SIDA swine" Volume 5 1972 — 1974 SIDA Contributions to Botany volume 5 (in 5 numbers) Copyright 1972, 1973, 1974, 1975 by Wm. F. Mahler SMU Herbarium Dallas, Texas 75275 DATES OF PUBLICATION No. 1, pp. 1—58: 24 Mar. 1972 No. 2, pp. 59—136: 30 May 1973 No. 3, pp. 137—190: 18 Jan. 1974 No. 4, pp. 191—300: 29 July 1974 No. 5, pp. 301—354: 11 Feb. 1975 Index, pp. 355—361 For contents, see the unnumbered pages forming the front covers of the separate issues. S| DA CONTRIBUTIONS TO BOTANY 2A ee ee eee VOLUME 5 NUMBER 1 MARCH 1972 2 ee A el Ss ie a CONTENTS A reconsideration of the correct name for the Hawaiian Gossypium, Paul A. Fryxell. 1 Chromosome numbers in Physalis and Solanum (Solanaceae). John E. Averett and A. Michael Powell, 3 A revision of the genus Poliomintha (Labiatae). Robert S. Irving. 8 Two new species of Isocoma (Compositae-Astereae) from north-central Mexico. B. L. Turner 23 Field observations of Cuthbertia (Commelinaceae) with description of a new form. Olga Lakela 26 Observations on some Upper Amazonian formicarial Melastomataceae. Trevor Whiffin. 33 NOTES. Galium pedemontanum (Bell.) All. (Rubiaceae) reported from Oklahoma. 42—Three grasses new to Georgia: Eriochloa gracilis, Setaria faberii, Leptochloa uninervia (Gramineae), 42—New combinations in Hoffmanseggia Cav. and Caesalpinia L. 43— Vernonia crinita Raf., a synonym of ¥. arkansana DC. (Compositae). 44—Rotala indica (Lythraceae) in Louisiana, 45—New combinations and a new species in Helenium (Compositae-Helenieae). 45—Adventive records of Eriochloa contracta (Gramineae) in the United States. 48—Two new species of Chamaesaracha (Solanaceae) from Texas. 48—A new variety of Carex bicknellii from Arkansas. DOCUMENTED PLANT CHROMOSOME NUMBERS 1972: 1, 2. 50 BOOK NOTICES 53 LOWELL DAVID FLYR, 1937-1971 54 SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. Mahler, SMU Herbarium, Dallas, Texas, 75222, U. S. A. Subscription price $8 (U.S.) per volume; parts issued at irregular intervals. ASSOCIATE EDITOR John W. Thieret REVIEWERS & CONSULTANTS (for this issue) Rupert C. Barneby Robert Kral Walter H. Lewis Lloyd H. Shinners (C) g SIDA Contributions to Botany, Volume 5 Number 1, pages 1—55 Copyright 1972 by Wm. F. Mahler A RECONSIDERATION OF THE CORRECT NAME OR THE HAWAIIAN GOSSYPIUM. PAUL A. FRYXELL Research Geneticist, Plant Science Research Division, Agricultural Re. search Service, U. S. Department of Agriculture, Texas A&M University, College Station, Texas 77843. Wilbur (1964) proposed rejection of the name Gossypium tomentosum utt. ex Seem. for the native Hawaiian cotton, and acceptance of the next oldest valid name, G. sandvicense Parl. His basis for the rejection of See- mann’s (1865) name is that Seemann’s original conception of the species was too broad, encompassing plants both from Hawaii, where the species in question occurs, and from Fiji, where it does not. Wilbur clearly documents the fact that Seemann’s description is hetero- geneous, as Watt (1907) had noted earlier, and that it was derived both from Hawaiian specimens of G. tomentosum and from Fijian specimens that have been described under the name G. taitense Parl., but which are now generally included in G. hirsutum L. Wilbur Heeupon attempted to lectotypify Seemann’s name by circumscription: he stated that “G. to- mentosum must be typified by the plants principally characterized by the publishing author. . . He then expressed the opinion that the description is more heavily weienied toward the Fijian element than the Hawaiian element, stating that “it is readily apparent that the greater portion of Seemann’s original diagnosis can apply only to the Fijian cot- ton.’ His opinion is surely subject to challenge. One could, for example, note that Seemann cited three specimens from Fiji (Smythe, Pritchard, and Seemann) and four specimens from Hawaii (Diell, Nelson, Menzies, and Nuttall) as well as a specimen of Trove “from Joynegau”’ which Watt (1907, p. 129) indicates is a specimen of Hove “‘from Joynegare’’? (=Juna- gadh, Kathiawar), located in western India. On such a basis, one might equally well conclude that the Hawaiian element rather than the Fijian element “predominated” in Seemann’s conception of the species. More to the point, perhaps, is a re-examination of Seemann’s descrip- tion and Wilbur’s analysis of it. Wilbur notes three items (stipules, bract teeth, and seed hairs) that favor the Fijian element in Seemann’s descrip- tion and only one (calyx form) that favors the Hawaiian. I agree with Wilbur’s interpretation of bract teeth, seed hairs, and calyx form, but not that of the stipules. (I do not believe any conclusion can be reached for this item, because of the variability in these plants of the stipules, which differ in vegetative and reproductive branches. Apparently neither Seemann nor Wilbur was aware of this fact, and the descriptive terminology employed does not permit any conclusion to be reached concerning the origin of the specimens whose stipules Seemann described.) Moreover, Wilbur overlooked two items (corolla color and indumentum) that tend SIDA 5(1): 1—2, 1972, to favor the Hawaiian element. Therefore, one cannot conclude that cither the Fijian or the Hawaiian clement ‘‘predominated” in Seemann’s descrip- tion, but only that the description is indeed composite. Wilbur also failed to consider the extent to which the description might have been based upon Hove’s plant from India. However, such attempts to express a value judgement are beside the point. The problem of the correct application of the name G. tomentosum is to be resolved on the basis of typification. Wilbur rightly emphasized the importance of lectotypification in resolving this question, but curiously failed to choose a lectotype I designate Nuttall’s specimen, bearing his manuscript name and kept at the British Museum (Natural History), as the lectotype of Gossypium tomentosum Nutt. ex Seem. Nuttall’s specimen is the one from which See- mann took the name, and it may be viewed as the ‘‘holotype’’ of Nuttall’s unpublished name, which Seemann subsequently published. It therefore seems the most appropriate choice as lectotype of Seemann’s name. Such a choice preserves established usage. I see no advantage to a lectotypifi- cation that upsets established usage and is therefore contrary to Recom- mendation 7B of the Code, as was suggested (although not done) by Wilbur. That Seemann included Fijian and Indian material in his citation of speci- mens and utilized them to a degree in drafting his description, was simply a taxonomic error on his part that need not deflect us from making a correct nomenclatural decision. The problem is indeed resolved by a satis- factory typification. The name that Wilbur concluded was the correct one for the Hawaiian endemic, G. sandvicense Parl. (Parlatore, 1866), was published one year later than G. tomentosum, and is thus to be relegated to synonymy under the latter name, as most authors have done. Parlatore’s name has evidently never been typified, and so I will take this opportunity to rectify that omission. Parlatore cites specimens of Forster, Menzies, and Nuttall (the last-named cited with a question mark) following his description of the species, all collected in the Sandwich Islands (i.e., Hawaii). Only Menzies’ specimen is cited with an exclamation point (the usual indication that a specimen was actually examined) and mentioned in the subsequent dis- cussion. Therefore, I designate Menzies’ specimen as lectotype of Gossy- pium sandvicense Parl. That the same specimen was cited by Seemann, underscores the synonymy noted above. Aliotta (1903) notes the lectotype to be in the ‘‘Erbario Centrale Italiano” in Florence (FI). REFERENCES ALIOTTA, A. 1903. Revista critica del genere Gossypium. Della Torre: Portici. [cf. G. fomentosume on pp. 92-94. PARLATORE, F. 1866. Le specie dei cotoni. Stamperia Reale: Firenze. SEEMANN, B. C. 1865-1873. Flora Vitiensis. London. WATT, SIR GEORGE. 1907. cee wees of the world. Longmans, Green & Co.: London [ef. G. Pig dares on pp. 69- ie _- L. 1964. The correct name for the Hawaiian Gossypium, Pacific Sci, 18:101- CHROMOSOME NUMBERS IN PHYSALIS AND SOLANUM (SOLANACEAE) ' JOHN E. AVERETT and A. MICHAEL POWELL Department of Botany, The University of Texas, Austin, 78712* and Department of Biology, Sul Ross State University, Alpine 79830 While conducting a chromosomal survey Of Chamaesaracha, meiotic materials of several other Solanaceous taxa were collected from a wide area of the southwestern United States and northern Mexico. Chromosome counts obtained from this material are reported in Table I. Techniques for producing pollen mother cell squashes essentially follow those presented by Turner and Johnston (1961). Voucher specimens are deposited in the herbaria of The University of Texas, Austin, and Sul Ross State University. We are grateful to Dr. U. T. Waterfall for the identifi- cation of many of the Mexican species of Physalis. Table I. Species of Physalis and Solanum examined for chromosome number, Saree Taxon Chromosome Vouchers number (n) PHYSALIS: P. crassifolia Benth. 12 MEXICO: B.C.S. 67 mi N of Villa Constitu- var. crassifolia tion. Sikes and Babcock 280a,b,c 12 MEXICO: B.C.S. 2 mi N of San Antonio. Sikes and Babcock 266. MEXICO: B.C.N. 27 mi NE of San Felipe- Rosario Junction. Sikes and Babcock 307. P. crassifolia Benth. 12 MEXICO: B.C.N. 7 mi NE of San Fe lipe- _ bo var. infundibularis Rosario Junction. Sikes and Babcock 305. I.M. Johnst. *P. glabra Benth. 12 MEXICO: B.C.S. mi W of La Palmilla. 7 Sikes and Babcock 242. a ea na Schlect. 12 MEXICO: Durango. 4 mi W of Durango. Sikes var. glutinosa and Babcock 375. P. he ie pen Gra 12) NEW MEXICO: Otero Co. 29 mi SE of Cloud- var. cordifolia Gay croft. Sikes 69. Waterfall 12 ARIZONA: Yavapai Co. 38 mi S of Flagstaff. Tomb 272 *“P. hederaefolia Gray 12 TEXAS: iesweios Co. 6 mi S of Alpine. Aver- var. hederaefolia ett 126. Aided in part by National Science Foundation Training Grant GB-69 * Present address, Dept. of Biology, University of Missouri, St. Louis, OL SIDA 5(1): 3—7. 1972. P. lobata Torrey *P_ sordida Fernald TEXAS: Brewster Co. 13 mi S of Alpine. Averett I11. TEXAS: Brewster Co. 6 mi S of Alpine. Boston 29. MEXICO: Durango. 15 mi N of La Zarca Junction on Mex 45. Sikes and Babcock 364. TEXAS: Brewster Co. 16 mi FE of Alpine. Averett 84. TEXAS: Ector Co. 10 mi N of Odessa. Watson 139 TEXAS: Ector Co. 14 mi S of Odessa. Wat- son 191. TEXAS: Pecos Co. 18-24 mi W of Bakersfield. Tomb 414. TEXAS: Pecos Co. 21 mi E of Ft. Stockton. Tomb and Bierner 430. TEXAS: Presidio Co. 11 mi N of Presidio. Sikes and Averett 139. TEXAS: Val Verde Co. 1 mi FE of Langtry. Averett 283. TEXAS: or Verde Co. E city limits of Del Rio. Tomb 402 TEXAS: aN Co. 3/4 mi W of Dryden. Averett 277. NEW MEXICO: Chaves Co. 13 mi W of Ros- well, Averett and Tomb 330. NEW MEXICO: Grant Co. 9 mi W of Hachita on N. Mex #9. Tomb and Bierner 4 MEXICO: Chihuahua. 26 mi NE of Carmaree. Sikes and Patterson 406. NEW MEXICO: Lea Co. 1 mi W of Broncho. Averett and Tomb 321. NE EXICO: Union Co. 18 mi W of Clay- ton. Tomb 538. OKLAHOMA: Cimarron Co. 12 mi S_ of Campo, Colorado. Tomb 342. TEXAS: Concho Co. 3 mi S of Runnels county line on hwy. 88. Averett and Bierner XAS: Hardeman Co. 3 mi S of Quanah. Averett and Bierner 477. EXAS: Jones Co. 3 mi S of Stamford. Aver- ett and Bierner 479 TEXAS: Lubbock Co. 11 mi N of Lubbock. Averett a Tomb 35a. TEXAS: Young Co. Olney, Texas. Seigler 1469. MEXICO: Coahuila. 43 mi NW of Muzquiz. Powell and Patterson 1586. Sinaloa. 3 mi N of Mazatlan. Sikes and Babcock 200. *“P. vestita Waterfall . viscosa L. var. cinerascens (Dunal) Waterfall P. wrightii Gray SOLANUM: *S. amazonicum Ker. S. eleagnilolium Cav. *S. hindsianum Benth. *S. madrense Fernald S. nigrum L, S. rostratum Dunal 12 -— bo H bo — bo 12 12 MEXICO: Sinaloa. 6 mi N of Mazatlan. Sikes and Babcock 202. TEXAS: Bee Co. N side of Beeville. Tomb 354. ra Ector Co. 14 mi S of Odessa. Watson TEXAS hk Davis Co. 26 mi W of Toyahville. Sikes ARIZONA: Pinal se - mi S of Phoenix. Averett and Watson : ARIZONA: Pinal - mi § of Phoenix. Averett and Watson 412. MEXICO: Sonora. 1 mi E of Navajoa. Sikes and Babcock 184. TEXAS: Brewster Co. 6 mi SW of Marathon. Watson 18 TEXAS: faa Co. 1 mi W of Sierra Blan- ca. Sike TEXAS: en Co. 1 mi W of Sierra Blan- ca. Sikes G60b. TEXAS: Jeff oo Co. 24 mi NW of Ft. Davis. Boston NEW eo. Hidalgo Co. 17 mi W of Lordsburg. Averett and Watson 383. MEXICO: B.C.S. 1 mi N of Villa Constitution. Stkes and Babcock 253. MEXICO: Sinaloa. 70 mi S of Mazatlan. Sikes and Babcock 208. ARIZONA: Cochise Co. 24 mi W of E entrance to Chiricahua Nat'l. Monument. Averett and Watson 392. ARIZONA: Greenlee Co. 12 mi W of Mule Creek, N. Mex. Averett and Watson 435. ARIZONA: Santa Cruz Co. Madera Canyon. Averett and Watson 408. NEW MEXICO: Chaves Co. 43 mi SE of Cloudcroft. Sikes 71. nage MEXICO: Hidalgo Co. 10 mi S of I-10 n U.S. 80. Averett and Watson 384. TEXAS: Jeff ae Co. 1 mi E of Boy Scout mp. Sikes oo ne Co. Pinto Canyon. Sikes 82. TEXAS: San Patricio . Lake Corpus Chris- ti State Park. Tomb 3 TEXAS: Brewster a > mi E of Alpine. Watson 63. *S. triflorum Nutt. 12 UTAH: Beaver Co. 12 mi E of Milford. Aver- ett and Watson 421. * First report for the taxon. DISCUSSION Excepting n = 11 in P. lobata (= Quincula lobata (Torr.) Raf.), chromo- some numbers presented for the 18 taxa of Physalis and Solanum are on a base of x — 12. Initial counts are reported for nine taxa which are denoted by an asterisk in Table I. Most of the species examined are diploid (n — 12,11), substantiating the rarity of polyploidy in Physalis, Solanum, and related genera. In the Solanae, Chamaesara ha alone displays con- siderable polyploidy (Powell and Averett, 1967). Some elaboration is appropriate with regard to the occurrence of diploid and tetraploid races in P. lobata (Table I.). Menzel (1950) observed that three seed classes (based on size) of this species are correlated with fairly distinct geographic ranges. As seen in Figure 1, class TY (2.7-3.0 mm) ex- tends from central Texas north into Kansas, Colorado, and New Mexico. Class IL (1.8-2.2 mm) is largely restricted to southern and western regions of Texas and adjacent Mexico. Class II (3.8-4.0 mm) is limited to southern Arizona. Based upon counts from one tetraploid and four diploid popula- tions, Menzel further correlated seed sizes with chromosome numbers in classes I and II, representing the 4n and 2n populations respectively. A still higher ploidy level was thought to characterize class Ill. The counts reported in Table I generally adhere to the geographic ranges for the diploid and tetraploid races suggested by Menzel (Figure 1). No chromo- some counts are available from southern Arizona (class Iq). Menzel (1950) also noted a few herbarium specimens with large seeds within the range of the small-seeded populations and vice versa, Chromo- some counts for these mixed populations are lacking, but a tetraploid count for P. lobata was obtained from specimens collected in northern Mexico, well within ma diploid range of the species (Figure 1 An oid origin is suspected for the tetraploid race of P. lobata. oe the tetraploids are nearly identical to the diploids, and there is no indication that other species have been involved in their origin. Ithough segmental allopolyploidy is possible, Lewis (1967) has pointed out that it is not necessary to postulate a hybrid origin between ecologi- cally distinct populations in order to account for the establishment and maintenance of discrete polyploid populations. In any case, even con- sidering alloploidy, the tetraploid race presumably arose in desert areas of the southwestern United States and northern Mexico, and subsequently spread into the grassland areas to the north. Menzel (1950) concluded that the nm = 11 in P. lobata was derived from n = 12 by aneuploid loss. The existence of n = 12 in this taxon could be offered to support this conjecture. However, the extra chromosome could have resulted through aneuploid gain. Fig. 1. Distribution of tetraploid and diploid Physalis lobata; triangles (n = 22), circles (n = 11), and squares (n = 12). The distribution of three classes of seed sizes recognized by Menzel (1950) is also shown; 1 (2.7-3.0 mm), 2 (1.8-2.2 mm), and 3 (3.8-4.0). Discussion in text. REFERENCES LEWIS, oe Ee The taxonomic significance of auropo ypoidy . Taxon 16:267-271 MENZE Cytotaxonomic observations in some genera of the Solanae: Mars cerntns, ‘Suri a ee 2 J. Bot. 37:2 and J. E POW ETT. 1967. Give ke Pets of Chamaesaracha Solnacene) in iba Pecos ae and adjacent regions. Sida 3:156-162. TURN B. L. and M. C. JOHNSTON. 1961. Chromosome ee in the Compositae. Il. aus Mexican species. Brittonia 13:62-69. A REVISION OF THE GENUS POLIOMINTHA (LABIATAE) ROBERT S. IRVING Department of Botany, University of Montana, Missoula 59801 Poliomintha (Labiatae) is a small North American genus whose four shrubby species inhabit the arid regions of the southwestern United States and northern Mexico. Bearing only two fertile stamens in combination with a tubular calyx, it is placed in the subtribe Melissinae, a heterogeneous group long recognized as taxonomically difficult. The two-stamen genera within the Melissinae, including Poliomintha, merge imperceptibly into one another and provide considerable difficulty in their definition. In addition, their morphological affinities with the four-stamen genera Melissa and Satureja open the question of unnaturalness within the subtribe and the taxonomic weight accorded stamen number. As pointed out by Epling & Stewart (1939) it is entirely possible that there has been extensive parallel- ism and convergence with the abortion of the upper stamen pair occurring repeatedly within the four-stamen groups. Indeed, one member of the Melissinae, Pogogyne, was revised to include both two- and four-stamen species (Howell, 1931). The present study, which stems from preparations for the Flora of Texas, attempts to examine closely the generic status and relationships of Polio- mintha as well as to bring needed insight into its morphology, distribution, and possibly phylogeny. It was originally hoped that the small number of species in the genus would permit breeding and chromosome studies. Unfortunately, repeated attempts at such studies over a number of years were successful with only one species, P. longiflora. HISTORICAL ACCOUNT The genus Poliomintha was created by Gray in 1870 to accommodate two species, P. longiflora and P. incana, the latter serving as the type. It should be noted, however, that informal recognition of the group came 11 years earlier with Torrey’s description of Hedeoma incanum in 1859. Regard- ing the disposition of his new species, he stated, ‘It may remain in Hedeoma for the present, but, if other species like it should be found, it may be the type of a new genus.’ Between 1870 and 1890 three species were added (P. bicolor, P. marifolia, and P. glabrescens). In 1897, with Briquet’s treat- ment of the Labiatae in Die Natiirlichen Pflanzenfamilien, Poliomintha was merged with Hedeoma, reflecting the inherent difficulties in the disposition of the taxa. With the work of Epling & Stewart (1939), generic status was again given Poliomintha; in addition, the sections Incanae and Saturejoides were informally established. Poliomintha conjunctrix was added in 1940, bringing the genus to its current number of four species. SIDA 5(1): 8—22. 1972. CHROMOSOME STUDY The chromosome numbers recorded to date from the subtribe Melissinae seem to reflect current concepts of monophyletic units; moreover, both number and morphology may be valuable criteria in segregating related genera (Irving, 1968). Unfortunately, only a few representative species of five of the 11 genera of Melissinae have been counted. The first count for Poliomintha, 2n = 36, was made from aceto-carmine squashes of seed root-tips of P. longiflora Gray: MEXICO. NUEVO LEON. Calcareous hills near Monterrey, Irving 698 (MONTU, TEX). The number suggests a separation of Poliomintha from several possibly related groups. For example, the single species counted for Hesperozygis, H. ee (2n = 44: Irving, unpublished), has been extremely difficult s it shows affinities to both Poliomintha and Hedeoma. The chromosome number 2n = 36 for Poliomintha tentatively suggests a separa- tion of Poliomintha and H. marifolia. The chromosome numbers of the four-stamen genera Melissa and Satureja are diverse (2n — 18, 20, 22, 30, 32, 42, 45, 48; Darlington & Wylie, 1955; Cave, 1965; Ornduff, 1967, 1968, 1969), but as yet 2n = 36 species have not been encountered. Needless to say, additional chromosome numbers are needed for all these groups before conclusive relationships can be ascer- tained. The chromosomes of P. longiflora, which average 3 microns in length, are large compared to those of Hedeoma or Hesperozygis (Irving, 1968). Moreover, they are primarily metacentric or submetacentric. With addi- tional work these size and structural differences may prove useful in dis- tinguishing and relating these taxa. GENERIC LIMITS AND RELATIONSHIPS On the basis of the present understanding of morphology, ecology, and distribution, Poliomintha is believed to be a natural group that is most likely monophyletic in origin. The question of its disposition remains. If it is argued that genera are to be recognized on the basis of clearcut and nonintergrading characters, then Poliomintha cannot be maintained as a distinct genus and should be merged with Hedeoma. With the application of this view, however, consistency in concept would demand that all 11 of the two-stamen genera of the Melissinae be brought together into one poly- morphic genus. The distinctness of the characters used to separate Polio- mintha from its closest ally, Hedeoma, are equal to those used to separate the remaining genera. This is approximately the rationale of Briquet (1897), who expanded Hedeoma to include most of these genera. This posi- tion, although logical, results in a taxon dissatisfying from several stand- points. Its resulting diversity of form would preclude meaningful generic characterization; indeed, there would remain but one character, the two stamens, to nd distinguish the group. Equally important, such a taxon would severely obscure many evolutionary trends; quite likely the resulting genus would be partly horizontal in aspect, cutting across several z. oo nN 10 phyletic lines. Fortunately, however, it is well understood that the course of evolution does not permit application of a priori rules in the disposition of taxonomic rank. Each group must evolve its own generic and_ specific criteria. Thus, I have used the combination of subtle characters which do distinguish Poliomintha to accord it generic status. Poliomintha is related to several sub-groups in the four-stamen genus Satureja, especially the shrubby section Guardoquia. The obvious difference between these groups is the number of stamens, but there are others. The canescent and shrubby habit and the symmetrical calyx can be found throughout the genus Satureja, but they occur together only in Poliomintha. The chromosome number of Poliomintha, may represent another possible difference Within the two-stamen when more completely known, groups, three genera need to be considered: Zizophora, Hesperozygis, and Hedeoma. For those seemingly more distantly related see Epling & Stewart (1939). Zizophora is a Mediterranean group whose morphological attributes are surprisingly similar to Poliomintha One of the latter’s most salient features is a radially symmetrical calyx whose subequal, deltoid teeth close the orifice at maturity. Another charac teristic is its canescent pubescence. All three of these features are found consistently in Zizophora. The slight differences include the abortion of the lower anther sacs and the non-shrubby habit of Zizophora. The continental separation of Poliomintha and Zizophora is, however, a clear-cut difference Unfortunately, the chromosome number(s) of Zizophora is unknown. Poliomintha and the primarily South American genus Hesperozygis are relatively distinct except for the somewhat ambiguous species H. marifolia (2n = 44) of Mexico. The latter possesses the habit of Poliomintha, a calyx intermediate between the two genera, and the calyx annulus of more typical Hesperozygis. The differences between Poliomintha and the re- mainder of the genus Hesperozygis are tabulated below. Hes perozygis Poliomintha Calyx either sore maae and cam- Calyx symmetrical, tubular. panulate or bilabia Calyx teeth subequal, erect or dis- Calyx teeth subequal, inwardly re- posed into upper and lower sets. flexed to close the orifice Calyx annulus well defined and Calyx annulus absent or irregular usually seated below the base of and seated at the base of the teeth. the teeth. Corolla tube hirtellous. Corolla tube annulate. Principally South American, Exclusively North American. 2n — 44 (one count). 2n = 36 (one count). The genera Hedeoma and Poliomintha are quite close and nearly ‘‘con- nected” by several transitional species: H. palmeri, H. molle, and H. mon- tanum. Although Epling & Stewart (1939) mention a few distinguishing 11 features, several more have been encountered which in combination sep- arate the two genera. These are tabulated below. Hedeoma Occasionally semishrubs, usually herbaceous perennials or annuals. Calyx tube gibbous or saccate. Calyx teeth usually acuminate and Poliomintha Shrubs. Calyx tubular and symmetrical. Calyx teeth subequal, convergent to close the orifice. or differentiated into upper and sets. lower Calyx annulus in a_ well-defined Calyx a absent or in an ir- ring. regular r Nutlets Po or orbicular in out- Nutlets eae over 1.5 mm long, 1.5m line, ca. 1. m long or less, usual- t becoming mucilaginous when ly becoming Saas when moistened. moistened 2n = 34, 36, 44, 72, 144. 2n = 36. PHYLOGENETIC CONSIDERATIONS To speculate on the intra- and inter-phyletic relationships of Poliomintha in the face of experimental ignorance is indeed tenuous. Yet as chromo- somal morphology, populational dynamics, and other clues into evolutionary behavior are likely to remain undetermined for some time, the presenta- tion of a tentative phyletic picture is, perhaps, not altogether out of place. Poliomintha appears to represent an older phylad whose populations have experienced dissection and extinction with evolutionary time habit, cylindrical, symmetrical calyx tube, undifferentiated calyx teeth, genera. Morphological transitions between taxa, often a salient feature of youthful and evolutionary active phylads, are conspicuously absent in Polio- mintha. Although interpopulational variation occurs, the specific criteria remain intact. The complete allopatrism of its species also hints at long evolutionary divergence. By comparison, the species of the related genus Hedeoma are often annuals with highly zygomorphic calyces and cleisto- gamy. Its species frequently intergrade one into another, and as many as five species may grow sympatrically. In addition, the frequent occurrence of aneuploidy reinforces the more derived status of Hedeoma. (Irving, 1968 Poliomintha has diverged along two lines, representing the subgeneric groupings Saturejoides and Poliomintha. Yet the two lines display interest- ing parallelisms following their divergence. Both reveal identical shifts in inflorescence, corolla morphology, habitat, and distribution accompanying speciation from the ‘‘primitive”’ types. Poliomintha longiflora (Saturejoides) ranges along the eastern periphery 12 of the Chihuahuan Desert from Monterrey southward to San Luis Potosi (Fig. 1). Its populations are found disjunctively on open exposed sites in the foothills flanking the Sierra Madre. It is most often associated with Juniperus monticola and Agave lecheguilla in an ecotone region between mesquite grassland and pine-oak forest. Morphologically it is well marked } S 1 Fig. 1. Distribution of Poliomintha longiflora var. longiflora (circles), P. longiflora var. congesta (squares), and P. glabrescens (triangles). 13 by solitary flowers and large red corollas. The allied species P. glabrescens is found to the north of P. longiflora in the Chihuahuan Desert and in low xeric habitats. Although P. glabrescens is very similar to P. longiflora it possesses an inflorescence of 6-flowered cymes and bears small white corollas. Poliomintha glabrescens thus may be viewed as derived. Section Poliomintha reveals a similar pattern with P. conjunctrix repre- senting the relatively ‘‘primitive’’ mesophytic element. Poliomintha con- northern Baja California (Fig. 2). It has a 3-flowered inflorescence with long red tubular corollas. In contrast, P. incana is a widespread, more northern species occupying extreme xerophytic sites in the lower Great Basin and western Chihuahuan Desert (Fig. 2). It possesses 6-flowered cymes and small white corollas. By superimposing knowledge of the distribution, ecology, and morphology of Poliomintha on the presently known events of the Tertiary, a chrono- logical evolutionary history can be formulated. Although this formulation is speculative, it provides the rationale for the existing relationships within the genus and hopefully the impetus for further study. As the events of the Tertiary have been reviewed in both botanical and geological literature they will not be reviewed here (Axelrod, 1950, 1958: Grant, 1959; Raven, 1962). By the close of the Oligocene, a large xeric floristic element, the Madro- tertiary flora, had developed between the tropical Neotertiary flora to the south and the temperate Arctotertiary flora to the north. As the primitive members of Poliomintha are today associated with the remnants of this flora, we can speculate that the Madrotertiary brought with it increased distribution and adaptive radiation in Poliomintha. Although it is possible that this period witnessed the origin of the group, it must be remembere that xeric sites, albeit limited in area, have been available to evolution since the Cretaceous (Axelrod, 1967). With the nee of the western mountain axes in e Pliocene and early Pleistocene, increased aridity resulted in a dissection of the Madrotertiary flora into a number of isolated woodlands and widespread desert regions (Axelrod, 1950, 1958). Popula- tions of Poliomintha perhaps at this time became contracted and frag- mented with widespread extinction. However, with the development of true deserts, an evolutionary impetus was created to which Poliomintha re- sponded, viz. P. glabrescens and P. incana. The role of the fluctuating en- vironments of the Pleistocene is difficult to assess. Undoubtedly, there were distributional shifts and populational fragmentations. SYSTEMATIC TREATMENT POLIOMINTHA A. Gray, Proc. Am. Acad. 8:295. 1870. EOMA sect. POLIOMINTHA Briq. in Engler & Prantl, Nat. Pflanzenf. IV. 3a. 298, 1897. Shrubs or semishrubs 1-3 m tall. Primary shoots numerous, narrowly Fig. 2. Distribution of Poliomintha conjunctrix (triangles) and P. incana (circles). or widely ascending, arising from a stout woody rootstock or caudex, puberulent, canescent, or tomentose with hairs spreading or antrorsely curling; branches slender and usually short, arising freely along the axes at wide or narrow angles. Leaves membranous, well spaced or crowded (internodes short), cernuous or spreading, linear, oblong, elliptical, ovate, or oval, 6-21 mm long, 1-6 mm wide, entire or serrulate, attenuated or 15 shortly petiolate, acute or obtuse; upper surfaces glabrate (Saturejoides) or tomentose (Poliomintha), lower surface strigillose or tomentose, hairs directed toward the apex; nervation inconspicuous, secondaries when prominent consisting of 2-3 alternating, straight pairs; both upper and lower surfaces uniformly glandular-punctate. Flowers solitary in the upper leaf axils or aggregated in 3—7-flowered cymes (cymules) in the upper 1/2-1/3 of the shoots; peduncles inconspicious or up to 3 mm long, pedicels 1-4 mm long, both peduncles and pedicels typically cinereous with retrorse- ly directed hairs; bracteoles (diminutive leaves subtending the pedicels) triangulate or subulate, shorter or longer than the pedicels. Calyx actino- morphic or nearly so, 4-12 mm long, 13—15-ribbed; the tube 3-9 mm long, cylindrical or pentagonal, the walls straight and not outwardly distended, cinereous or pilose, the antrorse or spreading hairs distributed uniformly or restricted to the ribs; calyx teeth 5, subequal or separable into an upper set of 3 and a lower of 2, variously pubescent on the inner face and con- vergent at some stage to close the calyx orifice; the upper teeth free or connate for 1/2 of their length, forming a small upper lip (P. conjunc- trix), triangulate, 1-3 mm long, 0.5—1 mm wide, equal to or shorter than the lower teeth; lower teeth never connate, narrowly or broadly triangu- late, 1-3 mm long, 0.5—1 mm wide at the base; pubescence within the tube either cinereous and restricted to the teeth faces (Saturejoides) or pilose and in an irregular ring or annulus seated at the base of the teeth (Poliomintha). Corolla strongly zygomorphic, with a pronounced tube (within the calyx) and throat (above the calyx), the limbs forming a con- spicuous subgaleate upper lip and a 3-lobed spreading lower lip, lavender, orange-red or red, 10-35 mm long, pubescent on the outer surface with spreading hairs, densely pubescent within at the juncture of the tube and throat to form an annulus 2-4 mm wide; the tube 4-12 mm long; the throat 4-16 mm long, tubular, arching upwardly, dilating dorsiventrally, occasion- ally iterally: distended (P. incana); the upper lip ligulate, emarginate, and straight, 3-7 mm long, 2.5—5 mm wide at the base; the lower lip spreading, the lobes equal or subequal, the central lobe again bilobed and exserted beyond the laterals, 3.5—8 mm long, 3.5—10 mm wide. Fertile stamens 2, seated on the abaxial surface and arched upwardly under the upper lip; filaments glabrous, anther sacs widely divergent on well-de- veloped connectives, usually well exserted at anthesis; sterile stamens 2, reduced to subulate, glabrous filaments 1-2 mm long, seated at about the mid-point of the tube. The style long, exserted, its apex arching down- wardly and unequally bifid, the lower division twice the length of the upper. Nutlets oblong in outline, oval or ovate in cross section, light to dark brown, 1-2 mm long, 0.5—1 mm wide, the surface smooth but glandu- lar at the apex, not becoming mucilaginous when moistened. Chromosome number, 2n = 36. Distribution (Fig. 1 & 2): Northern Mexico and southwestern United States. Center of distribution appears to be in the northern Sierra Madre 16 Mountains of Mexico. Type species: Poliomintha incana (Torr.) Gray KEY TO THE TAXA OF POLIOMINTHA Leaves and stems canescent to glabrate; leaves oval or elliptical: calyx 5-9 mm long, cinereous, 13-ribbed; calyx annulus absent (Sect. Saturejoides). 2. Flowers solitary in the axils; corolla orange-red, ca. 25 mm long. Leaves typically elliptical and well spaced; calyx teeth clearly converging at maturity; plants of Mexico. —_ 0.5 2 2 oe we oS P: inno var lenge seaves oval or rhombic, very crowded; calyx teeth slightly con- verging at maturity; plants of Haiti. . lb P. longiflora var congesta pay) la. PC} we — . Flowers in axillary, 6-flowered cymes; corolla white-lavender, ca. 15 mm long. ek ee eS Bm 2. P. glabrescens Leaves and stems lenenieee: leaves linear-oblong; ie 3-4 mm long pilose, often 15-ribbed, calyx annulus present (Sect. Poliomintha). 4. Corolla tubular, orange-red, ca. 25 mm _ long; endemic to Baja —_ California. an en a Gees 3. P. conjunctrix 4. Corolla nics inflated, lavender, up to 15 mm long; widespread throughout the southwest. . 2. . 2... 2 ee . 4. P. incana POLIOMINTHA sect. Saturejoides Irving, sect nov. Frutices canescentes; folia ovalia vel elliptica; calyx 5.0-12.0 mm. lacs 13-costatus, annulo nullo. Type species Poliomintha longiflora Gra Canescent shrubs; leaves oval to elliptical; | 5-12 mm long, 18-ribbed, without an annulus. la. POLIOMINTHA LONGIFLORA Gray var. LONGIFLORA, Proc. Am Acad. 8: 296. 1870. Holotype GH!: Mexico. COAHUILA. ‘‘Saltillo, moun- tains,”’ 30 July 1848, Gregg 313. Isotype K! Poliomintha bicolor Wats., Proce. Am. Acad. 25: 160. 1890. Sete pa UEVO LEON. ‘Summit ledges, ae x la Silla,’ 5 1889, Watson 2536. Isotypes F!, NY!, PH!, Us Pande see (Gray) Briq. in Engler & Prantl, Nat. Pflanzenf. IV . 294, 1897. Based on P. longiflora Gray. Hedeoma bicolor (Wats.) aR Ann. Conserv. Jard. Bot. Genéve 2: 185. 1898. Based on P. bicolor Wats Poliomintha greggii Gray in Watson, Proc. Am. Acad. 25: 160. 1890 (nom. nud.) Hedeoma greggii (Gray) Brig., Ann. Conserv. Jard. Bot. Geneve 2: 185. 1898 (nom. nud.). Robust shrubs or semi-shrubs 13-35 em high. Shoots slender, ascending or somewhat decumbent, puberulent or glabrous below, pubescent or canescent above, hairs spreading, antrorsely curling, or appressed; branches slender, 17 brittle, and diverging at close angles, Leaves well spaced, spreading, or cernuous, oval to broadly or narrowly elliptical, 7-15 mm (11.3 mm) long, 2.5—6 mm (3.9 mm) wide, entire, attenuate and shortly petiolate, the petiole 1-2 mm long, apex obtuse, surfaces pubescent to tomentose below, glabrate above; nervation inconspicuous, the secondaries consisting of 2—3 alternate, straight pairs. Flowers solitary in the axils of the upper leaves, rarely in 2-flowered axillary cymes; peduncles prominent, 1.5—2 mm long; pedicels 1.5—4 mm long, cinereous with short antrorsely directed hairs; bracteoles lanceolate, exceeding the pedicels they subtend. Calyx 7-12 mm (9 mm) long, 13-ribbed; the tube 5.5—9 mm (6.9 mm) long, pen- tagonal, the sides straight and paralle! below, becoming constricted above, evenly cinereous with appressed hairs; calyx teeth only scarcely differen- tiated into upper and lower sets, converging to close the orifice, densely and evenly pubescent within; the upper teeth very slightly connate (less than 1/2 their length), the lobes triangulate, 1.5—3 mm long, ca. 0.7 mm wide at the base; the lower teeth triangulate, 1.5—3 mm long, ca. 1 mm wide at the base; annulus absent, replaced by pubescence on the inner face of the teeth. Corolla orange-red, 27-35 mm (32 mm) long, pubescent within at the juncture of the tube and throat, the trichomes forming an annulus ca. 4 mm wide; the tube 7-12 mm long: the throat 15—16 mm long, expanded above, ca. 5 mm wide dorsiventrally; the upper lip broadly ligu- late, emarginate, 5-7 mm long, 3.5—5 mm wide, straight (not upwardly reflexed), and flat (not concave); the lower lip 5-8 mm long, 6-10 mm wide, the lobes subequal with the central lobe slightly exserted beyond the later- in o a, 2mm long, 1 mm wide, the surface smooth but glandular distally, ih number, 2n Distribution (Fig. 1). Open exposed sites in the foothills of the Sierra adre Oriental and outlying ranges in Mexico; Monterrey, N.L., southwest to Catorce, S.L.P., 5,000-7,000 ft. June—September. Despite the poor representation of P. longiflora in herbaria it is quite common throughout its range. On the somewhat inaccessible ridge tops and mesas where it is found it builds extensive populations. One of the most salient features of this species is its production of carvacol as the major constituent of its essential oil (Irving, unpublished). It is the same com- pound which gives the spice oregano (Origanum spp.) its distinctive odor and flavor. Because of this and the large size of its populations P. longiflora is collected in large ae for sale locally or export to the United States as ‘““Mexican oregan Representative specimens: MEXICO. COAHUILA: 6 mi SE of Saltillo, 21-23 Sep 1904, Palmer 417 (GH, UC, US); 4 km E of Fraile, 11 Jul 1941, Standford, Retherford & Northcraft 308 (DS, GH, UC); 3 mi SW of Saltillo, 3 Aug 1966, Irving 703 (TEX); NUEVO LEON: Sierra de la Silla, 5 Jun 1889, Pringle 2536 (F, GH, K, NY, PH, UC, US): SAN LUIS POTOSI: 12 km al W de Estacion Berrendo, municipio de Charcas, 11 Sep 1955, Rzedow- ski 662 (GH, MICH, US); ca 30 mi W of Matchuala at Catorce, 1 Aug 18 1966, Irving 695 (TEX); ZACATECAS: Aranzazu, 19 Jul 1934, Pennell 17457 (MICH, PH); 2 mi above Maypil (W of conception del Oro), 2 Aug 1966, Irving 698 (TEX). lb. POLIOMINTHA LONGIFLORA Gray var. congesta Irving var. nov. Holotype MICH!: HAITI. Morne Vincent, M. des Commissaires, 14 Dee 1944, Holdridge 2060. Frutices glabrati; folia congesta, ovata, ovalia, vel rhombica, 6.0—9.5 (8.5) mm longa, 4.0—5.0 (4.7) mm_ lata, marginibus serrulata, basi decurrentia; calyx 9.0—10.0 mm longus, dentibus erectis; corolla rosea ca. 20.0 mm longa, annulata. Robust shrubs up to 1 m in height. Shoots woody, ascending, with numer- ous short branches, glabrous and woody below, tomentose in the upper herba- ccous stem regions. Leaves crowded, spreading, with margins becoming revolute on drying, ovate or rhombic, 6-9.5 mm (8.5 mm) long, 4-5 mm (4.7 mm) wide, serrulate, base attenuate, apex acute, petiole ca. 1 mm long, decurrent; both surfaces felty tomentose, nervation inconspicuous, the secondaries consisting of 2—3 subopposite, curved pairs. Flowers soli- tary in the uppermost leaf axils; peduncles ca. 0.7 mm long; pedicels ca. 3 mm long, cinereous; bracteoles linear-subulate, ca. 2.55 mm long. Calyx 9-10 mm long, 13-ribbed; the tube ca. 7 mm long, cylindrical but expanding upwardly, cinereous, the hairs antrorsely appressed, restricted to the nerves: calyx teeth slightly differentiated into upper and lower sets, pubes- cent on the inner face; the upper teeth connate for slightly less than 1/2 of their length, forming an upper lip, the lobes triangulate, ca. 1 mm long and 0.7 mm wide at the base, erect; lower teeth shorter than the upper, triangulate, ca. 1.5 mm long, ca. 0.5 mm wide at the base, erect or slightly converging with the upper; annulus absent. Corolla pink, ca. 20 mm long, densely pubescent in the tube, the trichomes forming an annulus ca. 7 mm wide: the tube ca. 9 mm long; the throat ca. 9 mm long, curved and ex- panded above, ca. 4 mm wide dorsiventrally; the upper lip broadly ligulate and cleft, ca. 2 mm long, 3 mm wide, straight and flat; the lower lip ca. 3 mm long, 5 mm wide. Nutlets not seen. Chromosome number, un- known. Distribution (Fig. 2). Known only from the type locality. Details of its habitat are absent from the collection data. 9 POLIOMINTHA GLABRESCENS Gray, in Hemsl. Biol. Centr. Am. Bot. 9: 549. 1882. Holotype K!: MEXICO. COAHUILA. ‘Soledad, 25 mi SW of Monclova,”’ Feb-Oct 1880, Palmer 1056. Isotypes F!, PH!, US! Hedeoma glabrescens (Gray) Briq. in Engler & Prantl, Nat. Pflanzenf. IV. 3a. 294. 1897 9» Semi-shrubs up to 50 em tall. Shoots slender, sharply ascending, arising from a stout woody rootstock, puberulent throughout with minutely spread- ing hairs, occasionally densely so; branches brittle, slender, and diverging at close angles. Leaves cernuous, oblong to elliptical, 7-16 (12 mm) long, 3-4 mm (3.6 mm) wide, entire, subsessile, apex acute or obtuse; strigillose below, glabrate above; nervation inconspicuous, the secondaries consist- ing of 2—8 indistinct, alternate pairs. Flowers in 5—7-flowered axillary 19 cymules along the upper 1/2 of the shoots; peduncles conspicuous, ca. 1 mm long; pedicels ca. 1 mm long, canescent; bracteoles linear-lanceolate, longer than the pedicels they subtend. Calyx ca. 5 mm long, 13-ribbed; the tube 3.8 mm long, tubular or slightly pentagonal, evenly canescent with short appressed hairs; the calyx teeth scarcely differentiated into upper and lower sets, densely tomentose on their inner borders and apex, tri- angulate; the upper ca. 1.2 mm long, ca. 0.5 mm wide at the base; the lower ca. 1.2 mm long, and 0.7 mm wide at the base; all 5 teeth tightly convergent to close the calyx orifice; annulus absent. Corolla lavende ca. 13 mm long, with a well-defined annulus at the juncture of the tube and throat, ca. 1.5 mm wide; the tube ca. 5 mm long; the throat ca. 5 mm long, dilated above, ca. 4 mm wide dorsiventrally; the upper lip ligulate, emarginate, ca. 4 mm long, ca. 3 mm wide, straight and somewhat con- cave; the lower lip ca. 4 mm long, ca. 5 mm wide, the lobes equal or subequal with the central lobe slightly exserted beyond the laterals. Nut- lets oblong, ca. 1.5 mm long, ca. 0.7 mm ones the surface smooth, glandu- lar at the apex. Chromosome number, unknow Distribution (Fig. 1). Dry calcareous ne in the Chihuahuan Desert; Texas to Coahuila, Mexico. 4,000—6,000 ft. August—September. Representative specimens: MEXICO. COAHUILA: Castanos, Puerta de San Lazaro, Sierra de San Lazaro, 30 Aug 1939, Muller 3038 (GH, MICH, UC); Cuatro Cienegas, Sierra de la Madera, Canon del Aqua, 10 sep 1939, Muller 3255 (GH, MICH, UC); UNITED STATES. TEXAS: BREWSTER CO., Mc- Rae Canyon on Cox Ranch, 31 Jul 1953, Warnock 11397 (LL); PRESIDIO CO., Solitario Peak area, 30 Jul 1957, Warnock 15710 (NY, T ) POLIOMINTHA sect. POLIOMINTHA. Type species Poliomintha incana (Torr.) Gray. Tomentose shrubs; leaves linear-oblong; calyx 4.0—7.0 mm long, cinereous or villous, 13—15-ribbed, annulate 3. POLIOMINTHA CONJUNCTRIX Epling & Wiggins, Contrib. Dudley Herb. 3:85. 1940. Holotype DS!: MEXICO. BAJA CALIFORNIA. “5 mi SE of Rancho San Antonio,’ Buenos Aires Plains, 20 Jun 1937, George Lindsay, S.n. (accession no. 259214). Isotype F! Tall slender shrubs over 1 m in height. Shoots widely ascending, arising from a woody caudex, tomentose with a felty tomentum of retrorsely ap- pressed hairs; branches short, brittle, diverging at wide angles. Leaves cinereous, cernuous, linear—oblong, 7-17 mm (13 mm) long, 2.5—3.5 mm (2.6 mm) wide, entire, attenuated and sessile, apex narrowly or broadly obtuse; surfaces tomentose, the lower densely so. Flowers in 3-flowered axillary cymules borne in the upper 1/3 of the shoots; peduncles 2-3 mm long; pedicels 1.5—2 mm long, tomentose with antrorsely appressed hairs: bracteoles subulate, shorter than the pedicels they subtend. Calyx 6-7 mm (6.5 mm) long, 13-ribbed; the tube 5.5—6.5 mm (6 mm) long, symmetrical and tubular, cinereous with short antrorsely curling hairs restricted to the nerves; calyx teeth subequal, slightly differentiated into upper and lower 20 sets, erect or slightly converging at maturity; the lower set slightly shorter than the upper, connate for ca. 1/2 its length, the lobes triangulate, 0.5 mm long and ca. 0.5 mm wide at the base; the upper teeth narrowly triangulate, ca. 1 mm long, ca. 0.7 mm wide at the base; annulus an ir- regular dense ring of long hairs (pilose) seated at the base of the teeth and extending upward onto the inner tooth faces, slightly exserted. Corolla orange-red, 20-21 mm long, densely pubescent at the juncture of the tube and throat, the trichomes forming an annulus ca. 3.5 mm _ wide; the tube 6-7 mm long; the throat ca. 11 mm long, expanded above, ca. 4 mm wide dorsiventrally; the upper lip ligulate, emarginate, ca. 7 mm long and 3 mm wide: the lower lip ca. 5 mm long, ca. 5 mm wide, the lobes sub- equal. Nutlets tan, oblong in outline, ca. 2 mm long, ca. 1 mm wide, the surface smooth or weakly areolate, glandular at the apex. Chromosome number, unknown. Distribution (Fig. 2). Known only from the type locality. 4. POLIOMINTHA INCANA (Torr.) Gray, Proc. Am. Acad. 8: 296. 1870. Holotype NY!: TEXAS. EL PASO CO., “near El Paso,” Parry, Wright, Bigelow (Wright 1523). Isotypes, GH!, K!, PH!, US! Hedeoma incanum Torr., U.S. Mex. Bound. Surv. 130. 1859. Robust canescent shrubs to 3 m high. Shoots slender, ascending, bearing eray-fissured bark below, tomentose with short, retrorsely appressed hairs ~— above: branches slender and arising at close angles. Leaves gray-green, spreading or cernuous, linear-oblong or occasionally obovate below, 11-21 mm (15 mm) long, 1.3—3 mm (2.1 mm) wide, entire, attenuated and ses- sile, apex acute or obtuse; surfaces equally tomentose or tomentulose; nervation indistinct. Flowers typically in 3-flowered axillary cymules borne in the upper 1/2 of the shoots; peduncles ca. 1 mm long; pedicels ca. 1.1 mm long, tomentose; bracteoles linear, longer than the pedicels they sub- tend. Calyx 4-5 mm long, 13—15-ribbed; the tube 3-4 mm long, cylindrical, the sides parallel, moderately to densely pilose with hairs up to 2 mm long strigose beneath the pilose coat, calyx teeth subequal, only scarcely dif. ferentiated into upper and lower sets, lower slightly longer than upper, narrowly triangulate, 1-2 mm _ long, ca. 1 mm wide at the base; teeth converging to close the calyx orifice; annulus an unequal dense ring of exserted long hairs (pilose) situated at the base of the teeth. Corolla laven- der, 10-14 mm long, pubescent at the base of the tube and throat, the trichomes forming an annulus; the tube 8-9 mm long; the throat both later- ally and dorsiventrally inflated (3-5 mm wide laterally and dorsiventrally); the upper lip ligulate, emarginate, 3.5—4 mm long, ca. 2.5 mm wide, straight and slightly concave; the lower lip ca. 3.5 mm long, 3.5—4 mm wide, the lobes unequal, the central lobe bilobed and exserted beyond the laterals. Nutlets oblong, ca. 2 mm long, ca. 1 mm wide, the surface smooth and glandular. Chromosome number, unknown. Distribution (Fig. 2). Sand hills frequently of gypsum origin; Mexico, 21 Texas, New Mexico, Arizona, Utah, Colorado, and California. 2,000—4,000 ft. April—October. Poliomintha incana is the most widespread member of the genus, although disjunctively so, and forms extensive populations where it occurs. It is apparently well adapted to the dune-sand habitat and is able to withstand repeated burial. Representative specimens: MEXICO. CHIHUAHUA: near El Paso del Norte, 9 Apr 1885, Pringle 275 (F, GH, PH, POM, US); Samalayuca, 28 Jan 1903, Coville 1700 (US); 5 mi S of Samalayuca, 19 Apr 1960, Cruch- field & Johnston 5265 (MICH, TEX): UNITED STATES, ARIZONA: APA- CHE CO., Holbrook, 20 Aug 1883, Rusby 784 (MICH, PH, UC, US); COCO- NINO CO., 10 mi W of Navajo Bridge, 24 May 1939, Barkley & Reed 4370 (DS, PH, POM, UC); NAVAJO CO., Monument Valley, 14 Sep 1938, East- wood & Howell 6656 (CAS, US); CALIFORNIA: SAN BERNARDINO CO., Cushenbury Springs, 12 Jul 1938, Jaeger s.n. (POM): COLORADO: MON- TEZUMA CO., E of Aneth, Utah, 19 Jun 1968, H. D. Harrington 10107 (CS); NEW MEXICO: DONA ANA CO., White Sands, 19 Jun 1899, Wooton s.n. (DS, MICH, NMC, POM, TEX, UC, US); OTERO CO., 13 mi W of Tularosa, > Jun 1938, Cutler 2016 (CAS, GH, UC); SAN JUAN CO., S of Bloomfield, 9 Jul 1929, Mathias 615 (GH, POM); TEXAS: CULBERSON CO., 29 mi W of Orla, 11 Jul 1965, Irving 613 (MONTU, TEX): EL PASO CO., 2 mi NE of Fabens, 5 Jul 1958, Correll & Johnston 19276 (LL); HUDSPETH CO., 3 mi E of McNary, 18 Apr 1930, Cory 3445 (POM); UTAH: EMERY CO., 20 mi SW of Greenriver, 3 Jun 1953, McVaugh 14421 (CAS, MICH, TEX); GARFIELD CO., 16 mi N of Hite & 35 mi S of Hanksville, 17 May 1961, Cronquist 9055 (DS, MICH, TEX, UC); GRAND CO., ca 1 mi N of Moab, 20 May 1957, Vickery, Jr. 801 (DS, GH, UC); SAN JUAN CO., 5 mi N of Mexican Hat, 24 Jul 1939, Cutler 2779 (DS, GH); WAYNE CO., 15 mi S of Hanksville, 16 May 1950, Harrison 11532 (UC). EXCLUDED NAMES Poliomintha marifolia (Briq.) Gray, Proc. Am. Acad. 8: 365, 1872. = Hesperozygis marifolia (Briq.) Epling in Rep. Spec. Nov. Beiheft 115: 13. 1939 Poliomintha mollis (Torr.) Gray, Proc. Am. Acad. 8: 365. 1872. = Hedeoma molle Torr. in Bot. U.S. & Mex. Boundary Surv. 2(1): 129. 1859. ACKNOWLEDGEMENTS Acknowledgement is made to all persons responsible for the loan of specimens. Abbreviations of herbaria from which specimens were borrowed are those standardized by Lanjouw & Stafleu (1964). Thanks are also given to Drs. D. Bilderback and C. Miller for their help with the manuscript. REFERENCES AXELROD, D. I. 1950. Perec ee = desert vegetation in western North America. Car- negie Inst. Ba sh. Publ. 215 i OS eee. of che W N hes Tertiary Geoflora. Bot. Rev. 24: 433-509 F var a 201- 209, EVs i ought, dia: ali and quantum evolution BRIQUE r, ,y 1897. Labiatae, in Engler & Prantl, Nat. Pflanzenf. TV. 3a: 183- . (ed.) 6 dex to plant chromosome numbers. Uniy. North Cupola Press. and A. P. WYLIE. 1955, Chromosome atlas of flowering plants mes) as Allen “iaal Unwin, Lonc EPLING, C. aed W. S. STE WART. 1939. A revision ¥ + a with Repert. Spec. Nov. Regni Veg. Beiheft 115: GRANT, V. 1959. Natural history of the phlox Foe The Hague. HOWELL, J. T. 1931. The genus Pogogyne. Proc. Calif. Acad. Sci, 20: 105-128. ee R. S. 1968 The systematics of Hedeoma, Ph.D. Thesis. University of Texas, a review of allied tin. oe J. and P. A. STAFLEU. 1964. Index herbariorum, 31. Se Rar R. (ed.) 1967. Index to plant chromosome numbers for 1965 sth ed. Regnum Vegetabile . Regnum Vege- tabile : 1-128. = _. 1968. Index to plant chromosome numbers for 1966, Regnum Vegetabile 55: 1969. Index to plant chromosome numbers for 1967. Regnum Vegetabile 59: 1-129. RAVEN, P. H. 1962. The systematics of Oenothera subgenus Chylismia. Univ. Calif. Publ. Bot. 34: 1-121. TORREY, J. 1859. Bot. of the Boundary. U.S. & Mex. Boundary Sury, 2(1): 130. TWO NEW SPECIES OF ISOCOMA (COMPOSITAE-ASTEREAE) FROM NORTH CENTRAL MEXICO B. L. TURNER Department of Botany, The University of Texas, Austin, 78712 ISOCOMA halophytica Turner sp. nov. Frutex ad 1 m altus; caudex distincte lignosus fide collectorum; caules fide collectorum decumbentes fragiles dense pubentes pilis brevibus ubi juvenes demum glabrescentes. Folia integra uniformiter lineario-oblanceo- lata dense pubentia pilis brevibus plerumque 1.5—2.5 cm longa 1.5—2.0 (—3.0) mm lata apice mucronibus albis. Caules secundarii 10—25 cm longi 5—8-capitulati capitulis dense confertis 15—18-floris. Involucra subturbinata 4—5 mm alta 4—5 mm lata 2—3-seriata. Exteriora phyllaria dense pubentia. Interiora phyllaria scariosa apicibus latis fuscatis ciliatis. Receptaculum planum alveolatum radiati flores nulli. Disci flores flavi corollis 7—8 mm longis tubulis 4—5 mm longis faucibus distincte obliquiis 2.0—2.5 mm longis lobulis acutis 1.0—1.5 mm longis. Styli rami ca 1 mm longi appendiculibus dense pubentibus acutis. Achenia ca 2 mm longa dense pubentia pilis antrorsis plerumque adpressis. Pappus 50—60-setosus setis 3—5 mm longis. Shrub up to 3 ft tall, the stems (according to label data) decumbent, arising from a decidedly woody rootstock. Stems brittle, at first densely short pubescent, becoming glabrate with age. Leaves entire, uniformly linear-oblanceolate, densely short pubescent, mostly 1.5—2.5 cm_ long, 1.5—2.0(3.0) mm wide, terminated by a small, white mucro. Heads 5—8, densely clustered at the apexes of secondary stems, the latter 10—25 cm long. Involucres subturbinate, 4—5 mm _ high, 4—5 mm broad. Bracts imbricated in about 2—3 lengths, the outer row densely pubescent, the inner scarious with broad, darkened, ciliate apices. Receptacle flat, alveo- late. Ray flowers absent. Disc florets yellow, 15—18; corollas 7—8 mm long, tube 4—5 mm long, throat decidedly oblique, 2.0—2.5 mm long, the lobes 1.0—1.5 mm long, acute. Style branches about 1 mm long, the appendage densely pubescent, acute. Achenes ca 2 mm long, densely pubescent with mostly appressed, ascending hairs. Pappus of ca 50—60 bristles, 3—d) mm long. MEXICO. Coahuila: Along road from San Vicente SW (about 7 mi) to southern end of Laguna de Jaco. ‘“‘Rocky saline soil at south end of lake, stems decumbent, up to 3 ft tall.’”’ 9 Sep 1940. I. M. Johnston & C. H. Muller 1090. (Holotype, MICH). Isocoma halophytica has non-fasciculate, entire leaves, deeply cleft corollas and, except for the former, would most readily key to I. pluriflora (= T. wrightii) in Hall’s (1928) treatment of the genus. However, it is SIDA 5(1): 23—25. 1972. 24 readily distinguished from that species by its densely pubescent, gray foliage, apparently sprawling habit, and short, broad involucres. The taxon is known only from the locality concerned where, I suspect, it occurs i eypseous-saline soils, 1 ISOCOMA gypsophilia Turner sp. nov. Herbae perennes 5—10 cm altae caulibus fohisque recumbentibus saepe eleganter tegetiformes 10—40 cm diametro laxae vel crebrae succulentae. Caules sparse vel dense pubentes pilis ascendentibus crispatis albis. Folia principalia 0.5—1.0 cm longa spathulata vel oblanceolata conspicuo glandu- lo-punctata sparse pubentia pilis ascendentibus crispatis dense conferta imbricata caulem occultantia plerumque 3—4-dentata apicem versus denti- bus distincte albomucronatis. Rami secundarii apice pedunculiformes uni- flori rare biflori 2—10 mm longi. Capitula 30—40-flora. Involucra late tur- binata vel hemisphaerica 7—9 mm alta 6—9 mm lata 4—5-seriata. Phyllaria ordinate imbricata viridia conspicuo glandulo-punctata ciliata apicem ver- sus. Receptaculum planum = conspicuo setaceo-alveolatum. Radiati flores nulli. Disci flores flavi corollis ca 5—6 mm longis tubulis 3—4 mm longis faucibus 2 mm longis lobulis ca 0.5 mm longis. Styli rami ca 1 mm longi appendiculibus ovatis dense papillatis. Achenia ca 2 mm longa dense pu- bentia pilis adpressis argenteis. Pappus 30—40-setosus setis 3—5 mm longis. Chromosomatum numerus n = 6. Perennial herb 5—10 cm tall, the stems and foliage more or less re- cumbent, often forming an attractive, loose to compact, succulent, dark green mat, 10 to 40 cm across. Stems sparsely to densely pubescent with crisp, white hairs; principal leaves, 0.5—1.0 em long, spatulate to oblanceo- late, densely crowded and overlapping so as to obscure the stem, mostly 3—4 dentate near the apex, each of the denticulations terminated by a distinct, white mucro, the blades conspicuously glandular-punctate and sparsely covered with a loose, crispy, crinkled pubescence. Heads single (or rarely double) on peduncles 2—10 mm long at the apices of secondary branches. Involucres broadly turbinate to hemispheric, 7—9 mm high, 6—9 mm broad. Bracts regularly imbricated in about 4-5 lengths, green and conspicuously glandular-punctate above, the margins ciliate, especially near the apices. Receptacle flat, conspicuously bristly alveolate. Ray flowers absent. Disc florets yellow, 30—40: corollas ca 5—6 mm long, tube 3—4 mm long, throat 2 mm long, the lobes ca 0.5 mm long: style branches about 1 mm long, the ovate appendages densely papillate. Achenes ca 2 mm long, densely pubescent with silvery, appressed hairs: pappus of 30 to 40 setae, 3—5 mm long. Chromosome number, n — MEXICO. Nuevo Leon: 15 mi S of San Roberto Junction. Growing on white, gypseous soils on what appears to be saline flats along the roadside. 26 Sep 1970 B. L. Turner 6213. (Holotype, TEX; isotypes to be distributed). The taxon is apparently most closely related to Isocoma veneta (HBK) Greene, a widespread species of central and northern Mexico where it occurs in mostly rocky limestone soils, or less frequently in mixed gypseous, 29 alluvial soils, as noted below. On the former soils it apparently assumes a more robust form; the latter sites characteristically producing a low, the vicinity of the type locality and both have diploid counts of n = 6 (Turner & Crutchfield 6313, robust form; Turner & Crutchfield 6289, Sinai form) as does I. gypsophila. The latter, however, grows only in gypsum, forming open stands with such endemic gypsophilous species as Frankenia gypsophila, Sartwellia mexicana, Atriplex reptans and a remarkable, as yet unpublished, new genus of the Compositae, tribe Tagetinae (Turner, in press Isocoma gypsophila is readily recognized from all other species of the genus by its habit; small, overlapping leaves; predominantly single-headed flowering stems; and broad involucres possessing conspicuously punctate involucral bracts. The species was found growing with or near I. veneta (Turner & Crutchfield 6303) which, while relatively small at this locality, has a decidedly woody, 3—5 headed, glutinous stem and subturbinate in- volucres, the latter composed of only faintly punctate, eciliate involucral bracts. Intermediates between the two taxa were not seen in the field, in spite of a search for these in a second visit to the site. (In my initial visit I was unaware that the material here described as I. gypsophila might even be a species of Isocoma!) The holotype consists of 5 plants. On an additional sheet I have included a plant of I. veneta (found along the roadside at the type locality) along with isotypic material of I. gypsophilia. I follow Shinners (1950), M. C. Johnston (1967) and perhaps others in recognizing the natural group Iso- coma as a genus, believing Hall's treatment of the taxon as part of Haplopappus to be highly artificial, in spite of his claims to the contrary. I am grateful to Dr. M. C. Johnston for the Latin descriptions. Ficld work was supported, in part, by NSF grant 5548X REFERENCES HALL, H. M. 1928. The genus Hab opanpes. Carnegie Pas Wash. Public. JOHNSTON, M. C. 1967. Ericameria anstrotexana M. Johnston iGompence’, NOMEN novum. Southwestern Nat. 12: ee ar L. H. 1950. Notes on Texas Compositae. TV. Field and Lab. 18: 25-32. FIELD OBSERVATIONS OF CUTHBERTIA (COMMELINACEAE) WITH DESCRIPTION OF A NEW FORM OLGA LAKELA! Herbarium, University of South Florida, Tampa, Florida, 33620 The genus Cuthbertia Small includes three closely related species that are endemic to the South Atlantic Coastal Plain. The combination Cuth- bertia rosea (Vent.) Small (1903) superseded Tradescantia rosea Vent. (Jard. Cels. pl. 24, 1800). In addition, Small described C. graminea and C. ornata as new species (1933). Later Anderson and Woodson (1935) re- duced Cuthbertia to synonomy under Tradescantia and made Small’s three species into varieties, thus Tradescantia rosea var. rosea, T. rosea var. graminea (Small) Anders. & Woods., and T. rosea var. ornata Anders. & Woods. The authors apparently did not recognize the morphological dis- tinctions of the involucral bracts that Small used to seperate the two genera. Woodson, after a comprehensive study of American and tropical genera of Commelinaceae, chose to emphasize the structure of the inflorescence in the delimitation of genera. Reevaluation of his earlier revision led him to transfer Tradescantia rosea var. rosea to Tripogandra rosea (Vent.) Woods. However, he did not make any transfers involving T. rosea var. graminea or T. rosea var. ornata because he was not certain of their proper taxonomic status. Woodson stated that none could be made. until “extensive field study’? had been completed. After several years of observation of the plants, both in the field and in cultivation in the botanical garden, I am convineed that Dr. Small’s delimitation of Cuthbertia is valid. Other workers (Giles, 1942, 1943: Tom- linson, 1966) concur with this position, using other lines of evidence to reach the same conclusion. The following taxonomic diagnosis is offered based on my field and garden observations. CUTHBERTIA Perennial herbs in small or large tufts. Stems and leaves glossy green or yellowish green, nodose, never hirsute, stems erect to spreading. Leaves fleshy, semiterete, or flat, folded when young, with a closed sheathing base; stomata abaxial; sheaths red-striate, or sometimes whitish-striate, with oblique orifice. Roots fibrous, lanate or glabrate, clustered or spread- ing and fistulose (even at extremities). Flowers ephemeral, bisexual, trimerous, radially symmetrical; sepals green or petaloid; petals free, roseate, or rarely white, crenulate on mar- gins; stamens six in two alternating cycles of three’s, all fertile and morpho- logically similar; anther connective trapezoidal; pollen yellow; stamen ' Contribution 61, Botanical Laboratories of University of South Florida. SIDA 5(1): 26—32. 1972 27 hairs roseate, rarely white, moniliform, in groups on the lower part of the filament, the upper part hairless; ovary superior, 3-locular, typically 6- ovulate; pistil white; style declinate, tubular, dilating to tubular stigma. Capsule loculicidal, valves coriaceous, persistent; seeds hemispherical, ribbed-reticulate, hilum punctiform. (Tradescantia L. of authors; Tripo- gandra Raf. of authors.) KEY TO SPECIES 1. Leaves loosely spreading, blades linear or linear-lanceolate, flat, as wide or wider than the sheaths; roots slender, yellow; capsules obovoid- trigonal. . . . 1. C. rosea 1. Leaves erect or ascending: pies ie narrower tian the sheaths; roots dilated at bases: capsules globose—oblong-trigonal. 2. Plants in small tufts with a few erect stems; roots copiously lanate, clustered. 2 4 2. C. ornata De lants cespitose, a. numerous - Sspaniching: ese: 4 stems; roots moderately lanate, glabrate. . . . . . . . . 3. C. graminea 1. CUTHBERTIA ROSEA (Vent.) Small. in Fl SE. U.S. 237. 1908. Tradescantia rosea Vent. Jard. Cels. pl. 24, 1800. Tripogandra rosea (Vent.) Woods. Ann. Mo. Bot. Gard. 29: 141-154. 1942. lants 2—5.5 dm tall: leaf blades 0.8—2.5 dm long, 4—15 mm _ wide, puberulent or glabrous; cymes 10—15-flowered; involucral bractlet 3—5 mm long, simple or cleft, rarely acuminate to 14 mm long. Sepals 3—9 mm long, suffused with rose; petals ovate, 8—12 mm long, shallowly crenu- late, bright rose; capsules 3—4 mm ignoe ee silvery gray, ribbed-reticu- late, 2 mm wide. Sandy banks, hammocks; Southampton Co., Virginia to vicinity of Jacksonville, Florida. 2. CUTHBERTIA ORNATA Small, Man. SE. FI. 259. 1933. Tradescantia rosea Vent. var. ornata (Small) Anders. & Woods. Contrib. Arn. Arb. 9: 111-116. 1935. Small in describing C. ornata did not designate a type specimen. The LECTOTYPE selected is Small and Winkler 9054, sandhills of Avon Park to Sebring, De Soto County, 1 May 1919 (NY). A later civil division of De Soto County placed the type locality in Highlands County. Plants 2.5—4.5 dm tall, yellowish green, essentially glabrous, branches few, internodes 2—4; leaf blades 1.7—2.5 dm long; sheaths red-striate, 0.3—1.7 mm long. Involucral bract 1—3 mm long; bracteoles 2 mm long, base greenish, whitish on margins. Cymes 3—15-flowered, pedicels 10—14 mm long; flowers opening two or three at a time; sepals 4—5 mm long, petaloid, apex slightly incurved; petals 10—15 mm long, prominently crenulate; stamen hairs often purplish; filament white, sometimes suffused with pink. Mature capsules 3—5 mm long; seeds 2 mm wide, typically compressed, silvery, ribbed. Distributed only in Florida; oak scrub and sandhills. 28 3. CUTHBERTIA GRAMINEA Small, Man. SE. FI. 259. 1933. Tradescantia rosea Vent. var. graminea (Small) Anders. & Woods. Con- trib. Arn, Arb. 9: 111-116. 1935. HOLOTYPE: J. kK. Small, June 27-July 1, 1895, about Augusta, Richmond Co., Georgia (NY). Plants 0.4—3.8 dm tall, glabrous with 6 or fewer nodes. Leaf blades 0.4—2 dm long, 1—5 mm wide; sheaths purple-striate, pilose about orifice, puberulent on surface. Roots from rhizomatous base crowns or in congested tufts from the lower nodes. Involucral bracts 3 mm or rarely 14 mm long, cleft or obscurely lobed. Flowers 3—13 or more; pedicels 8—12 mm long; sepals green, suffused with rose; petals 8—10 mm long, bright rose. Mature capsules 3.5 mm long; seeds 1.5—1.8 mm wide, ribbed-reticulate, typically silvery. 2n = 12, 24, 36. Sandy woods; Coastal Plain, adjacent to the Fall Line, and in the interior formations throughout the range of the genus. CUTHBERTIA GRAMINEA Small f. leueantha Lakela, f. nov. HOLO- TYPE: Lakela 32048 (USF). A typo different cum pctalis, staminibus et pistillo albidis. Grown from offshoots of plant collected 1 April 1969, sandhill, vicinity of University of South Florida campus, 18 May 1970, Plates 1, 2, 3. Differs principally from forma graminea in absence of anthocyanin pigments, hence bracteoles, sepals, and leaves margined with white; stem bases, nodes, sheath striations suffused with white: petals, stamen filaments, and hairs white; anthers and pollen yellow; pistil wholly white with lustrous stigma. 2n = 12. After several years of field acquaintance with pink-flowered Cuthbertia, the appearance of a color variant was more than a passing surprise. A luxuriant plant with many flowering stems and white budding cymes was discovered in an opening of a sandhill vegetation April 1, 1969. To facilitate continued observations, two stems were detached and transferred to the University Botanical Garden. Later, upon returning to the site for flower- ing and fruiting specimens, the plant was not there and others could not be found, Perpetuation of the novelty depended on the two plants in culti- vation. The attractive white flowers opened at dawn and closed in the early afternoon. After ripening of the pollen, petals with stamens rumpled over the ovary and the style remained exserted beyond the closed sepals. Fruit did not form, however, and the flowers one after another fell with withered pedicels. The flowerless rachis with persistent bracteoles did not wither. In due time, tips of green leaves appeared in axils of proximal bracteoles below, tips of rootlets were clongating. One of the plants on September 11, 1969, had an offshoot of five leaves 3—8 em long. The stem was cut above the roots and pressed as a herbarium specimen. The other plant was more prolific by producing two offshoots of similar size. The reclining stem tip was secured in soil until the plants were independent. During the cooler months, the plants were placed in the lath house where they freely ie Sa gga 8 2 att. 88 Plate 1. Cuthbertia graminea f. leucantha: plant grown from offshoots for type specimens, Lakela 32048. 30 PLATE 2 Plate 2. Cuthbertia graminea f. leucantha: flower with dehisced anthers, the other in post anthesis with mature stigma. flowered and fruited through the spring of 1970. The plants, developing from the two offshoots since September 1969 comprise twenty flowering stems. Seeking parallelism of rachial propagation in nature, at least one prone stem of C. ornata was found with a rooting propagulum in sand of the topo- site of forma leucantha. This is an undeveloped wooded area, west side North 30th Street, North Tampa. Although spoiled and destined for destruc- tion by urban development its vegetation still features many native species typical to Pinus palustris-Quercus laevis association. In the undergrowth on mats of pine needles and among grasses, thrive fourteen Fabaceous genera, and various Florida endemics such as Cacalia a loridana, Baptisia Lecontei, Phoebanthus grandiflora, Cuthbertia ornata, Cuthbertia graminea forma graminea and forma leucantha. The white-flowered variant reap- PLATE 3 | Plate 3. Cuthbertia graminea f. leucantha: declining style with unripe stigma among mature stamens. peared in 1971. At present its known distribution is extended to Pinellas County. The author is indebted to the late Dr. Lloyd H. Shinners for his advice in literature, for the xeroxed references received from the SMU Science Library; to the University of Michigan for the type plate of Cuthbertia rosea; to Dr. Harold Moore, for sharing his concept of the generic status of Cuthbertia; to Dr. Wilbur Duncan for sending living plants of C. rosea; to Mrs. Martha Meagher for the chromosome number; to Curators of the New York Botanical Garden and the University of North Carolina for loans of specimens; and to Dr. Long for consultation, editing, and for the Latin description. REFERENCES ANDERSON, E. and WOODSON, R. FE. 1935. The species of Tradescantia indigenous to the United States. Contrib. Arn. Arb. 9: 111-116, pl. 1- ARISTEGUIETA, LEANDRO. 1965. Notas sobre la familia ee eer en Venezuela. Bol. Acad. Cienc. Fis., Mat. y Nat. 25 (68): 94-142 sera cared hal M. 1964, Estudio sobre las Conmnelinacent Argentinas, I. Darwiniana (1):°8 BRENAN, - 1966. The classification of Commelinaceae. J. Linn. Soc. (Bot.) 59: 349-370. Nae - L. 1950. Gray’s anual of botany, 8th ed. American Book Co., Boston. GILES, N. Jr. 1942. Aexopolypleid and eee ae distribution in Cuthbertia gra- mined Sl. poser J. Bot. 29: 37- Studies of ee oo dies of Cuthbertia graminea in the Carolinas. ehehy Mechel Cee 59: 73-80. ee E ree N, H. A 57. The new Britton and Brown illustrated flora of northeastern United States and ee ee: N.Y. Bot. Gard., Bronx, N. Y. MOORE, H. E., seg 1960. Tripogandra grandiflora ( (Commelinaceae) Baileya 8(3): 77-83. SMALL, J. K. 190 Fler m the southeastern United States. N. Bot. Gard., New York. . 1933, ae of the southeastern flora. Reprint. Univ. of North Carolina Press, TOMLINSON, P. iB. wee Anatomical data in the classification of Commelinaceae. J. Linn, a a Y fan ht YY wr a — ~ ee oi ‘ Cee on the North American genera of Commelinaceae. Ann. Mo. Bot. Gard. 29: 141-154 OBSERVATIONS ON SOME UPPER AMAZONIAN FORMICARIAL MELASTOMATACEAE TREVOR WHIFFIN Department of Botany, University of Texas, Austin, Texas 78712. The observations reported here were made in the area around Leticia, Amazonas, Colombia during July and August 1968, and have subsequently been extended by further study in the herbarium. Nevertheless, apart from the section on taxonomic notes, the main emphasis is on the field observa- tions, and the extent to which they support (or not) some current ideas related to formicaria and to speciation in the tropical rain forest The author citations for all names used in the first two sections of the paper (unless given) are provided in the section on taxonomic notes. Speci- mens collected from the various populations studied in the field are listed at the end of the paper. FORMICARIA A large proportion of upper Amazonian melastomes have formicaria. Among these are Maieta guianensis var. guianensis, M. guianensis var. leticiana, M. poeppigii, Ossaea bullifera, Tococa juruensis, T. ulei and T. aff. stephanotricha, which were studied in the field. Quantitative observa- tions of the disposition of formicaria on a number of plants were made for M. guianensis var. leticiana, M. poeppigii, T. ulei, and T. aff. cic tricha, while qualitative observations were made for the other pa tax In Maieta guianensis (Fig. 2A,B) and M. poeppigii (Fig. 2C), the leaves of a pair are usually markedly unequal; the larger leaf has a eee oped formicarium, the smaller leaf not. The general habit of Maieta is illustrated in Fig. 1, showing the ao tive positions of the larger and smaller h leaves at a node are more or less equal, and in this case both have a formicarium. Also, the rare intermediate leaves have formicaria of cor- poupenanes intermediate size. There is no significant difference between e two spec In Ossaea ee (Fig. 2D), the leaves are usually unequal, the larger having a formicarium and the smaller not. In Tococa juruensis (Fig. 2E), the leaves are more or less equal, both having a formicarium; occasionally one leaf of a pair will be smaller, and with a small or no formicarium. In Tococa ulei (Fig. 2F), a young seedling, at the first few nodes, has only one or sometimes neither of a pair of leaves bearing a formicarium. In larger plants, the situation is variable, even on the same plant. The leaves are not markedly anisomorphic, but usually one leaf of a pair is slightly smaller. The larger leaf has a large well-developed formicarium; the opposite, and usually slightly smaller leaf, has either a small formi- SIDA 5(1): 33—41. 1972, me NN ri MON ti aN i ACY ms we wo Fig. 1. General habit of Maieta guianensis var. leticiana (Whiffin 68). Fig. 2. Details of formicaria. A. Maieta guianensis var. leticiana (Whiffin 68); B. M. guianensis var. guianensis (Whiffin 21); C. M. poeppigit (Whiffin 56); D. Ossaea bullifera (Whiffin 18); FE. Tococa juruen- sis (Whiffin 8); F. T. ulei (Whiffin 15); G. T. aff. stephanotricha (Whiffin 52). All x4. 36 carium or none, in about equal frequencies. Occasionally this opposiic leaf will have a large formicarium, and at other times a minute, scarcely developed formicarium. The disposition of these various forms on the ma- ture plant seems to follow no set sequence. However, the size of the formi- carium seems to vary proportionately with the size of the leaf (though not so markedly as in Maieta, where the range in leaf size is greater). In an unidentified Tococa species (aff. stephanotricha) (Fig. 2G), the leaves at the lower two or three nodes are markedly unequal, with only the larger leaf of each pair having a formicarium; the leaves at higher nodes are more or less equal, and both have a subspherical formicarium. The utility of the disposition of formicaria as taxonomic characters has not been extensively examined. Genera such as Clidemia, Ossaea and Tococa contain both formicarial and non-formicarial species; all three species of Maieta are formicarial. The formicaria appear to be a genetic feature, developing in Tococa guianensis, for example, even in the absence of ants (Wurdack, pers. comm.). Observations on living plants indicate that the degree of development of the formicarium varies with the size of the leaf, as might be expected. Species which are markedly anisophyllous (e.g. in Maieta) have a formi- carium usually only on the larger leaf of a pair, but if the opposing leaves are subequal to equal then both have a formicarium. Species in which the leaves are variably subequal to somewhat unequal (e.g. Tococa ulei) usual- ly have a well-developed formicarium on the larger leaf, and an intermedi- ate to no formicarium on the opposing leaf. Species in which the leaves are equal to subequal (e.g. Tococa aff. stephanotricha) usually have a well- developed formicarium on both leaves of a pair; often, however, there is some slight anisophylly even in these species, and the size of the formi- carium will vary correspondingly. Any attempt to use disposition of formi- caria as a taxonomic character would have to consider this problem of allometry. The shape and probably also the placement of the formicaria is often, however, a useful taxonomic character. For example, the shape of the formicarium helps to distinguish Tococa ulei from T. guianensis; indeed, several Tococa species have formicaria with distinctive shapes. There have been a number of observations on the function and evolution of formicaria (often also called myrmecodomatia). Various different types of relationship exist between ants and plants, as has been outlined by van der Pijl (1955). As regards formicarial melastomes, it appears that the plants provide shelter for the ants (a myrmecodomic relationship); there is no evidence that the plants provide a source of nutrition for the ants (Melin, 1931). It is often postulated that the ants protect the plant from attack by phytophagous insects; this proved to be so in bull-horn acacias, where the removal of the ant colonies was followed by severe defoliation of that acacia (Jansen, 1967). In addition, Jansen (1967, 1969) has noted an allelopathic effect of the ants, maintaining a cleared area 37 around Acacia and Cecropia by selectively killing the shoot and lateral branch tips of neighboring plants. However, there is no evidence of such an interaction between ants and formicarial melastomes. Melin (1931) could find no evidence that ants had any effect on the extent of leaf damage caused by phytophagous insects on several Tococa plants in middle Amazonia. Field observations in upper Amazonia would, on the whole, support Melin’s conclusion. Although the question of allelopathy has not been investigated in formicarial melastomes, the advantage of this would be much less to melastomes growing as small shrubs in tropical rain forest than it is to Acacia and Cecropia trees in dense young secondary regrowth. Spruce (1908) postulated a correlation between myrmecophily and inun- dation; this may have been due to the high frequency of Cecropia trees, which show myrmecophily, in the disturbed and often periodically inun- dated areas alongside the rivers. In the melastomes, however, there ap- pears to be no such correlation. There are few melastomes in the varzea forest (periodically or seasonally inundated forest) and, although the latter are often, but not always, formicarial, there is also a high frequency of formicarial melastomes in the neighboring terra firme forest (upland). Melin (1931) also indicates that the formicarial melastomes which he studied were on the edge of the inundated areas, and in the terra firme forest. As noted earlier, formicaria are probably produced even in the absence of ants; thus they are part of the genetic make-up of the taxon, and have presumably arisen and been fixed during the course of evolution. From the taxonomy of formicarial melastomes, it would appear that formicaria and the ant-plant relationship have arisen independently on a (small) num- ber of separate occasions. Spruce (1908) attempted to explain this evolu- tion in neo-Lamarckian terms; Melin (1931) could find no explanation in relationship. At the moment, therefore, the question is not settled, at least in the melastomes. Some authors (Schnell, 1967, and others cited there) have indicated that formicaria may have arisen during evolution from acarodomatia, but the evidence at the moment is not very satisfactory. For selection to have favored the formation and evolution of formicaria, there must have been some selective advantage of the ants to the plant, even if the nature of this advantage is not known at present. Within the variation of the taxon, those forms which allowed or increased this associ- ation with ants would have been favored, thus facilitating the fixation of this trait in the population, and later in the taxon. This is similar to the idea of pre-adaptation (Jansen, 66). probable outline of the further development of the ant-plant relationship through co-evolution is well de- tailed by Jansen (1966) SPECIATION The modes of speciation in the tropics are probably not different in quality from those in temperate areas (Cain, 1969; Mayr, 1969), although 38 they may vary in the relative importance of the various different modes. Discussions of speciation in the tropics variously emphasize genetic drift in small populations (Federov, 1966) or natural selection (Ashton, 1969); sometimes other modes such as saltatory non-adaptive evolution (van Steenis, 1969) are postulated, but generally these are unsatisfactory. The general assumptions concerning the rain forest environment are that it has been relatively stable over geological time, and that as far as plant survival is concerned, it is optimal, thus leading to the idea of a low survival level (van Steenis, 1969) and a consequent lack of selective force. The rain forest environment is complex, not only hort- zontally (area) but also vertically (different height levels). The greater diversity of niches is part, but probably not all, of the explanation for the greater diversity of species. Among the rain forest trees, especially those which are emergent or which form the canopy, fairly large, though comparatively widespread, populations occur which are probably inter- breeding. Ashton (1969) has shown that selection appears to have been a factor in speciation in such trees. However, in understory shrubs, the populations in many cases appear to be rather small and isolated. Thus in the area around Leticia, only one small population of Ossaea_ bullifera and one of Tococa juruensis were found. Populations of Maieta guianensis var. guianensis, M. guianensis var. leticiana, and M. poeppigit were found more often, but each population consisted of one or, at most, a few indi- viduals, and was some distance from any other population. Often, however, two or even all three taxa were found together, indicating a similar eco- logical preference, although no intermediates were found. The question of how these taxa diverged is less easy to answer. However, some insight may be gained from recent studies by Haffer (1969), indicating that the Ama- zonian rain forest has not remained stable over a long period of time. He presents evidence that, during the Pleistocene dry periods, rain forest vegetation was restricted to certain forest refugia, which probably corre- sponded with the present areas of higher rainfall; this theory is useful in explaining speciation and present distribution of Amazonian birds. His theory applies equally well to speciation in rain forest plants. Maieta guianensis var. guianensis and M. poeppigii are widespread across Ama- zonia; M. neblinensis is restricted to Cerro de Neblina, Amazonas, Vene- guela, and presumably, after becoming isolated there, has diverged and become distinct. M. guianensis var. leticiana is found only in Amazonian Colombia and adjacent Pertti and Brasil (Whiffin, 1971). Its distribution corresponds to the Napo forest refugium of Haffer (1969) or, perhaps more correctly, to the riverside refugia to the east of the main Napo refugium. This taxon could well have arisen in the forest refugium during one of the Pleistocene dry periods. A number of other melastomes have a very similar known distribution to Maieta guianensis var. leticiana; these include Adel- obotrys macrophylla Pilger, Clidemia ulei Pilger, Ossaea bullifera, Tococa juruensis, and Tococa ulei. TAXONOMIC NOTES The notes given here refer specifically to those formicarial melastomes mentioned above, indicating a number of points of interest found while studying these taxa in the herbarium; notes on other melastomes collected are given by Whiffin (1970). Maieta Aubl. Maieta guianensis Aubl., which is widespread from the Guianas across Amazonia to eastern Colombia, Pert, Ecuador, and Bolivia, is fairly com- mon in upper Amazonia. The variation in this latter area is greater than in other parts of its range, and proves to be of interest (Whiffin, 1971). Two varieties of M. guianensis are recognized: var. guianensis and var. leticiana Whiffin; both occur in the area around Leticia. Maieta poeppigii Mart. ex Triana is similarly widespread, though less co only collected throughout its range. The two species are easily ane ane Paes 1931a; Whiffin, 1971). Ossaea bullifera (Pilger) Gleason. This, a distinct species from a restricted area in upper Amazonia, has been very rarely collected. It may be recognized by the following charac- ters: unequal leaves, the larger with a formicarium, the smaller not; young branches densely long-setose, and leaves long-setose on nerves, margin and formicarium, with long, white to golden, setae; inflorescence lax, sparsely long-setose, and hypanthium more densely long-setose, with some setae glandular; calyx lobes long and narrow, green; petals long and narrow, white to translucent; fruit a comparatively large, long-setose berry. This species, originally described in Leandra, was later transferred to Ossaea by Gleason (1931b); its lax, lateral or axillary inflorescence with a few four- merous flowers may be better placed in Ossaea, although its large, fleshy, blue fruit seems a little out of place. Tococa juruensis Pilger. his, one of the more easily recognized of the upper Amazonian species of Tococa, has only rarely been collected. It appears to have a distribution including upper Amazonian Brasil, Colombia and Pert. It may be recog- nized by the following characters: young branches, larger formicaria, peduncles, pedicels and hypanthia long-setose pubescent; leaves variously unequal to subequal, most having a formicarium of a size varying with the leaf size: hypanthium terete, the calyx more or less truncate. Tococa ulei Pilger. This species is common in the area around Leticia. It has sometimes been confused with Tococa guianensis Aubl., which has perhaps slowed down appreciation of its specific limits. Tococa ulei is an upper Amazonian species, mostly in Colombia, Peru, and adjacent Brasil. The shape of the formicarium in T. ulei is variable; typically it is longer than broad, but sometimes it is aberrant and more or less hemispherical. The inflorescence is typically racemose, but may be a simple panicle. Some collections of T. ulei may have been misidentified as T. guianensis, 40 which is usually paniculate but may also be racemose; in any case, there is no sharp distinction on this character. As presently understood, T. gui- anensis covers a wide undefined variation, and needs re-definition. Gleason (1931b) has some useful comments on this point. T. ulei is a distinct spe- cies, however, and may be separated from T. guianensis by the following characters: formicarium usually elongate on the larger leaves; leaves often elliptic, sometimes more elongate; hypanthium with ten broad veins; hypanthium at most sparsely hirsute, and usually scurfy or stellate pubes- cent to glabrous; calyx teeth with a terminal black bristle, which may be entire or divided into three (or rarely more). Tococa ulei approaches, but is distinct from, the typical form of T. fera Mart. & Schrank ex DC.; however, some of the varieties placed under T. bullifera may be better considered as T. ulei. In some respects T. ulei is similar to T. occidentalis Naud., but is surely distinet; T. occidentalis is also similar to T. guianensis, but again they are probably distinct. In fact, T. occidentalis, a rather neglected species, appears to be somewhat inter- mediate between T. guianensis and T. bullifera, the latter in turn connect- ing with T. ulei. Tococa ulei also approaches T. discolor Pilger, but in the latter the leaves are generally broader in shape, and less attenuate at the base. It is also similar to T. loretensis Ule, which does not have the black setae on the calyx lobes, but is otherwise very similar, especially in inflor- escence form and hypanthium shape. These three upper Amazonian spe- cies, T. discolor, T. loretensis, and T. ulei, seem to form a very close group. It has not been possible to examine the types, but type photos are available. At the moment, no valid specific differences between the three can be found. Tococa aff. stephanotricha Naud. This plant, collected sterile, was not matched, but in vegetative details, especially pubescence, it shows affinities with Tococa stephanotricha Naud. ed yulli- and its allies. COLLECTIONS Below are listed the specimens collected from the various populations of formicarial melastomes studied in the field; a full set of these is de- posited at Kew (KX), nearly complete sets at New York (NY) and at Wash- ington (US), and a partial set at Bogota (COL). Maieta guianensis var. guianensis : Whiffin 19, 21, Maieta guianensis var. leticiana : Whiffin 83 (type), 3, 22, 49, 51, 57, 60, 61, 6S. Maieta poeppigti : Whiffin 23, 24, 56, 82. Ossaea bullifera : Whiffin 18. Tococa juruensis : Whiffin 8. Tococa ulei : Whiffin 4, 15, 25, 50, 62. Tococa aff. stephanotricha : Whiffin 52. For the herbarium study, further material was examined in the herbaria 41 of the New York Botanical Garden (NY) and of the U.S. National Museum (US). ACKNOWLEDGEMENTS The field observations and collections were made while a botanist on the Cambridge Medical Expedition to Colombia, 1968; I am grateful to all those who made that expedition possible (full details are given in Gundry et al., 1970). The taxonomic studies were undertaken while at the New York Botanical Garden, and I am very grateful for the assistance that I re- ceived there. I am grateful also to the curators and staff of the U. S. Na- tional Herbarium (US) for allowing me to examine the collections there. I am grateful to Dr. B. L. Turner for a number of helpful comments on the manuscript, and to Mr. Geza Knipfer for preparing the illustrations. I am especially grateful to Dr. John J. Wurdack for help with the identi- fications, for reading the manuscript, and for his help and encouragement. REFERENCES ASHTON, P. S. 1969. seo ee tropical rain — trees: some deductions in the light of recent evidence. Biol. Soc. 1: 155- aaa A. J. 1969. Speciation in | environments: summing up. Biol. J. Linn. Soc. 1: 233-236. FEDEROV, An. A. 1966. The structure of the tropical rain forest and speciation in the humid tropics. J. Ecol. 54(1): 1-11. GLEASON, H. A. 1931a. The relationships of certain myrmecophilous me Torr. Bot. Club 73-85, “ 1931b. Studies in the flora of northern South America. nis ae Collections of Mclastomavieete from Peru and Amazonian Brazil. Bull. Torr. Club 5 15-262. GUNDRY, D. R. T., ef al. 1970. Report of the Cambridge va aie aa to oe bia, 1968. Published privately by the authors. HAFFER, J. 1969. Speciation in Amazonian forest birds. Science 165 See 131-137. JANSEN, D. H. 1966. Co-evolution of mutualism between ants and acacias in Central America, Evolution 20: 249-275, 9 ay vegetation structure, and the ant x acacia interaction in Central astomes. Bull. America. eas 48 (1): 26-3 969. Allelopathy Be myrmecoph yes: the ant Azfeca as an allelopathic agent ta Ecology eee -153 MAYR, E. 1969. Bird speciation in nee mebpies Biol. J. Linn. Soc. 1: 1-17. MELIN, D. 1931. pees aie to the study of the theory of selection 1. Zool. Bidr. Upp- i 0 I Pl hol 5: 190-200. PI. van dee: Some remarks on phy SouNEEL. R. se cn ution a l’étude des genres Gu uyano- ie owen Tococa Aubl. et Maicta Aubl. (Melastomacées) et de leurs poches foliaires. Adansonia sér. 2, 6(4): 2 25-35 ace. R. 1908. . - a ra on the . ane Andes. — STEE ee C. G. G. J. Plant speciation in Malesia, with cial reference to the theory of non- sdaptive ioe Gee Biol. J. oe Soc. 1: WHIFE IN, T. 1970. Botany. In D. R. T. Gundry ef al., Report of “the ‘Cambridge Medical Expedition to cate nbia, 1968. 1 ry 971. Studies in the genus Malefa Aubl. (Melastomataceae). Brittonia 23: 325- os 329, NOTES GALIUM PEDEMONTANUM (BELL.) ALL, (RUBIACEAE) REPORTED FROM OKLAHOMA.—A specimen on loan to the Bebb Herbarium from the Southeastern State College Herbarium, Durant, Oklahoma, proved to be this rare European introduction, first reported in the United States from Kentucky and West Virginia (Bartholomew in Castanea 6:141-142, 1941). This specimen, John and Connie Taylor 6419, was collected along a roadside 6 miles southeast of Eagletown, McCurtain County, April 26, 1970.—Cheryl A. Lawson, Bebb Herbarium, Department of Botany and Microbiology, University of Oklahoma, Norman, Oklahoma 73069. THREE GRASSES NEW TO GEORGIA: ERIOCHLOA GRACILIS, SE- TARIA FABERU, LEPTOCHLOA UNINERVIA (GRAMINEAE).—Three grasses, probably not native to Georgia, seem firmly established at separate localities within the state. Specimens of Eriochloa gracilis (Fourn.) Hitche. were sent to the Uni- versity of Georgia Herbarium for naming. They were from the southwestern Coastal Plain of Georgia, where the species is described as slowly spread- ing but a vigorous weed in ditches and in and around fields of cotton, corn, and peanuts. Hitchcock (1950) reports this species as ‘‘often a weed in fields’? and shows a distribution of southern California to western Texas and Oklahoma. Small (1933), Fernald (1950), Gleason and Cronquist (1963), and Radford, ect al. (1968) do not report it for their areas, which includes all of the United States cast of the Mississippi River. Setaria faberii Herrm. is known as a vigorous weed, now widely distri- buted (Gleason and Cronquist, 1963), mostly in the central part of the eastern half of the United States (Hitchcock, 1950). Previously it has been reported southward only into Arkansas (Hitchcock, 1950), and into Tennes- see, throughout most of North Carolina, and in central north South Caro- lina (Radford, ct al., 1968). The new locality is in the southeastern Pied- mont of Georgia, where it is abundant in abandoned fields and roadside cuts and ditches. Leptochloa uninervia (Presl.) Hitche. & Chase has been found near the seacoast, distributed in a broad, low, open area between a woods and the main highway across White Marsh Island east of Savannah, Georgia. Many plants had been mowed, but had grown vigorously afterward and were flowering and fruiting. The nearest localities reported by Hitchcock (1950) are in Mississippi and North Carolina. Radford, et al. (1968) record the species from Mississippi and from the northern part of the Piedmont of North Carolina. The Eriochloa and Setaria are likely to become important weeds over large areas of the Southeast. Specimens of the three species are on file in the University of Georgia Herbarium: ERIOCHLOA GRACILIS (Det. by SIDA 5(1): 42. 1972. 43 W. H. Duncan), Hull Andrews, ca. 26 June 1970, Lee Co., Ga.; SETARIA FABERII, W. H. Duncan, 23456, 1 Aug. 1970, just east of Washington, Wilkes Co., Ga.; LEPTOCHLOA UNINERVIA, W. H. Duncan 23451, 31 July 1970, Chatham Co., Ga.—Wilbur H. Duncan, Department of Botany, Uni- versity of Georgia, Athens, Georgia 30601. REFERENCE GLEASON, H. A. and ae THs CRONOQUIST. 1963. Manual of vascular plants of north- eastern United States and Canada. D. Van Nostrand Co., Inc., New ues 8 pp. FE oo M. L. 1950. ca s manual of botany. 8th ed. Aenea Book Co., N. Y. 1632 HITCHCOCK, A. S. 1951. Manual of grasses - the United States. Revised by Agnes Chase. U. §. Dept. ge none Publ. No. 200. 1051 pp. RADFORD. A. E., et al. . Manual of the v ee flora of the Carolinas. Univ. of North Carolina Press, ae a 1183 pp. SMALL, K, 1933, ie of the southeastern flora. Reprint. Uniy. of North Carolina Press, C Shapel Hill. pp. NEW cake Geel IN HOFFMANSEGGIA CAV. AND CAESALPINIA L.—l. HOFFMANSEGGIA glauea (Ort.) Ejifert, comb. nov sarrea i ca Ort., N. Hort. Matrit. Decad. 15, Hoffmanseggia falearia Cav., Icones Plant. 4:63, a 392. 1798 (‘1797’). Caesalpinia glauca (Ort.) an, Rev. Gen. Plant. 3(2):53. 1898. C. Seles nec ee & Macbr., Publ. Field Mus. Nat. Hist., Chicago, Bot. r. 18(3):191. 1948. a ee described Hoffmanseggia falcaria, he clearly stated that it is the same species as Larrea glauca Ort., but his remark was overlooked by all subsequent workers, with the exception of Kuntze, who did not con- sider Hoffmanseggia Cav. distinct enough from Caesalpinia L. and made the new combination in Caesalpinia. After studying the genus Hoffmanseggia Cav., I concluded that it is sufficiently distinct from Caesalpinia L. to warrant generic recognition. Because my revisionary work can not appear in print for some time, I make this new combination available for this species which is widely distributed in the arid and semiarid regions of the New World. A list of synonyms is given by Britton in the North Ameri- can Flora (vol. 23, p. 311. 1930; under Larrea densiflora), to which list the above synonyms have to be added. 2 ESALPINIA snag (Britton) Ejifert, comb. nov. Larrea peninsularis Britton, N. Am. FI. 23:311. 1930. Hoffmanseggia aden yee Wiggins, Contrib. Dudley Herb., ord Univ 0. This is a species Be Caesalpinia, which begins to flower at a very early stage while still herbaceous. Older specimens are, however, definitely woody, a character which occurs only occasionally, and then only at the base of the shoot, in two species of Hoffmanseggia. Also, the lunate pods are unknown in the latter genus, but they are characteristic of some parvi- foliolate species of Caesalpinia, most of which were formerly included in Hoffmanseggia. These species, the closest relatives of C. peninsularis, are SIDA 5(1): 43. 1972. 44 the following: C. drummondii (Torr. & Gray) Fisher, in North America, and C. pumilio Griseb., C. ternata (Phil.) Macbr., C. viscosa (Ruiz & Pav.) Macbr., C. egena (Macbr.) Macbr., and a still unnamed Peruvian species in south America.—Imre J. Eifert, Department of Botany, The University of Texas, Austin, Texas, 78712. VERNONIA CRINITA RAF., A SYNONYM OF V. ARKANSANA DC. (COMPOSITAE)—The well known Vernonia of the Ozarks of Missouri and Arkansas, V. crinita Raf. in the regional and state manuals of the area (Fernald, 1950; Gleason and Cronquist, 1963; and Steyermark 1963), is incorrectly named. Gleason (1922), in his revision of the genus Vernonia in North America, appeared to indicate that V. crinita Raf., New Flora Am. 4:77, was published in October with the notation ‘‘(?0) 1838,’’ and that the name V. arkansana DC., Prodr. 7:264, appeared in the literature in December, 1838 with the symbol ‘‘(?D) 1838.’ After studying some of Rafinesque’s correspondence, Barnhart (1907) had earlier determined that part 4 of Rafinesque’s New Flora N. Am. was actually available for distribution late in 1838 but before December and not in 1836 as listed on the frontispiece of the entire work. Shinners (pers. com.) suspected that Gleason may have consulted Barnhart about the valid publication dates and that these were Barnhart’s best opinion at that time. The International Code of Botanical Nomenclature provides that the title page date may not be accepted if it can be shown to be incorrect. It is therefore necessary to determine the date of publication for Vernonia arkansana DC. Volume 7 of DeCandolle’s Prodromus appeared in two parts: pages 1-330, late April 1838 and pages 331-801 late December 1839 (Staf- leu, 1967). Gleason’s ‘‘(?D)’? was correct but the year was not, as the December was for part 2 which was not relevant for V. arkansana DC. (7:264 There can be no doubt that both DeCandolle’s and Rafinesque’s descrip- tion refer to the same Vernonia. It would not be possible for their descrip- tions to fit any other species of Vernonia and no one familiar with the genus has ever suggested this. In conclusion, Vernonia arkansana DC., Prodro- mus 7:264, late April 1838 is earlier than V. crinita Raf., New Flora N, Am. 4:77, late in 1838 and possibly October. Therefore, V. arkansana DC. is the earliest legitimate name. It is unfortunate when a well-known name must be changed but it is necessary for nomenclatural stability. I want to thank the late Dr. L. H. Shinners for his assistance with this nomenclatural problem. This study was supported by National Science Foun- dation Grant GB 7208.—S. B. Jones, Jr., Department of Botany, University of Georgia, Athens, Georgia 30601. REFERENCES BARNHART, J. Hl. 1907. The dates of Rafinesque’s New Flora and Flora 'Telluriana. Vorreya 7:177-181. SIDA 5(1): 44. 1972. 45 DE CANDOLLE, A. a oe 1839. Prodromus systematis naturalis regni vegetabilis. 7 CL 195 0y P23 30-8 FERNALD, M. L. 1950. ae s manual of botany, 8th ed. American Book Co., N. Y. p. 1359. GLEASON, - 1922. Vernonieae. fa Am. Fl. 33:85. Se er . and A. CRONQUIST. 1963. Manual of vas ae ee of northeastern nited — hie Canada. a Nostrand Reinhold Co., N. : eae E, -C; 836. New flora and botany of North America. Ph ‘de adelphia. STAFLEU, F. A. i Taxonomic literature. IBPT Utrecht, Netherlands. Regnum Vege- tabile 52:75, 373. STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Press. