characters and leaf dimensions for the P. thermale complex are variable across its range. Given this range of variablilty, the D. thermale complex was interpreted more from the raw data and a review of specimens than from the analysis (see notes under D. thermale for details). This resulted in the recognition of D. thermale subsp. themale and subsp. sericeum, which is similar to the treatment of Freckmann and Lelong (2003).
While the PCA, phenetic analysis and t-tests demonstrate strong differences between D. lanuginosum and D. lindheimeri, the application of morphological characters to specimens can be challenging. Much of the separation generated in the analyses above originates from pubescence characters, where D. lindheimeri is mostly glabrous and D. lanuginosum is always pubescent. The two taxa are very similar especially when sparsely pubescent specimens of D. lanuginosum are encountered (see notes under D. lindheimeri for details on distinguishing these taxa). This similarity has led some authors to lump theses taxa or treat one as a subspecific taxon of the other. Because of its glabrosity, D. lindheimeri is more closely linked with D. spretum in the PCA and phenetic analysis. While this may seem wholly artificial, plants from calcareous lake
10 shores in the Great Lakes region that are here treated as D. lindheimeri are morphologically similar to D. spretum. Said plants have the overall stature of D. spretum coupled with pubescence and floral characters more aligned with D. lindheimeri (for details see notes under D. lindheimeri). In the phenetic analysis the more sparsely pubescent specimens of D. lanuginosum sorted together in a branch of D. lanuginosum. These specimens represent what has been called P. tennesseense and P. lindheimeri var. septentrionale. Following a review of these OTUs, as well as numerous additional herbarium specimens, no clear distinction could be found to justify maintaining this cluster as an infraspecific taxon. The phenetic analysis likewise demonstrated other small, seemingly distinct, groups in the D. lanuginosum and D. praecocius branches. Upon investigation of these groups, no justification for subspecific separation could be found, based on morphology.
In the Short Ligule Group, the results of the phenetic analysis (Fig. 6) and PCA (Fig 7 and Table 6) demonstrate the segregation of seven entities that are here interpreted as distinct species. One outlier within the D. columbianum cluster in Figure 7 represents the type specimen of P. scoparioides. This specimen possesses all the traits of typical D. columbianum except for its larger spikelets. This is interpreted as the extreme maximum of the phenotypic plasticity found within D. columbianum rather than a distinct taxon. Gould and Clark (1978) placed P. scoparioides into synonymy under their broad interpretation of D. acuminatum. However, using their key and descriptions, the type specimen better fits their D. sabulorum var. thinium which is here treated as a synonym of D. columbianum (typical D. sabulorum and var. patulum are now under D. portoricense in section Lancearia). Freckmann and Lelong (2003) refer to P. scoparioides as being a sterile hybrid between D. oligosanthes and their broad interpretation of D. acuminatum (which includes D. columbianum of the present treatment). The possibility of P. scoparioides being a hybrid does not seem unreasonable. Given the numerous specimens reviewed throughout the range of D. columbianum it is abundantly clear that the spikelet length demonstrated by P. scoparioides is a rare exception to the otherwise narrow morphological range of D. columbianum for this character.
11
Similarly, one outlier within the D. acuminatum cluster (Figure 7) represents the type specimen of P. comophyllum. It falls outside the main cluster due to its wider leaves and more densely pubescent adaxial leaf surfaces. Hitchcock and Chase (1910) and Gould and Clark (1978) treated it as a synonym of P. acuminatum, while Freckmann and Lelong (2002, 2003) did not include any mention of its placement within the section. Given this, it is here interpreted as being either an aberrant extreme within D. acuminatum or an unrecognized hybrid with another taxon. Either way, it is in no way characteristic of the more uniform morphology expressed by D. acuminatum. The phenetic analysis of the Short Ligule Group demonstrated some small groups within the larger branches of D. acuminatum and D. villosissimum. A detailed investigation of the data pertaining to these branches revealed no strong morphological or geographical associations.
According to the phenetic analysis and PCA, what is here treated as D. subvillosum is closely associated with D. thermale and D. acuminatum. According to the phenetic analysis, D. subvillosum is most similar to D. thermale, but their morphology and habitats differ strikingly. Dichanthelium thermale has a double vestiture of pubescence on the basal and lower leaf sheaths of the vernal culm, shorter pubescence on vernal sheaths and only occurs in moist geothermal soils of the Cascade and Rocky Mountains. Dichanthelium subvillosum has a single vestiture throughout the vernal culm, a longer pubescence on the vernal sheaths and does not occur in the habitat or near the altitudes of D. thermale. The geographical ranges of both D. thermale and D. subvillosum are restricted to northern North America while the range of D. acuminatum is limited to the coastal regions of the southeastern United States, Gulf of Mexico and into South America (for details see notes under D. thermale and D. subvillosum).
Relevance to Past Studies The present study confirms that the treatment of Hitchcock and Chase (1910) and their recognition of 33 species and two varieties within what is now section Lanuginosa over-emphasize the measurable morphological diversity found within the section. The
12 plethora of names associated with the section is demonstrative of the inherent morphological variability within the group, coupled with the slow rates of communication between 19th century botanists as they gradually described the diversity of the New World. Subsequent treatments, such as those of Gould and Clark (1978) and Freckmann and Lelong (2002, 2003), have attempted to narrow the taxonomic limits within the section. Furthermore, regional treatments demonstrate numerous intermediate alternatives to the treatments mentioned above. Because these secondary treatments are too numerous to discuss in detail and because they do not explore the full geographical range of the section, they will only be addressed where pertinent to the present treatment.
The most obvious difference between the current treatment and those of Gould and Clark (1978) and Freckmann and Lelong (2002, 2003) is the circumscription of D. acuminatum. In the treatment of Gould and Clark (1978), 13 of the 15 species in the present treatment are placed into eight varieties under D. acuminatum. They qualify this treatment with the statement “It may be that the taxa retained as varieties do not adequately delimit the recognizable populations of this extremely widely distributed and morphologically variable species”. The treatment of Freckmann and Lelong (2003) places 10 of the 15 species recognized in the present treatment into 10 subspecies of D. acuminatum. They state that morphologically intermediate specimens derived from hybridization and autogamy produce “a reticulate pattern of intergradation between members of the section [Lanuginosa]”. While this could offer a mechanism for intermediate morphologies, the present study failed to find a significant number of intermediate specimens. Of the numerous specimens reviewed during the present treatment, the vast majority were easily placed into the taxa as outlined below. In direct reference to their broad circumscription of D. acuminatum, Freckmann and Lelong (2003) state that “there appears to be widespread introgression from other Dichanthelium species, such as D. dichotomum, D. sphaerocarpon, D. ovale and D. aciculare”. While such introgression may exist, the present study did not find it to be “widespread”. Rather, the few exceptions were no more common than one would expect from other complex genera such as Carex or Quercus.
13 Gould and Clark (1978) initiated, and Freckmann and Lelong (2002, 2003) maintained, a broad interpretation of D. ovale, though the two treatments differ significantly. Gould and Clark (1978) placed what is here treated as D. commonsianum as a variety of D. ovale (var. addisonii, which is placed in synonymy with D. columbianum in the present treatment) and maintained the typical variety to represent what had traditionally been D. ovale. The basis of that treatment, as gleaned from their keys, seems to be the presence of a double vestiture on the vernal sheaths, longer spikelets and the distribution of pubescence along the adaxial surface of vernal leaves. While the present study failed to confirm any differences in adaxial leaf pubescence or spikelet length, the presence and extent of a double vestiture along the vernal sheaths did prove informative. Freckmann and Lelong (2002, 2003) proposed an even broader interpretation of D. ovale that includes D. villosissimum and D. praecocius (as subsp. villosissimum and praecocius) and incorporated D. commonsianum into subsp. pseudopubescens (see notes under D. ovale for details). In their treatment of D. ovale, Freckmann and Lelong (2002, 2003) and Lelong (1984) are the only authors to combine taxa from the informal groups Columbiana and Lanuginosa of Hitchcock and Chase (1910).
Like the present treatment, Freckmann and Lelong (2003) maintained D. wrightianum as a distinct species. However, Gould and Clark (1978) chose to place D. wrightianum as a variety of D. acuminatum. Given its diminutive stature and consistently tiny spikelets (0.8 to 1.0 mm long), there should be little debate regarding the distinct nature of this taxon.
While the treatments of Gould and Clark (1978) and Freckmann and Lelong (2002, 2003) do provide infraspecific recognition for some taxa here treated as species, such placement does not well represent the morphological, ecological or geographical diversity of the section. Their broad interpretation of D. acuminatum renders the defining characters to ambiguities such as spikelets 0.7-3mm long, plants glabrous or puberulent to villose, and ligules 0.3-5.0 mm long. Additionally, this broad interpretation fails to capture the ecological amplitude, habitat fidelity and geographical distinctness of the taxa
14 involved. If the broad interpretation of D. acuminatum is maintained, it would constitute one of the most ecologically variable and morphologically plastic species known to science. Such an inclusive treatment also warrants concern regarding the conservation status of the entities involved. When a species as habitat-specific and morphologically distinct as D. thermale is placed under a broad concept of D. acuminatum, its conservation status could be lost. Furthermore, otherwise newly discovered populations of rare taxa may remain unreported if they are subsumed under more common entities.
Morphological Themes within the Section Though all of the taxa in the present study are maintained as members of section Lanuginosa, there appear to be some morphological themes and species groups, within the section, worthy of note. These groups are reminiscent of the informal groups of Hitchcock and Chase (1910, 1951). Dichanthelium ovale, D. columbianum and D. commonsianum are the only taxa to exhibit a restriction of pubescence to near the margins of the adaxial surfaces of vernal leaves (central portions glabrate). They also exhibit a double vestiture along vernal sheaths with puberulent pubescence often below a pilose or villose pubescence and ligules that are usually composed of intermeshed hairs of varying length. Dichanthelium lanuginosum, D. lindheimeri, D. praecocius and D. implicatum appear to form an association based on an overall similarity in size and proportions, a similar range of spikelet lengths, ligules uniformly longer than in other taxa and less fidelity to acidic soils. Dichanthelium acuminatum, D. villosissimum, D. thermale and D. subvillosum seem to form a natural group based on their uniformly short ligules, similar spikelet length, lack of double vestiture on sheaths, and densely pubescent vernal culms. Dichanthelium wrightianum, D. leucothrix, D. longiligulatum and D. spretum also exhibit morphological similarities in terms spikelet shape, habitat requirements, and overall stature. Between each of these groups there exists too much overlap to justify any taxonomic separation from section Lanuginosa as it is currently defined. Enlisting the techniques of molecular taxonomy for a further study of the section would prove to be interesting and informative.
Sources of Confusion
15 Much of the past confusion within section Lanuginosa originates from the varying interpretations of taxonomically useful characters. Most treatments have relied heavily on spikelet length, the length of the ligule, and characters involving the indument. The present study has also found that these traits provide clarity, but only for certain species and in conjunction with other characters. Furthermore, when applied to the section, characters like spikelet length, double ligule and double vestiture require a more detailed characterization than most treatments provide. Spikelet length in the present treatment is defined as the length of the spikelet from the base of the first glume to the tip of the second glume. Where possible the most mature spikelets of the vernal inflorescence should be used. Immature spikelets and those of the autumnal inflorescence are often highly variable can result in consistent measurements.
Characterization of the diverse expressions in the ligule length, shape, and uniformity can also be problematic. Such taxa as D. lanuginosum and D. implicatum possess a “single ligule”, which means there is no pseudoligule or that the hairs of the ligule and the pseudoligule are of the same length and thereby not discriminable (Fig 8a). This is in contrast to D. ovale and D. columbianum which often have a “double ligule” or a distinct pseudoligule, where the true ligule is composed of short hairs and the pseudoligule is composed of longer hairs behind the ligule. The hairs of the pseudoligule often originate from the immediate adaxial leaf surface (Figure 8b). Adding to the confusion, some taxa that typically have a longer pseudoligule occasionally lack it and thus appear to possess a “single” ligule (Figure 8c). In herbarium specimens, distinguishing the ligule and pseudoligule can often be problematic because the ligule and pseudoligule (when both are present) may become intermeshed during the drying process (Figure 8d). One would think this a simple matter, but several of the type specimens reviewed for this project possessed a ligule type that differs significantly from that in the author’s original description (included ligule and pseudoligule in original measurements for “ligule”, for example). In the present treatment, the key to taxa breaks from this traditional terminology by referring only to the first row of ligule hairs. This avoids the confusion associated with ligule and pseudoligule and focuses attention on the relevant issue of whether the immediate hairs (first row) are long or short.
16
Another character state that has caused confusion in the group is the presence or absence of a “double vestiture” in reference to sheaths, internodes, peduncles and the axes of the inflorescences. This is problematic because two types of double vestiture are possible for taxa in section Lanuginosa. First, taxa that typically have a single vestiture of long pilose hairs (Figure 8a) can occasionally have an additional indument of short pilose hairs (Figure 8b). Second, some taxa have long pilose hairs above short puberulent hairs (Figure 8d). In the latter case, some specimens can lack the longer pilosity (Figure 8c) and thus do not truly have a “double vestiture”. All previous treatments that have utilized the "double vestiture" character to distiguish taxa have failed to define these subtlties. To avoid this confusion in the present treatment, any reference to “double vestiture” includes a description of whether the subordinate pubescence is pilose or puberulent. Because the distinguishing features of the ligule, spikelet and sheath pubescence can be minute, a minimum of 30X magnification is required to fully discern the differences in taxa. Furthermore, all measurements regarding these characters should be taken with calipers or a sub-millimeter scale while under a dissecting scope.
Because the morphology of the autumnal branches and inflorescences often differ strikingly from that of the vernal culms and inflorescences, all key characters in the present treatment are in reference to the vernal culm. The autumnal branches, leaves and inflorescences of most taxa in the section become highly fascicled and finely pubescent late in the summer. This is often accompanied by the senescence of the vernal stem leaves. Thus, plants collected late in the season become increasingly difficult to identify. As the collector of Carex must be mindful of the shattering of perigynia from the spike, the collector of Dichanthelium should seek specimens that possess fresh components of the vernal stem and thus the full complement of morphological expression.
Currently there are 160 names associated with section Lanuginosa. Such lengthy nomenclature has undoubtedly added an element of confusion to the section. The most problematic source of confusion in the section is the relationship of D. lanuginosum and D. acuminatum. Much of this confusion seems to stem from the regional misapplication
17 of names. Dichanthelium acuminatum, long considered a species of the Caribbean, was not included in the flora of North America until the treatment of Gould and Clark (1978), though it certainly occurred throughout the southern coastal plain. The name Panicum lanuginosum (now referred to as Dichanthelium lanuginosum) was often misapplied to these coastal plants (Hitchcock and Chase 1910; Small 1903, 1933). The name P. lanuginosum was also being applied, and correctly so, to plants of the interior of North America. In 1910, Hitchcock and Chase restricted the name P. lanuginosum to plants of the southern coastal plain and relegated the plants of the interior to P. huachucae. Finding little to no difference between specimens of P. huachucae and earlier published names such as P. lindheimeri and P. implicatum (Dichanthelium lindheimeri and D. implicatum of the present treatment), the regional treatments of Fernald (1922), Pohl (1947), Shinners (1944), Voss (1972) and Swink and Wilhelm (1994) utilized these earlier names for plants in their respective regions (since P. lanuginosum was considered to be restricted to the coastal plain). Fernald (1934, 1950), attempted to correct this discreprency and realigned these taxa with P. lanuginosum by placing P. lindheimeri, P. implicatum, as well as other species, as varieties of P. lanuginosum while restricting the typical variety to the southern coastal plain. In Fernald’s realignment, P. lanuginosum var. fasciculatum (of which P. huachucae is a synonym) came to represent the most wide-ranging taxon of the section in the interior of North America. In 1978, Gould and Clark became the first authors to include D. acuminatum into the flora of North America. However, instead of maintaining D. lanuginosum as distinct from D. acuminatum (as does the present treatment) they placed it, all the above mentioned taxa, plus several others, into synonymy with D. acuminatum. A similar arrangement was followed by Freckmann and Lelong (2002, 2003).
While the inclusion of D. acuminatum in the flora of North America helped clarify this issue, the placement of D. lanuginosum in synonymy with D. acuminatum does not seem justified. In fact, it only adds more confusion and conflict. The persisting confusion between D. lanuginosum and D. acuminatum hinges on the subtle characters of the type specimen of D. lanuginosum. Though the said type specimen is velvety pubescent like D. acuminatum, it lacks the short ligule and ciliate leaf margins that define
18 D. acuminatum. Since the type specimen of P. lanuginosum has a longer and uniformly single ligule and lacks extensive marginal cilia on the vernal leaves, separating it from D. acuminatum adds clarity to the section. In this case it becomes better aligned with P. huachucae of Hitchcock and Chase (1910, 1951), P. lindheimeri of Farwell (1928) and Fernald (1922), P. implicatum of Pohl (1947), Shinners (1944) and Voss (1972), and P. lanuginosum var. fasciculatum of Fernald (1934, 1950) and Gleason and Cronquist (1991). Though the type specimen of P. lanuginosum represents a more velvety pubescent variation found along the coastal plain of North America, it does not differ significantly from those specimens of the interior. Should future studies find a strong difference between the coastal and interior manifestation of P. lanuginosum, those of the interior would be relegated either to Fernald’s var. fasciculatum or the species “huachucae”.
Confusion has also arisen from taxa outside section Lanuginosa. Specimens of D. consanguineum (section Angustifolia) and D. laxiflorum (section Strigosa) are commonly encountered in the herbarium folders of taxa from section Lanuginosa as misidentifications. Dichanthelium consanguineum differs from all members of section Lanuginosa in having attenuate spikelets and thin primary glumes. Because it often has a light puberulence on the culm, it is often confused with D. ovale or D. commonsianum. Dichanthelium laxiflorum differs in having longer winter rosette leaves, and culm leaves that are more basally disposed. It often has more of a yellow-green coloration, a smaller ligule and more extensive cilia on the leaf margins than members of section Lanuginosa. It most resembles D. acuminatum and D. villosissimum.
Conclusion The data generated and subsequent analyses made in the present study support the view that section Lanuginosa comprises of 16 species and two subspecies. The morphological boundaries of each taxon are well defined and a key to their identification is provided below. Though occasional plants are difficult to interpret due to the subtleties of distinguishing characters, the majority of specimens encountered are easily placed. It is the occasional morphological extremes within section Lanuginosa, which already
19 possesses subtle character differentiation, that make the group confusing. These exceptions do not represent taxonomic entities or intermediates but rather the plasticity of isolated populations and should not detract from the notions of these entities as clear species across their range. Though frustrating, once the range of morphological variation for these taxa is understood, they do indeed make sense.
20 TAXONOMIC TREATMENT
Dichanthelium sect. Lanuginosa (Hitchc.) Freckmann & Lelong Plants cespitose, 5-120 cm, glabrous to densely pilose, villose or puberulent. Basal rosettes conspicuous and over-wintering. Culms erect early in the season, often sprawling later. Leaves evenly distributed along the culm, almost always less than 1 cm wide. Ligules a dense ring of hairs, 0.2-5.0 mm long, often backed by a pseudoligule. Spikelets 0.8-3.0 mm long, elliptic to ovate-elliptic, pubescent, the first glume ¼ to ½ the spikelet length, the second glume and sterile lemma 5-9 nerved.
1. The first row of ligule hairs ≥ 1.7 mm long (key questionable specimens here) 2. The surfaces of terminal and midstem leaf sheaths of vernal culms glabrous or glabrate 3. Inflorescence < 1/3 as wide as long; spikelets acute to attenuate at the apex and ovate in outline; leaves and sheaths rigid with a stiff, smooth texture; inflorescences densely flowered; collar region of vernal leaves lacking cilia…………………………………….………………………...……D. spretum 3. Inflorescences at least ½ as wide as long; spikelets obtuse at the apex and elliptic to elliptic-ovate in outline; leaves and sheath thin with a papery texture; inflorescences sparsely flowered; collar region of vernal leaves ciliate 4. Under surfaces of vernal leaf blades glabrous to glabrate; largest leaves of the vernal culm mostly > 60 mm long and 5 mm wide...... D. lindheimeri 4. Under surfaces of vernal leaf blades puberulent; largest leaves of the vernal culm mostly < 60 mm long and 5 mm wide……………...... D. longiligulatum 2. The surfaces of terminal and usually midstem leaf sheaths of vernal culms conspicuously pubescent with a uniform indument of variable density 5. More than 30 cilia on each margin of the middle and lower leaves of the vernal culm; marginal cilia extending from the base of the leaves to approximately half way up the blade………………………...... D. acuminatum 5. Less than 20 cilia on each margin of the middle and lower leaves of the vernal culm; marginal cilia restricted to the basal ¼ of the blade 6. Plants restricted to hot geothermal soils of the Cascade and Rocky Mountains 7. The largest vernal stem leaves < 8 mm wide; pubescence on the vernal sheaths < 2.0 mm long.………….…D. thermale subsp. thermale 7. The largest vernal stem leaves > 8 mm wide; pubescence on the vernal sheaths > 2.0 mm long…………..….D. thermale subsp. sericeum 6. Plants never growing in hot geothermal soils 8. The upper surface of mid-stem leaves of vernal culms lacking pubescence along the central portions of the blade (lacking pubescence along the central one third of the blade from base to
21 tip)…………………………………………………..……D. columbianum 8. The upper surface of mid-stem leaves of vernal culms uniformly pubescent across the surface (margin to margin) or glabrous 9. Plants with a puberulent pubescence (less than 0.3 mm long) on the under surfaces of vernal stem leaves; main axis of the inflorescence possessing a puberulent pubescence often beneath longer pilose hairs; largest vernal leaves 22-65 mm long 10. Spikelets ≤ 1.1 mm long and 0.5 mm wide……….D. wrightianum 10. Spikelets ≥ 1.2 mm long and 0.5 mm wide 11. Longer hairs of the mid-stem sheaths and the upper leaf surfaces of vernal stem leaves ≤ 2 mm long; other than occasional puberulence, upper surface of vernal stem leaves with sparse pilosity or lacking pilosity; almost exclusively a species of coastal regions from New Jersey south through the Gulf of Mexico to northern South America and the Caribbean ...... D. leucothrix 11. Longer hairs of the mid-stem sheaths and the upper leaf surfaces of vernal stem leaves mostly > 2 mm long; the upper surface of vernal leaves conspicuously pilose; a species of southern Canada and the northern United States south to Georgia and Alabama including coastal and inland regions…………...... ……………….…..D. meridionale 9. Plants lacking hairs or with a velutinous, pilose or hispid pubescence on the under surfaces of vernal stem leaves; main axis of the inflorescence lacking a puberulent pubescence; largest vernal leaves 40-110 mm long 12. Spikelets 1.8-2.1 mm long; first glume 0.7-1.0 mm long; longest hairs of the mid-stem vernal sheaths mostly 2.7-4.5mm long; often 10 or fewer nodes along the central axis of the inflorescence; ligule a mixture of long and short hairs with shorter hairs more abundant in the center thereby creating a U-shaped ligule in general outline...... D. praecocius 12. Spikelets 1.3-1.8 mm long; first glume 0.3-0.7 mm long; longest hairs of the mid-stem vernal sheaths usually < 2.7 mm long; usually 12 or more nodes along the central axis of the inflorescence; ligule hairs approximately the same length throughout 13. Largest vernal leaves typically ≤ 65 mm long; vernal stem leaves typically < 6.0 mm wide; plants growing in acidic substrates and usually at sites with significantly high native floristic quality; typical habitats include bogs, cedar swamps, tamarack swamps and sandy or cherty woodlands, prairies and glades……………....…...D. implicatum 13. Largest vernal leaves typically > 65 mm long; vernal stem leaves typically > 6.0 mm wide; plants more often associated with neutral or alkaline soils and not indicative of high native floristic quality; typical habitats include old fields, woodlands, prairies and glades ...... D. lanuginosum
22 1. First row of ligule hairs ≤ 1.5 mm long 14. Upper surfaces of vernal stem leaves glabrous or glabrate along entire blade 15. Lower and basal sheaths possessing a single indument of long villose pubescence and lacking an additional short puberulence (occasional specimens may have pilose hairs under the villose hairs)…....D. villosissimum 15. Lower and basal sheaths possessing a double vestiture of long pilose or villose pubescence above a short puberulence or only possessing a short puberulence 16. Spikelets ≤ 2.1 (rarely to 2.4) mm long; ligule usually > l mm long; first glume ≤ 1 mm long...... D. columbianum 16. Spikelets > 2.1 mm long; ligule usually ≤ 1 mm long; first glume typically > 1 mm long 17. Terminal sheath of vernal culms with a double vestiture of short puberulence beneath longer pilose to villose hairs; ligule of vernal stem leaves > 0.7 mm long...... D. commonsianum 17. Terminal sheath of vernal culms with a single vestiture of longer pilose or villose pubescence; ligule of vernal stem leaves < 0.7 mm long...... D. ovale 14. Upper surfaces of vernal stem leaves conspicuously pubescent 18. The upper surface of mid-stem leaves of vernal culms uniformly pubescent across the surface (margin to margin) 19. More than 30 cilia on each margin of the middle and lower leaves of the vernal culm; marginal cilia extending from the base of the leaves to approximately half way up the blade…………………...... D. acuminatum 19. Less than 20 cilia on each margin of the middle and lower leaves of the vernal culm; marginal cilia restricted to the basal ¼ of the blade 20. Lower and basal sheaths possessing a double vestiture of long pilose or villose hairs above shorter pilose or puberulent hairs 21. Plants restricted to hot geothermal soils of the Cascade, Sierra Nevada and Rocky Mountain ranges...D. thermale subsp. thermale 21. Plants never growing in hot geothermal soils……D. columbianum 20. Lower and basal sheaths possessing a single indument solely of villose hairs 22. Indument of vernal stem sheaths > 2 mm long; spikelets > 1.9 mm long...... D. villosissimum 22. Indument of vernal stem sheaths < 2 mm long; spikelets < 1.9 mm long...... D. subvillosum 18. The upper surface of mid-stem leaves of vernal culms lacking pubescence along the central portions of the blade (lacking pubescence along the central one third of the blade from base to tip) 23. Spikelets ≤ 2.1 (rarely to 2.4) mm long; ligule usually > l.0 mm long; first glume ≤ 1.0 mm long...... D. columbianum 23. Spikelets > 2.1 mm long; ligule usually ≤ 1.0 mm long; first glume typically > 1.0 mm long 24. Terminal sheath of vernal culms with a double vestiture of short puberulent hairs below pilose to villose hairs; ligule of vernal stem
23 leaves > 0.7 mm long...... D. commonsianum 24. Terminal sheath of vernal culms with a single vestiture of pilose or villose hairs; ligule of vernal stem leaves < 0.7 mm long...... D. ovale
Dichanthelium acuminatum (Sw.) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1121. 1978. Panicum acuminatum Sw., Prodr. 23. 1788. Panicum dichotomum var. acuminatum (Sw.) Griseb., Fl. Brit. W.I. 553. 1864. TYPE: JAMAICA: locality unknown, Swartz s.n. (holotype: S; isotype: BM, US!)
Panicum ciliosum Nash, Bull. Torrey Bot. Club 26(11):568. 1899. TYPE: USA. Mississippi: Harrison Co., Biloxi, 1 Sep 1898, Tracy 4580 (holotype: NY!; isotype: NCU, US!)
Panicum orangense Ashe, J. Elisha Mitchell Sci. Soc. 15:113. 1899. TYPE: USA. North Carolina: Orange Co., Chapel Hill, 29 Jun 1898, Ashe s.n. (lectotype: US!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 220. 1910)
Panicum thurowii Scribn. & J.G. Sm., Circ. Div. Agrostol. U.S.D.A. 16:5. 1899. Dichanthelium acuminatum var. thurowii Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1125. 1978. Panicum acuminatum var. thurowii (Scribn. & Sm.) Reed, Phytologia 67(6):452. 1989. TYPE: USA. Texas: Waller Co., locality unknown, 5 Jun 1898, Thurow 9 (holotype: US!; isotype: US!)
Panicum auburne Ashe, North Carolina Agric. Res. Serv. Bull. 175:115. 1900. Dichanthelium auburne (Ashe) Mohlenbr., Vasc. Fl. Illinois 419. 2002. TYPE: USA. Alabama: Lee Co., Auburn, 7 May 1898, Earle & Baker 1527 (holotype: NCU!; isotypes: NY!, US!)
Panicum comophyllum Nash, Bull. Torrey Bot. Club 30(7):380. 1903. TYPE: PUERTO RICO. San Juan: collected in rich soil at Santurce, 9 Jan 1899, Heller 12 (holotype: NY!; isotype: fragment US)
Panicum olivaceum Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:225. 1910. TYPE: GUATEMALA. Alta Verapaz: Coban, 1400m, Feb 1888, Tuerckheim 428 (holotype: US!; isotype: ISC!, US!)
Panicum glutinoscabrum Fernald, Rhodora 49(581):122. 1947. TYPE: USA. Virginia: Nansemond Co., sphagnous and peaty bog by Norfolk and Western Railway, Fernald, Long & Clement 15186 (holotype: GH!; isotype: PH, US!)
Plants erect or ascending from a geniculate base, 20-70 cm tall; lower internodes double invested with short, often sparse, pilose hairs below longer villose hairs (Figure 8b, for
24 pubescence type), mid-stem and upper-stem internodes single invested with velutinous or villose hairs (Figure 8a, for pubscence type); sheaths conspicuously velutinous to villose (Figure 8a) with hairs 0.2–2.5 mm long, the lower stem sheaths double invested with short, pilose hairs and longer villose hairs (Figure 8b); ligule 0.4-1.3 mm long; pseudoligule distinct, 2-4 mm long; leaf blades spreading to ascending, 3-12 cm long, 4- 13 mm wide, margins ciliate on at least the lower half, adaxial leaf surface glabrate to villous, abaxial surface velutinous; panicles 3-11 cm long, half as wide to as wide as long, pilose to velutinous; spikelets 1.6-2.1 mm long, approximately 1 mm wide, elliptic, pilose; first glume 0.4-0.9 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 9). Dichanthelium acuminatum is found in coastal regions from Georgia to Texas, south to Mexico, Central America and northern South America. It is also found in the islands of the Caribbean. Its habitat consists of prairies, pine woodlands and dry open woods with sandy soil.
NOTES. Given the distinct morphology found in the present study as well as a primarily coastal distribution, the present treatment of D. acuminatum is much more restrictive than that of Gould and Clark (1978) and Freckmann and Lelong (2002, 2003). These past treatments place several taxa, here recognized as distinct species, as infraspecific taxa of D. acuminatum. Much of this confusion can be traced to the nomenclatural association of D. acuminatum with D. lanuginosum (see discussion). Dichanthelium acuminatum is readily identified by its short ligule, longer pseudoligule and the presence of marginal cilia on the vernal leaf blades which extend nearly half the blade length or more. These marginal hairs often have pustulate bases which can be detected even in older specimens that have lost the hairs. Typically, the whole plant is velvety-pilose and the lower blade surfaces are soft to the touch. It also produces an abundance of rosette leaves which are proportionately larger than the leaves of other species in the section. Hitchcock and Chase (1910) maintained the separation of D. olivaceum and D. thurowii from D. acuminatum based solely on plant color. While variation does exist, the abundance of intermediates and the many factors that can influence plant color suggest
25 that this character lacks taxonomic value. Gould and Clark (1978) maintained var. thurowii in reference to more robust specimens, but no justification for this could be found in the present treatment.
REPRESENTITIVE SPECIMENS.
BELIZE. Orange Walk District: 25 Jul 1980, J. Dwyer & C. Berry 15016 (MO); COLOMBIA. CAUCA: 1889, Lehmann 974 (GH); COSTA RICA. SAN JOSE: 26 Dec 1925, P. Standley & J. Valerio 43377 (GH); DOMINICAN REPUBLIC. SANTO DOMINGO: 13 Feb 1929, E. Ekman 11547 (GH); GUATEMALA. Alta Verapaz, Feb 1888, H. Turckheim 428 (MO,US); HAITI. 28 Sep 1943, L. Holdridge 1727 (MO); HONDURAS. FRANCISCO MORAZAN: 23 Oct 1951, J. Swallen 11012 (MO); JAMAICA. 24 Jan 1913, W. Harris 11597 (GH); MEXICO. CHIAPAS: 15 Nov 1984, G. Davidse 29845 (MO); OAXACA: 14 Aug 1975, G. Davidse & J. Davidse 9688 (MO); TABASCO: 8 Aug 1975, G. Davidse 9368 (MO); VERA CRUZ: 2 Sep 1910, A. Hitchcock 134 (MO); VERA CRUZ: Oct 1927, C. Purpus 7879 (MO); NICARAGUA. Nueva Segovia, 10-13 Aug 1977, W. Stevens & B. Krukoff 3271 (MO); PANAMA. CHIRIQUI: 25 Jun 1938, M. Davidson 766 (GH); UNITED STATES. ALABAMA: Lee Co., 7 May 1898, F. Earle & C. Baker 1527 (NY); FLORIDA: Franklin Co., Chapman 7189 (MO); GEORGIA: Burke Co., 4 Jun 1964, R. Freckmann 1180 (MO); LOUISIANA: Acadia Parish, 17 Apr 1913, C. Chambliss s.n. (MO); Jeff Davis Parish, 15 May 1915, E. Palmer 7610 (MO); MISSISSIPPI: Harrison Co., 1 Sep 1898, S. Tracy 4580 (US); NORTH CAROLINA: Orange Co., 29 Jun 1898, W. Ashe s.n. (US); PUERTO RICO: 17 Jan 1886, Hackel 5908 (GH); PUERTO RICO: 9 Jan 1899, A. Heller 12 (NY); TEXAS: Montgomery Co., 11 May 1980, C. Morden 29 (MO); Polk Co., 10 May 1973, F. Gould, S. Hatch & F. Waller 14233A (MO); Upshur Co., 28 May 1901, J. Reverchorn 2355 (MO); Waller Co., 5 Jun 1898, F. Thurow 9 (US); VIRGINIA: Nansemond Co., 2 & 12 Sep 1946, M. Fernald, B. Long & I. Clement 15186 (GH)
Dichanthelium columbianum (Scribn.) Freckmann, Phytologia 39(4):270. 1978. Panicum columbianum Scribn., Bull. Div. Agrostol. U.S.D.A. 7:78. 1897. Panicum acuminatum var. columbianum (Scribn.) Lelong, Brittonia 36(3):270: 1984. Dichanthelium acuminatum subsp. columbianum (Scribn.) Freckmann & Lelong, Sida 20(1):167. 2002. TYPE: USA. District of Columbia: Brookland, 14 Jul 1894, Scribner s.n. (holotype: US!; isotype: NCU, NY!)
Panicum addisonii Nash, Bull. Torrey Bot. Club 25(2):83-84. 1898. Panicum commonsianum subsp. addisonii (Nash) W. Stone, New Jersey State Mus. Annual Rep. 1910:205. 1911. Panicum commonianum var. addisonii (Nash) Fernald, Rhodora 37:391. 1935. Dichanthelium ovale var. addisonii (Nash) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1115. 1978. Panicum ovale var. addisonii (Nash) C.F. Reed, Phytologia 67(6):452. 1989. TYPE: USA. New Jersey: Cape May Co., in sandy soil at Wildwood, 30 & 31 May 1897, Bicknell s.n. (holotype: NY!; isotype: NY!, US)
26
Panicum scoparioides Ashe, J. Elisha Mitchell Sci. Soc. 15:53. 1898. Panicum villosissimum var. scoparioides (Ashe) Fernald, Rhodora 36(423):79. 1934. Dichanthelium scoparioides (Ashe) Mohlenbr., Erigenia 6:26. 1985. TYPE: USA. Delaware: New Castle Co., Centerville, dry soil, 25 Jun 1873, A. Commons (lectotype: US!; isotype: NY!)
Panicum tsugetorum Nash, Bull. Torrey Bot. Club 25(2):86. 1898. TYPE: USA. New York: Bronx Co., locality unknown, G. Nash 287 (holotype: NY!; isotype: US!)
Panicum lanuginosum var. siccanum Hitchc. & Chase, Rhodora 8(95):207-208. 1906. Panicum columbianum var. siccanum (Hitchc. & Chase) Boivin, Le Naturaliste Canadien 95:526. 1967. TYPE: USA. Illinois: La Salle Co., Starved Rock, dry hot sand of sandstone cliff, 13 Jul 1901, A. Chase 1602 (holotype: US!)
Panicum unciphyllum var. thinium Hitchc. & Chase, Rhodora 8(95):209. 1906. Panicum columbianum var. thinium (Hitchc. & Chase) Hitchc. & Chase, Rhodora 10(112):64. 1908. Panicum heterophyllum var. thinium (Hitchc. & Chase) Hubb, Rhodora 14(164):172. 1912. Dichanthelium sabulorum var. thinium (Hitchc. & Chase) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1113. 1978. TYPE: USA. New Jersey: Tom’s River, in mats, sandy open ground, 28 Jul 1906, A. Chase 3577 (holotype: US!)
Panicum owenae E.P. Bicknell, Bull. Torrey Bot. Club 35(4):185. 1908. TYPE: USA. Massachusetts: Nantucket Co., Nantucket, on the sandy commons west of town, 20 Sep 1907, E.P. Bicknell s.n. (holotype: NY!; isotype: US)
Plants erect or ascending from a geniculate base, 10-50 cm tall; internodes and sheaths pubescence of short puberulent to pilose hairs and long villose hairs (Figure 8d), occasionally lacking the longer villose hairs on the middle and upper internodes and sheaths (Figure 8c),longer hairs 0.3-2.0 mm long when present; ligule 0.9-1.5 mm long; pseudoligule sparse, intermeshed, or commonly absent (see Figs. 7b, c or d); leaf blades spreading to ascending, 3-9 cm long, 3-8 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface villose to glabrate, hairs absent along the central third of the blade, abaxial surface with appressed puberulence; panicles 2-8 cm long, half as wide to as wide as long, puberulent and often pilose; spikelets (1.3-) 1.5-1.9 (-2.4) mm long, approximately 1 mm wide, elliptic, pilose; first glume 0.5-1.0 mm long, typically 1/3 to 1/2 as long as the spikelet, obtuse to broadly acute.
27 DISTRIBUTION AND HABITAT (Figure 10). Dichanthelium columbianum is found in scattered counties throughout eastern North America ranging from Nova Scotia and western Ontario south to Texas and Florida. It is more commonly collected from the northeast portion of its range as well as from the Great Lakes region. The habitat of D. columbianum typically consists of dunes, pine woodlands and open woods with sandy soils.
NOTES. Dichanthelium columbianum varies widely in degree of pubescence but consistently possesses a short puberulence on all the internodes and sheaths of the vernal culm. This character, as well as spikelet, first glume and ligule length are useful in separating it from D. ovale. The spikelet lengths of D. columbianum can approach those of D. commonsianum, in which case the ligule and glume length can be used to separate the two. When a pseudoligule is present in D. columbianum, it is sparse and weakly differentiated, but can be easily detected with 30x magnification or higher. Plants possessing a pseudoligule are typically more pubescent overall than those lacking a pseudoligule. While all plants possess an extensive puberulence, there is much variation in the degree of longer pilose or villose pubescence (double vestiture). Plants with more dense pilosity or villosity often possess pubescence all across the adaxial leaf surfaces while more sparsely pubescent plants may be nearly glabrous along the adaxial leaf surfaces.
Panicum addisonii is here placed in synonymy. This is a unique placement, in that previous treatments have never directly associated P. addisonii with D. columbianum. Rather, it has often been treated under D. commonsianum or D. ovale on the basis of having spikelets equal to or greater than 2 mm (Deam 1940; Fernald 1950; Freckmann and Lelong 2002; Freckmann and Lelong 2003; Gleason and Cronquist 199; Gould and Clark 1978; Pohl 1947; Radford et al. 1968; Stone 1911). However, the holotype of P. addisonii has spikelets that range from 1.8-2.0 mm long which is well within the range of typical D. columbianum and outside the range for D. commonsianum or D. ovale. Furthermore, the length of its ligule and first glume better align it with D. columbianum. This confusion is thought to stem from the treatments of Hitchcock and
28 Chase (1910, 1951) wherein the vernal culms of P. addisonii are referred to as being “similar to that of P. commonsianum”. Stone (1911) was the first to subsume addisonii under P. commonsianum, referring to it as being a diminutive of the typical variety.
In the treatments of Gould and Clark (1978) and Hitchcock and Chase (1910), specimens with smaller spikelets (1.3-1.4 mm) have been referred to as var. thinium and specimens with larger spikelets (2.2-2.4 mm) have been referred to as P. scoparioides. These, as well as var. siccanum, are better interpreted as being within the range of variation of the species rather than as infraspecific taxa.
REPRESENTITIVE SPECIMENS.
UNITED STATES. DELAWARE: Eussex Co., 16 Jun 1926, J. Fogg Jr. s.n. (MO); DELAWARE: New Castle Co., 25 Jun 1823, A. Commons 283 (US); ILLINOIS: LaSalle Co., 13 Jul 1901, A. Chase 1602 (US); MARYLAND: Calvert Co., 8 Jul 1934, A. Chase 12270 (MU); MASSACHUSETTS: Nantucket Co., 20 Sep 1907, P. Bicknell s.n. (NY); Plymouth Co., 15 Sep 1916, H. St. John 11755 (MU); NEW JERSEY: Cape May Co., 30 May 1897, E. Bicknell s.n. (NY); NEW YORK: Clinton Co., 3 Jul 1902, W. Eggleston 2843 (MU); Oneida Co., 3 Jul 1945, R. Crockett 7577 (MO); OHIO: Hocking Co., 15 Jun 1982, A. Cusick 21692 (MU); Lucas Co., 19 Jun 1981, N. Easterly 12688 (MU)
Dichanthelium commonsianum (Ashe) Freckmann, Phytologia 39(4):271. 1978. Panicum commonsianum Ashe, J. Elisha Mitchell Sci. Soc. 15:55. 1898. Panicum columbianum var. commonsianum (Ashe) Dore, Le Naturaliste Canadien 103(6):562. 1972. TYPE: USA. New Jersey: Cape May Co., Cape May, collected in drifting sands along the coast, 29 Jun 1898, Commons 341 (holotype: unknown; isotypes: NY!, US!)
Panicum deamii Hitchc. & Chase, Indiana Dept. Cons., Publ. 82:284. 1929. TYPE: USA. Indiana: Lake Co., collected on crest of low dune a quarter mile south of Pine, 21 Jun 1926, C.C. Deam 43287 (holotype: US!)
Panicum euchlamydeum Shinners, Amer. Midl. Naturalist 32:170. 1944. Panicum commonsianum var. euchlamydeum (Shinners) Pohl, Amer. Midl. Naturalist 38:507. 1947. Dichanthelium commonsianum var. euchlamydeum (Shinners) Freckmann, Phytologia 39(4):271. 1978. TYPE: USA. Wisconsin: Adams Co., sandy slope, PH 7.5, 12 miles southeast of Adams, 28 Jun 1942, L.H. Shinners & C.G. Shaw 4415 (holotype: WIS!; isotypes: GH, US!)
29 Plants erect or ascending from a geniculate base, 10-60 cm tall; internodes double invested with short puberulent to pilose hairs below a villous pubescence (Figure 8d) or occasionally lacking the longer villous hairs on the middle and upper internodes (Figure 8c); sheaths double invested with short puberulent hairs below a pilose or villous pubescence (Figure 8d) or occasionally lacking the longer villous hairs on the middle and upper sheaths (Figure 8c), longer hairs 0.5-3.7 mm long when present; ligule 0.5-1.3 mm long; pseudoligule sparse or lacking (Figs. 7b or c); leaf blades spreading to ascending, 3- 11 cm long, 3-7 (-10) mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface glabrate to villous, hairs absent along the longitudinal central third of the blade, abaxial surface with appressed puberulence; panicles 3-9 cm long, half as wide to as wide as long, puberulent; spikelets 2.2-2.7 mm long, approximately 1 mm wide, elliptic, pilose; first glume 0.9-1.8 mm long, typically 2/5 to 1/2 as long as the spikelet, obtuse to broadly acute.
DISTRIBUTION AND HABITAT (Figure 11). Dichanthelium commonsianum is found along the coastal plain of the eastern and southeastern United States from Massachussets to Texas. Its distribution within the interior of North America is scattered and poorly understood. It is reported from every state east of and including Arkansas, Kansas and Iowa as well as throughout the Great Lakes region. The present study has verified interior North American collections from Missouri, Wisconsin, Indiana, Kansas and Ohio. Where found, D. commonsianum is associated with dry sandy soils of open woods, dunes and prairies.
NOTES. Dichanthelium commonsianum is similar in morphology to D. columbianum and D. ovale. It can be distinguished from D. columbianum by its larger spikelets, shorter ligule and longer first glume. It differs from D. ovale by the presence of puberulence on internodes and sheaths of the middle and upper culm which D. ovale lacks, as well as a longer ligule. There is also a strong tendency for the first glume of D. commonsianum to be slightly greater than half the overall spikelet length while the first glume of D. ovale is typically less than half the spikelet length. Some authors have included P. addisonii as synonymous or infra specific to D. commonsianum. For a
30 discussion of this see notes under D. columbianum. Panicum euchlamydeum and P. commonsianum var. euchlamydeum are names once used in reference to Midwestern specimens of D. commonsianum. However, as is evident in Pohl’s (1947) description of the variety, there is little value in maintaining var. euchlamydeum other than as representing the inland populations and there does not appear to be any morphological distinction other than a tendency for slightly more pubescence in Midwestern specimens. This is very much in agreement with the findings of the present treatment.
REPRESENTITIVE SPECIMENS. UNITED STATES. DELAWARE: Sussex Co., 18 Jun 1905, A. Hitchcock 143 (MO); Sussex Co., 24 Jun 1942, R. McVaugh 6512 (MO); INDIANA: Jasper Co., 15 Jun 1947, R. Friesner 21490 (MO); MARYLAND: Prince George’s Co., 25 Jul 1907, A. Chase 146 (MO); MASSACHUSETTS: Barnstable Co., 23 Jun 1918, M. Fernald 16118 (MO); Barnstable Co., 4 Aug 1918, M. Fernald 16119 (MO); Dukes Co., 5 Sep 1928, J. Fogg Jr. 3875 (MO); MISSOURI: Scott Co., 7 Sep 1993, P. McKenzie 1294 (MO); NEW JERSEY: Cape May Co., 29 Jun 1899, A. Commons 341 (US); Mercer Co., 18 Aug 1866, E.D. s.n. (MO); Middlesex Co., 9 Jul 1905, K. MacKenzie 1485 (MO); Ocean Co., 28 Jul 1907, K. MacKenzie 2779 (MO); NEW YORK: Suffolk Co., 19 Jul 1927, R. Latham 4169 (MO); Suffolk Co., 4 Aug 1956, G. Van Schaak 3550 (MO); Suffolk Co., 4 Aug 1956, G. Van Schaak 3551 (MO); TEXAS: Robertson Co., 30 Apr 1951, J. Launchbaugh 397 (MO); WISCONSIN: Adams Co., 28 Jun 1942, L. Shinners & C. Shaw 4415 (UW); La Crosse Co., 18 Jun 1925, L. Pammel 1334 (MO)
Dichanthelium implicatum (Scribn.) Kerguélen, Fl. Descript. France, Suppl. 5:544. 1979. Panicum implicatum Scribn., Bull. Div. Agrostol. U.S.D.A. 11:43. 1898. Panicum unciphyllum var. implicatum (Scribn.) Scribn. & Merr., Rhodora 3(29):123. 1901. Panicum lindheimeri var. implicatum (Scribn.) Fernald, Rhodora 23(274):228. 1921. [1922]. Panicum lanuginosum var. implicatum (Scribn.) Fernald, Rhodora 36(423):77. 1934. Dichanthelium acuminatum var. implicatum (Scribn.) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1126. 1978. 1979. Panicum acuminatum var. implicatum (Scribn.) Beetle, Phytologia 48(2):192. 1981. Dichanthelium acuminatum subsp. implicatum (Scribn.) Freckmann & Lelong, Sida 20(1):167. 2002. TYPE: USA. Maine: Cumberland Co., Cape Elizabeth, on salt marshes, 26 Jul 1895, F.L. Scribner s.n. (holotype: US!; isotype: NY!)
Panicum occidentale Scribn., Annual Rep. Missouri Bot. Gard. 10:48. 1899. TYPE: CANADA. British Columbia: Nootka Sound, no date, T. Haenke s.n. (holotype: PR; isotype: MO!, US!)
Panicum pacificum Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:229. 1910. TYPE: USA. California: Shasta Co., Castle Crag, one-fourth miles west of hotel,
31 in moist places in woods, 3 Aug 1908, A.S. Hitchcock 3070 (holotype: US!; isotype: NY!)
Plants erect or ascending from a geniculate base, 15-55 cm tall; internodes conspicuously to sparsely pilose to villose (Figure 8a); sheaths conspicuously to sparsely pilose to villose (Figure 8a), hairs 0.9-4.9 mm long; ligule 2.0-5.0 mm long (Figure 8a); pseudoligule lacking or indistinguishable from ligule; leaf blades spreading to ascending, 2.0-6.5 (-8) cm long, 2.0-6.0 (-8) mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface villous with the longest hairs often 3.0-6.0 mm long, abaxial surface pilose with the longest hairs often 0.8-2.0 mm long ; panicles 3-6 cm long, half as wide to as wide as long, densely to sparsely pilose; spikelets 1.3-1.7 mm long, approximately 0.7-1.0 mm wide, elliptic, pilose; first glume 0.3-0.7 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 12). Dichanthelium implicatum is found across southern Canada from British Columbia to Nova Scotia. In the United States it is found from Kansas north to Minnesota and east to Tennessee and Maine, plus the states of California, Oregon and Washington. It is also listed as naturalized in southern Europe (Coste 1979). The habitat of D. implicatum is highly variable and includes dry, open habitats with sandy soils such as dunes, sand prairies and sandy or cherty woods. It is also occasionally found in bogs with acidic soils.
NOTES. Some specimens of D. implicatum and D. lanuginosum can be morphologically similar (see discussion). This is espceially true when dealing with small D. lanuginosum and large D. implicatum. However, most specimens of D. implicatum are very distinct. It also inhabits a much narrower ecological niche than D. lanuginosum. Compared to D. lanuginosum, it is smaller in stature and the larger vernal leaves are less than 7.0 cm long. The longer hairs on the adaxial surface of vernal leaves average 3.2 mm in D. implicatum compared to 2.3 mm in D. lanuginosum. The longer hairs of the abaxial surface of vernal leaves average 0.9 mm long in D. implicatum, while in D. lanuginosum they average 0.6 mm. Furthermore, upon drying naturally, the vernal leaves of D. implicatum often become strongly reflexed.
32 The similarities of the two species have prompted several authors (Braun 1967; Gleason and Cronquist 1991; Fernald 1950; Radford et al. 1968; Scoggan 1978; Steyermark 1963) to treat D. implicatum as a variety of D. lanuginosum. Pohl (1947), in his treatment of Pennsylvania’s grasses, followed the treatment of Hitcock and Chase (1910) in which P. lanuginosum (often representing the nomenclatural misapplication of what is here treated as D. acuminatum) was restricted to the southeastern coastal plain. He then applied the name P. implicatum to what is here treated as D. lanuginosum. This treatment has since found its way into several floras in the Great Lakes Region (Swink and Wilhelm 1994; Voss 1972) and no doubt added to the confusion of species concepts. It is worth noting that D. implicatum is limited to the northern half of the geographical range of D. lanuginosum.
The type specimens of two species, P. pacificum and P. occidentale, described from western North America, are treated as synonymous with D. implicatum. However, there are notable differences in the western populations of D. implicatum, including wider leaves and differences in pubescence density that may warrant further investigation. Many “non-type” specimens labeled “P. pacificum” and “P. occidentale” clearly belong to D. lanuginosum rather than D. implicatum. This is most likely due to erroneous identification rather than actual taxonomic confusion. Dichanthelium implicatum can superficially resemble D. meridionale but differs in its lack of a double vestiture (or at least no puberulence below the villose and pilose pubescence).
REPRESENTITIVE SPECIMENS.
CANADA. ONTARIO: Prince Edward Co., 10 Jul 1947, J. Gillett 1366 (MO); UNITED STATES. ARKANSAS: Fulton Co., 19 May 1909, B. Bush 5637 (MO); CALIFORNIA: Shasta Co., 3 Aug 1908, A. Hitchcock 3070 (US); IOWA: Johnson Co., B. Shimek s.n. (MO); OHIO: Lucas Co., 1 Jul 1997, J. McCormac 5971 (MO); OREGON: 1871, E. Hall s.n. (MO); Curry Co., 11 Jul 1919, M. Peck 8795 (MO); MAINE: Lincoln Co., 13 Jul 1922, F. Grover s.n. (MO); Cumberland Co., 25 Jul 1895, F. Scribner (US); impl MASSACHUSETTS: Berkshire Co., 16 Jul 1917, J. Churchill s.n. (MO); Middlesex Co., 27 Jun 1915, F. Forbes s.n. (MO); MINNESOTA: Anoka Co., 7 Jul 1926, A. Stoltz s.n. (MO); Anoka Co., 10 Aug 1926, A. Stoltz s.n. (MO); Chippewa Co., 9 Jul 1982, R. Gereau 1014 (MO); Itasca Co., 30 Jun 1935, C. Rosendahl & J. Maple 707 (MO); NEW YORK: Tompkins Co., 18 Jul 1917, K. Wiegand, A. Bechtel & A. Gershoy 7504 (MO); RHODE ISLAND: Providence Co., Olney s.n. (MO); TENNESSEE: Coffee
33 Co., 20 Jun 1938, H. Svenson 9230 (MO); VERMONT: Caledonia Co., F. Seymour 18632 (MO); WASHINGTON: Presl 3978 (MO); Chelan Co., 25 Jun 1931, W. Thompson 6928 (MO); WISCONSIN: Bayfield Co., 10 Jul 1938, N. Fassett, J. Curtiss & L. Knowlton 19571 (MO)
Dichanthelium lanuginosum (Ell.) Gould, Brittonia 26(1): 60. 1974. Panicum lanuginosum Ell., Sketch Bot. S. Carolina 1(2):123. 1816. Panicum dichotomum var. lanuginosum (Ell.) Wood, A Class-book of Botany 786. 1861. TYPE: USA. Georgia: unknown locality, unknown date, Baldwin s.n. (holotype: CHARL; fragment and photo, US!)
Panicum dichotomum var. fasciculatum Torr., Fl. N. Middle United States 145. 1824. Panicum huachucae var. fasciculatum (Torr.) F.T. Hubb., Rhodora 14(164):171. 1912. Panicum lindheimeri var. fasciculatum (Torr.) Fernald, Rhodora 23(274):228. 1921 [1922]. Panicum lanuginosum var. fasciculatum (Torr.) Fernald, Rhodora 36(423):77. 1934. Dichanthelium lanuginosum var. fasciculatum (Torr.) Spellenb., Madrono 23(3):145. 1975. Dichanthelium acuminatum var. fasciculatum (Torr.) Freckmann, Phytologia 48(1):108. 1981. Panicum acuminatum var. fasciculatum (Torr.) Beetle, Phytologia 49(1):40. 1981. Dichanthelium acuminatum subsp. fasciculatum (Torr.) Freckmann & Lelong, Sida 20(1):167. 2002. TYPE: USA. New Jersey: unknown locality, in sandy fields, unknown date, Torrey s.n. (holotype: NY)
Panicum huachucae Ashe, J. Elisha Mitchell Sci. Soc. 15:51. 1898. Panicum lanuginosum var. huachucae (Ashe) Hitchc., Rhodora 8(95):208. 1906. TYPE: USA. Arizona: Huachuca Mountains, 1882, J.G. Lemmon s.n. (lectotype: US!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 214. 1910)
Panicum tennesseense Ashe, J. Elisha Mitchell Sci. Soc. 15:52. 1898. Panicum lindheimeri var. tennesseense (Ashe) Farw., Amer. Midl. Naturalist 11:45. 1928. Panicum lanuginosum var. tennesseense (Ashe) Gleason, Phytologia 4(1):21. 1952. TYPE: USA. Tennessee: La Vergne Co., cedar glades, 7 Aug 1897, Biltmore 7087 (holotype: location unknown; isotypes: NY!, US!)
Panicum nitidum var. pilosum Torr., Fl. N. Middle United States 146. 1824. Panicum lindheimeri subvar. pilosum (Torr.) Farw., Amer. Midl. Naturalist 11:45. 1928. Panicum lanuginosum subvar. pilosum (Torr.) Farw., Pap. Michigan Acad. Sci. 26:5. 1941. TYPE: USA. New York: unknown locality, in dry woods & c. New York, no date, unknown collector (holotype: NY?)
Panicum nitidum var. ciliatum Torr., Fl. N. Middle United States 146. 1824. TYPE: USA. New Jersey: unknown locality, no date, unknown collector (holotype: NY?)
34 Panicum funstonii Scribn. & Merr., Circ. Div. Agrostol. U.S.D.A. 35:4. 1901. TYPE: USA. California: Tulare Co., collected on the bank of the Kaweak River at Three Rivers, 26 Jul 1891, Coville & Funston 1286 (holotype: US!)
Panicum unciphyllum f. prostratum Scribn. & Merr., Rhodora 3(29):124. 1901. TYPE: USA. Maine: York Co., unknown locality, 26 Sep 1897, M.L. Fernald s.n. (holotype: US!)
Panicum huachucae var. silvicola Hitchc. & Chase, Rhodora 10(112):64-65. 1908. TYPE: USA. District of Columbia: Reno, 28 Jun 1904, A. Chase 2400 (holotype: US!)
Panicum lindheimeri var. septentrionale Fernald, Rhodora 23(274):227-228. 1921 [1922]. Panicum lanuginosum var. septentrionale (Fernald) Fernald, Rhodora 36(423):77. 1934. Dichanthelium acuminatum var. septentrionale (Fernald) Mohlenbr., Ill. Fl. Illinois (ed. 2) Grasses: Panicum to Danthonia: 383. 2001. TYPE: CANADA. New Brunswick, Woodstock, river gravels and shingly border of thicket by the St. John River, 14 Jul 1916, M.L. Fernald & Long 12527 (holotype: GH!)
Panicum brodiei H. St. John, Fl. SE Washington 51. 1937. TYPE: USA. Washington: Wawawai, Snake River, 1898, D.A. Brodie s.n. (holotype: WS!; isotype: US)
Plants erect or ascending from a geniculate base, 15-75 cm tall; internodes conspicuously to sparsely pilose to villose (occasionally velutinose) (Figure 8a); sheaths copiously to sparsely pilose to villose (Figure 8a), hairs (0.7-) 1.2–2.5 (-3.9) mm long, sheath margins ciliate; ligule (2.0-) 2.4-4.3 (-5.0) mm long (Figure 8a); pseudoligule lacking or indistinguishable from ligule; leaf blades spreading to ascending, 3-12 cm long, 3-12 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface glabrate to villous with the longest hairs typically 1.3-3.0 mm long, abaxial surface short pilose with the longest hairs typically 0.1-0.7 mm long; panicles 3-12 cm long, half as wide to as wide as long, densely to sparsely pilose; spikelets 1.3-1.9 mm long, approximately 0.7- 1.1 mm wide, elliptic, pilose; first glume 0.3-0.7 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 13). Dichanthelium lanuginosum is found throughout North America but is most common in the central and eastern United States. Being the most generalist species within the section, D. lanuginosum inhabits a wide
35 range of habitat types and is by far the most frequently encountered taxon in the section. It is most commonly associated with dry open habitats such as prairies, open woods, old fields and forest edges.
NOTES. Dichanthelium lanuginosum exhibits some morphological variation across its range and in different habitats. In western North America the spikelets typically range from 1.7-1.9 mm long. In deeply shaded environments plants are more sparsely pubescent, the stems often become diffusely branched and the plants tend to sprawl. Also, the leaves often become thinner, wider and darker green and the inflorescences are smaller and fewer flowered. In dry habitats with high light the plants are smaller, more densely pubescent, more erect, and produce larger inflorescences with many spikelets. It is these latter specimens that most closely approach and are occasionally difficult to discern from D. implicatum (see notes under D. implicatum). In moist, well-drained habitats plants are often sparsely pubescent on the sheaths and lack pubescence on the adaxial leaf surfaces. These plants have previously been called P. tennesseense, P. lanuginosum var. tennesseense, and P. lanuginosum var. septentrionale. However, given that the spectrum of pubescence is so closely associated with light and moisture, there seems to be little justification for any taxonomic recognition (see notes under D. lindheimeri).
Much of the taxonomic confusion surrounding D. lanuginosum began with the regional misapplication of the name and its association with D. acuminatum (see discussion). Lelong (1984) claimed that the type specimens of P. acuminatum and P. lanuginosum are identical. However, the present study found several meaningful morphological differences between these types as well as the plants they represent. The long, clearly single ligule and the absence of extensive marginal cilia on the leaf blades of D. lanuginosum easily distinguishes it from D. acuminatum which has a short ligule (backed by a longer pseudoligule) and conspicuous marginal cilia along the leaf blades. While the geographical ranges of these two species do overlap along the coastal plain, D. acuminatum does not typically occur in the interior of North America.
36 REPRESENTITIVE SPECIMENS. CANADA. BRITISH COLUMBIA: Nelson Co., 20 Jul 1939, J. Thompson 14433 (MO); Vancouver Island, 26 Jun 1907, C. Rosendahl 1931 (MO); ONTARIO: Pipineau Co., 19 Jul 1947, W. Dore 1051 (MO); QUEBEC: Missisquoi Co., 10 Aug 1923, C. Knowlton s.n. (MO); UNITED STATES. ARKANSAS: Benton Co., 18 Jun 2002, J. Thomas 1502 (private); ARIZONA: Mt. Huachuca, 1882, J. Lemmon 2907 (US); CALIFORNIA: Shasta Co., 3 Aug 1908, A. Hitchcock 136 (MO); Tulare Co., 26 Jul 1891, Coville & Funston 1286 (US); DISTRICT OF COLUMBIA, 28 Jun 1904, A. Chase 2400 (US); ILLINOIS: Macon Co., 12 Jun 1915, I. Clokey 2440 (MO); Morgan Co., 16 Jun 1999, S. Hill 31708 (MO); Tazewell Co., 23 Jun 1919, V. Chase 3146 (MO); INDIANA: Morgan Co., 11 Jun 1934, R. Friesner 7497 (MO); IOWA: Wapello Co., 23 Jun 1939, M. Hayden 925 (MO); KANSAS: Douglas Co., 15 Jun 1992, R. McGregor 40533 (MO); KENTUCKY: Ballard Co., 2 Jun 1923, W. Anderson s.n. (MO); LOUISIANA: Lake Charles Parish., 21 Apr 1906, A. Hitchcock 1147 (MO); MAINE. York Co., 26 Sep 1897, M. Fernald s.n. (US); MARYLAND: Balto Co., 21 Jun 1940, C. Reed 2165 (MO); MASSACHUSETTS: Bristol Co., 28 Jun 1961, F. Seymour 19272 (MO); MISSOURI: Jasper Co., 2 Jun 1912, E. Palmer 3714 (MO); Jasper Co., 9 Jun 1912, E. Palmer 3753 (MO); NEBRASKA: Kearney Co., 8 Jul 1918, H. Hapeman s.n. (MO); Nemaha Co., 22 Jun 1972, S. Churchill 200 (MO); NEW JERSEY: Hunterdon Co., 12 Jul 1915, H. Fisher s.n. (MO); NEW YORK: Tompkins Co., 12 Jul 1913, A. James 88 (MO); OHIO: Huron Co., 20 Jun 1887, F. Leonard 14 (MO); Scioto Co., 27 Jun 1934, D. Demaree 10756 (MO); OKLAHOMA: Creek Co., 11 Jun 1939, L. Harvey 808 (MO); LeFlore Co., 17 May 1936, D. Demaree 12712 (MO); OREGON: Lane Co., 14 Jun 1928, M. Peck 15950 (MO); PENNSYLVANIA: Montgomery Co., 15 Jun 1899, A. MacElwee 544 (MO); Philadelphia Co., no date, C. Bigyn s.n .(MO); SOUTH CAROLINA: Anderson Co., May 1924, J. Davis s.n. (MO); TEXAS: Wheeler Co., 25 Jun 1957, D. Correll & I. Johnston 17045 (MO); UTAH: San Juan Co., 2 Jul 1933, B. Maguire 5910 (MO); VIRGINIA: Augusta Co., 5 Jul 1937, H. Allard 3170 (MO); WASHINGTON: Kittilas Co., July 1898, Sandberg & Leiberg 425 (MO); Okanogan Co., Jul 1897, A. Elmer 489 (MO); Whitman Co., Jun 1898, D. Brodie s.n. (WS); WISCONSIN: La Crosse Co., 8 Jul 1975, S. Ziegler 1656 (MO)
Dichanthelium leucothrix (Nash) Freckmann, Phytologia 48(1):101. 1981. Panicum leucothrix Nash, Bull. Torrey Bot. Club 24(1):41-42. 1897. Panicum acuminatum var. leucothrix (Nash) Lelong, Brittonia 36(3):271. 1984. Dichanthelium acuminatum subsp. leucothrix (Nash) Freckmann & Lelong, SIDA 20(1):167. 2002. TYPE: USA. Florida: Lake Co., low pine land at Eustis,16-31 Jul 1894, Nash 1338 (holotype: NY!; isotypes: MO!, NCU, US)
Panicum parvispiculum Nash, Bull. Torrey Bot. Club 24(7):347-348. 1897. TYPE: USA. Georgia: McIntosh Co., collected at Darien Junction, 25-27 Jun 1895, Small s.n. (holotype: NY!; isotype: NY!, US)
Plants erect or ascending from a geniculate base, 10-50 cm tall; internodes short puberulent, occasionally also sparsely pilose on lower culms (Figs. 7c or d); sheaths
37 puberulent, occasionally also pilose (Figs. 7c or d), longer hairs 0.3-1.6 mm long when present; ligule 2.2-4.0 mm long (Figure 8a); pseudoligule absent or indistinguishable from ligule; leaf blades spreading to ascending, 2-7 cm long, 2.0-5.0 (-7.0) mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface puberulent to glabrate, abaxial surface puberulent; panicles 3-8 cm long, half as wide to as wide as long, puberulent and often pilose; spikelets 1.2-1.5 mm long, approximately 0.5-0.8 mm wide, elliptic, puberulent to pilose; first glume 0.3-0.5 mm long, typically 1/3 as long as the spikelet, obtuse to acute.
