Breeding Science 59: 561–570 (2009)

Morphological identification of genomic genera in the

Mary E. Barkworth*1), D. Richard Cutler2), Jeffrey S. Rollo1), Surrey W.L. Jacobs3) and Abdur Rashid4)

1) Intermountain Herbarium, Department of Biology, Utah State University, Logan, Utah 84322-5305, USA 2) Department of Mathematics and Statistics, Utah State University, Logan, Utah 84322-3900, USA 3) National Herbarium of , Mrs Macquarries Road, Sydney, N.S.W. 2000, 4) University of Peshawar Botanic Garden, University of Peshawar, Peshawar, Northwest Frontier Province, Pakistan

Our goal was to determine whether the genomic groups of perennial species Triticeae having solitary spikelets could be identified morphologically and, if so, to construct identification keys that could be used for this purpose. If so, it would strengthen the argument for recognizing such groups as genera. We conducted Dis- criminant and Random Forest® analyses of 61 characters scored on 218 herbarium specimens representing 13 genomic groups. In addition, we closely examined some additional characters that came to our attention, evaluating our findings on specimens not scored for the two kinds of analysis. Random Forest® analysis was almost always more successful in distinguishing the genomic groups, whether separating all 13 groups or a subset of the 13. The results suggest that it is usually possible to identify the genomic group to which a spec- imen of perennial Triticeae with solitary spikelets belongs on the basis of its morphology but that doing so will require examination of characters that have not been considered particularly important in the past. Among these are the length of the middle inflorescence internodes, the width of the palea tip, and the morphology of the glumes. Generic descriptions and keys have been posted to the web (see http://herbarium.usu.edu/ triticeae). They include all the genera that we recognize in the tribe, not just those included in the analyses, and will be improved as additional information becomes available.

Key Words: Genomic group, solitary spikelets, inflorescence, palea tip, glume.

Introduction Triticeae, species with different genomic constitutions being placed in different genera. They argued that these genomic Botanical names are an important and conspicuous mecha- genera would better reflect phylogenetic relationships with- nism for communicating information. The amount of infor- in the tribe; Dewey added that they would be a useful guide mation they convey is limited by their brevity, existing for breeders. The response from taxonomists was knowledge, and the taxonomic philosophy of the person pro- largely negative (see, for example, Baum et al. 1987, moting a particular treatment. The International Code of Kellogg 1989, Seberg and Petersen 1998). In this study, we Botanical Nomenclature (McNeill et al. 2007, referred to after focused on one of the objections expressed: that the genomic this as the “Code”) states only that the name of a species is a genera could not be distinguished morphologically, a factor binomial, with the first word being the name of a . that makes their adoption impractical, if not impossible, for Most of us also learned that species in the same genera are, collectors and other field-oriented scientists. Our goal was to in some sense, more closely related to each other than to identify morphological criteria for distinguishing different members of another genus, a statement that reflects a philo- genomic groups within the perennial Triticeae with solitary sophical position, not a requirement of the Code. It is this spikelets. apparently simple philosophical statement that is behind There have been two previous studies of the relationship be- most nomenclatural disagreements, including those in the tween morphology and genomic constitution in the Triticeae. Triticeae. The problem is deciding how best to determine Salomon and Lu (1992) studied seven taxa representing which taxa are “more closely related”. Another area of dis- two genomic groups, StY and StH tetraploids (genome sym- agreement is the relative importance that should be attached bols from the International Triticeae Consortium; http:// to having a system that reflects relationships and one that is herbarium.usu.edu/Triticeae/genmsymb.htm) and found a easy to use. correlation between genome constitution and both palea apex In 1984, Löve and Dewey independently proposed using shape and the shape of the teeth on the palea keels. They cau- genomic constitution to determine generic boundaries in the tioned, however, that more species needed to be examined before it could be concluded that these features were a reli- Communicated by H. Tsujimoto able means of distinguishing between the two groups. Baum Received August 20, 2009. Accepted October 30, 2009. et al. (1995) conducted discriminant analysis of nine charac- *Corresponding author (e-mail: [email protected]) ters scored on 290 specimens representing 100 species in 562 Barkworth, Cutler, Rollo, Jacobs and Rashid four genomic groups, P, StH, StYP, and StY. The resulting Materials and Methods functions correctly identified the genomic group of the spec- imens 73% of the time in their jackknife analyses. We examined herbarium specimens of all the Australian Our study began as an investigation of the Australasian species of perennial Triticeae and representatives of the Triticeae, superficial observation of which suggested that Asian and American genomic groups that include perennial they could be distinguished morphologically from other species with one spikelet per node. The species we included members of the tribe. All the Australasian species are peren- are those with known genomic constitution (see http:// nials with one spikelet per node and all, so far as is known, herbarium.usu.edu/Triticeae/genomes.htm). We excluded have the W genome. In the polyploids, it is combined with Secale, Dasypyrum, and Leymus because those of their one or more of the St, Y and H other genomes. Löve (1984), perennial species that have solitary spikelets are rarely who restricted to the crested wheat grasses, placed in the wrong genus. The specimens used came from placed the diploid Australian species in and CANB, CHR, MO, NSW, and UTC (herbarium codes from the polyploid species in because the diversity of Index herbariorum; http://sweetgum.nybg.org/ih/). The New their genomic constitution was not known in 1984. Connor Zealand specimens were identified by Dr. Henry Connor, (1994) argued for recognition of , a morpholog- the remaining Australasian specimens by Jacobs. The non- ically distinct genus endemic to New Zealand. Its species Australasian specimens came from UTC and MO. Many of have since been shown to be HW tetraploids. The remaining the Asian specimens had been identified by Drs. Bjorn Australasian species are usually included, together with Salomon and Bao-Rong Lu. The remaining specimens were some Asian species, in Elymus sect. Anthosachne (Löve 1984, identified by Rollo and Barkworth. Several references were Connor 1994, Jacobs et al. 2009, but see Yen et al. 2006). used for this purpose, notably Bor (1970), Tutin (1980), Thus our interest in providing a better generic treatment for Cope (1982), Tsvelev (1983), Koyama (1987), Salomon the Australasian Triticeae required that we identify mor- (1994), Lu (1995), Assadi (1996), Edgar and Connor (2000), phological characteristics for distinguishing all the genomic Wu et al. (2006), Barkworth et al. 2007 and Jacobs et al. groups of Triticeae that include perennial species with soli- (2009). Voucher information is presented in the appendix. tary spikelets. Almost all these species were included in Seventy nine characters (Table 2) were chosen for the ini- Elymus sensu lato by Löve (1984), the exceptions being tial examination. They were selected based on previous the diploid genera Agropyron P genome), Australopyrum work, floristic treatments, and our own observations. Of the (W genome), and (L genome). For convenience, 79 characters: 52 were continuous, nine meristic or ordered, we refer to the groups by the appropriate generic name and 18 qualitative. Only the continuous and ordered charac- (Table 1) even though some of the species we examined do ters were used in the discriminant analyses. not have combinations based on such names. The Asian spe- Our intent was to examine five specimens per species, but cies that have been included in Anthosachne are StY tetra- in some instances we were unable to obtain that many. Rec- ploids, i.e., members of Roegneria. ognizing that our samples probably did not include the ex- tremes of variation for the species involved, we attempted to create an artificial maximum and minimum for each species,

Table 1. Genomic groups that include perennial species having one spikelet per node and the applicable generic name. In asterisked groups, most species have more than one spikelet per node. The last three groups, , Leymus and Pascopyrum, were not included in this study. Genomic composition Generic name Distribution P Agropyron Eurasia St Pseudoroegneria Eurasia and North America W Australopyrum Australia, New Zealand, and New Guinea StH* Elymus Eurasia, North America, South America StY Roegneria Eurasia HW Stenostachys New Zealand StYW Anthosachne Australasia StYP Kengyilia Asia (Tibetan Plateau) StYH Campeiostachys Eurasia StYHW Connorochloa New Zealand StP Douglasdeweya and western Asia E+ Thinopyrum Eurasia (a genomically mixed and poorly understood group, united by their possession of the E genome). L Festucopsis Albania Xp Peridictyon Southwestern Bulgaria and northern Greece (not included) StHNsXm Pascopyrum North America (not included) NsXm* Leymus Eurasia, North and South America (not included) Morphological identification of perennial genomic genera 563

Table 2. Characters and method of scoring. If a feature such as an awn was not present, it was scored “not applicable”. VEGETATIVE CHARACTERS: Growth habit (not rhizomatous, rhizomatous). CULM: height (cm); thickness at middle of first internode (mm); peduncle length (cm); nodes (number); trichome length middle of first internode (mm); trichome length top of first internode (mm); trichome length on first culm node (mm); trichome at top of peduncle. BASAL LEAVES: trichome length on sheath (mm); auricle length (mm); auricle frequency on (0, 1, 2, 3, 4); ligule length (mm); ligule shape (truncate, rounded, obtuse, acute, dentate); ligule edge (entire, irregular, denticulate); ligule ciliate (yes, no); blade length (cm); blade width (mm); blade vein number; trichome length lower surface (mm); trichome length upper surface (mm). TOP LEAF: ligule length (mm); blade length (cm); angle (declined, perpendicular, ascending, erect). REPRODUCTIVE CHARACTERS: INFLORESCENCE: length (cm); rachis extension length (mm); posture (drooping, nodding, inclined, erect); nodes (number); disarticulation (in rachis, below spikelet, below florets); lowest internode length (cm); middle internode length (cm); spikelets per node, lowest node (number); spikelets per node, mid-spike (number); pedicel length (mm); rachis cross-section shape (lunate, semi- circular, trapezoidal); trichome length rachis surface (mm); trichome length rachis edge (mm). SPIKELET: length (mm); florets (number); orien- tation to rachis (radial, tangential). LOWER GLUME: length (mm); shape (not applicable, rectangular, lanceolate, linear, horn-shaped); tip (not applicable, aristate, acute, obtuse); width (mm); hyaline margin width (mm); awn length (mm); veins (number); midvein trichome length (mm); surface trichome length (mm); surface trichome density (0, 1, 2, 3, 4). UPPER GLUME: length (mm); width (mm) ; shape (not applicable, rectan- gular, lanceolate, linear, horn-shaped); symmetry (symmetric, asymmetric); tip (not applicable, aristate, acute, obtuse); hyaline margin width (mm); awn length (mm); veins (number); keel (number); keel prominence (not prominent, prominent); midvein trichome length (mm); midvein scabrous (yes, no); surface trichome length (mm). LEMMA: length (mm); veins (number); awn length (mm); midvein trichome length (mm); back trichome length (mm); margin trichome length (mm); awn curvature (not applicable, straight, up to 90 degrees, more than 90 degrees). PALEA: apex width (mm); width at midlength (mm); keel cilia length (mm); palea tip shape (acute, rounded, truncate). Anther length (mm). using the lowest and highest values, respectively, from de- each sample, 63% of the records occur at least once. The scriptions in the references cited above, but they contained trees are used to predict the membership of the excluded too few of the characters in our list to permit use of such ar- records with the final disposition reflecting the majority tificial records in our analyses. rule, ties being split randomly. RF supersedes classification The species studied represented 13 of the 16 genomic trees, being a more accurate classifying procedure and one groups of perennial Triticeae (Table 1). Three groups are uni- that is minimally impacted by small perturbations in the data specific: Festucopsis, Peridictyon (Seberg et al. 1991), and (Cutler et al. 2007). Connorochloa (Zhang et al. 2009, Barkworth et al. 2009). Because of their different approaches, DA and RF also Two groups, Elymus and Thinopyrum, are known to include differ in their identification of the most important characters variants of the basic constitution for their group: Elymus, for distinguishing groups. In DA, only one of a set of highly which consists primarily of StH tetraploids also includes a correlated characters will be given high weight because the few hexaploids that are either StStH (e.g., E. repens) or other characters add only minimally to the discriminating StHH (e.g., E. patagonicus; Dewey 1972). Thinopyrum, as power of a function. RF ranks all the characters that are use- treated here, includes species with just the E genome or both ful for classifying the groups without regard for their corre- the E and the St genome. The latter species are sometimes lation. placed in Trichopyrum (Löve 1986, Yen et al. 2005). We used both methods to examine whether the records The ability of individual quantitative characters to distin- could be placed into the appropriate genomic groups and guish the groups was explored by examining box and whis- which characters were most useful for achieving this goal. In ker plots for each character. These have been posted to the addition, we looked at the ability of the methods to separate web at http://herbarium.usu.edu/triticeae/idgenomicgenera. the species containing a particular genome from the species We used both Discriminant Analysis (DA) and Random not containing that genome and at two subsets of the records, Forests® (RF) to determine whether some combination of those of the northern hemisphere St containing species and the characters could be used to distinguish the genomic those of the four groups examined by Baum et al. (1995). groups and, if so, to identify the relative importance of the We used SYSTAT 12 (Systat Software Inc. 2007) for devel- individual characters in making such distinctions. Only the oping the box and whisker plots and DA. Software for RF is quantitative and meristic characters were used in these anal- freely available at http://www.math.usu.edu/˜adele/forests. yses. DA uses a succession of linear combinations of the In practice, taxonomists rely on morphological keys for character values to classify the groups, with each successive plant identification. To create such keys, we first developed function weighting the characters in such a way as to maxi- descriptions of the genomic groups in this study, drawing on mize the additional separation of the groups. RF builds on knowledge gained while scoring the specimens, our previ- classification trees (CT). CTs are built by recursive binary ous experience, and descriptions in the references used for division of the records until the groups are separated or no identification. In addition, we entered information on the improvement in separation can be achieved. RF fits many distribution of individual species, as reported in various flo- classification trees to a data set and then combines the pre- ras. This information was then used to generate maps show- dictions from all the trees. It uses many (e.g., 500) bootstrap ing the species density of each genomic group by country, samples of the data to create multiple classification trees. In or, for the former Soviet Union, by the floristic regions used 564 Barkworth, Cutler, Rollo, Jacobs and Rashid by Tsvelev (1983); by state for the USA; and by province or matous. In Stenostachys and Connorochloa the rachis was territory for Canada. The resulting maps were stored as consistently lunate in cross-section whereas in Thinopyrum JPEG files. We used Fact Sheet Fusion (CBIT 2009), to it was consistently trapezoidal but both shapes are also create a web page for each genus. This program aids in cre- present in other genera. ation of linked pages that include thumbnail images linked Stenostachys and Festucopsis are both reported to have to larger versions of the image resources. The pages were spikelets that are radial, rather than tangential to the rachis posted to http://herbarium.usu.edu/triticeae/genera. (Tutin 1980, Connor 1994). In Stenostachys, the glumes are The descriptive information was used to construct both a tangential to the rachis but the florets are radial or almost so. dichotomous and multi-access key. Dichotomous keys dic- In Festucopsis, only some of the spikelets appeared to be ra- tate which characters are used for identification and the or- dial to the rachis, others appearing tangential. Examination der in which they are used. Multi-access keys enable users to of fresh material is needed to clarify the situation. select the characters they wish to use and the order in which Attempts to score the palea tips according to the terms they use them. In Lucid, the user may also click on the used in existing descriptions (e.g., acute, obtuse, emarginate, “wand” icon to determine which of the remaining characters truncate) were unsatisfactory. It seemed to depend on a com- would narrow down the possibilities most. If this character is bination of factors, including caryopsis maturity and width. not available or is one the user does not feel comfortable us- ing, the next best characters can be determined by succes- Quantitative characters sively selecting the “wand plus arrow” icon. In both the The box and whisker plots have been posted to the web dichotomous and multi-access key, the names of the taxa are at http://herbarium.usu.edu/triticeae/idgenomicgenera. They linked to the descriptive pages. show that a few genera were almost distinct from all others We used Phoenix (CBIT 2004) to create an interactive with respect to a single character. For example, Australopyrum version of the dichotomous key and Lucid (CBIT 2008) to differs from almost all other taxa in having long culm hairs develop the interactive multi-access key. Phoenix permits a and Stenostachys and Connorochloa have glumes that are user to skip one couplet in the key and includes links to the unusually narrow distally. In general, however, the range of descriptive pages for the genera but does not provide the values among the genera exhibits considerable overlap. flexibility of true multi-access keys such as Lucid. The keys incorporate images designed to aid in interpreting the char- Discriminant analyses acters and links to the descriptive pages. They are ongoing Of the 208 records for which there were complete data, projects. In addition, we have posted a PDF version of the 98% were assigned to the correct genus when all the data text material in the dichotomous key to the web. All three were used in the analysis. With Jackknife analysis, however, keys can be accessed via http://herbarium.usu.edu/triticeae/ the success rate was only 80%. Eliminating the poorly repre- keys. sented genera had almost no impact on these results. The first three discriminant functions accounted for 28.1%, Results 18.2%, and 16.2% of the total dispersion, respectively, for a total of 62.5%. The first and second functions separated Two hundred and eighteen specimens, representing 78 taxa Australopyrum and Stenostachys from the remaining genera. and 13 genomic groups, were formally scored for the numeri- Connorochloa was distinguished by the first and third func- cal analyses. Several additional specimens from the listed tions and Anthosachne was almost completely separated by herbaria were examined more informally in the course of de- the second and third functions. Thus the Australasian genera veloping the descriptions and keys. were distinguished from all the remaining genera on the first three functions. Qualitative characters When the four groups examined by Baum et al. (1995) were Most of the qualitative characters examined were either analyzed (Kengyilia, Roegneria, Elymus, and Agropyron), so constant among the groups or so similarly variable within our jackknifed classification success rate was only 69%, each group as to be of little value by themselves for identifi- 4% lower than the 73% that they obtained. As in their study, cation. For instance, all the specimens had truncate ligules. the greatest confusion was between Elymus (StH) and In most taxa, the ligule edges were irregular, although some Roegneria (StY). specimens of Australopyrum calcis had denticulate ligules and one species of Anthosachne had ciliate ligules. The in- Random Forest Analysis florescences almost always appeared to be erect but in some Because RF can use records with some missing data, the genera a minority of the specimens had drooping inflores- number of records in the complete analysis was 218, ten cences. This is, however, a character that it is difficult to more than for DA. Of these, 85% were placed in the correct score reliably from herbarium specimens. Three of the genome group (Table 3). All the specimens of Stenostachys, genera (Elymus, Pseudoroegneria, and Thinopyrum) include a Festucopsis, Anthosachne, and Australopyrum, were correct- few rhizomatous species or species that are sometimes rhizo- ly classified and 98% of the specimens of Elymus. The most matous but even in these genera most species are not rhizo- poorly classified were Douglasdeweya (80% misclassified), Morphological identification of perennial genomic genera 565

Table 3. Classification results from Random Forests Analysis. E+—Thinopyrum; HW—Stenostachys; L—Festucopsis; P—Agropyron; St— Pseudoroegneria; StH—Elymus; StHYW—Connnorochloa; StP—Douglasdeweya; StY—Roegneria; StYH—Campeiostachys; StYP—Kenyilia; StYW—Anthosachne; W—ustralopyrum. E+ HW L P St StH StHYW StP StY StYH StYP StYW W class error E+ 10 00000001001016.7% HW 0 19 00000000000 0.0% L002 0000000000 0.0% P0003 00000010025.0% St00004 0000002033.3% StH0000050 0001000 2.0% StHYW0000005 00001016.7% StP00001201 0001080.0% StY0000020020 104025.9% StYH0000060036 01062.5% StYP00000100202 1171.4% StYW0000000000043 00.0% W 00000000000020 0.0%

Kengyilia (71% misclassified), and Campeiostachys 63% mis- able species groups, but it has also suggested new ways of classified), all of which contain the St genome and were repre- looking at the taxa. In this study, we examined the hypothe- sented in this study by few specimens. Misclassified speci- sis that groups of genomically similar species are morpho- mens of Douglasdeweya were placed in Pseudoroegneria logically distinguishable. (1), Elymus (2), and Anthosachne (1); misclassified speci- Our results provide more support for this hypothesis than mens of Kengyilia were placed in Elymus (1), Roegneria suggested by current taxonomic treatments. Some groups, (2), Anthosachne (1), and Autralopyrum (1); and misclassi- such as the Australasian taxa, were relatively easy to distin- fied specimens of Campeiostachys in Elymus (6), Roegneria guish. Others, such as Kengyilia, Campeiostachys, and (3), and Anthosachne (1). Douglasdeweya, were not well sorted out when all the records Restricting the analysis to the 112 records from the six were included in the analysis. This is partly attributable to northern hemisphere St-containing groups (Pseudoroegneria, their low representation in the analyses. Both DA and RF Elymus, Douglasdeweya, Roegneria, Kengyilia, and tend to favor increasing the accuracy of identification for the Campeiostachys) resulted in 79.5% successful classifica- larger groups in an analysis in order to improve the overall tion, an increase of 8 correctly classified records over that success rate. Because there are only two known species of obtained from the complete analysis. For the four groups ex- Douglasdeweya and few species of Campeiostachys with amined by Baum et al. (1995), the success rate was 89.9%, a solitary spikelets, having equal, or nearly equal, species notable improvement over the results obtained with DA. samples for each group is impossible. Table 4 presents the results of the analyses when each of The success rate for Kengyilia would undoubtedly be im- the P-, H-, St-, Y- and W-containing species was compared proved if we included a larger sample of the species because with all the rest of the records. None of the P-containing speci- our selection included an almost equal number of typical and mens were correctly identified as such. The classification atypical species. We did not have access to enough reliably success rate for the other genomes ranged from 86.9–97.5% identified material to increase the number of species includ- when only five characters were used and from 85.96–99.38% ed. The species of Campeiostachys included had solitary when all characters were used. The important characters spikelets although the majority of species in the genus have varied with each analysis. Only the length of the middle inter- two or more spikelets per node. Inclusion of species with 2 node was one of the top ten characters in all five analyses; or more spikelets per nodes from both Elymus and palea tip width was in the top ten for three of the analyses. Campeiostachys might increase the confusion between these two genera, both of which include the St and H genomes. Discussion Douglasdeweya currently has only two species, both of which were described on the basis of grown in experimental Within the Triticeae, most species are perennial and have plots. Discovery of wild-grown material might enable it solitary spikelets. Ten different genomes have been identi- to be more effectively distinguished from other genomic fied among these species, eight of which are present in vary- genera. ing combinations in the numerous polyploid taxa. It is not Our inability to identify suites of characters that always surprising, therefore, that their taxonomic treatment is con- placed all the specimens in the correct genomic group could troversial. The genetic and cytogenetic information accumu- be used to argue for adopting a broad interpretation of lated during the last century furthered our understanding of Elymus. In our opinion, this would be counter-productive why it has been so hard to delimit morphologically recogniz- because it would conceal what is known about the genetic 566 Barkworth, Cutler, Rollo, Jacobs and Rashid

Table 4. Classification success of RandomForest Analysis for individual genomes with all, ten, or five characters respectively. The ten most im- portant characters for each analysis are listed in descending order of importance. H not HPnot PStnot St Y not YWnot W Records 92 126 16 202 161 57 99 119 88 130 All Chars 95.65% 95.24% 0.00% 100.00% 99.38% 85.96% 90.91% 94.12% 95.45% 96.92% 10 Chars 93.48% 93.65% 56.25% 99.50% 97.52% 87.72% 88.89% 93.28% 89.77% 96.15% 5 Chars 93.48% 90.48% 31.25% 99.01% 97.52% 87.72% 86.87% 89.92% 87.50% 90.77% Char 1 palea tip width palea length rachis extension length lemma awn length upper glume width Char 2 upper glume awn lemma awn length palea cilia length palea tip width number of inflorescence nodes Char 3 number of inflorescence width of hyaline margin lemma awn length length of middle inflo- culm thickness nodes on lower glume rescence internode Char 4 length of middle inflo- lemma margin hair lower glume width spikelet length length of middle inflores- rescence internode length cence internode Char 5 lower glume awn lemma length palea length length of lowest inflores- lower glume width cence internode Char 6 upper glume vein num auricle frequency lemma length number of inflorescence rachis extension length nodes Char 7 lemma length length of middle inflo- length of middle inflo- rachis extension length lower glume length rescence internode rescence internode Char 8 lower glume hair dens lemma back hair length lower glume hair density lemma tip hair length anther length Char 9 florets per spikelet lemma vein number upper glume width palea cilia length culm height Char 10 spikelet length Inflorescence length upper glume vein num- anther length palea tip width ber structure of the tribe, potentially impeding the development phrase, for instance, whether “densely hairy” meant that the of new insights into its morphology, ecology, and biology by hairs concealed all or about half of the underlying surface. the next generation of field-oriented botanists and plant Even measurements are somewhat ambiguous, for instance, breeders. where does the lemma body end and the awn start? These Of greater interest, in many respects, was the identifica- problems are not peculiar to the Triticeae. They can be ad- tion by RF analysis of the characters that are most useful in dressed, in part, by using images. distinguishing the genomic genera and of the characters that In building the generic descriptions, we attempted to take tend to characterize species that have a particular genome in into account all available information, not just that gained common. These analyses confirmed Salomon and Lu’s ob- through this study. Thus Elymus is described as usually hav- servation that palea tip width is a useful character and sug- ing two or more spikelets per node, but with only one in gested several other characters that should be given closer some species although, for this study, we examined only spe- attention such as middle inflorescence internode length and cies with one spikelet per node. Moreover, the disposition glume shape. of some of these species, notably those treated in Hystrix, Users of taxonomic classifications, however, do not wish is problematic. At present, it appears that some belong in to run a computationally demanding analysis in order to Elymus, others in Leymus. Because they do not have solitary identify their plants. Consequently, one of our goals was to spikelets, such species were not included in this study. Once develop practical identification keys to the genomic groups they have been examined and their appropriate generic based on morphological features. Nowadays, such keys are membership decided, it may be necessary to modify the ex- usually presented in one of two forms, multi-access keys, in isting generic descriptions. which the user determines which information to provide and For the sake of those working with the tribe, we posted the order in which to provide it, or dichotomous keys, in generic descriptions for all the genera of the tribe to the web which the author of the key decides the order in which the (http://herbarium.usu.edu/triticeae/genera), not just those in- characters are used. Before any keys can be constructed, cluded in this study. All the descriptions will be modified as however, good descriptions are needed. more information and images become available. Our origi- Descriptions of genera should be developed from de- nal intent was to provide keys solely to the perennial species scriptions of the species they include. In the absence of with solitary spikelets, but we eventually decided to expand detailed and parallel species descriptions, we developed ge- them to include all the genera that we currently recognize. neric descriptions by combining information from our The keys are intended to permit generic identification of observations with information from published descriptions non-hybrid Triticeae from anywhere in the world. This in- and identification keys. This process often required examin- creases their value to plant breeders and gene bank curators ing specimens to clarify what an author meant by a given who often hold accessions from distant locations; it makes Morphological identification of perennial genomic genera 567 them less useful to regional botanists because some of the olution images, effectively providing the viewer with a genera will be absent from their region. With the web-based magnifying lens (for examples, see http://www.herbarium. keys, however, it is possible to specify that the specimen is usu.edu/triticeae/zoom). CoolIris (2009) facilitates viewing from Europe, Asia, the Americas, Australasia, or Africa. multiple images, thereby enabling visitors to have overview Like the descriptions, the keys reflect our current knowledge of the species in a genus, or of variation in a species. Prior to of the genera. the web, such an overview could only have been obtained by In developing the keys to the genomic genera, we drew borrowing specimens from multiple herbaria. The only re- on information gained from the DA and RF. It was, howev- striction on developing such resources, and it is a significant er, only after conducting additional examination of the spec- restriction, is the time and effort involved in obtaining the imens and the leads used in other works that we were able to images and, in the case of images of living plants, preparing develop the keys that we have posted to the web. The dichot- the specimens that document them. The greatest restriction omous key was easier to construct than the multi-access key on using web-based resources is internet access. In some because we were able to decide in what order to bring out parts of the world, this is inexpensive and reliable; in others each group. This meant we could bring out the distinctive it is non-existent, expensive, unreliable, or some combina- groups first, using the characteristics that set them apart, and tion of these three. A partial solution to these problems is to did not need to know about their other characteristics. We make the resources available on a flash drive or compact are, however, aware that users will find many of the leads in disk and develop a mechanism for updating the files that the dichotomous key difficult. Multi-access keys are gener- does not require a complete download. Those interested in ally thought to be easier to use but constructing them re- obtaining such copies of the descriptions and keys devel- quires having information about all the characters for all the oped for this paper should write to Barkworth. genera. Because we currently lack such information, the The greatest need now for taxonomists working in the multi-access key is, as yet, incomplete. Triticeae is to develop detailed, parallel species descriptions The dichotomous key relies primarily on qualitative char- and documented distribution maps. In globally distributed acters because the range of values for all the quantitative taxa such as the Triticeae, this requires collaboration in iden- characters overlapped among genera. For instance, most tifying the characters to examine and how to record their species of Kengyilia have short rachis internodes at mid- variation. Digital technology can help in this regard, but it inflorescence, but a few species have long internodes. Similar will not, by itself, provide the solution. exceptions exist in all the genera having more than a few In the last few paragraphs, we emphasize the role that species. digital technology can play in the creation of taxonomic re- Unlike the dichotomous key, the multi-access key fea- sources. It does not, however, address our primary question, tures many quantitative characters, because users often feel whether or not the genomic genera can be identified mor- more confident in using such characters. Their inclusion in phologically. Our conclusion is that they can be, but that it the key will enable users to narrow down the possibilities us- will require examination and clearer description of features ing the characters they prefer but a final determination may such as palea shape, middle inflorescence internode length, require use of the qualitative characters. The multi-access and glume shape. We also found that, as is to be expected, key also includes a distributional character. It should be used genera that have some genomes in common are more diffi- with caution because many species of Triticeae have become cult to distinguish than those with no genomes in common. established beyond their native range. Both the dichotomous and multi-access keys are available at http://www.herbarium. Acknowledgements usu.edu/Triticeae/keys. The absence of detailed, comparable species descriptions We thank the curators of the herbaria from which we have means that both the generic descriptions and the keys must borrowed specimens (CANB, CHR, MO) and Drs. B.R. be regarded as preliminary. They will undoubtedly require Baum, Henry Connor, B.R. Lu, B. Salomon, Chi Yen, and modification as more information becomes available. Modi- H.-Q. Zhang for many discussions concerning the fying multi-access keys is relatively simple; modifying di- of the Triticeae and for sharing information with us. chotomous keys, including those presented on the web is harder, because it may require rethinking all the leads. But Literature Cited providing web-based keys and descriptions makes it easier to address another stumbling block for users, that of under- Assadi,M. (1996) A taxonomic revision of Elymus sect. Caespitosae standing what an author meant by a particular term or and sect. Elytrigia (, Triticeae) in Iran. Willdenowia 26: 251–271. phrase. Digital files can incorporate illustrative images and Barkworth, M.E., K.M. Capels, S. Long, L.K. Anderton and M.B. Piep, drawings at minimal expense. The programs that we used re- editors. (2007) Flora of North America, volume 24. Poaceae, part strict the number and size of the files that they can incorpo- 1 Oxford University Press, New York, p. 911. rate, but they can include links to other web sites where such Barkworth, M.E., S.W.L. Jacobs and H.Q. Zhang (2009) Connorochloa: resources are made available in a different style. For in- a new genus in Triticeae. Breed. Sci. 59: 685–686. stance, Zoomify (2009) permits zooming in on high res- Baum, B.R., J.R. Estes and P.K. Gupta (1987) Assessment of the 568 Barkworth, Cutler, Rollo, Jacobs and Rashid

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APPENDIX. List of voucher specimens, grouped according to their genomic constitution. Not all species have names that reflect their genomic constitution. [Most of the specimens of Roegneria and Kengyilia were collected in China and have labels that are written in Chinese. I shall try to have them translated, at least to the point where we can list the province in which they were collected, by the time this paper is back from review].

AGROPYRON Gaertn. (P): A. fragile (Roth) Candargy: (1) U.S.S.R., T. Stockton & D.R. Dewey PI-3146043, UTC188505; (2)U.S.S.R., T. Stockton & D.R. Dewey PI-440089, UTC 181143; (3) UTC 188515; A. dasyanthum (Ledeb. ex Spreng.) Ledeb.: (1) MO 4983310. ANTHOSACHNE Steud. (StWY): A. aprica: (Á. Löve & Connor) C. Yen & J.L. Yang: (1) Rexburgh, Otago, New Zealand, V.D. Zotov, CHR 19791; (2) East Roxburgh, Central Otago, New Zealand, H.E. Connor, CHR 370823; (3) Lindis Pass, Otago, New Zealand, V.D. Zotov, CHR487198; Elymus falcis Connor: (1) Mt. Ida, Otago, New Zealand, V.D. Zotov, CHR 19684; (2) Mt. John, Mackenzie County, New Zealand, H.E. Connor, CHR402714; Elymus fertilis Song Wang ex S.W.L. Jacobs & Barkworth (1) Brisbane River, Moreton District, Queensland, Australia, C.E. Hubbard 4811, 3654, 5311, CANB 497897, CANB 497853, CANB 299342; (2) Darling Downs district Queensland, Australia, C.E. Hubbard 5628, CANB 304216; (3) Leichardt District, Queensland, Australia, C.E. Hubbard 4981, CANB 304217; Elymus longisetus (Hitchc.) Veldkamp: (1) Mt. Suckling, Abau Subdistrict, Central District, Papua, K. Paijmans 973, CANB 219409; (2) Isuani Basin, SE flank of Mt. , Central District, Papua, L. A. Craven 2864, CANB 268673.1; (3) Sugarloaf grasslands, South of Wapenamanda, Wabag Area, western Highlands District, New Guinea, R. G. Robbins 2757, CANB 87785; (4) Pompameri, Kuraglumba track, Chimbu Valley, New Guinea, J.M.B. Smith 15508, CANB 240648; (5) 10 km SE of Keglsugl, Kundiawa Subdistrict, Chimbu District, New Guinea, K. Paijmans 1264, CANB 222948.1; Elymus multifloris (Hook.f.) Á. Löve & Connor: (1) North Island, New Zealand, T. Kirk 956, CHR 333058; (2) Coast of Hobson Bay (cliffs of), New Zealand, D. Petrie, CHR 335237; (3) Hamurama, Lake Rotorua, New Zealand, D. Petrie, CHR 1599; (4) Tauroa, New Zealand, H. Carse 32513, CHR 333064; (5) Whanearva, New Zealand, H. Carse 32511, CHR 333062; A. multiflora (Banks & Sol. ex Hook.) C. Yen & J.L. Yang: (1) Stonehenge, New England, New South Wales, Australia, R. W. Jeasup & M. Gray, CANB 93078; (2) “Aughamore” Biddeston, Australia, W. J. Bisset 5752, CANB 22846; (3) Between “Harrow” & Cambooya, Queensland, Australia, W.J. Bisset 5733, CANB 22064; (4) One mile E. of Bunnan, Hunter Valley, New South Wales, Australia, R. Story 6943, CANB 73412; (5) 32.3 km from South on Tilpa Road, East of Darling Road, Australia, C.W.E. Moore 7836, CANB 270686.1; Elymus sacandros Connor: (1) Maxborough, New Zealand, A. P. Druce, CHR 279243, CHR 279211, CHR 279245; A. scabra (R.Br.) Nevski: (1) Alexandra, Central Otago, New Zealand, H.E. Connor, CHR 402716; (2) Bank’s Peninsula, New Zealand, E.A. Madden, CHR 56231; (3) North Canterbury, New Zealand, H.E. Connor, CHR 402719, CHR 402720; (4) Bankside Reserve, Ellesmere Co., Canterbury Plns., B.P.J. Molloy, CHR 201486; Elymus solandri (Steud.) Connor: (1) Mt. Edwards, Canterbury, New Zealand, V.D. Zotov, CHR 19686; (2) Portabello, Ortago Harbour, New Zealand, E. Edgar, CHR 95442; (3) Whararchni Bead, MN Nelsa, New Zealand, A.P. Druce, CHR 355260; (4) Mt. Edward, Mackenzie Country, New Zealand, H.E. Connor, CHR 402744; Kaweka Range, Hawkes Bay, New Zealand, B.C. Aslin & D. Petrie 32614, CHR 333081. AUSTRALOPYRUM (Tzvelev) Á. Löve (W): A. calcis (Connot et Molloy): (1) Leatham River, Marlborough, New Zealand, B.P.J. Molloy & K.W. Ryan L4, CHR 468519; (2) Leatham River, Marlborough, New Zealand, A.P. Druce, CHR 282637; (3) Leatham River, Marlborough, New Zealand, M.I. Dawson 19352, CHR 475216C; (4) Castle Hill Basin, Canterbury, New Zealand, B.P.J. Molloy PH 3, CHR 468523; (5) Flock Hill, Canterbury, New Zealand, B.P.J. Molloy FH1c, CHR 468528; A. pectinatum (Labill.) Á. Löve: (1) Kybean Range, New South Wales, Australia, L.G. Adams & M.P. Austin 2905, CANB 291092; (2) Mt. Lowden, New South Wales, Australia, L.G. Adams 3273, CANB 381607; (3) Between Pikes Saddle and the Badja Sawmill, New South Wales, R. Pullen 11,078, CANB 345596; (4) On trail to Mt. Cowangerong, New South Wales, Australia, R. Pullen 11,076, CANB 345594; (5) Along Badja track, New South Wales, Australia, R. Pullen 11,077, CANB 345595; A. retrofractum (Vickery) Á. Löve: (1) Mt. Cooper, New South Wales, Australia, P. Michael, CANB 91023; (2) Near Coopers Lake, New South Wales, Australia, R. Pullen 11,079, CANB 345597; (3) Southern Tablelands, New South Wales, Australia, I. Crawford 3976, CANB 604507.1; (4) Rock Flat, 15 km south of Cooma, New South Wales, Australia, F.A. Zich 18, CANB 393532; A. uncinatum (Veldkamp): (1) Mt. Kenive (Nisbet), Subdist Kokoda, Papua, J.R. Croft et al. 65112, CANB 391072.1; A. velutinum (Nees) B.K. Simon: (1) Victoria, Australia, M.J. Henwood 223& 226, CANB 393525, CANB 393527; (2) Kiandra Plain, New South Wales, A.V. Slee 2370, CANB 470504; (3) Blue Lake walking track from Charlottes Pass, New South Wales, Australia, J. Palmer 447, CANB 466333; (4) Path from Charlottes Pass toward Blue Lake, New South Wales, Australia, F.A. Zich 40, CANB 402916. CAMPEIOSTACHYS Drobow (StYH): Elymus drobovii (Nevski) Tzvelev: UTC 177461, UTC164446, UTC 179277; E. tsukushiensis: 5 specimen-UTC 179277, UTC 188352, UTC 188188, UTC 188081, UTC 188319; E. dahuricus: MO 4755744, MO 4725062, MO 4550628, MO 4482821, MO 4550623. DOUGLASDEWEYA C. Yen, J.L. Yang & B.R. Baum (StP): D. deweyi (K.B. Jensen, S.L. Hatch & Wipff) C. Yen, J.L. Yang, & B.R. Baum: (1) Stavropol Botanical Gardens, U.S.S.R., K. Asay & M. Rumbaugh PI-502264, UTC 233615; D. wangii C. Yen, J.L. Yang, B.R. Baum: (1) Iran, C. Nowak & D.R. Dewey PI 401328, UTC 177418; (2) Iran, T. Stockton & D.R. Dewey PI-401330 & PI-380646, UTC 186398 & UTC 186387; (3) Iran, C. Nowak & D.R. Dewey PI 401328, UTC 177678. ELYMUS L. (StH or StStH): E. agropyroides Presl.: (1) Argentina, Dean Corey CS-13-70, UTC 151117, (2) Argentina, S.L. Hatch 656, UTC 164523; (3) R.N. Hurst PI 204183, UTC 128449; (4) Argentina, C. Nowak & D.R. Dewey PI 269646, UTC 177823; E. alaskanus (Scribn. & Merr.) Á. Löve: (1) Northwest Territories, Canada, W.J. Cody & D.H. Ferguson 10461, UTC 202093; (2) Alaska, U.S.A., L.A. Viereck & J. Foote, UTC 223520; (3) Yukon Territory, Canada, J.M. Gillett & J.A. Calder et al. 4217, UTC 225724; (4) British Columbia, Canada, M.E. Barkworth & P. Hostin 9190, UTC 209001; E. arizonicus (Scribner & J.G. Smith) Gould: (1) New Mexico, U.S.A., C. Nowak & D.R. Dewey He-4-5, , UTC 179626; (2) Texas, U.S.A., R.D. Worthington 23759, UTC 213267; (3) Arizona, U.S.A., F.W. Gould 2600, UTC 67542; E. bakerii (E. Nelson) Á. Löve: (1) Colorado, U.S.A., R.K. Gierisch I283, UTC 121086; (2) Colorado, U.S.A., A.S. Hitchcock 1467, UTC 228101; E. caninus (L.)L: (1) U.S.S.R., T. Stockton & D.R. Dewey PI-314612, UTC 181013; E. dentatus (Hook f.)Tzvelev: (1) U.S.S.R., T. Stockton & D.R. Dewey PI-314614, UTC 188488; E. fibrosus (Shrenk) Tzvelev: (1) U.S.S.R., C. Nowak & D.R. Dewey HE-1-86, UTC 177811; (2) U.S.S.R., C. Nowak & D.R. Dewey PI 315493, UTC 179517; E. glaucus Buckley: UTC 138789, UTC217481; E. lanceolatus (Scribn. & Merr.) Gould: (1) Utah, U.S.A., B.F. Harrison & E. Larsen 7967, UTC 17457; (2) Nevada, U.S.A., G.P. Hallsten 2175, UTC 210575; (3) Nevada, U.S.A., C. Nowak & D.R. Dewey PI 232115, UTC 222838; E. macrourus (Turcz.) Tzvelev: (1) Alaska, U.S.A., A.S. Hitchcock 1466, UTC 228102; E. mutabilis (Drobow) Tzvelev: (1) Khirgiz, U.S.S.R., C. Nowak & D.R. Dewey Jaaska 20-73, UTC 177806; (2) , Steve Larson PI 564954, UTC 235940; E. repens (L.) Gould:-UTC 179288, UTC 20305 E. riparius Wiegand: (1) Kentucky, U.S.A., M.E. Barkworth & J.J.N. 570 Barkworth, Cutler, Rollo, Jacobs and Rashid

Campbell 94.115, UTC 216092; (2) Kentucky, U.S.A., M.E. Barkworth, J.J.N. Campbell, & M. Hines 94.111, UTC 216081; E. scabriglumis (Hack.) Á. Löve: (1) Argentina, C. Nowak & D.R. Dewey PI 202256 & PI 202147, UTC 244194 & UTC 244191; (2) Argentina, USDA-ARS PI- 331168, UTC 244190; (3) Argentina, E.E. Terrell 4760, UTC 244193; (4) Argentina, Dean Corey CS-12-66, UTC 244192; E. scribneri (Vasey) M.E. Jones: (1) Utah, U.S.A., A. Taye 2805, UTC 195077; (2) Nevada, U.S.A., N.H. Holmgren & J.L. Reveal 1585, UTC 108241; E. sibircus L.: (1) China, C. Nowak & D.R. Dewey D-2715, UTC 179678; (2) China, S. Larson PI 499464, UTC 235953; (3) Yukon Territory, Canada, W.J. Cody & J.H. Ginns 28365, UTC 225706; E. sierrae (Scrib. & Smith) Hitchc.: (1) California, U.S.A., C. Nowak & D.R. Dewey CS-10-15, UTC 179563; E. ternatus: (1) Bolivia, R.L. & K. Parker 25-a, UTC 136673; E. trachycaulus (Link) Gould ex Shinners: (1) Montana, U.S.A., C.L. Hitchcock & C.V. Muhlick 12928, UTC 71787; (2) Ontario, Canada, M.J. Oldham & W.D. Bakowsky 26648, UTC 234183; (3) Wyoming, U.S.A., A. Nelson & R.A. Nelson 1113, UTC 24888; (4) Colorado, U.S.A., J.L. Reveal 1866, UTC 163498; (5) Idaho, U.S.A., M.E. Barkworth & C. Annable 94.071, UTC 215834; E. transhyrcanus (Nevski)Tzvelev: (1) Iran, C. Nowak & D.R. Dewey PI 229912, UTC 177318; (2) U.S.S.R., T. Stockton & D.R. Dewey PI 314208, UTC 181051; E. vaillantianus (Wulf. & Schreb.) K. Jensen: (1) Coahuila, Mexico, J. Valdes & E. Demesa VR-1665, UTC 205834; (2) Coahuila, Mexico, P.S. Hoge, M.E. Barkworth & M.A. Carranza 294, UTC 207070; E. violaceus (Hornem.) Feilberg: (1) Northwest Territories, Canada, W.J. Cody & F.M. Brigham 20643, UTC 204826; E. wawawaiensis J. Carlson & Barkworth: (1) Washington, U.S.A., S. Larson PI 285272, UTC 235945; (2) Washington, U.S.A., T. Stockton & D.R. Dewey PI 285272, UTC 222854. FESTUCOPSIS Melderis (L): F. serpentini (C.E. Hubb.) Melderis: MO 1622253, MO 2256852. KENGYILIA C. Yen & J.L. Yang (StPY): K. alatavica (Drobow) J.L. Yang, C. Yen & B.R. Baum: (1) Khirgiz, U.S.S.R., T. Stockton & D.R. Dewey Jaaska 54-73, UTC 188330; (2) Khirgiz , U.S.S.R., C. Nowak 7 D.R. Dewey Jaaska 54-73, UTC 179175; K. hirsuta (Keng) J.L. Yang, C. Yen & B.R. Baum: (1) China, T. Stockton & D.R. Dewey D-2548, UTC 188356: (2) , China, C. Nowak & D.R. Dewey D-2588, UTC 179284; K. laxiflora (Keng) J.L. Yang, C. Yen & B.R. Baum: MO 5329378; K. melanthera (Keng) J.L. Yang, C. Yen & B.R. Baum: MO 5331440; K. thoroldiana (Oliv.) J.L. Yang, C. Yen & B.R. Baum: MO 5329389. PSEUDOROEGNERIA (Nevski) Á. Löve (St): P. spicata (Pursh) Á. Löve : (1) North Fork of Twin Creek,Wyoming, U.S.A., B. Neely & A. Carpenter 1334, UTC 242532; (2) City Creek Canyon, Utah, U.S.A., J. Ludwig 599, UTC 130194; (3) Washington, U.S.A., F.O. Kreager 419, UTC 50120; (4) Meadow Creek, Idaho, U.S.A., A. Cronquist 3675, UTC 49992; (5) Montana, U.S.A., H.T. Rogers 829, UTC 65271; P. tauri (Boiss. & Bal.) Á. Löve: (1) Iran, S.L. Hatch 637, UTC 164506. ROEGNERIA: K. Koch (StY): R. abolinii (Drob.) Nevski: (1) U.S.S.R., T. Stockton & D.R. Dewey PI-440104, UTC 188493; (2) U.S.S.R., T. Stockton & D.R. Dewey, UTC 188363; (3) U.S.S.R., S.L. Hatch 650, UTC 164506; R. antique (Nevski) B.S. Sun: MO 4365622; R. brevipes Keng: MO 4096873; R. burchan-buddhae (Nevski) B.S. Sun: MO 4482560; R. caucasica K. Koch: (1) U.S.S.R., T. Stockton & D.R. Dewey CS-7-11, UTC 188358; R. ciliaris (Trin.) Nevski (1) Japan, C. Nowak & D.R. Dewey D-2712, UTC 179614; (2) Japan, T. Stockton & D.R. Dewey PI-276395, UTC 181060; R. gmelinii (Ledeb.) Kitag.: (1) , China, C. Nowak & D.R. Dewey D-2660, UTC 179173; (2) China, C. Nowak & D.R. Dewey D-2670, UTC 179295; R. longearistata (Boiss.) Drobow: (1) Iran, T. Stockton & D.R. Dewey PI-401277, UTC 181101; (2) Iran, C. Nowak & D.R. Dewey PI 401276, UTC 177434; R. panormitana (Parl.) Nevski: (1) , T. Stockton & D.R. Dewey PI-206402, UTC 181048; (2) Iraq, C. Nowak & D.R. Dewey CS-9-39, UTC 179611; R. pendulina Nevski: MO 4497323; R. semicostata (Nees ex Steud.) Kitag.: (1) , T. Stockton & D.R. Dewey PI-207452, UTC 181041; (2) West Pakistan, C. Nowak & D.R. Dewey PI 275305, UTC177799; R. tschimganica (Drobow) Nevski: (1) Lake Issyk-kul, U.S.S.R., C. Nowak, D.R. Dewey CS-31-7, UTC 177432; (2) U.S.S.R., C. Nowak & D.R. Dewey CS-10-61, UTC 177403. STENOSTACHYS Turcz. (WH): S. deceptorix (Connor): (1) Upper Maruia River, New Zealand, A.P. Druce, CHR 358102A, CHR 358102B; (2) Taylor Basin, New Zealand, A.P. Druce, CHR 273750; (3) Owen Range, New Zealand, A.P. Druce, CHR 249770; Australopyrum ensyii (Kirk) Connor: (1) Mt. Torlesse, New Zealand, H. Talbot, CHR 92018; (2) Porters Pass, New Zealand, H.E. Connor, CHR 115530; (3) Mt. Peel, New Zealand, A.P. Druce, CHR 252248; (4) Owen Range, New Zealand, A.P. Druce, CHR 249773; (5) Mt. St. Patrick, New Zealand, Canterbury Botanical Society, CHR 225704; S. gracilis (Hook.f.) Connor: (1) Stewart Island, New Zealand, I.A. McNeur, CHR 72644; (2) Ruahine Mts., New Zealand, A.P. Druce DEG774, CHR 74357; (3) Eyre Mountains, New Zealand, A.P. Druce APD73, CHR 468998; (4) Marlborough, New Zealand, A.P. Druce, CHR 279248; (5) Bealey River, New Zealand, H. Talbot, CHR 301426; S. laevis (Petrie) Connor: (1) Landsborough River, New Zealand, P. Wardle, CHR 223892; (2) Black Bereh Range, New Zealand, A.P. Druce, CHR 165513; (3) Lake Coleridge Station, New Zealand, H.E. Connor, CHR 92011; (4) N.W. Ruahines, New Zealand, A.P. Druce DEG745, CHR 74358; (5) Copland Pass, New Zealand, A. Wall 620, CHR 333118. THINOPYRUM (J): T. caespitosum (Koch) Á. Löve: (1) Utah, U.S.A., D.R. Dewey 228389, UTC 121208; (2) Turkey, T. Stockton & D.R. Dewey PI-383583, UTC 188145; (3) Crimea, U.S.S.R., C. Nowak & D.R. Dewey Jaaska-5, UTC 188166; T. intermedium (Host) Barkworth & D.R. Dewey: (1) Utah, U.S.A., L.C. Anderson 790, UTC 93591; (2) Oregon, U.S.A., B.L. Wilson & R. Brainerd 11404, UTC 248635; (3) Savournon, France, Bureau of Genetic Resources 59, UTC 205378; T. junceum (Simonet ex Guin.) Á. Löve: (1) France, C. Nowak & D.R. Dewey TS-1-31, UTC177404; Lophopyrum nodosum (Nevski) Á. Löve: (1) U.S.S.R., C. Nowak & D.R. Dewey A-62-15, UTC 177320; T. obtusiflorum (Podp.) Barkworth: (1) Nova Scotia, Canada, A. Kepinsky, M.E. Barkworth, & J. McNeill 6215, UTC 165109; (2) British Columbia, Canada, M.E. Barkworth & P. Hosten 91104, UTC 208995; (3) Utah, U.S.A., N.H. Holmgren 188, UTC 100014; (4) Luganskiensis, Ukraine, M. Kotov, UTC 154494; (5) U.S.S.R., C. Nowak 7 d.R. Dewey A-62-15, UTC 177320. Connorochloa Barkworth, S.W.L. Jacobs, & H.Q. Zhang (StYHW). Connorochloas tenuis (Buch.) Barkworth, S.W.L. Jacobs, & H.Q. Zhang: (1) N. of Lake Clearwater, New Zealand, R. & E.F. Melville & H.E. Connor 6221, CHR 143003; (2) Kaimanawa Mts., New Zealand, A.P. Druce, CHR 402410; (3) Oreti River, New Zealand, V.D. Zotov, CHR 19708; (4) Central Otago, New Zealand, P.N. Johnson, CHR 364502; (5) Waeourii, New Zealand, A. Wall, CHR 333077.