DISTRIBUTION AND HABITAT (Figure 14). Dichanthelium leucothrix is found along the coastal plain of North America from New Jersey to eastern Mexico and inland to Tennessee. It also occurs in Central America, northern South America and the Caribbean. The habitat of D. leucothrix consists of moist, often sandy, pine savannas and bogs.
NOTES. Dichanthelium leucothrix is one of the smaller members of the section and is similar to D. wrightianum, D. longiligulatum and D. meridionale in leaf dimensions, presence of puberulence and ligule length. Gould and Clark (1978) treated both D. longiligulatum and D. wrightianum as varieties of D. acuminatum and lumped D. leucothrix and D. meridionale with D. acuminatum var. implicatum (D. implicatum in present treatment). Given the absence of puberulence on the culms in D. implicatum and its larger spikelets, Gould and Clark’s treatment lacks the resolution that is attained by keeping these taxa as separate species. Lelong (1984) and Freckmann and Lelong (2002, 2003) designated D. leucothrix as a subspecies of D. acuminatum (or P. acuminatum). Zuloaga et al. (1993b) placed D. leucothrix into synonymy under P. acuminatum var. longiligulatum. Compared to D. acuminatum, D. leucothrix has a distinctly different ligule (longer and lacking pseudoligule versus short ligule backed by pseudoligule in D. acuminatum), does not possess extensive marginal cilia on vernal leaves, is much smaller in stature, has a close puberulence that D. acuminatum lacks and has much smaller spikelets (averaging 1.3 mm versus 1.8 mm). Dichanthelium leucothrix differs from D. longiligulatum in possessing conspicuous pubescence on the sheaths and adaxial leaf
38 surfaces of vernal culms. Given the considerable morphological differences between D. acuminatum, D. longiligulatum and D. leucothrix, the present work finds that D. leucothrix is best recognized as a distinct species.
REPRESENTITIVE SPECIMENS. UNITED STATES. FLORIDA: Fauquier Co., 26 May 1903, S. Tracy 8410 (MO); Hillsborough Co., 2 Apr 1927, H. O’Neill 2513 (MO); Lake Co., 16-31 Jul 1894, G. Nash 1338 (NY); GEORGIA: McIntosh Co., 25-27 Jun 1895, J. Small s.n. (NY); SOUTH CAROLINA: Orangeburg Co., 19 Aug 1905, A. Hitchcock 118 (MO); TENNESSEE: Coffee Co., 20 Jun 1938, H. Svenson 9225 (MO); TEXAS: Robertson Co., 30 Apr 1951, J. Launchbaugh 394 (MO)
Dichanthelium lindheimeri (Nash) Gould, Brittonia 26(1): 60. 1974. Panicum lindheimeri Nash, Bull. Torrey Bot. Club 24(4):196-197. 1897. Dichanthelium acuminatum var. lindheimeri (Nash) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1127. 1978. Panicum lanuginosum var. lindheimeri (Nash) Fernald, Rhodora 36(423):77. 1934. Dichanthelium lanuginosum var. lindheimeri (Nash) Harvill, Castanea 42(2):177. 1977. Panicum acuminatum var. lindheimeri (Nash) Beetle, Phytologia 48(2):193. 1981. Dichanthelium acuminatum subsp. lindheimeri (Nash) Freckmann & Lelong, SIDA 20(1):168. 2002. TYPE: USA. Texas: Comal Co., springs, banks of Guadeloupe, near New Braunfels, May 1946, F. Lindheimer 565 (holotype: NY! isotype: MO!, NY!, US!)
Plants erect or ascending from a geniculate base, 15-100 cm tall; internodes glabrous to sparsely pilose; sheaths glabrous or lowermost sparsely pilose, margins ciliate; ligule 2.0- 3.2 mm long (Figure 8a); pseudoligule absent; leaf blades spreading to ascending, 3-10 cm long, 3-12 mm wide, margins ciliate at the base of blade or eciliate, adaxial leaf surface glabrous to glabrate, abaxial surface glabrous; panicles 3-12 cm long, half as wide to as wide as long, glabrous to sparsely pilose; spikelets 1.3-1.7 mm long, approximately 0.7-1.0 mm wide, elliptic, pilose; first glume 0.3-0.6 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 15). Dichanthelium lindheimeri is found throughout central and eastern North America as well as California, Oregon and New Mexico. In its southern range and in the Great Lakes region, D. lindheimeri is most commonly associated with sandy, ephemerally wet, soils. In the central and northern
39 regions of its range it is more commonly associated with floodplains, stream sides and lake shores.
NOTES. Dichanthelium lindheimeri can be a difficult taxon to interpret. Superficially, it appears to be a glabrous variety or even a form of D. lanuginosum. In fact, many authors (Braun 1967; Gleason and Cronquist 1991; Fernald 1950; Radford et al. 1968; Scoggan 1978; Steyermark 1963) have placed it in this context. However, D. lindheimeri differs significantly from D. lanuginosum in several characters. First, the middle and upper internodes and sheaths of the vernal culms, the adaxial leaf surfaces (except for marginal cilia) and the central axis of the inflorescences are glabrous in D. lindheimeri versus densely to sparsely pilose in D. lanuginosum. Second, the margins at the base of vernal stem leaves in D. lindheimeri possess long hairs that are thin and crooked versus hairs that are rigid and straight in D. lanuginosum. Third, the ligule of D. lindheimeri averages 2.4 mm long versus 3.1 mm long in D. lanuginosum. Although some overlap does occur, t-test for ligule length between these two species demonstrated a significant difference (P<0.005). Fourth, the typical habitats of D. lindheimeri are much wetter than D. lanuginosum and include stream and lake margins, sandy coastal marshes and floodplains versus dry to mesic fields, prairies, open woods and forest edges for D. lanuginosum. In all, D. lindheimeri and D. lanuginosum demonstrate a statistical difference (P ≤ 0.05) in 15 of the 24 morphological characters sampled. Hitchcock and Chase (1910 and 1951) found these differences striking enough to align D. lindheimeri in a separate group (Spreta) than D. lanuginosum (Lanuginosa). Gould and Clark (1978) and Freckmann and Lelong (2002, 2003) treated D. lindheimeri as infraspecific in their widely encompassing D. acuminatum, but neither mentions the distinctiveness of habitat as evidence for separation. To illustrate this point, the present study collected habitat information from 70 herbarium labels of D. lindheimeri. Of these, two labels referred to the habitat as “dry”, six labels provided no indication of site moisture, and 62 labels (89 percent) indicated or implied saturated soil conditions. A similar review of D. lanuginosum labels indicated a predominance of dry soils.
40 Voss (1972) and Tony Reznicek of the Univeristy of Michigan (pers. comm.) note that along calcareous lake shores in the Great Lakes region, the morphological expression of what is here treated as D. lindheimeri approximates that of D. spretum (a known acidophile). These plants have the overall habit of D. spretum coupled with the pubescence characters of D. lindheimeri. Furthermore, their vernal inflorescences are open panicles and their spikelets are not pointed, in contrast to D. spretum which has narrow inflorescences and pointed spikelets. The inflorescences of D. spretum are conspicuously more densely flowered. These plants may represent an undescribed taxon within the group that is worthy of further investigation and help explain why Hitchcock and Chase (1910, 1951) placed D. lindheimeri in the informal group “Spreta” rather than “Lanuginosa”. The more typical morphology of D. lindheimeri in Michigan seems to be restricted to the southern portion of the lower peninsula.
REPRESENTITIVE SPECIMENS. CANADA. NEW BRUNSWICK: Carleton Co., 14 Jul 1910, M. Fernald & B. Long 12527 (GH); UNITED STATES. ARKANSAS: St. Francis Co., 19 Jun 1956, S. McDaniel 568 (MO); CALIFORNIA: Humbolt Co., 13 Jul 1915, A. Hitchcock 527 (MO); KENTUCKY: Caldwell Co., 27 May 1970, J. Conrad 1008 (MO); MICHIGAN: Emmet Co., 28 Jul 1922, F. Gates 13354 (MU); Washtenaw Co., 5 Jul 1935, F. Hermann 6839 (MO); MISSOURI: Carter Co., 16 May 2001, J. Thomas 1416 (private); Howell Co., 5 Jun 1999, J. Thomas 678 (private); NEW YORK: Albany Co., 23 Jun 1951, G. Van Schaak 3230 (MO); Cayuga Co., 25 Jul 1916, F. Metcalf 5539 (MO); TEXAS: F. Lindheimer 5651846 (NY); Gregg Co., 18 Jun 1901, J. Reverchon 2357 (MO)
Dichanthelium longiligulatum (Nash) Freckmann, Phytologia 48(1): 102. 1981. Panicum longiligulatum Nash, Bull. Torrey Bot. Club 26(11):574. 1899. Dichanthelium acuminatum var. longiligulatum (Nash) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1127. 1978. Panicum acuminatum var. longiligulatum (Nash) Lelong, Brittonia 36(3):270. 1984. Dichanthelium acuminatum subsp. longiligulatum (Nash) Freckmann & Lelong, SIDA 20(1):168. 2002. TYPE: USA. Florida: Franklin Co., Apalachicola, 1892, G.R. Vasey s.n. (holotype: NY!; isotype: US!)
Plants erect or ascending from a geniculate base, 30-70 cm tall; internodes glabrous; sheaths glabrous to glabrate; ligule 1.8-3.1 mm long (Figure 8a); pseudoligule absent; leaf blades spreading to ascending, 2-8 cm long, 2.0-6.0 mm wide, adaxial surface glabrous, abaxial surface puberulent; panicles 3-8 cm long, half as wide to as wide as
41 long, puberulent and often pilose; spikelets 1.1-1.5 mm long, approximately 0.5-0.8 mm wide, elliptic, puberulent to pilose; first glume 0.2-0.5 mm long, typically 1/3 as long as the spikelet, obtuse to acute.
DISTRIBUTION AND HABITAT (Figure 16). Dichanthelium longiligulatum is found in coastal regions from New Jersey to Texas, inland to Tennessee, south to Mexico, Central America and northern South America. It is also found in the islands of the Caribbean. The habitat of D. longiligulatum consists of pine savannas, sandy woodlands and various wetland community types.
NOTES. Dichanthelium longiligulatum can be distinguished from similar members of section Lanuginosa by its small stature, glabrous sheaths and the absence of puberulent hairs on the abaxial surfaces of vernal stem leaves. Its overall stature is reminiscent of D. spretum with which it was long aligned in the treatment of Hitchcock and Chase (1910, 1951). However, the spikelets of D. longiligulatum are smaller than those of D. spretum and are not as acute at the apex, the inflorescence is not narrowly paniculate as in D. spretum and D. longiligulatum has smaller leaf dimensions. Dichanthelium longiligulatum also resembles D. leucothrix, D. meridionale and D. wrightianum but differs from these three in its lack of puberulence on the vernal sheaths, internodes and adaxial leaf surfaces. Several authors (Freckmann and Lelong 2002; Freckmann and Lelong 2003; Gould and Clark 1978; Lelong 1984; Zuloaga et al. 1993b) have attempted to place D. longiligulatum as a variety or subspecies of D. acuminatum. However, D. longiligulatum is strikingly different from D. acuminatum. Dichanthelium longiligulatum has significantly smaller spikelets (1.1-1.5 mm vs. 1.6-2.1 mm), a longer ligule (1.8-3.1 mm vs. 0.4-1.3 mm), lacks the extensive marginal cilia that D. acuminatum has on its leaf blades, has glabrous sheaths, and possesses a light puberulence on the abaxial surfaces of vernal leaves. Furthermore, D. longiligulatum is generally smaller and much more diffusely branched in the autumnal form than D. acuminatum. In herbarium specimens D. longiligulatum often has a grey-green coloration.
42 REPRESENTITIVE SPECIMENS. NICARAGUA. Comarca Del Cabo, 12 Mar 1971, F. Seymour 4576 (MO); UNITED STATES. FLORIDA: Franklin Co., 1892, G. Vasey s.n. (NY); GEORGIA: Bulloch Co., 10 Jun 1901, R. Harper 839 (MO); Charlton Co., 26 Aug 1902, R. Harper 1575 (MO); NORTH CAROLINA: Henderson Co., July 1924, J. Davis s.n. (MO); TENNESSEE: Grundy Co. 23 Jun 1938, H. Svenson 9119 (MO)
Dichanthelium meridionale (Ashe) Freckmann, Phytologia 39(4): 270-271. 1978. Panicum meridionale Ashe, J. Elisha Mitchell Sci. Soc. 15:49. 1898. Panicum unciphyllum var. meridionale (Ashe) Scribn. & Merr., Rhodora 3(29):123. 1901. Panicum lindheimeri subvar. meridionale (Ashe) Farw., Amer. Midl. Naturalist 11(2):45. 1928. Panicum lanuginosum subvar. meridionale (Ashe) Farw., Pap. Michigan Acad. Sci. 26:5 1941. TYPE: USA. North Carolina: Burke Co., Blue Ridge, Jun 1893, W.W. Ashe s.n. (lectotype: US!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 210. 1910)
Panicum albemarlense Ashe, J. Elisha Mitchell Sci. Soc. 16(2):84. 1900. Panicum meridionale var. albemarlense (Ashe) Fernald, Rhodora 36(423):76. 1934. Dichanthelium meridionale var. albemarlense (Ashe) Mohlenbr., Ill. Fl. Illinois (ed. 2):381. 2001. TYPE: USA. North Carolina: Beaufort or Hyde Co., unknown locality, in open woods, well drained, 26 May 1899, W.W. Ashe s.n. (lectotype: US!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 220. 1910)
Panicum microphyllum Ashe, J. Elisha Mitchell Sci. Soc. 15:61. 1898. TYPE: USA. North Carolina: Orange Co., W.W. Ashe s.n. (lectotype: unknown, included in P. meridionale by Hitchcock and Chase, 1910, based on description; isolectotype: US!, NY!)
Plants erect or ascending from a geniculate base, 5-45 cm tall; internodes double invested with both short puberulent and long pilose pubescence (Figure 8d); sheaths double invested with short puberulent hairs below a pilose or villous pubescence (Figure 8d), longer hairs 1.7-3.8 mm long; ligule 1.7-4.0 mm long (Figs. 7a or d); pseudoligule absent or intermeshed with ligule; leaf blades spreading to ascending, 1.5-7.0 cm long, 2.0-6.0 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface long pilose, abaxial surface puberulent and occasionally pilose; panicles 1.5-5.0 cm long, half as wide to as wide as long, puberulent and pilose; spikelets 1.3-1.6 mm long, approximately 0.7-0.9 mm wide, elliptic, puberulent to pilose; first glume 0.3-0.8 mm long, typically 1/3 as long as the spikelet, obtuse to acute.
43 DISTRIBUTION AND HABITAT (Figure 17). Dichanthelium meridionale ranges from Nova Scotia to Minnesota and south to Alabama and Georgia. Its habitat consists of open woods and clearings, typically on sandy soils or shores.
NOTES. Gould and Clark (1978) placed D. meridionale under synonymy with D. acuminatum var. implicatum. Freckmann and Lelong (2002, 2003) make no mention of where D. meridionale fits into their revisions to section Lanuginosa, but the “Names and Synonyms” section of volume 25 of the “Flora of North America” lists it as a synonym of D. acuminatum subsp. implicatum. Neither treatment has provided any justification for placing D. meridionale into synonymy under D. acuminatum. Dichanthelium meridionale differs from D. implicatum in having a pronounced double vestiture of puberulent hairs beneath a longer pilosity. This is best demonstrated on the sheaths, internodes and abaxial leaf surfaces of the vernal stem. Because of its extensive double vestiture, D. meridionale is often confused with D. columbianum. The two can easily be distiguished by the smaller ligule of D. columbianum (0.7-1.5 versus 1.7-4.0 mm long).
REPRESENTITIVE SPECIMENS. UNITED STATES. MASSACHUSETTS: Barnstable Co., 3 Jul 1931, J. Churchill s.n. (MO); NEW YORK: Suffolk Co., 22 Jun 1916, A. Chase 526 (MO); NORTH CAROLINA: Beaufort Co., W. Ashe s.n. (US); Burke Co., W. Ashe s.n. (US); Orange Co., W. Ashe s.n. (NCU); Orange Co., W. Ashe s.n. (MO); TENNESSEE: Cannon Co., 20 May 1974, R. Kral 52796 (MO); Cannon Co., 20 May 1974, R. Kral 52800 (MO); VIRGINIA: Prince George Co., 30 May 1965, R. Freckmann 1565 (MO); Greensville Co., 18 Jun 1941, M. Fernald & B. Long 12940 (MO)
Dichanthelium ovale (Ell.) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1114. 1978. Panicum ovale Ell., Sketch Bot. S. Carolina 1(2):123. 1816. TYPE: USA. Georgia: Camden Co., St. Mary’s, no date, W. Baldwin s.n. (holotype: CHARL; fragment and photo, US!)
Panicum ciliiferum Nash, Bull Torrey Bot. Club 24(4):195-196. 1897. TYPE: USA. Florida: Lake Co., collected in the vicinity of Eustis, high pine land, 12-31 Mar 1894, G.V. Nash 147 (holotype: NY!; isotypes: US! MO!)
Panicum erythrocarpon Ashe, J. Elisha Mitchell Sci. Soc. 16(2):90. 1900. TYPE: USA. North Carolina: New Hanover Co., shady slopes on the sand hills one mile north of Wilmington, 17 May 1899, W.W. Ashe s.n. (holotype: NCU!; isotypes: NCU, US!)
44
Panicum wilmingtonense Ashe, J. Elisha Mitchell Sci. Soc. 16(2):86. 1900. TYPE: USA. North Carolina: New Hanover Co., shady slopes on the sand hills one mile north of Wilmington, 17 May 1899, W.W. Ashe s.n. (holotype: NCU!; isotype: US!)
Panicum benneri Fernald, Rhodora 46(541):2-3. 1944. TYPE: USA. New Jersey: Hunterdon Co., old field along the Delaware River ca. 1.5 miles east of Raven Rock, 7 Jun 1941, W.M. Benner 9635 (holotype: GH!; isotype: GH!, US, PH)
Plants erect or ascending from a geniculate base, 20-60 cm tall; lower-most internodes double invested with short puberulent to pilose hairs below a villous pubescence or occasionally lacking the longer villous hairs, the middle and upper internodes ascending to appressed pilose or villose, lacking the short puberulence of the lower internodes; lowermost sheaths double invested with short puberulent hairs below a pilose or villous pubescence (Figure 8d), upper sheaths pilose to villose and lacking puberulent hairs (Figure 8a), longer hairs 0.2-2.7 mm long; ligule 0.2-0.7 (-1.0) mm long (Figs. 7b or c); pseudoligule 2.0-4.0 mm long when present; leaf blades spreading to ascending, 3-11 cm long, 4-10 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface villose to glabrate, hairs absent along the cental third of the blade, abaxial surface with appressed puberulence or glabrate; panicles 4-9 cm long, half as wide to as wide as long, puberulent; spikelets 2.1-2.9 mm long, approximately 1.0 mm wide, elliptic, pilose; first glume 0.7-1.6 mm long, typically 2/5 to 1/2 as long as the spikelet, obtuse to broadly acute.
DISTRIBUTION AND HABITAT (Figure 18). Dichanthelium ovale is found along the coastal plain of North America from Massachusetts to Texas and south into Central America. Its habitat consists of sandy, open woods and clearings.
NOTES. All treatments involving what is now section Lanuginosa have kept D. ovale as a distinct species. However, other taxa within the group have often been assigned as infraspecific in D. ovale. Panicum addisonii (treated under D. columbianum in the present work) has often been treated as a variety of D. ovale. The treatment of Freckmann and Lelong (2002, 2003) relegated what is here treated as D. villosissimum
45 and D. praecocius to subspecies of D. ovale. Lelong (1984) also treated D. villosissimum under D. ovale (as Panicum ovale var. villosum). Interestingly, these are the only two treatments to bridge the gap between the traditional groups Lanuginosa and Columbiana (now both section Lanuginosa) of Hitchcock and Chase (1910, 1951) by combining taxa from each group. Lelong (1984) justifies this based on Gould and Clark’s (1978) placement of P. addisonii (including D. commonsianum) as a variety of D. ovale by stating that D. villosissimum is more like P. addisonii than D. ovale therefore D. villosissimum should probably also be a variety of D. ovale. However, the present study shows that P. addisonii is better placed under D. columbianum (for discussion of this see D. columbianum) and that D. villosissimum and D. praecocius demonstrate ample distinction for recognition as separate species.
Dichanthelium ovale differs from both D. villosissimum and D. praecocius in having larger spikelets, larger first glumes and a puberulent pubescence on the lower sheaths and internodes of the vernal culms. As with many members of the traditional group Columbiana, the hairs on the adaxial surfaces of vernal stem leaves of D. ovale are restricted to the blade edges, thus the central third of the adaxial leaf surface is glabrous. This is in stark contrast with both D. villosissimum and D. praecocius which possess a uniform pubescence across the adaxial leaf surfaces of vernal stem leaves. Furthermore, the true ligule of D. praecocius is significantly larger (typically 2.5-4.0 mm) than either D. villosissimum (0.3-1.3 mm) or D. ovale (0.2-0.7 mm). Freckmann and Lelong (2002) also saw fit to designate P. pseudopubescens as a subspecies of D. ovale. While this taxon has a convoluted nomenclature, it is best placed in the synonymy with D. villosissimum. For a discussion of these see notes under D. villosissimum. Nomenclatural confusion aside, the only taxon that is likely to be confused with D. ovale is D. commonsianum. For a discussion on the differences between these two closely related taxa see notes under D. commonsianum.
Panicum malacon has often been associated with D. ovale (Freckmann and Lelong 2003; Gould and Clark 1978; Hansen and Wunderlin 1988). The present treatment finds that P. malacon possesses little morphological resemblance to any
46 members of section Lanuginosa. Upon review of the type specimen, it seems better placed with members of section Angustifolia.
REPRESENTITIVE SPECIMENS. UNITED STATES. ALABAMA: Dekalb Co., 27 May 1974, R. Kral 53036 (MO); Henry Co., Oct 1972, R. Kral 48982 (MO); Monroe Co., 31 May 1972, R. Kral 46998 (MO); FLORIDA: May 1909, A. Chapman s.n. (MO); Duval Co., 24 Apr 1897, A. Curtiss 5866 (MO); Franklin Co., 16 May 1969, R. Godfrey 69494 (MO); Lake Co., 17-31 March 1894, G. Nash 147 (US); Lake Co., 13 May 1900, A. Curtiss 6616 (MO); Lee Co., Jul- Aug 1900, A. Hitchcock 474 (MO); Liberty Co., 19 Oct 1977, R. Kral 61184 (MO); GEORGIA: Romer s.n. (MO); Lee Co., 11 May 1940, W. Duncan 2317 (MO); MASSACHUSETTS: A. Chapman s.n. (MO); NEW JERSEY: Burlington Co., 16 Jun 1900, Parker s.n. (MO); Burlington Co.: 18 Jun 1933, F. Herman 4465 (MO); Hunterdon Co., 7 Jun 1941, W. Benner 9635 (GH); Monmouth Co., 26 Jun 1929, J. Churchill s.n. (MO); NORTH CAROLINA: Cumberland Co., 16 May 1976, J. Solomon s.n. (MO); New Hanover Co., 17 May 1899, W. Ashe s.n. (US); Sampson Co., 17 May 1966, R. Freckmann 2114 (MO); SOUTH CAROLINA: Aiken Co., 1 Jun 1867, H. Ravenel s.n. (MO); Aiken Co., 11 Jun 1867, H. Ravenel (MO); Aiken Co., 20 May 1899, H. Eggert s.n. (MO); Aiken Co., 21 May 1899, H. Eggert s.n. (MO); Aiken Co., Oct 1967, H. Ravenel s.n. (MO); Beaufort Co., 27 May 1916, A. Chase 7115 (MO); Beaufort Co., 6 May 1917, J. Churchill s.n. (MO); TENNESSEE: Wilson Co., Jun 1881, A. Gattinger 7387 (MO); VIRGINIA: Nansemond Co., 26-28 Jul 1939, M. Fernald & B. Long 10505 (MO); Northumberland Co., 1 Jun 1946, C. Reed 5136 (MO)
Dichanthelium praecocius (Hitchc. & Chase) Mohlenbr., Ill. Fl. Illinois (ed. 2) Grasses: Panicum to Danthonia: 383. 2001. Panicum praecocius Hitchc. & Chase, Rhodora 8(95):206-207. 1906. Panicum lanuginosum var. praecocius (Hitchc. & Chase) Dore, Le Naturaliste Canadien 103(6):562. 1976. Dichanthelium villosissimum var. praecocius (Hitchc. & Chase) Freckmann, Phytologia 39(4):270. 1978. Dichanthelium ovale subsp. praecocius (Hitchc. & Chase) Freckmann & Lelong, Sida 20(1):170. 2002. Panicum ovale var. praecocius (Hitchc. & Chase) Wipff, Sida 20(3):1044. 2003. TYPE: USA. Illinois: Stark Co., near Wady Petra, dry bank, 30 Jun 1900, V.H. Chase 649 (holotype: US!; isotype: NY!, GH!)
Plants erect or ascending from a geniculate base, 10-50 cm tall; internodes conspicuously to sparsely pilose to villose (Figure 8a), hairs ascending to appressed; sheaths copiously pilose to villose (Figure 8a), the hairs often darkened in hue, erect to ascending and (1.8-) 2.8–4.0 (-4.8) mm long; ligule (1.8-) 2.5-4.0 (-4.7) mm long (Figure 8a); pseudoligule lacking or indistinguishable from ligule; leaf blades ascending, 5-9 cm long, 3-7 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface villose often with dark hued hairs, abaxial surface pilose; panicles 4-6 cm long, half as wide to
47 as wide as long, densely to sparsely pilose, axillary inflorescences produced soon after terminal inflorescences; spikelets (1.6-) 1.8-1.9 (-2.1) mm long, approximately 0.8-1.2 mm wide, elliptic, pilose; first glume 0.6-1.0 mm long, typically 1/3 to 1/2 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 19). Dichanthelium praecocius ranges from southwestern Ontario to North Dakota in the north, east to Ohio, and south to Texas and Arkansas. The habitat of D. praecocius primarily consists of prairies but also includes glades, barrens and open woodlands. It becomes increasingly less common outside the Tallgrass Prairie Ecoregion. In contrast to D. lanuginosum, D. praecocius exhibits high fidelity to intact, undisturbed natural communities.
NOTES. Dichanthelium praecocius is unique in the timing of its secondary inflorescence production. Where other Dichanthelium produce a terminal inflorescences early in the growing season followed by axillary inflorescences late in the season, D. praecocius produces both terminal and axillary inflorescences early and continues to produce axilary inflorescences throughout the growing season. In overall morphology it is most similar to D. lanuginosum and D. implicatum. It differs from both in having larger spikelets, a longer first glume, longer hairs on the vernal sheaths, and a unique distribution of ligule hairs. Dichanthelium praecocius also tends to have narrower vernal leaves which are often more ascending or appressed than either D. lanuginosum or D. implicatum. The hairs on the sheaths and adaxial surfaces of D. praecocius often have a dark coloration and seem to be more brittle than other species. Because of this, late in the growing season the plants take on a “messy” appearance with numerous dark, broken and bent hairs. Dichanthelium praecocius tends to have fewer nodes on the central axis of the inflorescence (average of 7.7) than other members of the section.
Gould and Clark (1978) combined D. praecocius and D. villosissimum and treated the combination as a variety of D. acuminatum (D. acuminatum var. villosum). Freckmann (1978) treated D. praecocius as a variety of D. villosissimum. Given the present morphological examination, there does not appear to be any infraspecific
48 relationship between any of these three taxa. Both D. acuminatum and D. villosissimum have ligules less than 1.4mm long while that of D. praecocius exceeds 1.8mm long. Although D. acuminatum and D. villosissimum have similar ligule lengths, the ciliate margins and smaller spikelets of D. acuminatum and the dramatically longer vernal sheath pubescence of D. villosissimum clearly set each apart. Freckmann and Lelong (2002, 2003) treated both D. praecocius and D. villosissimum as subspecies of D. ovale. For discussion involving these taxa see notes under D. ovale. Populations at the northern and eastern edges of the range of D. praecocius are composed of larger, more densely pubescent plants that approximate D. villosissimum in terms of gross morphology. Said plants can be distinguished by their longer ligules and smaller spikelets. Plants in the central and western portions of the range have narrower, more appressed leaves and are less densely pubescent overall. These plants more closely resemble D. implicatum from which they can be distinguished by having larger spikelets and glumes and longer vernal sheath pubescence (see key for details).
REPRESENTITIVE SPECIMENS. UNITED STATES. ARKANSAS: Benton Co., Plank s.n. (MO); Benton Co., 18 Jun 2002, J. Thomas 1501 (MO); ILLINOIS: Champaign Co., 20 Jun 1906, H. Gleason s.n. (MO); Stark Co., 30 Jun 1900, V. Chase 649 (US); Stark Co., 10 Jun 1906, V. Chase 671 (MO); IOWA: Story Co., 18 Jun 1865, R. Freckmann 1927 (MO); KANSAS: Cloud Co., 15 Jun 1937, S. Fraser 796 (MO); MINNESOTA: Renville Co., 19 Jun 1940, G. Moore 13054 (MO); Sherburne Co., 6 Jul 1999, G. Swanson 1383 (MO); MISSOURI: Barton Co., 16 Jun 1990, G. Yatskievych & K. Yatskievych 90-147 (MO); Callaway Co., 12 Sep 1937, J. Steyermark 26220 (MO); Clark Co., 8 Jul 1909, B. Bush 5909A (MO); Dade Co., 9 Aug 2000, J. Thomas 1307 (MO); Dent Co., 30 Jun 1929, J. Kellogg 15078 (MO); Jasper Co., 5 Jun 1909, E. Palmer 2157B (MO); Jasper Co., 5 Jun 1909, E. Palmer 2144 (MO); NEBRASKA: Cuming Co., 31 May 1975, S. Churchill 5528 (MO); Lancaster Co., 15 Jun 1900, G. Hedgcock s.n. (MO); Lancaster Co., 4 Jun 1973, S. Churchill 941 (MO); Saunders Co., 25 Jun 1974, S. Churchill 3588 (MO); Stanton Co., 15 Jun 1976, S. Churchill 7709 (MO); Winneskiek Co., 12 Jun 1933, W. Tolstead s.n. (MO); OKLAHOMA: Creek Co., 11 May 1895, B. Bush 1220 (MO); Murray Co., U. Waterfall 6021 (MO); WISCONSIN: Columbia Co., 14 Jun 1942, L. Shinners & L. Shaw 4361 (MO); TEXAS: Harrison Co., 8 Aug 1907, B. Bush 632 (MO)
Dichanthelium spretum (Schult.) Freckmann, Phytologia 48(1): 102. 1981. Panicum spretum Schult., Mantissa 2:248. 1824. Dichanthelium lanuginosum var. spretum (Schult.) Harvill, Castanea 42(2):177. 1977. Dichanthelium acuminatum subsp. spretum (Schult.) Freckmann & Lelong, Sida 20(1):168. 2002. TYPE: USA. New
49 England: unknown locality, moist ground, no date, H.C. Muhlenberg 184 (holotype: PH; fragment and photo, US!)
Panicum nitidum var. densiflorum Rand & Redfield, Fl. Mt. Desert Isl. 174. 1894. Dichanthelium acuminatum var. densiflorum (Rand & Redfield) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1127. 1978. Panicum acuminatum var. densiflorum (Rand & Redfield) Lelong, Brittonia 36(3):270. 1984. TYPE: USA. Maine: Hancock Co., Mount Desert, shores of Ripples Pond, 28 Jul 1892, E.L. Rand s.n. (holotype: GH!; fragment, US)
Panicum eatoni Nash, Bull. Torrey Bot. Club 25:84. 1898. TYPE: USA. New Hampshire: Seabrook, unknown locality, 1897, A.A. Eaton s.n. (holotype: NY!; fragment and photo, US!)
Panicum paucipilum Nash, Bull Torrey Bot. Club 26:573. 1899. TYPE: USA. New Jersey: Cape May Co., at Wildwood, 30 & 31 May 1897, E.P. Bicknell s.n. (holotype: NY!; isotypes: NY!, US!)
Panicum octonodum Smith, U.S.D.A. Div. Agrost. Bull. 17:73. 1899. Panicum nitidum var. octonodum Scribn. & Merr., U.S.D.A. Div. Agrost. Bull. 24:34. 1901. TYPE: USA. Texas: Waller Co., Waller County line near Harris County, Jun 1898, F.W. Thurow 6 (holotype: US!)
DESCRIPTION. Plants erect or ascending from a geniculate base, 30-120 cm tall; internodes glabrous; sheaths glabrous; ligule 1.7-3.0 mm long (Figure 8a); pseudoligule absent; leaf blades spreading to ascending, 3-14 cm long, 3.0-9.0 mm wide, glabrous adaxially and abaxially; panicles 6-12 cm long, less than half as wide as long, glabrous to sparsely pilose; spikelets 1.3-1.9 mm long, approximately 0.7-0.9 mm wide, elliptic to narrowly ovate, pubescent or rarely glabrous; first glume 0.3-0.7 mm long, typically 1/4 to 1/3 as long as the spikelet, obtuse to acute.
DISTRIBUTION AND HABITAT (Figure 20). Dichanthelium spretum is found on the coastal plain of North America from Nova Scotia to Texas. It is also found within the Great Lakes Region. It is an acidophile and primarily inhabits saturated to seasonally saturated sandy to peaty soils along lake and bog margins or within coastal marshes and savannas. It is intolerant of natural community disturbance and may be of conservation concern where it is found.
50 NOTES. Once the cornerstone of Hitchcock and Chase’s (1910, 1951) informal group “Spreta”, D. spretum seems a bit out of place within section Lanuginosa. It differs from all other members of the section in having more acute spikelets, longer sheaths and internodes and strongly ascending panicle branches (narrow inflorescences). The most morphologically similar species is D. longiligulatum which is smaller in stature, has smaller spikelets and has a wider, more spreading inflorescence. Occasional specimens of D. spretum possess glabrous spikelets but this character does not appear to warrant taxonomic recognition. Along calcareous lake shores of the Great Lakes region, D. lindheimeri can loosely resemble D. spretum (see notes under D. lindheimeri for details).
REPRESENTITIVE SPECIMENS. UNITED STATES. TEXAS: Waller Co., Jun 1898, F. Thurow 6 (US); NEW JERSEY: Cape May Co., 30-31 1897, E. Bicknell s.n. (NY); NEW HAMPSHIRE: Rockingham Co., 1897, A. Eaton s.n. (NY); MASSACHUSETTS: Middlesex Co., 29 Jun 1896, J. Blankinship 109 (MO); CANADA, NOVA SCOTIA: Yarmouth Co., 23 Jul 1921, M. Fernald, B. Long & E. Bartram 23201 (MO); INDIANA: Porter Co., 27 Jun 1935, R. Tyron Jr. 3718½ (MO); Jasper Co., 12 Jul 1947, R. Friesner 21662 (MO)
Dichanthelium subvillosum (Ashe) Mohlenbr., Ill. Fl. Illinois (ed. 2) Grasses: Panicum to Danthonia: 383. 2001. Panicum subvillosum Ashe, J. Elisha Mitchell Sci. Soc. 16(2):86-87. 1900. TYPE: USA. Minnesota: Carlton Co., unknown locality, Aug 1899, W.W. Ashe s.n. (holotype: location unknown; fragment and photo US!; isotype: NCU!)
Panicum unciphyllum f. pilosum Scribn. & Merr., Rhodora 3(29):124. 1901. TYPE: USA. Maine: Penobscot Co., 7 Jul 1891, M.L. Fernald 501 (holotype: US!)
Plants erect or ascending from a geniculate base, 10-50 cm tall; lowermost internodes pilose to villose and often double invested with short puberulent to pilose hairs (Figure 8b); the middle and upper internodes pilose to villose (Figure 8a); lowermost sheaths copiously pilose to villose and usually double invested with short puberulent to pilose hairs (Figure 8b), upper sheaths pilose to villose and lacking puberulent hairs (Figure 8a), longer hairs 1.0-2.0 mm long; ligule 0.7-1.5 mm long (Figs. 7b or d); pseudoligule 1.5- 4.0 mm long, distinct or intermeshed with ligule; leaf blades spreading to ascending, 2-7 cm long, 3-7 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface villose to pilose, hairs uniformly distributed, abaxial surface pubescent to pilose;
51 panicles 2-6 cm long, half as wide to as wide as long, pilose; spikelets 1.6-2.0 mm long, approximately 1.0 mm wide, elliptic, pilose; first glume 0.7-1.0 mm long, typically 1/3 to 1/2 as long as the spikelet, obtuse to broadly acute.
DISTRIBUTION AND HABITAT (Figure 21). Dichanthelium subvillosum is sparsely distributed from Nova Scotia to Saskatchewan in the north, south to New York and Iowa. Its habitat consists of sandy prairies and open woods.
NOTES. Though they provided no explanation, Gould and Clark (1978) placed D. subvillosum in synonymy with D. acuminatum var. acuminatum. One must assume this placement is based on the superficial similarity of D. subvillosum to P. lanuginosum var. fasciculatum (D. lanuginosum in the present treatment), which was also placed in D. acuminatum, rather than any direct similarity to actual D. acuminatum. Freckmann and Lelong (2002, 2003) took a more conservative approach and elevated var. fasciculatum to subsp. fasciculatum under D. acuminatum and placed D. subvillosum as a synonym of this new combination. The present study demonstrates that D. subvillosum possesses several morphological characters that warrant specific recognition for this taxon. It differs from D. acuminatum var. fasciculatum (D. lanuginosum in the present treatment) in having a shorter true ligule (0.7-1.5 mm versus 2.0-5.0 mm long), the presence of a distinct pseudoligule, longer first glume (0.7-1.0 mm versus 0.3-0.8 mm long), and less autumnal branching which is more limited to the base and lower sheaths in D. subvillosum. Furthermore, D. subvillosum differs strikingly from D. acuminatum in that D. subvillosum lacks ciliate leaf margins, is more sparsely pubescent and has longer first glumes in relation to overall spikelet length (first glumes typically 1/3 to 1/2 the spikelet length in D. subvillosum versus first glumes typically 1/4 to 1/3 the spikelet length in D. acuminatum). The ranges of D. subvillosum and D. acuminatum are also distinct with D. subvillosum occuring in northern North America and D. acuminatum occurring in extreme southern North America and southward.
52 REPRESENTITIVE SPECIMENS. UNITED STATES. OHIO: Lucas Co., 17 Jun 1940, R. Kriebel 8731 (MO); MAINE: Oxford Co., 14 Jun 1931, F. Hunnewell 11973 (MO); MICHIGAN: Keweenaw Co., 14 Jul 1956, H. Bennett s.n. (MO); MINNESOTA: Anoka Co., 30 Jun 1888, J. Schuette s.n. (MO); Carlton Co., 6 Aug 1899, W. Ashe s.n. (NCU); NEW YORK: Franklin Co., 22 Jun 1930, W. Muenscher & B. Maguire 933 (MO); WISCONSIN: Ashland Co., 1 Jul 1976, B. Coffin 41 (MO); Oneida Co., 29 Aug 1918, R. Coffmann s.n. (MO)
Dichanthelium thermale (Bol.) Thomas & Vincent, comb. nov. in ed., Panicum thermale Bol., Proc. Calif. Acad. Sci. 2:181. 1862. Dichanthelium lanuginosum var. thermale (Bol.) Spellenb., Madrono 23(4):151. 1975. Dichanthelium acuminatum var. thermale (Bol.) Freckmann, Phytologia 48(1):107. 1981. Dichanthelium acuminatum subsp. thermale (Bol.) Freckmann & Lelong, Sida 20(1):168. 2002. TYPE: USA. California: Sonoma Co., unknown locality, [on hot rocks and in hot water flowing from Geyser springs and Geyser mountains, in the northern part of Sonoma County], no date, H.N. Bolander 3941 (holotype: GH!; isotypes: MO, US)
Plants erect or ascending from a geniculate base, 6.5-42 cm tall; internodes of the lower stem double invested with short, often sparse, pilose hairs below a villose pubescence or conspicuously to sparsely pilose (Figs. 7a or b), mid-stem and upper-stem internodes single invested with velutinose to villose pubescence (Figure 8a); sheaths conspicuously velutinose to villose, hairs 0.4–3.5 mm long (Figure 8a), the lower stem sheaths singly or double invested with short, pilose hairs (Figure 8b); ligule 0.6-3.5 mm long (Figs. 7a, b or d); pseudoligule distinct, lacking or intermeshed with ligule, 2-4 mm long; leaf blades spreading to ascending, 2-12 cm long, 4-12 mm wide, margins typically ciliate for no more than ¼ the blade length, adaxial leaf surface pilose to densely villose or velutinose, abaxial surface velutinose; panicles 4-9 cm long, half as wide to as wide as long, densely to sparely pilose to velutinose; spikelets 1.4-2.0 mm long, approximately 0.5-1.0 mm wide, elliptic, pilose; first glume 0.4-1.0 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
Dichanthelium thermale subsp. thermale
Panicum lassenianum Schmoll, Madrono 5(3):95-96. 1936. TYPE: USA. California: Plumas Co., Hot Spring Valley, Devil’s Kitchen, 6200 ft., 6 Jun 1910, W.L. Jepson 4082 (holotype: US!; isotype: UC)
Plants erect or ascending from a geniculate base, 12-42 cm tall; internodes of the lower stem double invested with short, often sparse, pilose hairs below a villose pubescence
53 (Figure 8b), mid-stem and upper-stem internodes single invested with velutinous to villose pubescence (Figure 8a); sheaths conspicuously velutinous to villose, hairs 0.4–2.4 mm long (Figure 8a), the lower stem sheaths double invested with short, pilose hairs (Figure 8b); ligule 0.6-1.5 mm long (Figs. 7b or d); pseudoligule usually distinct, 2-4 mm long; leaf blades spreading to ascending, 2-12 cm long, 4-8 mm wide, margins typically ciliate for no more than ¼ the blade length, adaxial leaf surface densely villose to velutinose, abaxial surface velutinose; panicles 4-9 cm long, half as wide to as wide as long, pilose to velutinose; spikelets 1.4-2.0 mm long, approximately 0.5-1.0 mm wide, elliptic, pilose; first glume 0.5-1.0 mm long, typically equal or less than 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 22). Dichanthelium thermale subsp. thermale is only found in the warm, mineral rich soils of isolated hot springs in the Cascade Mountains of northern California.
NOTES. Gould and Clark (1978) placed D. thermale in synonymy with D. acuminatum, while Freckmann and Lelong (2003) treated D. thermale as a subspecies of D. acuminatum (subsp. thermale). Given the disjunct range, specialized habitat and morphological autonomy of D. thermale, these treatments prove unsatisfying. Morphologically, D. thermale differs from D. acuminatum in having limited marginal cilia on the vernal leaf blades, and by possessing a double vestiture on the culms and sheaths of the vernal stem that D. acuminatum lacks. There is significant morphological variation in leaf size, pubescence density and spikelet length throughout the range of D. thermale (Hitchcock and Chase 1910). This could be attributed to inbreeding or genetic drift associated with their isolated populations and the island-like nature of their habitats. Schmoll (1939) attempted to sort out the variation in the D. thermale complex. In so doing, she limited D. thermale to the hot springs in Sonoma County California, described a new species (P. lassenianum) for plants occurring in Plumas County (California), described a new species (P. ferventicola) from Yellowstone National Park (Wyoming) and described two new varieties of P. ferventicola (vars. papillosum and sericeum); one in the vicinity of Banff, Alberta and another in Yellowstone National Park. The present
54 work could find no meaningful difference between what she called P. thermale and P. lassenianum, nor was any appreciable difference noted between what she called P. ferventicola and P. ferventicola var. papillosum and var. sericeum. Thus, P. lassenianum of Schmoll (1939) is here relegated to D. thermale. Panicum ferventicola and varieties are here treated as D. thermale subsp. sericeum. However, due to the scarcity of collections and the authors admitted unfamiliarity with the region, the present treatment is not entirely satisfying. The type specimen of P. ferventicola seems rather intermediate between the D. thermale of California and subsp. sericeum of the Rocky Mountains. It is because of this similarity in morphology and habitat that subsp. sericeum is linked to D. thermale instead of D. acuminatum as other authors have contended (Gould and Clark 1978; Freckmann and Lelong 2003). It is tempting to follow the treatment of Hitchcock and Chase (1951) that lumped those taxa described by Schmoll (1939) into D. thermale (then P. thermale), but this would be significantly less satisfying given the obvious differences in D. thermale and D. thermale subsp. sericeum. Essentially, the present work serves to provide a basic taxonomic framework that well represents the larger trends in morphological variation for the D. thermale complex. In order to more fully elucidate the ultimate taxonomy of these scattered and isolated groups, detailed population, morphological or molecular investigations will be necessary.
Recent research (Redman et al., 2002) has demonstrated that D. thermale possesses a fungal endophyte of the genus Curvularia. This endophyte equips D. thermale with the heat tolerance proteins and enzymes that allow it to tolerate soil temperatures in excess of 50ºC (Redman et al., 2002). Dichanthelium thermale closely resembles D. subvillosum; a species of northern North America.
REPRESENITIVE SPECIMENS. UNITED STATES. CALIFORNIA: Plumas Co., 6 Jun 1910, W. Jepson 4082 (US); Shasta Co., Jun 1899, L. Greata s.n. (US); Sonoma Co., 1865, H. Bolander 3991 (MO); Sonoma Co., 1866, H. Bolander 3941 (GH); Sonoma Co., 27 Aug 1872, J. Redfield 9792 (MO); Sonoma Co., 5 Jul 1931, M. Jones 29210 (MO)
Dichanthelium thermale subsp. sericeum (Schmoll) Thomas & Vincent, comb. nov. in ed.. Panicum ferventicola var. sericeum Schmoll, Madrono 5(3):93-94.
55 1939. Dichanthelium lanuginosum var. sericeum (Schmoll) Spellenb., Madrono 23(4):150. 1975. Dichanthelium acuminatum subsp. sericeum (Schmoll) Freckmann & Lelong, Sida 20(1):168. 2002. TYPE: USA. Wyoming: Yellowstone National Park, Mammoth Hot Springs, in wet ground about hot springs, 21 Jul 1889, A. Nelson & E. Nelson 6037 (holotype: RM!; isotypes: NY, US!)
Panicum ferventicola Schmoll, Madrono 5(3):92-95: 1939. TYPE: USA. Wyoming: Park Co., Yellowstone National Park, 5 miles north of Norris Geyser Basin, 10 Aug 1908, A. Chase 5252 (holotype: GH!; isotypes: F!, NY, US!)
Panicum ferventicola var. papillosum Schmoll, Madrono 5(3):94-95. 1939. TYPE: CANADA. Alberta: Banff, on a declivity wet with water from hot spring, 31 Jul 1914, A.S. Hitchcock 11511 (holotype: GH!; isotypes: F!, NY, US!)
Plants erect to ascending from a geniculate base or forming prostrate mats, 6.5-32 cm tall; internodes conspicuously to sparsely pilose (Figure 8a); sheaths copiously pilose to velvety villose (Figure 8a), hairs 2.0-3.5 mm long; ligule 2.0-3.5 mm long (Figs. 7a or d); pseudoligule lacking or intermeshed with ligule; leaf blades spreading to ascending, 3-6 cm long, 5-12 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface pilose to villous with the longest hairs typically 1.5-4.0 mm long, abaxial surface short pilose with the longest hairs typically 0.5-1.5 mm long; panicles 3-6 cm long, as wide as long, densely to sparsely pilose; spikelets 1.6-1.8 mm long, 0.7-1.0 mm wide, elliptic, pilose; first glume 0.4-0.7 mm long, typically 1/4 to 1/5 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 23). Dichanthelium thermale subsp. sericeum is restricted to the margins of hot springs and geysers in the Rocky Mountains. Verified populations occur in the vicinity of Banff, Alberta and Yellowstone National Park, Wyoming.
NOTES. A general lack of specimens and a high degree of morphological variation due to isolated populations makes this a difficult taxon to define. However, variations in morphology from the typical subspecies warrant some level of taxonomic recognition (see notes under D. thermale). Dichanthelium thermale subsp. sericeum differs from the typical subspecies in having a long pilose pubescence, a single ligule and unusually short and wide leaves. Further study involving population level analyses from multiple
56 locations within the range of D. thermale will be necessary to fully understand the morphological expression of the D. thermale complex.
REPRESENTITVE SPECIMENS. CANADA, ALBERTA: Banff, 31 Jul 1914, A. Hitchcock 220 (GH); UNITED STATES. WYOMING: Teton Co., 21 Jul 1899, A. Nelson & E. Nelson 6037 (GH); Teton Co., 10 Aug 1908, A. Chase 137 (GH)
Dichanthelium villosissimum (Nash) Freckmann, Phytologia 39(4): 270. 1978. Panicum villosissimum Nash, Bull. Torrey Bot. Club 23:149. 1896. Dichanthelium lanuginosum var. villosissimum (Nash) Gould, Brittonia 26(1):60. 1974. Dichanthelium ovale subsp. villosissimum (Nash) Freckmann & Lelong, SIDA 20(1):170. 2002. TYPE: USA. Georgia: Bibb Co., in the Ocmulgee River swamp, below Macon, 18-24 May 1895, J.K. Small s.n. (holotype: NY!; isotypes: NY!, US)
Panicum nitidum var. villosum A. Gray, N. Amer. Gram. 2:111. 1835. Panicum dichotomum var. villosum (A. Gray) Vasey, Dept. Agric. Bot. Div. Bull. 8:31. 1889. Dichanthelium acuminatum var. villosum (A. Gray) Gould & Clark, Ann. Missouri Bot. Gard. 65(4):1124. 1978. Panicum acuminatum var. villosum (A. Gray) Beetle, Phytologia 48(2):192. 1981. Panicum ovale var. villosum (A. Gray) Lelong, Brittonia 36(3):272. 1984. TYPE: USA. District of Columbia: Rock Creek, near Pierce’s Mill, 1 Jul 1883, G.R. Vasey s.n. (lectotype: US!)
Panicum pseudopubescens Nash, Bull. Torrey Bot. Club 26(11):577. 1899. Panicum villosissimum var. pseudopubescens (Nash) Fernald, Rhodora 36(423):79. 1934. Panicum ovale var. pseudopubescens (Nash) Lelong, Brittonia 36(3):271. 1984. Dichanthelium villosissimum var. pseudopubescens (Nash) Mohlenbr., Erigenia 6:26. 1985. Dichanthelium ovale subsp. pseudopubescens (Nash) Freckmann & Lelong, Sida 20(1):170. 2002. TYPE: USA. Alabama: Lee Co., collected at Auburn, 7 May 1898, F.S. Earle & C.F. Baker 1537 (holotype: NY!; isotype: US!)
Panicum atlanticum Nash, Bull. Torrey Bot. Club 24(7):346-347. 1897. TYPE: USA. New York: Kings Co., on dry somewhat shaded knolls in the grounds of the New York Botanical Garden, 24 Jun 1897, G.V. Nash s.n. (holotype: NY!; isotype: US!)
Panicum haemacarpon Ashe, J. Elisha Mitchell Sci. Soc. 15:55. 1898. TYPE: USA. District of Columbia: in the vicinity of Washington D.C., 6 Jun 1897, T.H. Kearney s.n. (lectotype: NY!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 233. 1910; isolectotypes: US!, NCU!)
57 Panicum xanthospermum Scribn. & Mohr, Contr. U.S. Natl. Herb. 6:348. 1901. TYPE: USA. Alabama: Butler Co., Greenville, collected in open sandy soil, 8 May 1898, C.T. Mohr s.n. (holotype: US!; isotype: NY)
Plants erect or ascending from a geniculate base, 10-60 cm tall; internodes conspicuously to sparsely villose to long pilose (Figure 8a), hairs ascending to appressed; sheaths copiously villose to long pilose, rarely double invested with short pilose pubescence below a long villose pubescence (Figs. 7a or b), erect to ascending and 1.4-3.8 mm long; ligule 0.3-1.3 mm long (Figure 8b); pseudoligule distinct, 2.0-5.0 mm long; leaf blades spreading to ascending, 3-11 cm long, 3-12 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface copiously villose to sparsely pilose with uniformly distributed hairs, abaxial surface pilose; panicles 4-10 cm long, half as wide to as wide as long, densely to sparsely pilose to villose; spikelets 1.8-2.5 mm long, approximately 0.9- 1.2 mm wide, elliptic, pilose; first glume (0.6-) 0.8-1.0 (-1.3) mm long, typically 1/4 to 1/3 the length of the spikelet, acute to broadly acute.
DISTRIBUTION AND HABITAT (Figure 24). Dichanthelium villosissimum is found throughout eastern North America from Massachusetts, southern Ontario and Minnesota, south to Texas and into Central America. It is typically found in acidic soils of sand or chert substrate. Throughout its range it is associated with dry pine or oak-hickory forests and woodlands.
NOTES. The most striking features of Dichanthelium villosissimum are its density and length of pubescence. This is especially noticeable on the sheaths and culms of the vernal stem where hairs are often in excess of 3.0 mm long. Past treatments (Ashe 1898; Braun 1967; Deam 1940; Fernald 1950; Gleason and Cronquist 1991; Hitchcock and Chase 1910; Hitchcock and Chase 1951; Jones 1945; Mohlenbrock 1986; Pohl 1947; Radford et al. 1968; Small 1933; Steyermark 1963; Stone 1911; Swink and Wilhelm 1994) involving section Lanuginosa have maintained D. villosissimum as a distinct species, subspecies or variety. Gould and Clark (1978) treated it as a variety of D. acuminatum (D. acuminatum var. villosum). Beetle (1981) and Zuloaga (1993b) treated it as a variety of P. acuminatum (P. acuminatum var. villosum). However, the present
58 work demonstrates ample morphological resolution to warrant distinction at the rank of species. Dichanthelium villosissimum differs from D. acuminatum in having significantly larger spikelets, longer sheath pubescence and a lack of extensive marginal cilia along the blades of the vernal leaves. Freckmann and Lelong (2002, 2003) placed D. villosissimum as a subspecies of D. ovale. However, the evidence provided in the present treatment suggests that it is best treated as a distinct species. For a discussion of the differences between D. ovale and D. villosissimum see the notes under D. ovale and D. praecocius. Freckmann and Lelong (2002, 2003) also placed P. pseudopubescens as a subspecies of D. ovale and differentiate it from subsp. villosissimum (D. villosissimum in the present treatment) as having shorter sheath pubescence that is ascending or appressed instead of spreading or retrose. They further state that subsp. pseudopubescens integrades morphologically with subsp. villosissimum. Though the type specimen of P. pseudopubescens does have shorter sheath pubescence than typical D. villosissimum, the present study could find no meaningful morphological separation between the two. Nor was any correlation between sheath pubescence length and orientation found. Thus P. pseudopubescens is here placed in synonymy with D. villosissimum. Deam’s (1929) treatment of Indiana grasses refers to the upper leaf surfaces of P. pseudopubescens as being pubescent only along the margins. However, the entity to which he is referring is P. pseudopubescens Deam (an illegitimate name) not P. pseudopubescens Nash. The type specimen of Deam’s P. pseudopubescens easily fits into D. commonsianum of the present treatment. This has led to numerous misidentification of Midwestern D. commonsianum as P. pseudopubescens. The leaf character to which Deam (1929) referred is limited to three taxa within the section (D. columbianum, D. ovale, and D. commonsianum). Morphologically, D. villosissimum is closely aligned with and easily confused with D. praecocius. For a discussion of the differences see notes under D. praecocius.
REPRESENTITIVE SPECIMENS. GUATAMALA. 13 Sep 1896, E. Seler 3235 (MO); HONDURAS. INTIBUCA: 21 Jun 1994, G. Davidse 35207 (MO); MEXICO. CHIAPAS: 5 Nov 1981, D. Breedlove 54707 (MO); UNITED STATES. ALABAMA. Butler Co., 8 May 1898, C. Mohr s.n. (US);
59 Dekalb Co., 27 May 1974, R. Kral 53030b (MO); Lee Co., 7 May 1898, F. Earl & C. Baker 1537 (NY); Pickens Co., 21 May 1960, S. McDaniel 1839 (MO); ARKANSAS: Garland Co., 18 Aug 1937, D. Demaree 15843 (MO); DISTRICT OF COLUMBIA: 6 Jun 1897, H. Kearney Jr. s.n. (US); 12 Jun 1894, T. Holmgren s.n. (MO); GEORGIA: Bibb Co., 18-24 May 1895, J. Small s.n. (NY); Dade Co., 8 May 1948, A. Cronquist 5164 (MO); Murray Co., 18 Jul 1973, R. Kral 50687 (MO); Screven Co., 17 Apr 1948, A. Cronquist 5006 (MO); ILLINOIS: Cook Co., 21 Jun 1891, W. Moffatt s.n. (MO); Henderson Co., H. Patterson s.n. (MO); IOWA: Jones 34 (MO); MISSISSIPPI: 1858, E. Higland s.n. (MO); MISSOURI. Benton Co., 24 May 1936, J. Steyermark 10744 (MO); Carter Co., 7 Jun 2001, J. Thomas 1399 (private); Ozark Co., 1 Jul 1937, J. Steyermark 22589 (MO); Barry Co., 6 Jun 1897, B. Bush 141 (MO); NEW JERSEY: Passaic Co., 14 Jul 1907, K. MacKenzie 2724 (MO); NEW YORK: Bronx Co., 24 Jun 1897, G. Nash s.n. (NY); Seneca Co., 24 Jun 1914, L. Randolf 11249 (MO); NORTH DAKOTA: Richland Co., 27 Jun 1940, O. Stevens 459 (MO); SOUTH CAROLINA: Bamberg Co., 17 May 1966, R. Freckmann 2133 (MO); Williamsburg Co., 10 Jul 1939, R. Godfrey 428 (MO); TENNESSEE: Hardeman Co., 14 May 1920, E. Palmer 19493 (MO); Knox Co., Jun 1896, A. Ruth s.n. (MO); Knox Co., Aug 1894, A. Ruth s.n. (MO); TEXAS: Gregg Co., 18 Jun 1907, J. Reverchon s.n. (MO); Polk Co., 10 May 1973, F. Gould & S. Hatch 14223 (MO); VIRGINIA: Alexandria Co., 9 Jun 1905, A. Chase 555 (MO); Princess Ann Co., 3 Sep 1905, K. MacKenzie 1710 (MO); Norfolk Co., 11 Jun 1940, M. Fernald & B. Long 11969 (MO)
Dichanthelium wrightianum (Scribn.) Freckmann, Phytologia 48(1): 101. 1981. Panicum wrightianum Scribn., U.S.D.A. Div. Agrostol. Bull 11:44. 1898. Dichanthelium acuminatum var. wrightianum (Scribn.) Gould & Clark., Ann. Missouri Bot. Gard. 65(4):1126. 1978. Panicum acuminatum var. wrightianum (Scribn.) Reed, Phytologia 80:284. 1996. TYPE: CUBA. unknown locality, 1865, C. Wright 3463, (lectotype: US!, designated by Hitchcock and Chase, Contr. U.S. Natl. Herb. 15: 210. 1910; isolectotypes: GH, MO!, NY, US)
Panicum deminutivum Peck, New York State Mus. Bull. 10:27. 1907. TYPE: USA. New York: Suffolk Co., Wading River, 20 Aug 1906, C.H. Peck s.n. (holotype: NY!; isotypes: NY!, US)
DESCRIPTION. Plants erect or ascending from a geniculate base, 5-40 cm tall; internodes puberulent and often pilose (Figs. 7c or d); sheaths double invested with short puberulent hairs below a pilose or villous pubescence (Figure 8d), the upper merely puberulent (Figure 8c), longer hairs 0.1-0.8 mm long; ligule 1.7-3.1 mm long (Figure 8a); pseudoligule absent or indistinguishable from ligule; leaf blades spreading to ascending, 1.5-4.5 cm long, 2.0-4.5 mm wide, margins ciliate only at the base of blade or eciliate, adaxial leaf surface puberulent, pilose or both, abaxial surface puberulent; panicles 1.5- 6.0 cm long, half as wide to as wide as long, puberulent; spikelets 0.8-1.0 mm long,
60 approximately 0.4-0.6 mm wide, elliptic, puberulent; first glume 0.2-0.4 mm long, typically 1/3 as long as the spikelet, obtuse to acute.
DISTRIBUTION AND HABITAT (Figure 25). Dichanthelium wrightianum ranges along the coastal plain of North America from Massachusetts to Florida and Texas, and south through Central America and northern South America. It is also found in the Caribbean Islands. Its habitat consists of saturated to seasonally saturated bogs, savannas and open woods with sandy, peaty or mucky soils.
NOTES. Given its tiny spikelets and diminuitive stature, there is little debate that D. wrightianum is a distinct species. Morphologically, it is closely aligned with D. leucothrix, D. meridionale and D. longiligulatum. These four taxa are smaller than others in the section, have relatively small spikelets and possess varying degrees of puberulence often under a longer pilosity. The extremely small spikelets of D. wrightianum easily distinguish it from the others. Gould and Clark (1978) placed D. wrightianum as a variety of D. acuminatum, yet provided no direct evidence for this lumping. Every treatment before and since Gould and Clark (1978) has maintained D. wrightianum as a distinct species.
REPRESENTITIVE SPECIMENS. CUBA. 1860-1864, C. Wright 3463 (MO); UNITED STATES. GEORGIA: Lowndes Co., 10 Jul 1968, W. Faircloth 5282 (MO); FLORIDA: Suwannee Co., 7 Jun 1900, A. Curtiss 6652 (MO); MASSACHUSETTS: Barnstable Co., 24 Jun 1909, C. Knowlton s.n. (MO); MISSISSIPPI: Jackson Co., 6 May 1898, S. Tracy 4885 (MO); NEW YORK: Delaware Co., 20 Aug 1906, C. Peck s.n. (US); NORTH CAROLINA: L. Bosc s.n. (US); VIRGINIA: Sussex Co., 6 Jul 1942, M. Fernald & B. Long 14267 (MO)
61 LITERATURE CITED
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62 Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 25, pp. 406-450. Giussani, L.M., J.H. Cota-Sanchez, F. Zuloaga, and E.A. Kellogg. 2001. A molecular phylogeny of the grass subfamily Panicoideae (Poaceae) shows multiple origins of C4 photosynthesis. American Journal of Botany 88: 1993-2012. Gleason, H.A. and A. Cronquist. 1991. Manual of the Vascular Plants of Northeastern United States and Adjacent Canada. 2nd ed. New York Botanical Garden, Bronx. Gould, F.W. 1974. Nomenclatorial changes in the Poaceae. Brittonia 26: 59-60. Gould, F.W. and C.A. Clark. 1978. Dichanthelium (Poaceae) in the United States and Canada. Annals of the Missouri Botanical Garden. 65: 1088-1132. Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas, Lawrence, Kansas. Hansen, B.F. and R.P. Wunderlin. 1988. Synopsis of Dichanthelium (Poaceae) in Florida. Annals of the Missouri Botanical Garden. 75: 1637-1657. Hitchcock, A.S. and M.A. Chase. 1910. Contributions from the U.S. Herbarium, vol 15, North American Species of Panicum. Government Printing Office, Washington, D.C.. Hitchcock, A.S. and M.A. Chase. 1951. Manual of the Grasses of the United States. United States Printing Office, Washington, D.C.. Jones, G.N. 1945. Flora of Illinois. The University Press, Notre Dame, Indiana. Lelong, M.G. 1984. New combinations for Panicum subgenus Panicum and Dichanthelium (Poaceae) of the southeastern United States. Brittonia 36: 262-273. Lelong, M.G. 1986. A taxonomic treatment of the genus Panicum (Poaceae) in Mississippi. Phytologia 61:251-269 Linnaeus, C. 1753. Species Plantarum. Vol. 1. Laurentius Salvius, Stockholm. Facsimile reprinted by the Ray Society. London. Mohlenbrock, R.H. 1986. Guide to the Vascular Flora of Illinois. Southern Illinois University, Carbondale, Illinois. Morrone, O. and F.O. Zuloaga. 1991. Estudios morfológicos en el subgénero Dichanthelium de Panicum (Poaceae: Panicoideae: Paniceae), con especial referencia a Panicum sabulorum. Annuals of the Missouri Botanical Garden. 78:915-927. Morrone, O., S.S. Denham, S.S. Aliscioni and F.O. Zuloaga. 2008. Parodiophyllochloa, a new genus segregated from Panicum (Paniceae, Poaceae) based on morphological and molecular data. Systematic Botany 33(1): 66-76. Naczi, R.F., A.A. Reznicek and B.A. Ford. 1998. Morphological, geographical, and ecological differentiation in the Carex willdenowii complex (Cyperaceae). American Journal of Botany 85(3): 434-447. Peterson, P.M., J. Cayouette, Y.S.N. Ferdinandez, B. Coulman, and R.E. Chapman. 2002. Recognition of Bromus richardsonii and B. ciliatus: evidence from morphology, cytology, and DNA fingerprinting (Poaceae: Bromeae). Aliso 20: 21-36. Pohl, R.W. 1947. A taxonomic study on the grasses of Pennsylvania. American Midland Naturalist 38: 513-600. Radford, A.E., H.E. Ahles, and C.R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. University of North Carolina Press, Chapel Hill, North Carolina. Rohlf, J.F. 2002. NTSYS-pc version 2.10t: Numerical Taxonomy and Multivariate
63 Analysis System. version 2.10t. Exeter Software, Setauket, NY. Rothrock, P.E. and A.A. Reznicek. 2001. The taxonomy of the Carex bicknellii group (Cyperaceae) and new species for central North America. Novon 11: 205-228. Schmoll, H.M. 1939. A realignment of the Panicum thermale group. Madroño 5(3): 90- 96. Scoggan, H.J., 1978. The Flora of Canada. Part 2. National Museum of Natural Sciences Publications in Botany 7, Ottawa. Shinners, L.H. 1944. Notes on Wisconsin grasses IV: Leptoloma and Panicum. American Midland Naturalist 32: 164-180. Small, J.K. 1903. Flora of the Southeastern United States. Published by Author, New York, New York. Small, J.K. 1933. Manual of the Southeastern Flora. Published by Author, New York, New York. Smith, B.N. and W.V. Brown. 1973. The Kranz syndrome in the Gramineae as indicated by carbon isotopic ratios. American Journal of Botany 60: 505-513. Steyermark, J.A. 1963. Flora of Missouri. Iowa State University Press, Ames, Iowa. Stone, W. 1911. The plants of southern New Jersey with especial reference to the flora of the pine barrens and the geographical distribution of the species. Annual Report of the New Jersey State Museum for 1910. 22-828. Strausbaugh, P.D. and E.L. Core. 1978. Flora of West Virginia. Seneca Books Inc., Morgantown, West Virginia. Swink, F. and G. Wilhelm. 1994. Plants of the Chicago Region. The Morton Arboretum, Lisle, Illinois. Webster, R.D. 1988. Genera of the North American Paniceae (Poaceae:Panicoideae). Systematic Botany 13: 576-609. Weishaupt, C.G. 1968. Vascular Plants of Ohio. Wm. C. Brown Company Publishers, Dubuque, Iowa. Yatskievych, G. 1999. Steyermark's Flora of Missouri. Vol. 1. Missouri Dept. of Conservation, Jefferson City, MO. Voss, E.G. 1972. Michigan Flora. Part 1. Cranebrook Institute of Science, Bloomfield Hills, Michigan Zuloaga, F.O., J. Dubcovshy and O. Morrone. 1993a. Infrageneric phenetic relations in New World Panicum (Poaceae: Panicoideae: Paniceae) numerical analysis. Canadian Journal of Botany 71. 1312-1327. Zuloaga, F.O., R.P. Ellis and O. Morrone. 1993b. A revision of Panicum subg. Dichanthelium sect. Dichanthelium (Poaceae: Panicoideae: Paniceae) in Meso America, the West Indies, and South America. Annals of the Missouri Botanical Garden 80: 119-190.
64 Table 1. Summary of past taxonomic treatments for all taxa involved in section Lanuginosa.
Hitchcock and Chase, 1910 Gould and Clark, 1978 Freckmann and Lelong, 2003
Group Columbiana No subgeneric rank given Section Lanuginosa Panicum addisonii D. sabulorum var. patulum Dichanthelium acuminatum subsp. acuminatum P. columbianum var. columbianum var. thinium subsp. columbianum var. thinium D. ovale var. ovale subsp. fasciculatum P. commonsianum var. addisonii subsp. implicatum P. malacon D. acuminatum var. acuminatum subsp. leucothrix P. oricola var. densiflorum subsp. lindheimeri P. tsugetorum var. implicatum subsp. longiligulatum P. wilmingtonense var. lindheimeri subsp. sericeum var. longiligulatum subsp. spretum Group Lanuginosa var. thurowii subsp. thermale P. acuminatum var. villosum D. ovale subsp. ovale P. albemarlense var. wrightianum subsp. praecocius P. auburne subsp. pseudopubescens P. huachucae var. huachucae subsp. villosissimum var. silvicola D. wrightianum P. implicatum P. languidum P. lanuginosum P. meridionale P. occidentale P. olivaceum P. ovale P. pacificum P. praecocius P. pseudopubescens P. scoparioides P. shastense P. subvillosum P. tennesseense P. thermale P. thurowii P. villosissimum
Group Spreta P. leucothrix P. lindheimeri P. longiligulatum P. spretum P. wrightianum
65 Table 2. Morphological characters used in the final phenetic analysis and principal components analysis. An “*” indicates characters pertaining to the Long Ligule Group. A “‡” indicates characters pertaining to the Short Ligule Group.
Character Abbreviation Character State 1. ligule pubescence maximum LigLen * ‡ (mm) length 2. ligule trichome uniformity LigTriUni* 0=no; 1=yes 3. mid-stem sheath pubescence MidShePubMax * (mm) maximum length 4. mid-stem sheath glabrous to MidSheGla * 0=no; 1=yes glabrate 5. mid-stem leaf adaxial pubescence AdaLeaPubMax * (mm) maximum length 6. adaxial surface of mid-stem leaf AdaSurMidLeaGla * 0=no; 1=yes glabrous to glabrate 7. terminal sheath pubescence TerShePubMax * (mm) maximum length 8. terminal sheath glabrous to glabrate TerSheGla* 0=no; 1=yes 9. terminal leaf abaxial pubescence TerLeaAbaPubMin * (mm) minimum length 10. terminal leaf abaxial pubescence TerLeaAbaPubMax * (mm) maximum length 11. spikelet length SpiLen*‡ (mm) 12. spikelet width SpiWid* (mm) 13. first glume length PriGluLen*‡ (mm) 14. longest leaf length LonLeaLen* (mm) 15. longest leaf width LonLeaWid*‡ (mm) 16. number of nodes in inflorescence NumInfNod* number 17. inflorescence axis with puberulent PubPubInf * 0=no; 1=yes pubescence 18. peduncle glabrous to glabrate PedGla* 0=no; 1=yes 19. peduncle pubescence double PedPubDou * (mm) invested 20. abaxial leaf surfaces with AbaLeaSurPub * 0=no; 1=yes puberulent pubescence 21. inflorescence < a third as long as wide InfNar* 0=no; 1=yes 22. fertile floret pointed FerFloPoi* 0=no; 1=yes 23. inflorescende w/ double vestiture InfDouVes* 0=no; 1=yes 24. spikelet pointed at apex SpiPoi* 0=no; 1=yes 25. double ligule conspicuous DouLigCon‡ 0=no; 1=yes 26. double vestiture on culm DouVes‡ 0=no; 1=yes 27. number of trichomes across NumTriAdaLeaSur ‡ number mid-stem adaxial leaf surface 28. mid-stem leaf glabrous along MidLeaGlaAda ‡ 0=no; 1=yes middle of adaxial surface 29. terminal leaf glabrous along TerLeaGlaAda ‡ 0=no; 1=yes middle of adaxial surface 30. mid-stem sheath indument length MidSheIndLen‡ (mm) 31. sheath pubescence orientation ShePubOri‡ scale; 0=appressed; 1=erect 32. mid-stem leaf marginal cilia >30 Cilia>30‡ 0=no; 1=yes
66 Table 3. Results of pair-wise comparisons of Long Ligule Group using two sample t-tests. Numbers correspond to characters listed on Table 2. The numbers not in bold are those characters that demonstrate a significant difference of P ≤ 0.005. Bold numbers are those characters that demonstrate P-values between 0.05 and 0.005.
D. wrightianum D. spretum D. meridionale D. longiligulatum D. leucothrix D. implicatum D. praecocius D. lindheimeri
1,3,5,7,10,11,12, 1,3,4,5,6,7,8,9, 1 ,2, 3 , 5 , 10 ,11, 1 ,4,3,5,6,7,8, 9, 1 ,2,3, 4 , 5 ,6, 5, 6 ,7,9,10, 2,3,5,6,7,9, 1,2,3,4,5,6, D. lanuginosum 13,14,15,16,17, 10, 12 ,15,16,18, 13 ,14,1 5,16,17, 10,11,12, 13 ,14,15, 7,10,11,12, 13, 11 , 12 ,14, 10,11,12,13, 7,8,9,10, 11, 19,20,22,23 21,22,24 19,20,23 17,18,19,20,22,23 14,15,20,22,23 15,16 14,15,16,17 13,14,15
2,4,6,7,8,9,10, 3,4,5,6,7,8,9,10, 4,7,8,9,10,11, 1 ,2,3,4,5, 2,3,4,5,6,7, 1,2,11,13,15, 2,9,10,11,12, 11,12,13,14,15, 13,14,15,16,17, 12,14,15,18, 6,7,8,9,10, 8.9,10,11,12, D. lindheimeri 16,21,22,24 14,15,20,22,23 17,18,19,20,22,23 18,19,20,23 20,22,23 13,14,18 13,16,17,18
1,2,3,5,7,9,10, 1 ,2,3,4,5,6,7,8, 2,3, 5 ,7,9, 10 ,11, 2,3,4,5,6,7,8,9,10, 2,3, 4 ,5,6,7,9,10, 2,3,5,7,9, D. praecocius 11,12,13,14,15, 9,10,11,12,13, 14 , 12,13,14,15 ,16, 11,12,13,14,15,16, 11,12,13,1 4, 15 , 10 ,11,12, 16,17,19,20,22,23 16,17,18,21,22,24 19,20,23 17,18,19,20,22,23 16,17,20,22,23 13,14,16,17
1,3,5,7,9,10,11, 1,3,4,5,6,7,8,9, 3,4,5,6,7,8,9,10, 11, 2,3,5,6,7,9,10, 2,10,13,14,15, 12,13,14,15,17, 10,14,16,18,21, 12, 13 ,15,17,18, 11,12,14,20,22, D. implicatum 16,17,19,20,23 19,20,22,23 22,24 19,20,22,23 23 1,2,3,4,5,7,8,9,10, 2,3,6,7,11,12, 3,5,6,10,11,12,13, 2,3,4,5, 7,8,9, 11,12,13,14,16, D. leucothrix 13,14,17,19 15,16,17,19,22 10,18 18,20,21,23,24 3,4,6,7,8,9,10, 1 , 9 , 10 ,11, 3,4,5,6, 7 ,8, 9 ,10, D. longiligulatum 11,12,13,14,17, 13, 14, 15 ,20, 11 , 12 ,13, 16 ,17, 18,19 21,23,24 18,19,22 1,2,3,4,5,6,7,8,9, 2,3,5,7,10,11, 10, 11 ,14,15,16,
D. meridionale 12,13,14,17,22 17,18,19,20,21, 22,23,24
3,4,6,7,8,9,10,11, D. spretum 12,13,14,15,16,17, 18,19,20,21,23,24
67 Table 4. Loadings for the first three principal components from the PCA of the Long Ligule Group. Bold numbers are characters greater than 0.60 (absolute value).
Variable PC1 PC2 PC3 (1) LigLen 0.470134 0.013531 0.311942 (2) LigTriUni 0.367315 -0.044885 -0.638994 (3) MidShePubMax 0.929179 -0.051885 -0.166881 (4) MidSheGla -0.702810 0.446930 -0.314728 (5) AdaLeaPubMax 0.841107 -0.131279 -0.102921 (6) AdaSurMidLeaGla -0.689265 0.401993 -0.077478 (7) TerShePubMax 0.882629 0.027453 -0.135609 (8) TerSheGla -0.695017 0.492033 -0.297461 (9) TerLeaAbaPubMin 0.805324 -0.174873 -0.234322 (10) TerLeaAbaPubMax 0.852446 -0.105936 -0.103876 (11) SpiLen 0.653451 0.549536 -0.320496 (12) SpiWid 0.690557 0.413550 -0.300294 (13) PriGluLen 0.677986 0.142037 -0.575833 (14) LonLeaLen 0.295223 0.698455 0.188139 (15) LonLeaWid 0.302144 0.469000 0.541051 (16) NumInfNod -0.519752 0.389019 0.330314 (17) PubPubInf -0.195249 -0.727720 -0.244033 (18) PedGla -0.651382 0.533550 -0.295013 (19) PedPubDou -0.271396 -0.774095 -0.179592 (20) AbaLeaSurPub -0.496088 -0.771455 -0.188159 (21) InfNar -0.455718 0.503179 -0.397369 (22) FerFloPoi -0.770360 -0.162314 -0.227806 (23) InfDouVes -0.471496 -0.781279 -0.181547 (24) SpiPoi -0.455718 0.503179 -0.397369
total variance explained 39.8226% 20.1040% 11.6452%
eigenvalues 9.5574 4.8249 2.7948
68 Table 5. Results of pair-wise comparisons of Short Ligule Group using two sample t-tests. Numbers correspond to characters listed on Table 2. The numbers not in bold are those characters that demonstrate a significant difference of P ≤ 0.005. Bold numbers are those characters that demonstrate P-values between 0.05 and 0.005.
D. villosissimum D. thermale D. subvillosum D. ovale D. commonsianum D. columbianum
1,13,15,25,26, 1,11,13,26,27, 1,11,13,15,25, 1,13,15,25,26,27, 1,11,13,30,31,32 13,25,26,31,32 D. acuminatum 30,31,32 28,29,30,31,32 26,27,28,29,31,32 28,29,30, 31,32
1,11,13,15,25,26, 1,25,26,27,28, 1,11,13,15,25, 1,15,26,27,28,29,31 1,11,13,27,30 D. columbianum 27,28, 29,30,31 29,30,31 26,27,30,31
1,11,13,15,25,26, 11,13,15,26,27, 11,13,25,26,27, 1,13,15,25,26,27 D. commonsianum 27,28, 29,30,31 28,29,31 28,29,31
1,11,13,26,27,28, 1,11,13,25, 26, 1,11,13,15,25,27,
D. ovale 29,30, 31 27,28,29,30,31 28,29,31
D. subvillosum 1,11,15,25,26,30 15,25,26,30
D. thermale 1,11,25,26,30
69 Table 6. Loadings for the first three principal components from the PCA of the Short Ligule Group. Bold numbers are characters greater than 0.60 (absolute value).
Variable PC1 PC2 PC3 (1) LigLen 0.178984 0.733770 0.409249 (11) SpiLen 0.551986 -0.754718 -0.028215 (13) PriGluLen 0.717599 -0.486356 0.150821 (15) LonLeaWid -0.501936 -0.259084 -0.311542 (25) DouLigCon 0.670133 0.481666 0.334808 (26) DouVes 0.825638 0.385092 0.202818 (27) NumTriAdaLeaSur -0.757211 -0.061297 0.413394 (28) MidLeaGlaAda 0.915842 -0.061084 -0.259726 (29) TerLeaGlaAda 0.915842 -0.061084 -0.259726 (30) MidSheIndLen -0.279299 -0.648330 0.569073 (31) ShePubOri -0.839872 -0.016393 0.045759 (32) Cilia>30 -0.593963 0.447627 -0.458003
total variance explained 46.8454% 20.2014% 10.6903%
eigenvalues 5.6214 2.4241 1.2828
70 ACUM-139-FL-BTHUR-205-TX-B T-OLIV-247-GUACUM-46-NI-BOLIV-228-MX-BTHUR-201-TX-B T-FERV-250-WYACUM-302-W.I.T-OLIV-284-GUATOLIV-226-MX-BACUM-140-GA-B T-GLUT-252-VAACUM-300-PROLIV-304-CLOLIV-305-CROLIV-224-MX-B THUR-203-LA-BTHUR-202-LA-BOLIV-225-HNACUM-301-JAACUM-47-HT-B OLIV-303-PNACUM-48-BE-BT-THUR-265-TXTHUR-229-TX-BT-ORAN-286-NC T-COMO-235-PR-BT-CILI-280-MSOLIV-227-MX-BT-AUBU-259-ALACUM-147-TX-B SUBV-221-MNT-THER-254-CAT-LASS-276-CASUBV-233-WISUBV-200-OH SUBV-311-MESUBV-312-MIT-SUBV-287-MNSUBV-222-NY T-THER-232-CATHER-231-CASUBV-313-WIVILL-06-ILTHER-230-CA-B VILL-105-TXT-VILL-242-GAVILL-22-TNVILL-41-MOT-ATLA-244-NY VILL-90-DCVILL-17-GAVILL-20-MSVILL-18-NYVILL-95-MX VILL-107-TNVILL-96-TNVILL-93-GAVILL-99-IAT-HAEM-270-DC VILL-92-VAVILL-115-ILVILL-113-ALVILL-97-NDT-XANT-268-AL VILL-111-MOVILL-109-GAVILL-35-TXVILL-116-IAShort VILL-106-TNVILL-27-MOVILL-91-MX?VILL-19-NJVILL-94-ALLigule T-PSEU-241-ALVILL-110-MOVILL-114-ARVILL-112-VAVILL-314-GT VILL-49-mexicoVILL-98-SCVILL-23-VACOLU-104-OHCOLU-100-MAGroup T-IMSI-275-ILCOLU-102-NYT-OWEN-234-MACOLU-101-MDCOLU-103-OH COMM-W-108-TXT-SCOP-283-DET-ADDI-238-NJCOLU-80-DECOLU-77-NY T-COMM-279-NJCOMM-79-NJCOMM-72-MDCOMM-66-NJT-EUCL-255-WI COMM-69-MACOMM-75-NJCOMM-70-DECOMM-73-DE COMM-74-NJCOMM-68-MACOMM-W-53-MOCOMM-76-NJCOMM-78-NY T-WILM-267-NCCOMM-W-89-WICOMM-W-59-INPSEU-60-TNOVAL-31-NC OVAL-81-VAOVAL-55-SCOVAL-63-NCOVAL-37-NJOVAL-34-NJ POVAL-84-SCOVAL-87-SCOVAL-82-SC SEU- 56- SC OVAL-29-FLOVAL-88-FLOVAL-64-FLPSEU-54-GAPSEU-58-VA OVAL-65-FLOVAL-33-FLPSEU-62-ALPSEU-61-ALOVAL-32-GA T-ERYT-277-NCT-CILI-256-FLOVAL-85-FLOVAL-36-SCOVAL-67-AL PSEU-57-FLOVAL-86-SCOVAL-83-MAOVAL-30-SCT-BENN-249-NJ T-OLIV-284-GUAT IMPL-177-MNIMPL-178-MIT-PACI-262-CAIMPL-02-MNCOMM-71-MA IMPL-169-TNIMPL-220-ORIMPL-134-ONT-OCCI-257-WAIMPL-04-MN IMPL-175-IAIMPL-173-RIIMPL-50-WILANU-137-NJLANU-132-LA IMPL-170-MAIMPL-130-ART-HUAC-272-AZLANU-144-ILLANU-145-IA IMPL-176-MAIMPL-171-NYIMPL-16-MEIMPL-51-VT IMPL-214-WAT-IMPL-274-MAIMPL-14-OHLANU-136-PALANU-10-IN IMPL-215-ORLANU-218-VANLANU-212-CALANU-213-WALANU-135-KY LANU-133-QULANU-131-KSLANU-141-MDLANU-12-ILLANU-11-OH T-FESE-269-WYT-FEPA-251-ALBT-UNPR-266-MALANU-146-VALANU-142-MA T-HUSI-273-DCLANU-143-ILLANU-217-WALANU-149-MOLANU-138-NE LANU-219-BCLANU-152-NYLANU-155-PALANU-42-ARLANU-211-OR TENN-154-SCT-BROD-248-WALANU-148-OKLANU-01-OHLong TENN-159-WITENN-150-MOTENN-158-UTTENN-157-NET-FUNS-271-CA PRAE-07-KSTENN-153-OKTENN-156-TXTENN-151-QULigule PRAE-117-MOPRAE-44-NEPRAE-40-MOPRAE-25-MNGroup PRAE-118-MOPRAE-315-?PRAE-21-ILT-PRAE-263-ILPRAE-45-NE PRAE-129-TXPRAE-124-NEPRAE-128-NEPRAE-119-MOPRAE-121-OK PRAE-125-NEPRAE-122-ILPRAE-126-IAPRAE-123-ARPRAE-26-NE PRAE-120-MOPRAE-167-WIPRAE-127-NEPRAE-38-MOPRAE-24-OK-A LEUC-197-SCLEUC-172-TXLEUC-194-FLPRAE-43-ARPRAE-39-MO T-PARV-240-GALEUC-199-TNLEUC-198-FLT-LEUC-243-FL WRIG-186-GAWRIG-184-MAWRIG-188-VAT-DEMI-278-NYT-WRIG-258-CUB MERI-181-MAMERI-174-VAWRIG-185-FLWRIG-187-MST-STRI-264-NC MERI-183-NCMERI-180-VAMERI-182-TNMERI-179-NYT-MERI-260-NC T-ALBE-281-NCLIND-163-MILIND-09-MIT-MICR-285-NCLIND-216-CA LIND-162-ARLIND-166-MOLIND-165-MOLIND-168-NYLIND-164-TX T-LISE-253-NBLONG-190-NCLIND-160-KYLIND-161-NY T-LONG-246-FLLONG-191-FLT-LIND-245-TXLONG-189-TNLONG-193-NIGU SPRE-208-NSSPRE-207-MASPRE-206-NJT-EATO-237-NHLONG-192-GA SPRE-210-INSPRE-209-INT-PAUC-239-NJT-OCTO-261-TX -0.38-0.38 -0.03-0.03 0.310.31 0.650.65 1.001.00
SimilarityCoefficient Coefficient
Figure 1. Phenogram for all 285 OTUs using 32 morphological characters. Cophenetic correlation coefficient (r) = 0.85.
71
Long Ligule
Short Ligule
0.0 1.0 2.0 3.0 4.0 5.0
Ligule Length (mm)
Figure 2. Box plot showing the lack of overlap in ligule length between the OTUs of the Long Ligule and Short Ligule groups.
72 IMPL-176-MAT-IMPL-274-MAIMPL-16-ME IMPL-130-ARIMPL-50-WI IMPL-171-NYIMPL-170-MAIMPL-51-VT IMPL-169-TNIMPL-178-MI IMPL-173-RIIMPL-175-IAD. implicatum IMPL-215-ORIMPL-220-ORIMPL-134-ON IMPL-177-MNT-OCCI-257-WA T-PACI-262-CAIMPL-02-MN IMPL-14-OHIMPL-04-MN T-FEPA-251-ALBIMPL-214-WAD. thermale T-FESE-269-WYPRAE-07-KS subp. seric eum PRAE-120-MOPRAE-44-NE PRAE-39-MOPRAE-38-MO PRAE-117-MOPRAE-25-MNPRAE-40-MO PRAE-118-MOPRAE-26-NE T-PRAE-263-ILPRAE-21-ILD. praecocius PRLA-167-WIPRAE-315-?PRAE-24-OK-A PRAE-119-MOPRAE-45-NE PRAE-123-ARPRAE-121-OK PRAE-128-NEPRAE-124-NE PRAE-127-NEPRAE-122-ILPRAE-126-IA PRAE-125-NEPRAE-129-TXPRAE-43-AR T-HUSI-273-DCLANU-01-OH LANU-42-ARLANU-143-ILLANU-138-NE LANU-12-ILLANU-211-OR LANU-217-WALANU-149-MO LANU-142-MALANU-133-QULANU-131-KS T-BROD-248-WALANU-146-VA LANU-10-INLANU-155-PALANU-152-NY LANU-218-VANLANU-135-KY T-FEPA-251-ALB LANU-148-OKLANU-136-PAD. lanuginosum LANU-141-MDLANU-219-BCLANU-137-NJ LANU-213-WALANU-212-CA LANU-145-IALANU-11-OHT-UNPR-266-MA LANU-144-ILLANU-132-LA TENN-150-MOT-HUAC-272-AZ TENN-151-QUTENN-158-UTTENN-153-OK TENN-157-NETENN-156-TX TENN-154-SCT-FUNS-271-CA LEUC-172-TXTENN-159-WI LEUC-197-SCT-LEUC-243-FLLEUC-194-FLD. leucothrix T-PARV-240-GALEUC-199-TNLEUC-198-FL T-DEMI-278-NY WRIG-184-MAWRIG-188-VAT-WRIG-258-CUBD. wrightianum WRIG-187-MSWRIG-186-GA WRIG-185-FLT-STRI-264-NC LONG-193-NIGULONG-190-NCD. longiligulatum LONG-191-FLLONG-189-TN T-LONG-246-FLLONG-192-GA MERI-181-MAMERI-174-VA MERI-182-TNMERI-179-NYD. meridionale MERI-183-NCMERI-180-VAT-MICR-285-NC T-ALBE-281-NCLIND-09-MIT-MERI-260-NC LIND-163-MI LIND-162-ARLIND-216-CAD. lindheimeri LIND-166-MOLIND-165-MO T-LIND-245-TXLIND-164-TXLIND-168-NY LIND-161-NYLIND-160-KY SPRE-206-NJT-LISE-253-NB SPRE-207-MAT-EATO-237-NHD. spretum SPRE-210-INSPRE-209-INT-PAUC-239-NJ SPRE-208-NST-OCTO-261-TX -0.30-0.31 0.000.01 0.300.34 0.700.66 0.991.00 Similarity Coefficient Coefficient
Figure 3. Phenogram of the Long Ligule Group based on 142 OTUs and 24 morphological characters. Cophenetic correlation coefficient (r) = 0.91
73 1.50
0.75 = D. implicatum (N=22) = D. lanuginosum (N=41) = D. leucothrix (N=7) = D. lindheimeri (N=12) = D. longiligulatum (N=6) PC2 0.00 = D. meridionale (N=9) = D. praecocius (N=25) = D. spretum (N=7) = D. wrightianum (N=7)
-0.75
-1.50 -1.50 -0.75 0.00 0.75 1.50 PC1
Figure 4. Scatter plot of the scores of principal component 1 (PC1) and principal component 2 (PC2) for all the standardized morphological characters of the 142 specimens in the Long Ligule Group.
74 1.15
0.61
= D. implicatum (N = 22) PC2R2 0.07 = D. lanuginosum (N = 41)
-0.46
-1.00 -1.00 -0.35 0.30 0.95 1.60 PC1R1
Figure 5. Scatter plots of the scores of principal component 1 (PC1) and principal component 2 (PC2) from PCA of Dichanthelium implicatum and D. lanuginosum specimens.
75 ACUM-139-FL-B OLIV-228-MX-BOLIV-226-MX-BTHUR-205-TX-B ACUM-140-GA-BTHUR-201-TX-B ACUM-302-W.I.ACUM-300-PROLIV-224-MX-B T-FERV-250-WYT-GLUT-252-VAOLIV-225-HN THUR-203-LA-BACUM-301-JAD. acuminatum ACUM-48-BE-BT-OLIV-284-GUATTHUR-202-LA-B OLIV-303-PNT-ORAN-286-NC OLIV-304-CLOLIV-305-CRACUM-46-NI-B T-OLIV-247-GUT-CILI-280-MS OLIV-227-MX-BACUM-47-HT-BACUM-147-TX-B T-COMO-235-PR-BT-THUR-265-TXTHUR-229-TX-B SUBV-200-OHT-AUBU-259-AL SUBV-311-MESUBV-222-NYD. subvillosum SUBV-233-WIT-SUBV-287-MN SUBV-221-MNSUBV-313-WISUBV-312-MI THER-231-CATHER-230-CA-BD. thermale T-THER-254-CAT-LASS-276-CAT-THER-232-CA VILL-99-IAT-ATLA-244-NY VILL-41-MOVILL-06-IL VILL-97-NDVILL-116-IA VILL-314-GTVILL-27-MOT-HAEM-270-DC VILL-105-TXT-VILL-242-GAT-XANT-268-AL VILL-90-DCVILL-17-GA VILL-22-TNVILL-18-NYVILL-95-MX VILL-49-mexicoVILL-20-MSD. villosissimum VILL-109-GAVILL-112-VAVILL-106-TN VILL-107-TNVILL-94-AL VILL-111-MOVILL-96-TNVILL-113-AL T-COMO-235-PR-B VILL-35-TXVILL-19-NJ T-PSEU-241-ALVILL-91-MX? VILL-110-MOVILL-98-SCVILL-23-VA VILL-114-ARVILL-115-IL VILL-93-GAVILL-92-VACOLU-100-MA COLU-101-MDCOLU-104-OH COLU-77-NYCOLU-103-OHCOLU-102-NYD. columbianum T-ADDI-238-NJCOLU-80-DE T-IMSI-275-ILT-OWEN-234-MA T-SCOP-283-DECOMM-66-NJ T-COMM-279-NJCOMM-74-NJCOMM-78-NY COMM-73-DECOMM-72-MDCOMM-70-DE COMM-69-MACOMM-79-NJD. commonsianum COMM-75-NJCOMM-71-MACOMM-76-NJ COMM-W-53-MOCOMM-68-MA COMM-W-89-WICOMM-W-59-IN COMM-W-108-TXT-EUCL-255-WE PSEU-61-ALOVAL-32-GAOVAL-29-FL OVAL-65-FLOVAL-33-FL OVAL-84-SCOVAL-36-SCOVAL-30-SC T-ERYT-277-NCPSEU-54-GA T-WILM-267-NCOVAL-31-NC OVAL-64-FLPSEU-60-TND. ovale OVAL-67-ALPSEU-62-ALOVAL-88-FL T-CILI-256-FLOVAL-37-NJOVAL-34-NJ OVAL-55-SCOVAL-63-NC OVAL-82-SCOVAL-87-SCOVAL-81-VA PPSEU-58-VA SEU- 56- SC PSEU-57-FLOVAL-86-SCOVAL-83-MA T-BENN-249-NJOVAL-85-FL --0.410.40 --0.060.10 0.300.29 0.600.64 1.001.00 Similarity Coefficient Coefficient
Figure 6. Phenogram of the Short Ligule Group based on 143 OTUs and 12 morphological characters. Cophenetic correlation coefficient (r) = 0.90.
76 1.50
0.75 = D. acuminatum (N=31) = D. columbianum (N=11) = D. commonsianum (N=20) = D. ovale (N=31) = D. subvillosum (N=8) PC2 0.00 = D. thermale (N=5) = D. villosissimum (N=39)
-0.75
-1.50 -1.50 -0.75 0.00 0.75 1.50 PC1
Figure 7. . Scatter plot of the scores of principal component 1 (PC1) and principal component 2 (PC2) for all the standardized morphological characters of the 143 specimens in the Short Ligule Group.
77 a. b.
c. d.
Figure 8. Examples of the four ligule types and the four pubescence types found within section Lanuginosa. All combinations of the eight illustrated characters are possible. Ligule illustration a.) occurs in D. implicatum, D. lanuginosum, D. leucothrix, D. lindheimeri, D. longiligulatum, D. meridionale, D. praecocius, D. spretum, D. thermale subsp. sericeum and D. wrightianum; b.) occurs in D. acuminatum, D. columbianum, D. commonsianum, D. ovale, D. subvillosum, D. thermale and D. villosissimum; c.) occurs in D. columbianum, D. commonsianum and D. ovale; d.) occurs in D. acuminatum, D. columbianum, D. meridionale, D. subvillosum, D. thermale subsp. sericeum and D. thermale subsp. thermale. Pubescence illustration a.) occurs in D. acuminatum, D. implicatum, D. lanuginosum, D. praecocius, D. thermale subsp. sericeum and D. villosissimum; b.) occurs in D. acuminatum, D. subvillosum, D. thermale subsp. thermale and D. villosissimum; c.) occurs in D. columbianum, D. commonsianum, D. leucothrix and D. wrightianum; d.) occurs in D. columbianum, D. commonsianum, D. leucothrix, D. meridionale, D. ovale, D. thermale subsp. thermale and D. wrightianum.
78
Figure 9. Distribution of Dichanthelium acuminatum based on specimens sampled.
Figure 10. Distribution of Dichanthelium columbianum based on specimens sampled.
79
Figure 11. Distribution of Dichanthelium commonsianum based on specimens sampled.
Figure 12. Distribution of Dichanthelium implicatum based on specimens sampled.
80
Figure 13. Distribution of Dichanthelium lanuginosum based on specimens sampled.
Figure 14. Distribution of Dichanthelium leucothrix based on specimens sampled.
81
Figure 15. Distribution of Dichanthelium lindheimeri based on specimens sampled.
Figure 16. Distribution of Dichanthelium longiligulatum based on specimens sampled.
82
Figure 17. Distribution of Dichanthelium meridionale based on specimens sampled.
Figure 18. Distribution of Dichanthelium ovale based on specimens sampled.
83
Figure 19. Distribution of Dichanthelium praecocius based on specimens sampled.
Figure 20. Distribution of Dichanthelium spretum based on specimens sampled.
84
Figure 21. Distribution of Dichanthelium subvillosum based on specimens sampled.
Figure 23. Distribution of Dichanthelium thermale based on specimens sampled. Open circles represent specimens pertaining to D. thermale subsp. sericeum.
85
Figure 23. Distribution of Dichanthelium villosissimum based on specimens sampled.
Figure 24. Distribution of Dichanthelium wrightianum based on specimens sampled.
86 Appendix A. All morphological characters sampled. An “*” indicates characters pertaining to the Long Ligule Group. A “‡” indicates characters pertaining to the Short Ligule Group.
Character Abbreviation Character State
Vegetative Characters plant height PlaHei (mm) geniculate stem Gen 0=no; 1=yes double vestiture on culm DouVes‡ 0=no; 1=yes node pubescence NodPub 0=absent; 1=present node pubescence length NodPubLen (mm) node width NodWid (mm) node length NodLen (mm) node length width ratio NodRat ratio (decimal to the hundredth) nodes with autumnal NodAutInf rank from bottom to top inflorescences second internode length SecIntLen (mm) second internode width SecInt Wid (mm) basal internode pubescence BasIntPubMax (mm) maximim length basal internode pubescence BasIntPubMin (mm) minimum length number of basal internode NumBasIntTri number trichomes per 0.5cm mid-stem internode pubescence MidIntPubMax (mm) maximum length mid-stem internode pubescence Mi dIntPubMin (mm) minimum length number of mid-stem internode NumMidIntTri number trichomes per 0.5cm terminal internode pubescence TerIntPubMax (mm) maximum length terminal internode pubescence TerIntPubMin (mm) minimum length number of terminal internode NumTerIntTri number trichomes per 0.5cm internode pubescence double IntPubDou 0=no; 1=yes invested internode pubescence orientation IntPubOri scale; 0=appressed; 1=erect internode pubescence pustulate IntPubPus 0=no; 1=yes Longest vernal leaf LonVerLea rank from bottom to top longest leaf length LonLeaLen* (mm) longest leaf width LonLeaWid*‡ (mm) longest leaf base width LonLeaBasWid (mm) leaf length width ratio LonRat ratio (decimal to the hundredth) terminal leaf length TerLeaLen (mm) terminal leaf width TerLeaWid (mm) terminal leaf length TerRat ratio (decimal to the hundredth) width ratio mid-stem leaf marginal cilia >30 Cilia>30‡ 0=no; 1=yes adaxial surface of mid-stem leaf AdaSurMidLeaGla * 0=no; 1=yes glabrous to glabrate mid-stem leaf glabrous along MidLeaGlaAda ‡ 0=no; 1=yes middle of adaxial surface
87 Character (vegetative con’t) Abbreviation Character State mid-stem leaf adaxial pubescence AdaLeaPubMax * (mm) maximum length mid-stem leaf adaxial pubescence AdaLeaPubMin (mm) minimum length number of trichomes across NumTriAdaLeaSur ‡ number mid-stem adaxial leaf surface abaxial leaf surfaces with AbaLeaSurPub * 0=no; 1=yes puberulent pubescence mid-stem leaf abaxial pubescence AbaLeaPubMax (mm) maximum length mid-stem leaf abaxial pubescence AbaLeaPubMin (mm) minimum length number of trichomes across NumTriAbaLeaSur number mid-stem abaxial leaf surface number of mid-stem leaf marginal NumMidLeaMarTri number trichomes length of mid-stem ciliate margins LenMidLeaCilMar (mm) terminal leaf glabrous along TerLeaGlaAda ‡ 0=no; 1=yes middle of adaxial surface terminal leaf adaxial pubescence TerLeaAdaMax (mm) maximum length terminal leaf adaxial pubescence TerLeaAdaMin (mm) minimum length number of trichomes across terminal NumTriTerAdaLeaSur number adaxial leaf surface terminal leaf abaxial pubescence TerLeaAbaPubMax * (mm) maximum length terminal leaf abaxial pubescence TerLeaAbaPubMin * (mm) minimum length number of trichomes across terminal NumTriTerAbaLeaSur number abaxial leaf surface number of terminal leaf marginal NumTerLeaCilMar number trichomes terminal leaf ciliate margin length LenTerLeaCilMar (mm) ligule pubescence maximum LigLen *‡ (mm) length ligule pubescence minimum LigPubMin (mm) length ligule double LigDou 0=no; 1=yes double ligule conspicuous DouLigCon‡ 0=no; 1=yes ligule intermeshed LigInt 0=no; 1=yes ligule trichome uniform LigTriUni* 1=lengths uniform; 2=lengths variable ligule edge maximum length LigEdgMaxLen (mm) ligule edge minimum length LigEdgMinLen (mm) ligule middle maximum length LigMidMaxLen (mm) ligule middle minimum length LigMidMinLen (mm) longest sheath length LonSheLen (mm) terminal sheath length TerSheLen (mm) basal sheath pubescence BasShePubMax (mm) maximum length basal sheath pubescence BasShePubMin (mm) minimum length number of basal sheath NumBasSheTri number trichomes per 0.5cm
88 Character (vegetative con’t) Abbreviation Character State mid-stem sheath glabrous to MidSheGla * 0=no; 1=yes glabrate mid-stem sheath indument length MidSheIndLen‡ (mm) mid-stem sheath pubescence MidShePubMax * (mm) maximum length mid-stem sheath pubescence MidShePubMin (mm) minimum length number of mid-stem sheath NumMidSheTri number trichomes per 0.5cm terminal sheath glabrous to glabrate TerSheGla* 0=no; 1=yes terminal sheath indument length TerSheIndLen (mm) terminal sheath pubescence TerShePubMax * (mm) maximum length terminal sheath pubescence TerShePubMin (mm) minimum length number of terminal sheath NumTerSheTri number trichomes per 0.5cm sheath pubescence double ShePubDou 0=no; 1=yes invested sheath pubescence orientation ShePubOri‡ scale; 0=appressed; 1=erect sheath pubescence pustulate ShePubPus 0=no; 1=yes peduncle exposed above PedExp (mm) subtending sheath peduncle length PedLen (mm) peduncle glabrous to glabrate PedGla* 0=no; 1=yes peduncle pubescence maximum PedPubMax (mm) length peduncle pubescence minimum PedPubMin (mm) length number of peduncle trichomes NumPedTri number per 0.5cm peduncle pubescence double PedPubDou * 0=no; 1=yes invested peduncle pubescence orientation PedPubOri scale; 0=appressed; 1=erect peduncle pubescence pustulate PedPubPus 0=no; 1=yes inflorescence < a third as long as wide InfNar* 0=no; 1=yes inflorescende w/ double vestiture InfDouVes* 0=no; 1=yes inflorescence pubescence InfPubMax (mm) maximum length inflorescence pubescence InfPubMin (mm) minimum length inflorescence with scabrous InfScaPub 0=no; 1=yes pubescence inflorescence axis with puberulent PubPubInf * 0=no; 1=yes pubescence number of spikelets in inflorescence NumSpi number number of nodes in inflorescence NumInfNod* number vernal inflorescence width VerInfWid (mm) vernal inflorescence length VerInfLen (mm) vernal inflorescence length InfRat ratio (decimal to the hundredth) width ratio number of trichomes along 0.5cm NumTriInf number of inflorescence axis pedicel maximum length PedLenMax (mm) pedicel minimum length PedLenMin (mm)
89 Character (vegetative con’t) Abbreviation Character State pedicel pubescence PedPub 0=no; 1=yes
Floral Characters spikelet length SpiLen*‡ (mm) spikelet width SpiWid* (mm) spikelet pubescence length SpiPubLen (mm) first glume length GluLen*‡ (mm) first glume width GluWid (mm) first glume length width ratio GluRat ratio (decimal to the hundredth) spikelet pointed at apex SpiPoi* 0=no; 1=yes number of veins on first glume NumVeiFirGlu number first glume pubescent GluPub 0=no; 1=yes number of veins on second glume NumVeiSecGlu number number of viens on sterile lemma NumVeiSteLem number fertile floret pointed FerFloPoi* 0=no; 1=yes
90 Appendix B. Histograms of the 116 morphological characters measured.
Plant Height Node Pubescence
16 45 14 40
12 35 30 10 25 8 20 Frequency Frequency 6 15 4 10
2 5 0 0 181.0 265.5 350.0 434.5 519.0 603.5 682.0 0 1 PlaHei (mm) NodPub
Geniculate Habit Node Pubescence Length
35 16
30 14 12 25 10 20 8 15 Frequency Frequency 6 10 4 5 2
0 0 0 1 0.0 0.6 1.1 1.7 2.2 2.8 3.2 Gen NodPubLen (mm)
Double Vestiture on Culm Node Wi dth
35 14
30 12
25 10
20 8
15 6 Frequency Frequency 10 4
5 2
0 0 0 1 1.0 1.2 1.4 1.6 1.8 2.0 2.2 Do uVes NodWid (mm)
91 Node Length Second Internode Width
16 16 14 14 12 12 10 10 8 8 Frequency Frequency 6 6 4 4 2 2 0 0 1.3 1.6 2.0 2.3 2.6 3.0 3.2 0.6 0.9 1.1 1.4 1.7 1.9 2.2 NodLen (mm) SecIntWid (mm)
Node Length to Width Ratio Basal Internode Pubescence Maximum Length
16 12 14 10 12 8 10
8 6 Frequency Frequency 6 4 4 2 2 0 0 1 1.2 1.4 1.6 1.8 2 2.2 0.7 1.4 2.0 2.7 3.3 4.0 4.6 NodRat BasInt PubMax (mm)
Number Nodes with Autumn Inflorescences Basal Internode Pubescence Minimum Length
20 18 18 16 16 14 14 12 12 10 10 8 Frequency Frequency 8 6 6 4 4 2 2 0 0 1 2 3 4 0.2 0.7 1.2 1.7 2.2 2.7 3.2 NodAut Inf BasInt PubMin (mm)
Second Internode Length Number of Basal Internode Trichomes per 0.5 cm
12 20 18 10 16 8 14 12 6 10 Frequency
Frequency 8 4 6 2 4 2 0 0 29.9 41.6 53.3 65.0 76.6 88.3 90.5 4.0 11.7 19.3 27.0 34.7 42.3 50.0 SecIntLen (mm) NumBasIntTri
92 Mid-Stem Internode Pubescence Maximum Length Terminal Internode Pubescence Minimum Length
14 18
12 16 14 10 12 8 10 6 8 Frequency Frequency 6 4 4 2 2 0 0 0.0 0.7 1.7 2.1 2.7 3.4 4.0 0.0 0.6 1.2 1.8 2.3 2.9 3.5 MidInt PubMax (mm) TerIntPubMin (mm)
Mid-Stem Internode Pubescence Minimum Length Number of Terminal Internode Trichomes per 0.5 cm
16 30 14 25 12 20 10 8 15 Frequency Frequency 6 10 4 5 2 0 0 0.0 0.6 1.2 1.8 2.3 2.9 3.4 0.0 6.7 13.3 20.0 26.7 33.3 40.0 MidInt PubMin (mm) NumTerIntTri
Number of Mid-Stem Internode Trichomes per 0.5 cm Internode Pubescence Double Invested
30 40 35 25 30 20 25
15 20
Frequency 15 Frequency 10 10 5 5 0 0 0 1 0.0 8.3 16.7 25.0 33.3 41.7 50.0 Int PubDou NumMidIntTri
Internode Pubescence Orientation Terminal Internode Pubescence Maximum Length
30 16 14 25 12 20 10 15 8 Frequency
Frequency 6 10 4 5 2 0 0 0 1 0.0 0.7 1.3 2.0 2.7 3.3 4.0 Int PubOri TerIntPubMax (mm)
93 Internode Pubescence Pustulate Longest Leaf Base Width
50 16 45 14 40 12 35 30 10 25 8
Frequency 20 Frequency 6 15 4 10 5 2 0 0 0 1 2.1 2.7 3.3 4.0 4.6 5.2 5.8 Int PubPus LonLeaBasWid (mm)
Longest Vernal Leaf Leaf Length-Width Ratio
25 18 16 20 14 12 15 10 8 Frequency Frequency 10 6 5 4 2 0 0 1 2 3 4 5 6.1 8.9 11.7 14.5 17.3 20.1 22.8 LonVerLea (rank) LonRat
Longest Leaf Length Terminal Leaf Length
14 18
12 16 14 10 12 8 10 6 8 Frequency Frequency 6 4 4 2 2 0 0 53.4 63.0 72.6 82.2 91.7 101.3 110.7 7.2 19.8 32.5 45.1 57.7 70.4 83.0 LonLeaLen (mm) TerLeaLen (mm)
Longest Leaf Width Terminal Leaf Width
14 12
12 10 10 8 8 6 6 Frequency Frequency 4 4 2 2
0 0 3.5 4.8 6.1 7.4 8.7 10.0 11.3 1.8 2.8 3.8 4.9 5.9 6.9 7.0 LonLeaWid (mm) TerLeaWid (mm)
94 Terminal leaf Length Width Ratio Mid-Stem Leaf Adaxial Pubescence Maximum Length
16 18 14 16 12 14 12 10 10 8 8
Frequency 6 Frequency 6 4 4 2 2 0 0 1.6 4.1 6.6 9.0 11.5 14.0 16.4 0.0 0.8 1.5 2.3 3.0 3.8 4.6 TerRat AdaLeaPubMax (mm)
Mid-Stem Leaf Marginal Cilia > 30 Mid-Stem Leaf Adaxial Pubescence Minimum Length
45 18 40 16 35 14 30 12 25 10 20 8 Frequency Frequency 15 6 10 4 5 2 0 0 0 1 0.0 0.6 1.2 1.8 2.3 2.9 3.5 Cilia>30 AdaLeaPubMin (mm)
Adaxial Surface of Mid-Stem Leaf Glabrous to Number of Trichomes Across Mid-Stem Adaxial Leaf Glabrate Surface
45 16 40 14 35 12 30 10 25 8 20 Frequency Frequency 6 15 10 4 5 2 0 0 0 1 0.0 7.3 14.7 22.0 29.3 36.7 44.0 AdaSurMidLeaGla NumT riAdaLeaSur
Mid-Stem Leaf Glabrous along Middle of Adaxial Abaxial Leaf Surfaces with Puberulent Pubescence surface 26 40 35 21 30 25 16 20
Frequency 11 15 Frequency 10 6 5 0 1 0 1 0 1 MidLeaGlaAda AbaLeaSurPub
95 Mid-Stem Leaf Abaxial Pubescence Maximum Length Length of Mid-Stem Leaf Marginal Trichomes
18 9 16 8 14 7 12 6 10 5 8 4 Frequency Frequency 6 3 4 2 2 1 0 0 0.0 0.3 0.6 1.0 1.3 1.6 2.0 1.1 1.5 1.9 2.3 2.7 3.1 3.5 AbaLeaPubMax (mm) LenMidLeaCilMar (mm)
Mid-Stem Leaf Abaxial Pubescence Minimum Length Terminal Leaf Glabrous Along Middle of Adaxial Surface 20 18 45 16 40 14 35 12 30 10 25
Frequency 8 20 6 Frequency 15 4 10 2 5 0 0 0.0 0.2 0.3 0.5 0.7 0.8 1.0 0 1 AbaLeaPubMin (mm) TerLeaGlaAda
Number of Trichomes Across Mid-Stem Abaxial Leaf Terminal Leaf Adaxial Pubescence Maximum Length Surface 20 18 18 16 16 14 14 12 12 10 10
8 Frequency 8 Frequency 6 6 4 4 2 2 0 0 0.0 13.2 26.3 39.5 52.7 65.8 79.0 0.0 1.0 2.0 3.0 4.0 5.0 6.0 NumT riAbaLeaSur (mm) TerLeaAdaMax (mm)
Number of Mid-Stem Leaf Marginal Trichomes Terminal Leaf Adaxial Pubescence Minimum Length
40 18 35 16 30 14 12 25 10 20 8 Frequency 15 Frequency 6 10 4 5 2 0 0 0.0 24.3 28.7 73.0 97.3 121.7 145.7 0.0 0.5 1.0 1.5 1.9 2.4 2.9 NumMidLeaMarTri TerLeaAdaMin (mm)
96 Number of Trichomes Across Terminal Adaxial Leaf Number of Terminal Leaf Marginal Trichomes Surface 25 18 16 20 14 12 15 10 8
Frequency 10 Frequency 6 4 5 2 0 0 0.0 6.8 13.7 20.5 27.3 34.2 41.0 0.0 5.8 11.7 17.5 23.3 29.2 35.0 NumTriTerAdaLeaSur NumTerLeaCilMar
Terminal Leaf Abaxial Pubescence Maximum Length Terminal Leaf Ciliate Margin Length
18 10 16 9 14 8 12 7 6 10 5 8 Frequency Frequency 4 6 3 4 2 2 1 0 0 0.0 0.5 0.9 1.4 1.9 2.3 2.8 1.1 1.5 1.9 2.4 2.8 3.2 3.6 TerLeaAbaPubMax (mm) LenTerLeaCilMar (mm)
Terminal Leaf Abaxial Pubescence Minimum Length Ligule Pubescence Maximum Length
16 18 14 16 12 14 12 10 10 8 8 Frequency 6 Frequency 6 4 4 2 2 0 0 0.0 0.2 0.4 0.6 0.8 1.0 1.2 0.2 1.0 1.8 2.7 3.5 4.3 5.0 TerLeaAbaPubMin (mm) LigLen (mm)
Number of Trichomes Across Terminal Abaxial Leaf Ligule Pubescence Minimum Length Surface 18 20 16 18 14 16 12 14 12 10 10 8
8 Frequency Frequency 6 6 4 4 2 2 0 0 0.0 8.7 17.3 26.0 34.7 43.3 52.0 0.4 0.6 0.7 0.9 1.1 1.2 1.3 NumTriTerAbaLeaSur LigPubMin (mm)
97 Ligule Double Ligule Edge Maximum Length
26 18 16 21 14 12 16 10 8
Frequency 11 Frequency 6 6 4 2 1 0 0 1 0.2 1.0 1.8 2.7 3.5 4.3 5.1 LigDou LigEdgMaxLen (mm)
Double Ligule Conspicuous Ligule Edge Minimum Length
30 18 16 25 14 20 12 10 15 8 Frequency Frequency 10 6 4 5 2 0 0 0 1 0.4 0.6 0.7 0.9 1.1 1.2 1.4 DouLigCon LigEdgMinLen (mm)
Ligule Intermeshed Ligule Middle Maximum Length
30 18 16 25 14 20 12 10 15 8 Frequency Frequency 10 6 4 5 2 0 0 0 1 0.2 1.0 1.8 2.7 3.5 4.3 5.1 LigInt LigMidMaxLen (mm)
Ligule Trichome Uniform Ligule Middle Minimum Length
40 18 35 16 30 14 12 25 10 20 8 Frequency 15 Frequency 6 10 4 5 2 0 0 0 1 0.4 0.6 0.7 0.9 1.1 1.2 1.4 LigTriUni LigMidMinLen (mm)
98 Longest Sheath Length Number of Basal Sheath Trichomes per 0.5 cm
18 18 16 16 14 14 12 12 10 10 8 8 Frequency Frequency 6 6 4 4 2 2 0 0 25.0 32.2 39.4 46.7 53.9 61.1 68.3 4.0 11.7 19.3 27.0 34.7 42.3 50.0 LonSheLen (mm) NumBasSheTri
Terminal Sheath Length Mid-Stem Sheath Glabrous to Glabrate
14 45
12 40 35 10 30 8 25 6 20 Frequency Frequency 4 15 10 2 5 0 0 14.3 26.6 38.8 51.1 63.3 75.6 87.9 0 1 TerSheLen (mm) MidSheGla
Basal Sheath Pubescence Maximum Length Mid-Stem Sheath Indument Length
16 9 14 8 12 7 10 6 5 8 4 Frequency
6 Frequency 3 4 2 2 1 0 0 1.0 1.6 2.2 2.8 3.3 3.9 4.5 0.0 0.6 1.2 1.9 2.5 3.1 3.7 BasShePubMax (mm) MidSheIndLen (mm)
Basal Sheath Pubescence Minimum Length Mid-Stem Sheath Pubescence Maximum Length
25 18 16 20 14 12 15 10 8
Frequency 10 Frequency 6 5 4 2 0 0 0.2 0.6 1.1 1.5 1.9 2.4 2.9 0.0 0.8 1.5 2.3 3.0 3.8 4.6 BasShePubMin (mm) MidShePubMax (mm)
99 Mid-Stem Sheath Pubescence Minimum Length Terminal Sheath Pubescence Maximum Length
18 14
16 12 14 10 12 10 8 8 6 Frequency Frequency 6 4 4 2 2 0 0 0 0.6 1.2 1.8 2.3 2.9 3.5 0.0 0.8 1.5 2.3 3.0 3.8 4.6 MidShePubMin (mm) TerShePubMax (mm)
Number of Mid-Stem Sheath Trichomes per 0.5 cm Terminal Sheath Pubescence Minimum Length
25 30
25 20
20 15 15 Frequency
Frequency 10 10
5 5
0 0 0.0 7.3 14.7 22.0 29.3 36.7 44.0 0.0 0.7 1.4 2.1 2.8 3.5 4.2 NumSheTriMid T erShePubMin (mm)
Terminal Sheath Glabrous to Glabrate Number of Terminal Sheath Trichomes per 0.5 cm
45 30 40 25 35 30 20 25 15 20 Frequency Frequency 15 10 10 5 5 0 0 0 1 0.0 8.3 16.7 25.0 33.3 41.7 50.0 TerSheGla NumTerSheTri
Terminal Sheath Indument Length Sheath Pubescence Double Invested
9 26 8 7 21 6 16 5 4
Frequency Frequency 11 3 2 6 1 0 1 0.0 0.6 1.2 1.9 2.5 3.1 3.7 0 1 TerSheIndLen (mm) ShePubDou
100 Sheath Pubescence Orientation Peduncle Glabrous to Glabrate
30 40 35 25 30 20 25 15 20
Frequency Frequency 15 10 10 5 5 0 0 0 1 0 1 ShePubOri P edGla
Sheath Pubescence Pustulate Peduncle Pubescence Maximum Length
50 14 45 12 40 35 10 30 8 25 6 Frequency
Frequency 20 15 4 10 2 5 0 0 0 1 0.0 0.6 1.2 1.8 2.4 3.0 3.6 ShePubPus PedPubMax (mm)
Peduncle Exposed Above Subtending Sheath Peduncle Pubescence Minimum Length
10 25 9 8 20 7 6 15 5
Frequency 10
Frequency 4 3 2 5 1 0 0 0.0 25.8 51.7 77.5 103.3 129.2 155.0 0.0 0.4 0.9 1.3 1.7 2.2 2.6 PedExp (mm) PedPubMin (mm)
Peduncle Length Number of Peduncle Trichomes per 0.5 cm
12 16 14 10 12 8 10
6 8 Frequency Frequency 6 4 4 2 2
0 0 6.6 45.2 83.7 122.3 160.9 199.4 238.0 0.0 6.3 12.7 19.0 25.3 31.7 38.0 PedLen (mm) NumPedT ri
101 Peduncle Pubescence Double Invested Inflorescence with Double Vestiture
35 40
30 35
25 30 25 20 20 15 Frequency Frequency 15 10 10 5 5 0 0 0 1 0 1 PedPubDou InfDouVes
Peduncle Pubescence Orientation Inflorescence Pubescence Maximum Length
35 12 30 10 25 8 20 6 15 Frequency Frequency 10 4
5 2
0 0 0 1 0.3 0.8 1.4 1.9 2.4 3.0 3.5 PedPubOri InfPubMax (mm)
Peduncle Pubescence Pustulate Inflorescence Pubescence Minimum Length
50 30 45 40 25 35 20 30 25 15
Frequency 20 Frequency 15 10 10 5 5 0 0 0 1 0.0 0.2 0.4 0.6 0.7 0.9 1.1 PedPubPus InfPubMin (mm)
Inflorescence < a third as Long as Wide Inflorescence with Scabrous Pubescence
45 50 40 45 35 40 30 35 25 30 25 20 Frequency
Frequency 20 15 15 10 10 5 5 0 0 0 1 0 1 InfNar InfScaPub
102 Inflorescence Axis with Puberulent Pubescence Vernal Inflorescence Length
40 18 35 16 30 14 12 25 10 20 8
Frequency 15 Frequency 6 10 4 5 2 0 0 0 1 28.3 42.6 56.9 71.3 85.6 99.9 114.2 PubPubInf VerInfLen (mm)
Number of Spikelets in Inflorescence Vernal Inflorescence Length-Width Ratio
25 20 18 20 16 14 15 12 10 Frequency 10 Frequency 8 6 5 4 2 0 0 26 57 88 119 150 181 211 0.7 1.0 1.4 1.7 2.0 2.3 2.6 Num Sp i InfRat
Number of Nodes in Inflorescence Number of Trichomes Along 0.5 cm of Inflorescence Axis 16 14 30 12 25
10 20 8 15
Frequency 6 Frequency 10 4 2 5 0 0 6 8 9 11 12 14 15 0.0 6.7 13.3 20.0 26.7 33.3 40.0 NumInfNod NumTriInf
Vernal Inflorescence Width Pedicel Maximum Length
16 16 14 14 12 12 10 10 8 8 Frequency Frequency 6 6 4 4 2 2 0 0 15.3 28.6 41.8 55.1 68.3 82.6 95.9 6.2 8.6 11 13.4 15.8 18.2 20.4 VerInfWid (mm) PedLenMax (mm)
103 Pedicel Minimum Length Spikelet Pubescence Length
18 16 16 14 14 12 12 10 10 8 8 Frequency Frequency 6 6 4 4 2 2 0 0 0.5 1.1 1.8 2.4 3.0 3.7 4.3 0.4 0.6 0.7 0.9 1.0 1.2 1.3 PedLenMin (mm) SpiPubLen (mm)
Pedicel Pubescence First Glume Length
40 18 35 16 30 14 12 25 10 20 8 Frequency 15 Frequency 6 10 4 5 2 0 0 0 1 0.3 0.5 0.7 0.9 1.1 1.4 1.6 PedPub GluLen (mm)
Spikelet Length First Glume Width
16 20 14 18 16 12 14 10 12 8 10 Frequency 6 Frequency 8 6 4 4 2 2 0 0 1.4 1.7 1.9 2.2 2.5 2.7 3.0 0.3 0.4 0.5 0.6 0.7 0.8 0.9 SpiLen (mm) GluWid (mm)
Spikelet Width First Glume Length-Width Ratio
16 20 14 18 16 12 14 10 12 8 10 Frequency 6 Frequency 8 6 4 4 2 2 0 0 0.6 0.7 0.8 0.9 1.0 1.1 1.2 0.6 1 1.3 1.6 1.9 2.3 2.7 SpiWid (mm) GluRat
104 Spikelet Pointed at Apex Number of Veins on Second Glume
45 25 40 35 20 30 15 25 20
Frequency Frequency 10 15 10 5 5 0 0 0 1 1 3 5 7 9 11 13 Sp iP o i NumVeiSecGlu
Number of Veins on First Glume Number of Veins on Sterile Lemma
45 30 40 25 35 30 20 25 15 20 Frequency Frequency 15 10 10 5 5 0 0 0 1 5 6 7 8 9 10 11 NumVeiFirGlu NumVeiSteLem
First Glume Pubescent Fertile Floret Pointed
40 45 35 40 30 35 30 25 25 20 20 Frequency 15 Frequency 15 10 10 5 5 0 0 0 1 0 1 GluPub FerFloPoi
105 Appendix C. Operational Taxonomic Units (OTU) and associated character data for the Long Ligule group. OTU codes assigned by acronym of the species group followed by a number that corresponds to the order sampled, and ending with the state or country of collection. The prefix “T” indicates a type specimen.
(mm)
LigLen MidShePubMax MidSheGla AdLfPubMax AdaSurMidLeaGla TerShePubMax TerSheGla TerLeaAbaPubmin TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonLeaWid NumInfNod PubPub Inf PedGla PedPubDou AbaLeaSurPub InfNar FerFloPoi InfDouVes SpiPoi OTU Code LigTriUni IMPL-02-MN 2.50 1.00 1.90 0.00 2.10 0.00 1.90 0.00 0.30 0.70 1.50 0.83 0.53 65.40 4.00 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-04-MN 2.30 1.00 1.10 0.00 1.50 0.00 1.00 0.00 0.30 0.60 1.50 0.80 0.60 56.00 5.25 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-130-AR 3.80 1.00 2.50 0.00 3.70 0.00 3.20 0.00 0.45 0.89 1.71 0.82 0.50 65.20 5.00 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-134-ON 3.00 1.00 2.10 0.00 3.50 0.00 1.10 0.00 0.40 0.32 1.65 0.85 0.45 63.30 5.20 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-14-OH 2.20 1.00 1.70 0.00 2.50 0.00 2.20 0.00 0.30 0.80 1.60 0.82 0.49 53.70 6.00 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-169-TN 4.30 1.00 1.90 0.00 4.00 0.00 2.00 0.00 0.35 0.87 1.40 0.80 0.32 49.20 5.50 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-16-ME 2.70 1.00 3.20 0.00 3.30 0.00 3.00 0.00 0.50 2.00 1.50 0.75 0.59 67.80 5.10 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-170-MA 3.40 1.00 2.30 0.00 4.30 0.00 3.80 0.00 0.60 1.10 1.30 0.79 0.50 49.20 4.80 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-171-NY 3.30 1.00 2.10 0.00 3.70 0.00 4.90 0.00 0.55 1.00 1.39 0.79 0.45 58.60 5.50 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-173-RI 4.00 1.00 1.80 0.00 2.60 0.00 1.20 0.00 0.30 0.65 1.50 0.75 0.32 64.20 4.60 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-175-IA 3.50 1.00 2.60 0.00 2.80 0.00 1.50 0.00 0.30 0.80 1.35 0.70 0.30 63.30 6.30 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-176-MA 2.50 1.00 2.80 0.00 3.00 0.00 3.40 0.00 0.45 1.40 1.35 0.70 0.50 78.20 5.30 16.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-177-MN 2.00 1.00 1.70 0.00 2.70 0.00 1.60 0.00 0.32 0.75 1.40 0.80 0.40 41.70 3.80 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-178-MI 2.80 1.00 2.10 0.00 4.10 0.00 2.60 0.00 0.50 1.20 1.40 0.72 0.50 42.70 3.80 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-214-WA 2.70 1.00 2.30 0.00 3.00 0.00 2.70 0.00 0.40 1.00 1.60 0.82 0.51 57.60 7.00 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-215-OR 3.20 1.00 1.80 0.00 3.30 0.00 1.30 0.00 0.30 0.80 1.60 1.00 0.40 46.40 7.40 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-220-OR 3.30 1.00 1.80 0.00 2.90 0.00 0.90 0.00 0.27 0.50 1.69 0.97 0.49 52.20 4.70 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-50-WI 4.40 1.00 2.00 0.00 3.60 0.00 1.40 0.00 0.60 1.00 1.65 0.80 0.65 56.50 4.80 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IMPL-51-VT 4.40 1.00 2.30 0.00 5.90 0.00 4.30 0.00 0.75 1.20 1.40 0.70 0.55 63.50 5.50 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-IMPL-274-MA 3.20 1.00 2.10 0.00 2.95 0.00 2.30 0.00 0.39 1.70 1.38 0.70 0.60 61.30 6.30 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
106
LfPubMax AdaSurMidLeaGla AbaLeaSurPub InfNar FerFloPoi InfDouVes SpiPoi TerShePubMax TerSheGla TerLeaAbaPubmin TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonLeaWid NumInfNod PubPubInf PedGla PedPubDou OTU Code LigLen LigTriUni MidShePubMax MidSheGla Ad T-OCCI-257-WA 2.70 1.00 2.20 0.00 2.90 0.00 1.60 0.00 0.25 0.50 1.50 0.87 0.39 52.00 5.00 9.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-PACI-262-CA 2.40 1.00 2.20 0.00 2.00 0.00 2.50 0.00 0.32 0.59 1.62 0.83 0.49 54.90 4.30 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-FEPA-251-ALB 2.45 1.00 3.15 0.00 3.80 0.00 4.30 0.00 0.35 1.10 1.88 0.83 0.69 46.70 12.00 18.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-FESE-269-WY 2.20 1.00 2.10 0.00 1.80 0.00 2.00 0.00 0.30 0.55 1.65 0.81 0.43 51.60 8.30 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-01-OH 2.80 1.00 2.60 0.00 2.10 0.00 2.70 0.00 0.35 0.60 1.60 0.75 0.67 95.30 7.20 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-10-IN 3.10 1.00 2.30 0.00 1.80 0.00 2.60 0.00 0.20 0.50 1.65 0.85 0.63 75.20 5.90 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-11-OH 2.90 1.00 2.10 0.00 3.30 0.00 0.90 0.00 0.34 0.50 1.55 0.75 0.70 90.30 6.20 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-12-IL 2.40 1.00 1.80 0.00 1.85 0.00 1.40 0.00 0.29 0.60 1.50 0.80 0.50 70.20 6.00 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-131-KS 3.85 1.00 2.40 0.00 2.33 0.00 1.20 0.00 0.30 0.48 1.60 0.82 0.55 90.70 7.50 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-132-LA 2.85 1.00 2.10 0.00 1.39 0.00 1.85 0.00 0.35 0.40 1.51 0.80 0.45 69.70 6.00 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-133-QU 3.30 1.00 1.80 0.00 2.10 0.00 0.80 0.00 0.30 0.40 1.50 0.75 0.48 81.30 7.70 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-135-KY 3.00 1.00 1.80 0.00 2.70 0.00 1.50 0.00 0.25 0.55 1.50 0.90 0.50 73.70 6.40 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-136-PA 3.30 1.00 2.40 0.00 3.20 0.00 2.90 0.00 0.25 0.52 1.60 0.89 0.45 84.50 6.80 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-137-NJ 3.30 1.00 2.00 0.00 2.60 0.00 2.10 0.00 0.30 0.49 1.68 0.90 0.57 82.30 6.20 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-138-NE 2.40 1.00 1.90 0.00 1.30 0.00 1.70 0.00 0.10 0.65 1.55 0.80 0.45 86.70 7.80 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-141-MD 3.80 1.00 1.80 0.00 4.10 0.00 1.80 0.00 0.25 0.60 1.60 0.80 0.52 82.00 6.60 9.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-142-MA 3.50 1.00 1.85 0.00 2.40 0.00 0.90 0.00 0.25 0.62 1.70 0.80 0.45 68.60 8.20 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-143-IL 2.90 1.00 1.89 0.00 1.85 0.00 2.00 0.00 0.29 0.70 1.67 0.90 0.60 110.20 9.30 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-144-IL 2.80 1.00 1.90 0.00 3.20 0.00 1.40 0.00 0.49 0.69 1.50 0.85 0.35 90.00 5.90 17.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-145-IA 3.10 1.00 1.60 0.00 2.60 0.00 1.00 0.00 0.35 0.51 1.50 0.80 0.60 78.20 5.70 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-146-VA 3.00 1.00 2.10 0.00 2.20 0.00 1.20 0.00 0.10 0.23 1.60 0.80 0.50 80.50 7.70 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-148-OK 3.40 1.00 2.60 0.00 3.90 0.00 3.85 0.00 0.37 0.95 1.60 0.90 0.50 87.50 8.40 19.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-149-MO 2.70 1.00 1.60 0.00 1.70 0.00 1.40 0.00 0.20 0.40 1.35 0.80 0.52 73.90 7.60 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-152-NY 2.60 1.00 2.30 0.00 2.30 0.00 1.30 0.00 0.00 0.67 1.36 0.79 0.29 78.70 9.10 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-155-PA 3.40 1.00 2.20 0.00 1.80 0.00 2.00 0.00 0.00 0.80 1.50 0.80 0.45 95.20 10.70 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-211-OR 2.70 1.00 2.20 0.00 3.30 0.00 2.00 0.00 0.20 0.60 1.80 0.90 0.55 111.40 9.90 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
107
AbaLeaSurPub InfNar FerFloPoi InfDouVes SpiPoi TerShePubMax TerSheGla TerLeaAbaPubmin TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonLeaWid NumInfNod PubPubInf PedGla PedPubDou OTU Code LigLen LigTriUni MidShePubMax MidSheGla AdLfPubMax AdaSurMidLeaGla LANU-212-CA 3.10 1.00 2.10 0.00 2.40 0.00 1.75 0.00 0.40 0.72 1.70 0.95 0.75 71.50 6.40 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-213-WA 3.20 1.00 1.70 0.00 1.40 0.00 1.10 0.00 0.33 0.40 1.55 0.98 0.60 63.30 6.65 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-217-WA 2.40 1.00 1.40 0.00 0.00 0.00 1.25 0.00 0.20 0.45 1.62 0.85 0.53 77.00 8.00 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-218-VAN 3.30 1.00 1.75 0.00 4.10 0.00 2.10 0.00 0.32 0.80 1.70 1.10 0.62 80.00 7.50 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-219-BC 3.80 1.00 2.40 0.00 2.50 0.00 3.50 0.00 0.80 1.30 1.62 0.98 0.43 86.80 9.20 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LANU-42-AR 2.90 1.00 1.90 0.00 3.40 0.00 1.50 0.00 0.20 0.70 1.65 0.80 0.53 108.20 10.00 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-HUSI-273-DC 2.70 1.00 2.00 0.00 1.60 0.00 2.40 0.00 0.35 0.70 1.50 0.85 0.50 86.30 7.30 14.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-BROD-248-WA 4.30 1.00 3.00 0.00 2.10 1.00 2.20 0.00 0.22 0.80 1.78 0.86 0.39 98.30 9.99 16.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-UNPR-266-MA 2.85 1.00 1.90 0.00 2.20 0.00 1.60 0.00 0.30 0.52 1.80 0.90 0.65 70.50 8.10 9.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-HUAC-272-AZ 3.30 1.00 1.90 0.00 1.50 0.00 2.00 0.00 0.30 0.90 1.70 0.82 0.45 90.50 4.90 17.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-FUNS-271-CA 3.00 1.00 1.20 0.00 0.00 1.00 0.90 0.00 0.10 0.15 1.52 0.81 0.50 81.40 5.90 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-150-MO 2.40 1.00 1.50 0.00 1.40 1.00 0.75 0.00 0.00 0.55 1.40 0.70 0.32 83.50 7.90 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-151-QU 3.00 1.00 1.30 0.00 1.85 1.00 0.80 0.00 0.18 0.12 1.60 0.82 0.40 69.70 6.40 17.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-153-OK 2.80 1.00 1.60 0.00 2.20 1.00 1.30 0.00 0.00 0.50 1.50 0.78 0.52 91.30 5.40 16.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-154-SC 3.90 1.00 1.90 0.00 4.20 1.00 1.50 0.00 0.00 0.45 1.67 0.90 0.40 81.30 7.80 10.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-156-TX 3.70 1.00 2.00 0.00 0.00 1.00 1.30 0.00 0.00 0.32 1.59 0.89 0.42 66.10 7.20 15.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-157-NE 3.70 1.00 1.00 0.00 1.85 1.00 1.10 0.00 0.00 0.49 1.65 0.98 0.55 67.00 5.15 13.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-158-UT 2.40 1.00 2.30 0.00 2.40 1.00 1.50 0.00 0.00 0.65 1.68 0.89 0.70 88.50 5.50 12.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TENN-159-WI 3.30 1.00 1.50 0.00 4.20 1.00 1.30 0.00 0.10 0.10 1.50 0.79 0.51 77.60 8.50 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LEUC-172-TX 2.50 0.00 0.75 0.00 2.00 0.00 0.41 0.00 0.25 0.25 1.32 0.72 0.48 40.00 4.80 13.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 LEUC-194-FL 2.60 0.00 0.72 0.00 1.20 1.00 0.30 0.00 0.25 0.10 1.40 0.69 0.49 38.20 3.90 14.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 LEUC-197-SC 2.20 0.00 0.63 0.00 0.49 1.00 0.50 0.00 0.15 0.25 1.20 0.60 0.43 35.20 4.80 12.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 LEUC-198-FL 2.60 0.00 0.75 1.00 0.40 1.00 0.37 0.00 0.10 0.10 1.25 0.68 0.40 47.40 4.60 11.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 LEUC-199-TN 2.60 1.00 1.40 0.00 1.00 1.00 1.60 0.00 0.10 0.10 1.25 0.67 0.35 39.40 3.30 15.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 T-LEUC-243-FL 2.45 0.00 0.50 0.00 1.50 1.00 0.44 0.00 0.10 0.10 1.20 0.58 0.32 37.50 4.73 12.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00 T-PARV-240-GA 3.10 1.00 1.50 0.00 1.40 1.00 1.10 0.00 0.10 0.10 1.26 0.62 0.49 63.50 5.30 15.00 0.00 0.00 0.00 1.00 0.00 1.00 1.00 0.00
108
eaWid AbaLeaSurPub InfNar FerFloPoi InfDouVes SpiPoi TerShePubMax TerSheGla TerLeaAbaPubmin TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonL NumInfNod PubPubInf PedGla PedPubDou OTU Code LigLen LigTriUni MidShePubMax MidSheGla AdLfPubMax AdaSurMidLeaGla LIND-09-MI 2.30 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.40 0.90 0.33 70.40 5.70 12.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-160-KY 3.10 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.32 0.79 0.38 68.50 4.20 11.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-161-NY 3.20 0.00 0.80 0.00 0.00 1.00 0.00 1.00 0.00 0.00 1.69 0.95 0.50 81.40 8.50 14.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-162-AR 2.60 0.00 1.50 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.40 0.80 0.40 61.50 3.50 13.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-163-MI 2.10 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.50 0.80 0.40 61.40 5.90 10.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-164-TX 2.10 0.00 0.00 1.00 4.00 1.00 0.00 1.00 0.00 0.00 1.42 0.75 0.40 84.80 7.70 14.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-165-MO 2.40 0.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.60 0.81 0.40 75.90 5.00 11.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-166-MO 2.50 0.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.50 0.80 0.35 50.80 6.10 9.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-168-NY 2.20 0.00 1.40 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.52 0.95 0.60 87.80 6.80 15.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LIND-216-CA 2.40 1.00 0.00 1.00 2.10 1.00 0.00 1.00 0.00 0.00 1.50 0.80 0.30 69.50 6.10 14.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 T-LIND-245-TX 2.00 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.42 0.75 0.31 72.10 8.50 15.00 0.00 1.00 0.00 0.00 0.00 1.00 0.00 0.00 T-LISE-253-NB 2.30 1.00 1.60 0.00 0.00 1.00 0.00 1.00 0.00 0.00 1.60 0.90 0.50 59.60 5.81 11.00 0.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 LONG-189-TN 1.80 1.00 0.00 1.00 0.00 1.00 0.30 1.00 0.10 0.00 1.30 0.60 0.50 62.60 4.20 13.00 0.00 1.00 0.00 1.00 0.00 1.00 1.00 0.00 LONG-190-NC 3.10 1.00 0.40 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.50 0.76 0.42 53.20 3.30 12.00 0.20 0.00 0.00 1.00 0.00 1.00 1.00 0.00 LONG-191-FL 2.60 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.10 1.26 0.69 0.25 45.20 4.90 12.00 0.00 1.00 0.00 1.00 0.00 1.00 1.00 0.00 LONG-192-GA 2.30 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.19 0.63 0.39 51.80 4.00 28.00 0.00 1.00 0.00 1.00 0.00 1.00 1.00 0.00 LONG-193-NIGU 2.70 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.10 0.10 1.50 0.81 0.40 52.00 3.50 10.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00 T-LONG-246-FL 2.60 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.10 0.10 1.30 0.68 0.40 22.20 2.90 15.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 MERI-174-VA 2.80 1.00 3.60 0.00 3.80 0.00 2.80 0.00 0.30 1.30 1.50 0.89 0.70 48.30 3.10 10.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 MERI-179-NY 2.60 1.00 2.60 0.00 2.50 0.00 0.30 0.00 0.25 0.80 1.50 0.80 0.69 47.40 2.80 9.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 MERI-180-VA 2.10 0.00 1.70 0.00 3.40 0.00 0.80 0.00 0.10 0.60 1.42 0.91 0.60 58.50 4.60 12.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 MERI-181-MA 3.00 1.00 1.80 0.00 3.70 0.00 2.40 0.00 0.45 1.20 1.50 0.81 0.60 48.50 3.80 9.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 MERI-182-TN 3.20 1.00 3.10 0.00 4.50 0.00 1.40 0.00 0.70 1.00 1.43 0.80 0.70 53.90 4.60 13.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 MERI-183-NC 2.80 0.00 1.90 0.00 2.80 0.00 0.30 0.00 0.20 0.52 1.40 0.82 0.72 30.10 3.10 7.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 T-MERI-260-NC 2.30 1.00 2.10 0.00 2.90 0.00 0.30 0.00 0.10 0.20 1.49 0.79 0.55 46.80 3.80 7.00 1.00 0.00 0.00 1.00 0.00 0.00 1.00 0.00 T-MICR-285-NC 2.20 0.00 1.90 0.00 2.00 0.00 0.30 0.00 0.20 0.20 1.60 0.80 0.55 35.50 2.80 8.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00
109
rFloPoi AbaLeaSurPub InfNar Fe InfDouVes SpiPoi TerShePubMax TerSheGla TerLeaAbaPubmin TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonLeaWid NumInfNod PubPubInf PedGla PedPubDou OTU Code LigLen LigTriUni MidShePubMax MidSheGla AdLfPubMax AdaSurMidLeaGla T-ALBE-281-NC 3.10 0.00 2.70 0.00 3.70 0.00 2.00 0.00 0.40 1.10 1.32 0.70 0.40 66.80 3.80 11.00 1.00 0.00 1.00 1.00 0.00 0.00 1.00 0.00 PRAE-07-KS 4.70 0.00 4.50 0.00 6.10 0.00 3.30 0.00 0.65 1.95 1.80 1.00 0.80 87.30 4.90 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-117-MO 3.60 0.00 3.60 0.00 4.80 0.00 4.80 0.00 0.80 0.95 1.70 0.90 0.90 68.80 5.25 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-118-MO 3.20 0.00 3.50 0.00 3.80 0.00 3.00 0.00 0.80 0.92 2.00 1.10 0.83 60.30 5.10 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-119-MO 2.00 0.00 3.90 0.00 5.30 0.00 4.00 0.00 0.40 0.95 1.85 1.10 0.96 76.00 4.90 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-120-MO 4.40 0.00 2.40 0.00 3.00 0.00 2.40 0.00 0.40 0.75 1.75 0.81 0.85 74.60 4.00 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-121-OK 2.10 0.00 4.10 0.00 5.40 0.00 4.70 0.00 0.60 1.20 1.95 1.10 1.00 61.20 5.50 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-122-IL 2.50 0.00 4.20 0.00 4.50 0.00 3.50 0.00 0.63 0.95 1.90 1.20 0.83 73.70 6.00 5.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-123-AR 2.40 0.00 4.40 0.00 5.80 0.00 3.70 0.00 0.50 1.20 1.90 1.05 0.73 76.10 5.85 9.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-124-NE 2.00 0.00 3.40 0.00 4.20 0.00 3.90 0.00 0.65 1.30 1.90 1.10 0.92 62.30 5.85 6.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-125-NE 2.00 0.00 4.40 0.00 3.70 0.00 4.30 0.00 0.60 1.10 1.83 0.85 0.70 70.00 5.25 6.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-126-IA 2.30 0.00 4.40 0.00 3.00 0.00 3.80 0.00 0.50 1.15 1.80 1.00 0.85 59.20 5.90 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-127-NE 1.80 0.00 3.40 0.00 3.20 0.00 3.80 0.00 0.75 0.80 1.87 1.05 0.87 66.40 6.80 9.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-128-NE 2.00 0.00 3.60 0.00 3.60 0.00 3.60 0.00 0.50 0.85 1.85 1.10 0.75 60.00 5.30 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-129-TX 2.00 0.00 3.40 0.00 4.20 0.00 3.40 0.00 0.80 0.82 1.85 0.95 0.89 74.70 5.30 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-21-IL 2.80 0.00 2.90 0.00 3.40 0.00 2.90 0.00 0.60 1.30 1.93 1.00 0.85 80.00 5.90 9.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-24-OK 3.10 0.00 3.80 0.00 4.20 0.00 4.00 0.00 0.50 1.20 2.10 1.10 0.73 75.70 6.20 12.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-25-MN 4.00 0.00 3.90 0.00 4.40 0.00 4.50 0.00 0.55 1.00 1.90 1.00 0.85 71.90 6.00 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-26-NE 2.70 0.00 4.10 0.00 2.70 0.00 4.10 0.00 0.80 1.40 1.90 1.00 0.77 64.40 5.00 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-315 2.80 0.00 3.60 0.00 4.00 0.00 3.80 0.00 0.50 1.30 1.80 0.90 0.80 75.60 6.00 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-38-MO 4.70 0.00 2.90 0.00 3.00 0.00 2.90 0.00 0.60 1.20 1.85 1.00 0.87 59.40 4.80 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-39-MO 3.40 0.00 2.30 0.00 2.40 0.00 1.80 0.00 0.30 0.70 1.83 0.96 0.60 57.50 3.50 6.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-40-MO 3.80 0.00 3.70 0.00 3.60 0.00 3.70 0.00 0.80 1.10 1.83 0.95 1.00 79.00 5.10 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-43-AR 2.30 0.00 2.70 0.00 3.70 0.00 2.50 0.00 0.30 0.61 1.79 0.90 0.70 82.30 3.60 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-44-NE 4.20 0.00 3.25 0.00 5.60 0.00 3.50 0.00 0.40 1.40 1.65 0.95 0.90 72.70 5.30 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 PRAE-45-NE 3.20 0.00 4.00 0.00 2.40 0.00 4.00 0.00 0.45 1.10 1.83 1.00 0.63 78.50 6.25 7.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
110
bMax
AbaLeaSurPub InfNar FerFloPoi InfDouVes SpiPoi TerLeaAbaPubMax SpiLen SpiWid PriGluLen LonLeaLen LonLeaWid NumInfNod PubPubInf PedGla PedPubDou TerSheGla TerLeaAbaPubmin TerShePu OTU Code LigLen LigTriUni MidShePubMax MidSheGla AdLfPubMax AdaSurMidLeaGla PRAE-167-WI 3.10 0.00 4.20 0.00 5.00 0.00 4.40 0.00 0.45 1.45 1.95 1.10 0.80 70.50 6.50 11.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 T-PRAE-263-IL 3.20 0.00 3.60 0.00 5.30 0.00 3.80 0.00 0.45 1.60 1.80 0.95 0.82 91.10 6.90 8.00 1.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 SPRE-206-NJ 2.50 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.79 0.72 0.60 93.10 5.50 21.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 SPRE-207-MA 2.10 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.76 0.81 0.52 79.00 4.60 19.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 SPRE-208-NS 2.10 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.80 0.87 0.70 56.50 4.60 18.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 SPRE-209-IN 1.70 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.60 0.72 0.50 78.90 4.40 15.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 SPRE-210-IN 2.40 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.60 0.79 0.53 80.50 4.30 16.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 T-EATO-237-NH 1.90 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.72 0.83 0.67 94.70 4.50 21.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 T-OCTO-261-TX 2.00 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.37 0.71 0.42 105.40 6.00 18.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 T-PAUC-239-NJ 2.00 1.00 0.00 1.00 0.00 1.00 0.00 1.00 0.00 0.00 1.40 0.72 0.40 70.80 5.32 18.00 0.00 1.00 0.00 0.00 1.00 1.00 0.00 1.00 T-WRIG-258-CUB 2.10 1.00 0.20 0.00 0.35 0.00 0.15 0.00 0.10 0.20 0.93 0.50 0.30 38.30 3.60 10.00 1.00 0.00 1.00 1.00 0.00 1.00 1.00 0.00 WRIG-184-MA 2.40 1.00 0.30 0.00 0.40 0.00 0.20 0.00 0.10 0.10 1.00 0.55 0.25 33.70 4.30 14.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00 WRIG-185-FL 2.90 1.00 0.30 0.00 0.50 0.00 0.20 0.00 0.30 0.20 0.90 0.50 0.27 36.60 4.50 11.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00 WRIG-186-GA 2.20 1.00 0.35 0.00 0.40 0.00 0.20 0.00 0.10 0.10 0.90 0.52 0.39 33.10 4.70 14.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00 WRIG-187-MS 2.00 1.00 0.80 0.00 0.40 0.00 0.20 0.00 0.20 0.20 0.90 0.45 0.31 39.60 4.80 15.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00 WRIG-188-VA 2.40 1.00 0.20 0.00 0.30 0.00 0.20 0.00 0.20 0.10 0.98 0.58 0.35 23.10 3.60 10.00 1.00 0.00 1.00 1.00 0.00 1.00 1.00 0.00 T-STRI-264-NC 2.10 1.00 0.15 0.00 3.20 0.00 0.15 0.00 0.15 0.15 0.85 0.41 0.27 32.00 3.20 4.00 0.00 0.00 1.00 1.00 0.00 1.00 1.00 0.00 T-DEMI-278-NY 1.90 1.00 0.10 0.00 0.20 0.00 0.10 0.00 0.15 0.10 0.88 0.49 0.30 23.10 2.20 8.00 0.50 0.00 1.00 1.00 0.00 1.00 1.00 0.00
111 Appendix D. Operational Taxonomic Units (OTU) and associated character data for the Short Ligule group. OTU codes assigned by acronym of the species group followed by a number that corresponds to the order sampled, and ending with the state or country of collection. The prefix “T” indicates a type specimen.
0
OTU Code LigLen DouLigCon DouVes NumTriAdaLeaSur LonLeaWid MidLeaGlaAda TerLeaGlaAda SpiLen PriGluLen MidSheIndLen ShePubOri Cilia>3 ACUM-139-FL 0.80 2.00 0.00 17.00 9.30 0.00 0.00 1.70 0.49 1.40 1.00 1.00 ACUM-140-GA 0.90 2.00 0.00 21.00 6.00 0.00 0.00 1.71 0.52 1.00 1.00 1.00 ACUM-147-TX 1.00 2.00 1.00 37.00 6.10 0.00 0.00 1.73 0.52 2.30 1.00 1.00 ACUM-300-PR 0.90 2.00 0.00 19.00 7.10 0.00 0.00 1.73 0.50 0.80 1.00 1.00 ACUM-301-JA 0.70 2.00 0.00 15.00 6.70 0.00 0.00 1.65 0.51 1.00 1.00 1.00 ACUM-302-W.I. 1.00 2.00 0.00 26.00 7.80 0.00 0.00 1.60 0.49 1.10 1.00 1.00 ACUM-46-NI 0.60 2.00 0.00 16.00 8.30 0.00 0.00 2.10 0.85 0.60 1.00 1.00 ACUM-47-HT 1.30 2.00 0.00 23.00 6.80 0.00 0.00 1.79 0.54 1.30 1.00 1.00 ACUM-48-BE 0.67 2.00 0.00 12.00 5.90 0.00 0.00 1.77 0.83 0.67 1.00 1.00 OLIV-224-MX 0.78 2.00 0.00 17.00 6.70 0.00 0.00 1.75 0.65 0.50 1.00 1.00 OLIV-225-HN 0.68 2.00 0.00 19.00 7.50 0.00 0.00 1.73 0.50 1.10 1.00 1.00 OLIV-226-MX 0.80 2.00 0.00 31.00 8.20 0.00 0.00 1.72 0.55 1.00 1.00 1.00 OLIV-227-MX 0.75 2.00 1.00 3.00 7.20 0.00 0.00 1.70 0.68 0.25 1.00 1.00 OLIV-228-MX 0.85 2.00 0.00 25.00 8.50 0.00 0.00 1.70 0.68 0.92 1.00 1.00 OLIV-303-PN 0.70 2.00 0.00 20.00 4.80 0.00 0.00 1.67 0.60 0.95 1.00 1.00 OLIV-304-CL 0.60 2.00 0.00 26.00 5.20 0.00 0.00 1.80 0.60 1.00 1.00 1.00 OLIV-305-CR 0.70 2.00 0.00 18.00 5.20 0.00 0.00 1.80 0.65 0.85 1.00 1.00 THUR-201-TX 1.10 2.00 0.20 33.00 8.50 0.00 0.00 1.85 0.57 1.80 1.00 1.00 THUR-202-LA 0.60 2.00 0.00 26.00 7.10 0.00 0.00 1.80 0.51 1.25 1.00 1.00 THUR-203-LA 0.50 2.00 0.00 13.00 6.70 0.00 0.00 1.70 0.47 1.30 1.00 1.00 THUR-205-TX 0.82 2.00 0.00 20.00 9.00 0.00 0.00 2.00 0.65 1.40 1.00 1.00 THUR-229-TX 0.95 2.00 0.00 9.00 4.80 0.00 0.00 1.90 0.70 1.80 1.00 1.00 T-GLUT-252-VA 0.72 2.00 0.00 18.00 6.40 0.00 0.00 1.79 0.55 0.65 1.00 1.00 T-FERV-250-WY 0.85 2.00 1.00 19.00 7.40 0.00 0.00 1.40 0.45 1.10 1.00 1.00 T-OLIV-247-GU 0.60 2.00 1.00 16.00 8.00 0.00 0.00 1.80 0.75 1.00 1.00 1.00 T-OLIV-284-GUAT 0.47 2.00 0.00 28.00 8.30 0.00 0.00 1.70 0.60 1.10 1.00 1.00 T-ORAN-286-NC 0.80 2.00 0.00 20.00 5.40 0.00 0.00 1.65 0.70 1.10 1.00 1.00 T-CILI-280-MS 0.50 2.00 0.00 5.00 9.20 0.00 0.00 1.65 0.52 0.80 1.00 1.00 T-COMO-235-PR 0.60 2.00 0.00 65.00 12.40 0.00 0.00 1.80 0.55 1.10 1.00 1.00 T-THUR-265-TX 0.62 2.00 0.00 6.00 5.00 0.00 0.00 1.90 0.50 1.70 1.00 1.00 T-AUBU-259-AL 0.60 2.00 0.00 6.00 5.90 0.00 0.00 1.85 0.50 1.60 1.00 0.00 COLU-100-MA 1.50 3.00 3.00 0.00 3.70 1.00 1.00 1.60 0.70 0.60 0.25 0.00 COLU-101-MD 1.20 3.00 3.00 0.00 3.50 1.00 1.00 1.65 0.65 0.30 0.25 0.00 COLU-102-NY 1.40 3.00 3.00 0.00 3.70 1.00 1.00 1.90 1.05 0.60 0.25 0.00 COLU-103-OH 1.50 3.00 3.00 2.00 5.50 1.00 1.00 1.50 0.60 0.80 0.00 0.00 COLU-104-OH 1.50 3.00 3.00 0.00 3.40 1.00 1.00 1.60 0.65 0.30 0.00 0.00 COLU-77-NY 1.10 3.00 3.00 2.00 4.80 1.00 1.00 1.90 0.93 0.80 0.00 0.00
112
laAda
OTU Code LigLen DouLigCon DouVes NumTriAdaLeaSur LonLeaWid MidLeaG TerLeaGlaAda SpiLen PriGluLen MidSheIndLen ShePubOri Cilia>30 COLU-80-DE 1.01 3.00 3.00 1.00 5.50 1.00 1.00 2.10 0.80 0.70 0.00 0.00 T-ADDI-238-NJ 1.20 3.00 3.00 0.00 6.00 1.00 1.00 1.88 0.80 0.30 0.25 0.00 T-IMSI-275-IL 1.00 3.00 3.00 5.00 3.40 1.00 1.00 1.60 0.58 1.00 0.25 0.00 T-OWEN-234-MA 0.95 3.00 3.00 2.00 2.40 1.00 1.00 1.90 0.75 0.80 0.25 0.00 T-SCOP-283-DE 1.10 3.00 3.00 3.00 6.90 1.00 1.00 2.45 1.00 1.30 0.50 0.00 COMM-66-NJ 0.80 3.00 3.00 4.00 5.10 1.00 1.00 2.40 1.50 0.80 0.00 0.00 COMM-68-MA 1.05 3.00 2.50 3.00 3.90 1.00 1.00 2.40 1.35 0.80 0.50 0.00 COMM-69-MA 0.50 3.00 3.00 5.00 4.40 1.00 1.00 2.60 1.30 1.40 0.00 0.00 COMM-70-DE 1.10 3.00 3.00 2.00 5.50 1.00 1.00 2.50 1.32 1.65 0.25 0.00 COMM-71-MA 0.70 3.00 3.00 2.00 4.50 1.00 1.00 2.70 1.80 1.60 0.00 0.00 COMM-72-MD 0.80 3.00 3.00 0.00 5.40 1.00 1.00 2.30 1.30 1.70 0.25 0.00 COMM-73-DE 0.90 3.00 3.00 0.00 6.40 1.00 1.00 2.30 1.30 1.20 0.00 0.00 COMM-74-NJ 0.80 3.00 3.00 2.00 5.00 1.00 1.00 2.30 1.50 0.50 0.00 0.00 COMM-75-NJ 0.70 3.00 3.00 2.00 5.80 1.00 1.00 2.55 1.50 1.00 0.50 0.00 COMM-76-NJ 0.70 3.00 3.00 8.00 4.90 1.00 1.00 2.65 1.40 1.30 0.25 0.00 COMM-78-NY 0.75 3.00 3.00 3.00 3.90 1.00 1.00 2.40 1.50 0.90 0.00 0.00 COMM-79-NJ 0.80 3.00 3.00 0.00 5.00 1.00 1.00 2.20 1.20 1.00 0.00 0.00 COMM-W-108-TX 1.35 3.00 3.00 9.00 6.15 1.00 1.00 2.35 1.20 3.70 0.75 0.00 COMM-W-53-MO 1.10 3.00 3.00 14.00 4.70 1.00 1.00 2.50 1.20 1.30 0.50 0.00 COMM-W-59-IN 1.00 3.00 3.00 9.00 6.10 1.00 1.00 2.20 1.20 1.90 0.50 0.00 COMM-W-89-WI 0.90 3.00 3.00 14.00 5.00 1.00 1.00 2.35 1.10 2.40 0.50 0.00 T-COMM-279-NJ 0.86 3.00 2.00 5.00 4.10 1.00 1.00 2.32 1.25 0.50 0.00 0.00 T-EUCL-255-WI 1.00 3.00 2.50 13.00 6.20 1.00 1.00 2.50 1.20 2.20 0.75 0.00 OVAL-29-FL 0.50 2.00 0.00 6.00 7.00 1.00 1.00 2.75 0.87 1.35 0.50 0.00 OVAL-30-SC 0.67 2.00 0.00 4.00 8.80 1.00 1.00 2.45 0.78 1.40 0.50 0.00 OVAL-31-NC 0.50 2.00 0.00 5.00 4.70 1.00 1.00 2.20 0.87 1.35 0.50 0.00 OVAL-32-GA 0.42 2.00 0.00 0.00 7.00 1.00 1.00 2.57 0.96 1.60 0.75 0.00 OVAL-33-FL 0.30 2.00 0.00 1.00 8.10 1.00 1.00 2.60 1.20 2.30 1.00 0.00 OVAL-34-NJ 0.80 2.00 2.00 3.00 5.30 1.00 1.00 2.40 1.08 0.40 0.00 0.00 OVAL-36-SC 0.35 2.00 0.00 1.00 8.30 1.00 1.00 2.20 0.80 1.40 0.75 0.00 OVAL-37-NJ 0.50 2.00 3.00 4.00 5.20 1.00 1.00 2.50 1.35 1.00 0.00 0.00 OVAL-55-SC 0.65 2.00 1.00 3.00 5.30 1.00 1.00 2.50 1.30 1.10 0.25 0.00 OVAL-63-NC 0.60 2.00 2.00 1.00 5.70 1.00 1.00 2.55 1.35 0.80 0.00 0.00 OVAL-64-FL 0.32 2.00 1.00 4.00 5.50 1.00 1.00 2.65 1.35 1.60 0.50 0.00 OVAL-65-FL 0.25 2.00 1.00 3.00 8.00 1.00 1.00 2.85 1.25 1.80 0.75 0.00 OVAL-67-AL 0.60 2.00 1.00 3.00 7.80 1.00 1.00 2.62 1.45 0.90 0.50 0.00 OVAL-81-VA 0.48 2.00 1.00 3.00 5.30 1.00 1.00 2.25 1.00 2.00 0.00 0.00 OVAL-82-SC 0.70 2.00 1.00 5.00 5.40 1.00 1.00 2.15 1.00 1.70 0.00 0.00 OVAL-83-MA 0.60 2.00 1.00 2.00 8.60 1.00 1.00 2.40 1.00 0.80 0.00 0.00 OVAL-84-SC 0.40 2.00 1.00 1.00 6.60 1.00 1.00 2.60 1.30 3.30 0.25 0.00 OVAL-85-FL 0.40 2.00 2.00 4.00 7.40 1.00 1.00 2.25 0.85 1.30 0.75 0.00
113
igCon
OTU Code LigLen DouL DouVes NumTriAdaLeaSur LonLeaWid MidLeaGlaAda TerLeaGlaAda SpiLen PriGluLen MidSheIndLen ShePubOri Cilia>30 OVAL-86-SC 0.62 2.00 1.00 2.00 7.80 1.00 1.00 2.32 1.00 1.70 0.00 0.00 OVAL-87-SC 0.50 2.00 1.00 0.00 5.00 1.00 1.00 2.15 1.00 1.30 0.00 0.00 OVAL-88-FL 0.40 2.00 1.00 4.00 4.00 1.00 1.00 2.90 1.50 1.50 0.75 0.00 PSEU-54-GA 0.60 2.00 0.00 3.00 6.90 1.00 1.00 2.30 1.00 2.30 0.25 0.00 PSEU-56-SC 0.85 2.00 1.00 5.00 5.20 1.00 1.00 2.30 1.10 1.60 0.00 0.00 PSEU-57-FL 0.50 2.00 1.00 1.00 8.50 1.00 1.00 2.25 1.00 1.30 0.25 0.00 PSEU-58-VA 0.60 2.00 1.00 5.00 6.10 1.00 1.00 2.40 1.30 2.00 0.00 0.00 PSEU-60-TN 0.60 2.00 1.00 3.00 5.60 1.00 1.00 2.50 0.85 1.00 0.50 0.00 PSEU-61-AL 0.50 2.00 0.00 1.00 6.20 1.00 1.00 2.25 0.90 1.60 0.75 0.00 PSEU-62-AL 0.50 2.00 1.00 3.00 5.80 1.00 1.00 2.40 1.20 1.80 1.00 0.00 T-ERYT-277-NC 0.46 2.00 0.00 3.00 7.40 1.00 1.00 2.33 1.03 1.80 0.50 0.00 T-WILM-267-NC 0.50 2.00 1.00 4.00 4.00 1.00 1.00 2.20 0.87 1.10 0.50 0.00 T-CILI-256-FL 0.45 2.00 1.00 3.00 7.20 1.00 1.00 2.60 1.12 0.60 0.75 0.00 T-BENN-249-NJ 0.90 2.00 2.00 3.00 7.00 1.00 1.00 2.20 0.89 1.70 0.75 0.00 SUBV-200-OH 1.30 2.00 0.00 23.00 6.50 0.00 0.00 1.80 0.73 1.50 1.00 0.00 SUBV-221-MN 0.90 2.50 0.00 24.00 5.00 0.00 0.00 1.89 1.00 1.90 0.75 0.00 SUBV-222-NY 1.00 2.00 1.00 23.00 4.50 0.00 0.00 1.80 0.77 1.15 0.75 0.00 SUBV-233-WI 0.88 2.00 1.00 23.00 6.20 0.00 0.00 1.75 0.67 1.45 1.00 0.00 SUBV-311-ME 1.00 2.00 0.00 16.00 6.00 0.00 0.00 1.85 0.85 1.40 0.50 0.00 SUBV-312-MI 0.90 2.00 1.00 16.00 5.00 0.00 0.00 1.78 0.95 1.50 0.75 0.00 SUBV-313-WI 0.80 2.00 1.00 15.00 4.50 0.00 0.00 1.89 0.97 1.50 0.50 0.00 T-SUBV-287-MN 0.80 2.00 1.00 21.00 4.40 0.00 0.00 1.63 0.73 1.50 0.75 0.00 THER-230-CA 1.20 3.00 2.00 17.00 8.00 0.00 0.00 2.00 1.02 0.60 0.75 0.00 THER-231-CA 0.70 3.00 2.00 19.00 7.40 0.00 0.00 1.80 0.70 0.40 0.75 0.00 T-LASS-276-CA 0.95 3.00 2.00 15.00 6.70 0.00 0.00 1.75 0.70 1.40 1.00 0.00 T-THER-232-CA 0.75 3.00 2.00 19.00 7.40 0.00 0.00 1.87 0.90 1.00 0.75 0.00 T-THER-254-CA 0.60 3.00 2.00 25.00 5.00 0.00 0.00 1.85 0.62 1.00 1.00 0.00 VILL-06-IL 0.68 2.00 0.00 22.00 6.60 0.00 0.00 1.80 0.67 2.80 1.00 0.00 VILL-105-TX 0.60 2.00 0.00 20.00 6.10 0.00 0.00 2.30 0.75 2.45 1.00 0.00 VILL-106-TN 0.60 2.00 0.00 48.00 7.40 0.00 0.00 2.05 0.80 3.10 0.75 0.00 VILL-107-TN 0.50 2.00 0.50 20.00 5.40 0.00 0.00 2.00 0.80 2.70 0.50 0.00 VILL-109-GA 0.65 2.00 1.00 48.00 5.80 0.00 0.00 2.05 0.95 2.20 1.00 0.00 VILL-110-MO 0.45 2.00 0.00 27.00 8.00 0.00 0.00 2.25 1.00 2.90 0.50 0.00 VILL-111-MO 0.90 2.00 1.00 30.00 6.90 0.00 0.00 2.30 1.10 3.20 1.00 0.00 VILL-112-VA 0.50 2.00 0.00 37.00 7.00 0.00 0.00 2.20 0.90 1.90 0.50 0.00 VILL-113-AL 0.55 2.00 0.00 18.00 5.00 0.00 0.00 2.15 0.90 2.00 0.50 0.00 VILL-114-AR 0.75 2.00 0.00 33.00 8.85 0.00 0.00 2.25 1.15 2.30 0.50 0.00 VILL-115-IL 0.40 2.00 0.00 17.00 7.00 0.00 0.00 2.10 1.00 2.10 0.50 0.00 VILL-116-IA 0.70 2.00 0.00 28.00 6.10 0.00 0.00 1.97 0.90 3.10 1.00 0.00 VILL-17-GA 0.55 2.00 0.00 11.00 7.40 0.00 0.00 2.40 1.00 2.60 1.00 0.00 VILL-18-NY 0.62 2.00 0.00 9.00 6.60 0.00 0.00 2.10 0.95 1.70 1.00 0.00
114
OTU Code LigLen DouLigCon DouVes NumTriAdaLeaSur LonLeaWid MidLeaGlaAda TerLeaGlaAda SpiLen PriGluLen MidSheIndLen ShePubOri Cilia>30 VILL-19-NJ 0.45 2.00 0.00 43.00 5.90 0.00 0.00 2.30 1.10 3.75 1.00 0.00 VILL-20-MS 0.70 2.00 0.00 18.00 5.80 0.00 0.00 2.20 0.87 1.45 1.00 0.00 VILL-22-TN 0.40 2.00 0.00 21.00 6.60 0.00 0.00 2.00 0.80 1.90 1.00 0.00 VILL-23-VA 0.63 2.00 0.00 17.00 9.40 0.00 0.00 2.25 0.75 2.60 1.00 0.00 VILL-27-MO 0.99 2.00 0.00 36.00 7.30 0.00 0.00 2.15 0.70 3.00 1.00 0.00 VILL-314-GT 1.10 2.00 0.00 35.00 8.60 0.00 0.00 2.05 1.00 2.70 1.00 0.00 VILL-35-TX 0.40 2.00 0.00 37.00 4.20 0.00 0.00 2.19 0.70 3.10 1.00 0.00 VILL-41-MO 0.72 2.00 0.00 21.00 7.70 0.00 0.00 2.05 0.70 3.00 1.00 0.00 VILL-49-MX 0.30 2.00 0.00 23.00 8.20 0.00 0.00 2.20 0.93 2.40 1.00 0.00 VILL-90-DC 0.55 2.00 0.00 22.00 7.10 0.00 0.00 2.35 1.00 2.40 1.00 0.00 VILL-91-MX 0.55 2.00 0.00 34.00 5.50 0.00 0.00 2.18 0.83 2.30 1.00 0.00 VILL-92-VA 0.45 2.00 0.00 14.00 6.30 0.00 0.00 2.08 1.28 2.50 0.50 0.00 VILL-93-GA 0.50 2.00 0.00 14.00 5.50 0.00 0.00 2.25 0.95 2.10 0.25 0.00 VILL-94-AL 0.70 2.00 1.00 29.00 5.90 0.00 0.00 2.30 1.12 3.70 0.75 0.00 VILL-95-MX 0.75 2.00 0.00 22.00 6.70 0.00 0.00 2.34 1.10 2.30 1.00 0.00 VILL-96-TN 0.45 2.00 0.50 24.00 4.70 0.00 0.00 2.20 0.70 2.30 0.50 0.00 VILL-97-ND 0.85 2.00 0.00 21.00 5.60 0.00 0.00 1.85 0.80 2.30 0.75 0.00 VILL-98-SC 0.75 2.00 0.00 22.00 8.80 0.00 0.00 2.32 0.85 3.45 0.75 0.00 VILL-99-IA 0.90 2.00 0.00 16.00 5.90 0.00 0.00 2.10 0.85 2.40 0.50 0.00 T-ATLA-244-NY 1.30 2.00 0.00 10.00 5.90 0.00 0.00 2.02 0.99 3.30 1.00 0.00 T-HAEM-270-DC 0.45 2.00 0.00 10.00 5.40 0.00 0.00 2.30 0.82 2.80 1.00 0.00 T-VILL-242-GA 0.49 2.00 0.00 20.00 5.90 0.00 0.00 2.33 0.89 2.70 1.00 0.00 T-XANT-268-AL 0.45 2.00 0.00 15.00 5.20 0.00 0.00 2.20 0.95 1.80 0.75 0.00 T-PSEU-241-AL 0.50 2.00 0.00 17.00 8.50 0.00 0.00 2.40 0.82 2.00 0.50 0.00
115 Appendix E. Specimens sampled for analysis (asterisk in the “Herbaria” column represents a type specimen).
Country State/Province County Collector Coll. No. Date OTU code Herbaria Belize - - Dwyer, J. & C. Berry 15016 July 25, 1980 ACUM-48-BE MO Canada - - Rosendahl, C. 1931 June 26, 1907 LANU-218-VAN MO Canada Alberta - Hitchcock A. 220 July 31, 1914 T-FEPA-251-ALB *GH Canada British Columbia Nelson Thompson, J. 14433 July 20, 1939 LANU-219-BC MO Canada New Brunswick Carleton Fernald, M. 12527 July 14, 1910 T-LISE-253-NB *GH Canada Nova Scotia Yarmouth Fernald, M., Long & Bartrum 23201 July 23, 1921 SPRE-208-NS MO Canada Ontario Pipineau Dore, W. 1051 July 19, 1947 TENN-151-QU MO Canada Ontario Prince Edward Gillett, J. 1366 July 10, 1947 IMPL-134-ON MO Canada Quebec Missiqubi Knowlton, C. s.n. August 10, 1923 LANU-133-QU MO Colombia Popayau - Lehmann 974 1889 OLIV-304-CL GH Costa Rica San Jose - Standley, P. & J. Valerio 43377 January, 1926 OLIV-305-CR GH Cuba - - Wright, C. 3463 1860-1864 T-WRIG-258-CB *MO Dom. Rep. Santa Domingo - Ekman, E. 11597 February 13, 1929 ACUM-302-DR GH Guatemala - - Seler, E. 3235 September 13, 1896 VILL-314-GT MO Guatemala - - Turckheim, H. 428 February, 1888 T-OLIV-247-GU *MO Guatemala - - Turckheim, H 428 February, 1888 T-OLIV-284-GT *US Haiti - - Holdridge, L. 1727 September 28, 1943 ACUM-47-HI MO Honduras - - Swallen, J. 11012 October 23, 1951 OLIV-225-HN MO Honduras Intibuca - Davidse, G. 35207 June 21, 1994 VILL-91-HN MO Jamaica - - Harris, W. 11597 January 24, 1913 ACUM-301-JA GH Mexico Chiapas - Breedlove, D. 54707 November 5, 1981 VILL-95-MX MO Mexico Chiapas - Davidse, G. 29845 November 15, 1915 OLIV-226-MX MO Mexico Oaxaca - Davidse, G. & J. Davidse 9688 August 14, 1975 ACUM-49-MX MO Mexico Tabasco - Davidse, G. 9368 August 8, 1975 OLIV-227-MX MO Mexico Vera Cruz - Hitchcock, A. 134 September 2, 1910 OLIV-224-MX MO Mexico Vera Cruz - Purpus, C. 7879 October, 1927 OLIV-228-MX MO Nicaragua - - Seymour, F. 4576 March 12, 1971 LONG-193-NIGU MO Nicaragua Nueva Segovia - Stevens, W. & B. Krukoff 3271 August, 1977 ACUM-46-NI MO Panama Chiriqui - Davidson, M. 766 June 25, 1978 OLIV-303-PN GH Puerto Rico - - Hackel 5908 January 17, 1886 ACUM-300-PR GH USA Alabama Butler Mohr, C. s.n. May 8, 1898 T-XANT-268-AL *US USA Alabama Dekalb Kral, R. 53036 May 27, 1974 PSEU-61-AL MO
116 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Alabama Dekalb Kral, R. 53030B May 27, 1974 VILL-94-AL MO USA Alabama Henry Kral, R. 48982 October, 1972 OVAL-67-AL MO USA Alabama Lee Earle, F. & C. Baker 1537 May 7, 1898 T-PSEU-241-AL *NY USA Alabama Lee Earle, F. & C. Baker 1527 May 7, 1898 T-AUBU-259-AL *NY USA Alabama Monroe Kral, R. 46998 May 31, 1972 PSEU-62-AL MO USA Alabama Pickens McDaniel, S. 1839 May 21, 1960 VILL-113-AL MO USA Arizona - Lemmon 2907 1882 T-HUAC-272-AZ *US USA Arkansas Benton Thomas, J. 1502 June 18, 2002 LANU-42-AR MO USA Arkansas Benton Thomas, J. 1501 June 18, 2002 PRAE-43-AR MO USA Arkansas Benton Plank s.n. no date PRAE-122-IL MO USA Arkansas Fulton Bush, B. 5637 May 19, 1909 IMPL-130-AR MO USA Arkansas Garland Demaree, D. 15843 August 18, 1937 VILL-114-AR MO USA Arkansas St. Francis Mcdaniel, S. 568 June 19, 1956 LIND-162-AR MO USA California Humbolt Hitchcock, A. 527 July 13, 1915 LIND-216-CA MO USA California Plummas Jepson, W. 4082 June 6, 1910 T-LASS-276-CA *US USA California Shasta Hitchcock, A. 136 August 3, 1908 LANU-212-CA MO USA California Shasta Hitchcock, A. s.n. August 3, 1908 T-PACI-262-CA *US USA California Shasta Greata, L. s.n. June 1899 T-LANU-282-CA MO USA California Sonoma Redfield, J. 9792 August 27, 1872 THER-230-CA MO USA California Sonoma Jones, M. 29210 July 5, 1931 THER-231-CA MO USA California Sonoma Bolander, H. 3991 1865 T-THER-232-CA *MO USA California Sonoma Bolander, H. 3941 1866 T-THER-254-CA *GH USA California Tulare Coville & Funston s.n. 1891 T-FUNS-271-CA *US USA Delaware Eussex Fogg, J. s.n. June 16, 1926 COLU-80-DE MO USA Delaware New Castle Commons, A. 283 June 25, 1823 T-SCOP-283-DE *US USA Delaware Sussex Hitchcock, A. 143 June 18, 1905 COMM-70-DE MO USA Delaware Sussex McVaugh, R. 6512 June 24, 1942 COMM-73-DE MO USA District of Columbia - Holmgren, T. s.n. June 12, 1894 VILL-90-DC MO USA District of Columbia - Kearney, H. s.n. June 6, 1897 T-HAEM-270-DC *US USA District of Columbia - Chase, A. 2400 June 28, 1904 T-HUSI-273-DC *US USA Florida - - - - OVAL-64-FL MO USA Florida Duval Curtiss, A. 5866 April 24, 1897 PSEU-57-FL MO USA Florida Fauquier Tracy, S. 8410 May 26, 1903 LEUC-194-FL MO USA Florida Franklin Chapman s.n. April, 1909 OVAL-29-FL MO
117 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Florida Franklin Godfrey, R. 69494 May 10, 1969 OVAL-85-FL MO USA Florida Franklin Chapman 7189 no date ACUM-139-FL MO USA Florida Franklin Vasey, G. s.n. 1892 T-LONG-246-FL *NY USA Florida Hillsborough O’neill, H. 2513 April 2, 1927 LEUC-198-FL MO USA Florida Lake Curtiss, A. 6616 May 13, 1900 OVAL-33-FL MO USA Florida Lake Nash, G. 1338 July 16-31, 1894 T-LEUC-243-FL *NY USA Florida Lake Nash, G. s.n. March 12-31, 1894 T-CILI-256-FL *NY USA Florida Lee Hitchcock, A. 474 July-August, 1900 OVAL-65-FL MO USA Florida Liberty Kral, R. 61184 October 19, 1977 OVAL-88-FL MO USA Florida Suwannee Curtiss, A. 6652 June 7, 1900 WRIG-185-FL MO USA Georgia - Georgia s.n. no date OVAL-32-GA MO USA Georgia Bibb Small, J. s.n. May 18-24, 1895 T-VILL-242-GA *NY USA Georgia Bulloch Harper, R. 839 June 10, 1901 LONG-192-GA MO USA Georgia Burke Freckmann, R. 1180 June 4, 1964 ACUM-140-GA MO USA Georgia Charlton Harper R. 1574 August 26, 1902 LONG-191-FL MO USA Georgia Dade Cronquist, A. 5164 May 8, 1948 VILL-109-GA MO USA Georgia Lee Duncan, W. 2317 April 11, 1940 PSEU-54-GA MO USA Georgia Lowndes Faircloth, W. 5282 July 10, 1968 WRIG-186-GA MO USA Georgia McIntosh Small, J. s.n. June 25-27, 1895 T-PARV-240-GA *NY USA Georgia Murray Kral, R. 50687 July 18, 1973 VILL-17-GA MO USA Georgia Screven Cronquist, A. 5006 April 17, 1948 VILL-93-GA MO USA Illinois Champaign Gleason, H. s.n. June 20, 1906 PRAE-121-OK MO USA Illinois Cook Moffatt, W. s.n. June 21, 1891 VILL-06-IL MO USA Illinois Henderson Patterson, H. s.n. no date VILL-115-IL MO USA Illinois LaSalle Chase, A. 1602 July 13, 1901 T-IMSI-275-IL *US USA Illinois Macon Clokey, I. 2440 June 12, 1915 LANU-143-IL MO USA Illinois Morgan Hill, S. 31708 June 16, 1999 LANU-12-IL MO USA Illinois Stark Chase, V. 671 June 10, 1906 PRAE-21-IL MO USA Illinois Stark Chase, V. 649 June 30, 1900 T-PRAE-263-IL *US USA Illinois Tazewell Chase, V. 3146 June 23, 1919 LANU-144-IL MO USA Indiana Jasper Friesner, R. 21490 June 15, 1947 COMM-W-59-IN MO USA Indiana Jasper Friesner, R. 21662 July 12, 1947 SPRE-210-IN MO USA Indiana Morgan Friesner, R. 7497 June 11, 1934 LANU-10-IN MO USA Indiana Porter Tyron, R 3718 ½ June 27, 1935 SPRE-209-IN MO
118 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Iowa - Jones 34 no date VILL-116-IA MO USA Iowa Johnson Shimek, B. s.n. no date IMPL-175-IA MO USA Iowa Story Freckmann, R. 1927 June 18, 1965 PRAE-125-NE MO USA Iowa Wapello Hayden, A. 9259 June 23, 1939 LANU-145-IA MO USA Kansas Cloud Fraser, S. 796 June 15, 1937 PRAE-07-KS MO USA Kansas Douglas McGregor, R. 40533 June 15, 1992 LANU-131-KS MO USA Kentucky Ballard Anderson, W. s.n. June 6, 1923 LANU-135-KY MO USA Kentucky Caldwell Conrad, J. 1008 May 27, 1970 LIND-160-KY MO USA Louisiana Acadia Chambliss, C. s.n. April 17, 1913 THUR-202-LA MO USA Louisiana Jeff Davis Palmer, E. 7610 May 15, 1915 THUR-203-LA MO USA Louisiana Lake Charles Hitchcock, A. 1147 April 21, 1906 LANU-132-LA MO USA Maine Cumberland Scribner, F. s.n. July T-IMPL-274-MA *US USA Maine Lincoln Grover, F. s.n. July 13, 1922 IMPL-16-ME MO USA Maine Oxford Hunnewell, F. 11973 June 14, 1931 SUBV-311-ME MO USA Maine York Fernald, M. s.n. September 26, 1897 T-UNPR-266-MA *US USA Maryland Balto Reed, C. 2165 July 21, 1940 LANU-141-MD MO USA Maryland Calvert Chase, A. 12270 July 8, 1934 COLU-101-MD MU USA Maryland Prince George Chase, A. 146 July 23, 1907 COMM-72-MD MO USA Massachusetts - Chapman, A. s.n. no date OVAL-83-MA MO USA Massachusetts Barnstable Fernald, M. 16118 July 23, 1918 COMM-68-MA MO USA Massachusetts Barnstable Fernald, M. 16119 August 4, 1918 COMM-69-MA MO USA Massachusetts Barnstable Churchill, J. s.n. July 3, 1931 MERI-181-MA MO USA Massachusetts Barnstable Knowlton, C. s.n. July 24, 1909 WRIG-184-MA MO USA Massachusetts Berkshire Churchill, J. s.n. July 16, 1917 IMPL-176-MA MO USA Massachusetts Bristol Seymour, F. 19272 June 28, 1961 LANU-142-MA MO USA Massachusetts Dukes Fogg, J. 3875 September 5, 1928 COMM-71-MA MO USA Massachusetts Middlesex Forbes, F. s.n. June 27, 1915 IMPL-170-MA MO USA Massachusetts Middlesex Blankinship, J. 109 June 29, 1896 SPRE-207-MA MO USA Massachusetts Nantucket Bicknell, P. s.n. September 20, 1907 T-OWEN-234-MA *NY USA Massachusetts Plymoth St. John, H. 11755 September 15, 1916 COLU-100-MA MU USA Michigan Emmet Gates, F. 13354 July 28, 1922 LIND-09-MI MU USA Michigan Keweenaw Bennett, H. s.n. July 14, 1956 SUBV-312-MI MO USA Michigan Washtnaw Hermann, F. 6893 July 5, 1935 LIND-163-MI MO USA Minnesota Anoka Stolz, A. s.n. July 7, 1926 IMPL-02-MN MO
119 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Minnesota Anoka Stoltz, A. s.n. August 10, 1926 IMPL-04-MN MO USA Minnesota Anoka Schuette, J. s.n. June 30, 1888 SUBV-221-MN MO USA Minnesota Carlton Ashe, W. s.n. August 6, 1899 T-SUBV-287-MN *NCU USA Minnesota Chippewa Gereau, R. 1014 July 9, 1982 IMPL-178-MI MO USA Minnesota Itasca Rosendahl, C. & J. Maple 707 June 30, 1935 IMPL-177-MN MO USA Minnesota Renville Moore, G. 13054 June 19, 1940 PRAE-25-MN MO USA Minnesota Sherburne Swanson, G. 1382 July 6, 1999 PRAE-315-MN MO USA Mississippi - Higland, E. s.n. 1858 VILL-20-MS MO USA Mississippi Harrison Tracy, S. 4580 September 1, 1898 T-CILI-280-MS *US USA Mississippi Jackson Tracy, S. 4885 April 6, 1898 WRIG-187-MS MO USA Missouri Barry Bush, B. 141 June 6, 1897 VILL-111-MO MO USA Missouri Barton Yatskievych, G & K. 90-147 June 16, 1990 PRAE-119-MO MO USA Missouri Benton Steyermark, J. 10744 May 24, 1936 VILL-27-MO MO USA Missouri Callaway Steyermark, J. 26220 September 12, 1937 PRAE-120-MO MO USA Missouri Carter Thomas, J. 1399 June 7, 2001 VILL-41-MO Private USA Missouri Carter Thomas, J. 1416 May 16, 2001 LIND-165-MO Private USA Missouri Clark Bush, B. 5909A July 8, 1909 PRAE-117-MO MO USA Missouri Dade Thomas, J. 1307 August 9, 2000 PRAE-40-MO MO USA Missouri Dent Kellogg, J. 15078 June 30, 1929 PRAE-39-MO MO USA Missouri Howell Thomas, J. 678 June 5, 1999 LIND-166-MO Private USA Missouri Jasper Palmer, E. 2157B June 5, 1909 PRAE-38-MO MO USA Missouri Jasper Palmer, E. 2144 June 5, 1909 PRAE-118-MO MO USA Missouri Jasper Palmer, E.J. 3753 June 9, 1912 LANU-149-MO MO USA Missouri Jasper Palmer, E.J. 3714 June 2, 1912 TENN-150-MO MO USA Missouri Ozark Steyermark, J. 22589 July 1, 1937 VILL-110-MO MO USA Missouri Scott Mckenzie, P. 1294 September 7, 1993 COMM-W-53-MO MO USA Nebraska Cuming Churchill, S. 5528 May 31, 1975 PRAE-127-NE MO USA Nebraska Kearney Hapeman, H. s.n. July 8, 1918 TENN-157-NE MO USA Nebraska Lancaster Hedgcock, G. s.n. June 15, 1900 PRAE-45-NE MO USA Nebraska Lancaster Churchill, S. 941 June 4, 1973 PRAE-123-AR MO USA Nebraska Lancaster Hedgcock, G. s.n. June 15, 1900 PRAE-124-NE MO USA Nebraska Nemaha Churchill, S. 200 June 22, 1972 LANU-138-NE MO USA Nebraska Saunders Churchill, S. 3588 June 25, 1974 PRAE-44-NE MO USA Nebraska Stanton Churchill, S. 7709 June 15, 1976 PRAE-26-NE MO
120 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Nebraska Winneskiek Tolstead, W. s.n. June 12, 1933 PRAE-126-IA MO USA New Hampshire Rockingham Eaton, P. s.n. 1897 T-EATO-237-NH *NY USA New Jersey Burlington Herman, F. 4465 June 18, 1933 OVAL-37-NJ MO USA New Jersey Cape May Bicknell, E. s.n. May 30-31 1897 T-ADDI-238-NJ *NY USA New Jersey Cape May Bicknell, E. s.n. May 30-31, 1897 T-PAUC-239-NJ *NY USA New Jersey Cape May Commons, A. 341 June 29, 1898 T-COMM-279-NJ *US USA New Jersey Hunterdon Fisher, H. s.n. July 12, 1915 LANU-137-NJ MO USA New Jersey Hunterdon Benner, W. 9635 June 7, 1941 T-BENN-249-NJ *GH USA New Jersey Mercer E.D. s.n. August 18, 1866 COMM-66-NJ MO USA New Jersey Middlesex MacKenzie, K. 1485 July 9, 1905 COMM-76-NJ MO USA New Jersey Monmouth Churchill, J. s.n. June 26, 1929 OVAL-34-NJ MO USA New Jersey Ocean MacKenzie, K. 2779 July 27, 1907 COMM-79-NJ MO USA New Jersey Passaic MacKenzie, K. 2724 July 14, 1907 VILL-19-NJ MO USA New York Albany Van Schaak, G. 3230 June 23, 1951 LIND-161-NY MO USA New York Bronx Nash, G. s.n. June 24, 1897 T-ATLA-244-NY *NY USA New York Burlington Parker s.n. June 16, 1900 SPRE-206-NJ MO USA New York Cayuga Metcalf, F. 5539 July 25, 1916 LIND-168-NY MO USA New York Clinton Eggleston, W. 2843 July 3, 1902 COLU-102-NY MU USA New York Delaware Peck, C. s.n. August 20, 1906 T-DEMI-278-NY *US USA New York Franklin Muenscher, W. & B. Maguire 933 June 22, 1930 SUBV-222-NY MO USA New York Oneida Crockett, R. 7577 July 3, 1945 COLU-77-NY MO USA New York Seneca Randolf, L. 11249 June 24, 1919 VILL-18-NY MO USA New York Suffolk Van Schaak, G. 3551 August 4, 1956 COMM-74-NJ MO USA New York Suffolk Van Schaak, G. 3550 August 4, 1956 COMM-75-NY MO USA New York Suffolk Latham, R. 4169 July 9, 1927 COMM-78-NY MO USA New York Suffolk Chase, A. 526 June 22, 1916 MERI-179-NY MO USA New York Tompkin Wiegand, K. & A. Betchtel 7504 July 18, 1917 IMPL-171-NY MO USA New York Tompkins James, A. 88 July 12, 1913 LANU-152-NY MO USA North Carolina - Bosc, L. s.n. no date T-STRI-264-NC *US USA North Carolina Beaufort Ashe, W. s.n. no date T-ALBE-281-NC *US USA North Carolina Burke Ashe, W. s.n. no date T-MERI-260-NC *US USA North Carolina Cumberland Solomon, J. s.n. May 16, 1976 OVAL-31-NC MO USA North Carolina Henderson Davis, J. s.n. July, 1924 LONG-190-NC MO USA North Carolina New Hanover Ashe, W. s.n. May 17, 1899 T-WILM-267-NC *US
121 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA North Carolina Orange Ashe, W. s.n. no date MERI-183-NC MO USA North Carolina Orange Ashe, W. s.n. May 17, 1899 T-ERYT-277-NC *MO USA North Carolina Orange Ashe, W. s.n. no date T-MICR-285-NC *NCU USA North Carolina Orange Ashe, W. s.n. June 29, 1898 T-ORAN-286-NC *US USA North Carolina Sampson Freckmann, R. 2114 May 17, 1966 OVAL-63-NC MO USA North Dakota Richland Stevens, O. 451 June 27, 1940 VILL-97-ND MO USA Ohio Hocking Cusick, A. 21692 June 15, 1982 COLU-104-OH MU USA Ohio Huron Leonard, F. 14 June 20, 1887 LANU-11-OH MO USA Ohio Lucas McCormac, J. 5971 July 1, 1997 IMPL-14-OH MO USA Ohio Lucas Easterly, N. 12688 June 19, 1981 COLU-103-OH MU USA Ohio Lucas Kriebel, R. 8731 June 17, 1940 SUBV-200-OH MO USA Ohio Scioto Demaree, D. 10756 June 27, 1934 LANU-01-OH MO USA Oklahoma Creek Harvey, L. 808 June 11, 1939 TENN-153-OK MO USA Oklahoma LeFlore Demaree, D. 12712 May 17, 1936 LANU-148-OK MO USA Oklahoma Murray Waterfall, U. 6021 no date PRAE-24-OK MO USA Oregon - Hall, E. s.n. 1871 IMPL-215-OR MO USA Oregon Curry Peck, M. 8795 July 11, 1919 IMPL-220-OR MO USA Oregon Lane Peck, M. 15950 June 14, 1928 LANU-211-OR MO USA Pennsylvania Montgomery MacElwee, A. 544 June 16, 1899 LANU-155-PA MO USA Pennsylvania Philadelphia Bigyn, C. s.n. no date LANU-136-PA MO USA Puerto Rico - Heller, A. 12 January 9, 1899 T-COMO-235-PR *NY USA Rhode Island Providence Olney s.n. no date IMPL-173-RI MO USA South Carolina Aiken Ravenel s.n. October, 1867 OVAL-30-SC MO USA South Carolina Aiken Eggert, H. s.n. May 21, 1899 OVAL-55-SC MO USA South Carolina Aiken Ravenel, H. s.n. June 1, 1867 PSEU-56-SC MO USA South Carolina Aikens Ravenel, H. s.n. June 11, 1867 OVAL-82-SC MO USA South Carolina Aikens Ravenel, H. s.n. June 1, 1867 OVAL-86-SC MO USA South Carolina Aikens Eggert, H. s.n. May 20, 1899 OVAL-87-SC MO USA South Carolina Anderson Davis, J. s.n. May, 1924 TENN-154-SC MO USA South Carolina Bamberg Freckmann, R. 2133 May 17, 1966 VILL-99-IA MO USA South Carolina Beaufort Churchill, J. s.n. May 6, 1917 OVAL-36-SC MO USA South Carolina Beaufort Chase, A. 7115 May 27, 1916 OVAL-84-SC MO USA South Carolina Orangeburg Hitchcock, A. 118 August 19, 1905 LEUC-197-SC MO USA South Carolina Williamsburg Godfrey, R. 428 July 10, 1939 VILL-98-SC MO
122 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Tennessee Cannon Kral, R. 52800 May 20, 1977 VILL-96-TN MO USA Tennessee Cannon Kral, R. 52796 May 20, 1974 MERI-182-TN MO USA Tennessee Coffee Svenson, H. 9230 June 20, 1938 IMPL-169-TN MO USA Tennessee Coffee Svenson, H. 9225 June 20, 1938 LEUC-199-TN MO USA Tennessee Grundy Svenson, H. 9119 June 23, 1938 LONG-189-TN MO USA Tennessee Hardinan Palmer, E. 19493 May 14, 1920 VILL-106-TN MO USA Tennessee Knox Ruth, A. s.n. August, 1894 VILL-22-TN MO USA Tennessee Knox Ruth, A. s.n. June, 1896 VILL-107-TN MO USA Tennessee Wilson Gattinger, A. 7387 June 1881 PSEU-60-TN MO USA Texas - - - - THUR-229-TX MO USA Texas - Lindheimer 5651846 no date T-LIND-245-TX *NY USA Texas Gregg Reverchon, J. s.n. June 18, 1907 VILL-35-TX MO USA Texas Gregg Reverchon, J. 2357 June 18, 1901 LIND-164-TX MO USA Texas Harrison Bush, B. 632 August 8, 1907 PRAE-128-NE MO USA Texas Montgomery Morden, C. 29 May 11, 1980 THUR-205-TX MO USA Texas Polk Gould, F. & S. Hatch 14223 May 10, 1973 VILL-105-TX MO USA Texas Polk Gould, F., Hatch & Waller 14233A May 10, 1973 ACUM-147-TX MO USA Texas Robertson Launchbaugh, J. 397 April 30, 1951 COMM-108-TX MO USA Texas Robertson Launchbaugh, J. 394 April 30, 1951 LEUC-172-TX MO USA Texas Upshur Reverchorn, J. 2355 May 28, 1901 THUR-201-TX MO USA Texas Waller Thurow, F. 6 June 1898 T-OCTO-261-TX *US USA Texas Waller Thurow, F. 9 June 5, 1898 T-THUR-265-TX *US USA Texas Wheeler Correll, D. & I. Johnston 17045 June 25, 1957 TENN-156-TX MO USA Utah San Juan Maguire, B. 5910 July 2, 1933 TENN-158-UT MO USA Vermont Caledonia Seyour, F. 18632 no date IMPL-51-VT MO USA Virginia Alexandria Chase, A. 555 June 9, 1905 VILL-112-VA MO USA Virginia Augusta Allard, H. 3170 July 5, 1937 LANU-146-VA MO USA Virginia Greeensville Fernald, M. & B. Long 12940 June 18, 1941 MERI-180-VA MO USA Virginia Nansemond Fernald, M. & B. Long 10505 July 26-28, 1939 OVAL-81-VA MO USA Virginia Nansemond Fernald, Long & Clement 15186 Septermber, 1946 T-GLUT-252-VA *GH USA Virginia Norfolk Fernald, M. & B. Long 11969 June 11, 1940 VILL-23-VA MO USA Virginia Northumberland Reed, C. 5136 June 1, 1946 PSEU-58-VA MO USA Virginia Prince George Freckmann, R. 1565 May 30, 1965 MERI-174-VA MO USA Virginia Princes Ann MacKenzie, K. 1710 September 3, 1905 VILL-92-VA MO
123 Country State/Province County Collector Coll. No. Date OTU code Herbaria USA Virginia Sussex Fernald, F. & B. Long 14267 July 6, 1942 WRIG-188-VA MO USA Washington - Presl 3978 no date T-OCCI-257-WA *MO USA Washington Chelan Thompson, W. 6928 June 25, 1931 IMPL-214-WA MO USA Washington Kittilas Sandberg & Leiberg 425 July, 1898 LANU-213-WA MO USA Washington Okanogan Elmer, A. 489 July, 1897 LANU-217-WA MO USA Washington Whitman Brodie, D. s.n. June 1898 T-BROD-248-WA *WS USA Wisconsin Adams Shinners, L. & C. Shaw 4415 June 28, 1942 T-EUCL-255-WI *WIS USA Wisconsin Ashland Coffin, B. 41 July 1, 1976 SUBV-313-WI MO USA Wisconsin Bayfield Fassett, N. 19571 July 10, 1938 IMPL-50-WI MO USA Wisconsin Columbia Shinners, L. & L. Shaw 4361 June 14, 1942 PRAE-167-WI MO USA Wisconsin LaCrosse Pammel, L. 1334 June 18, 1925 COMM-89-WI MO USA Wisconsin LaCrosse Ziegler, S. 1656 July 8, 1975 TENN-159-WI MO USA Wisconsin Oneida Coffmann, R. s.n. August 29, 1918 SUBV-233-WI MO USA Wyoming Teton Chase, A. 137 August 10, 1908 T-FERV-250-WY *GH USA Wyoming Teton Nelson, A & E. Nelson 6037 July 21, 1899 T-FESE-269-WY *GH
124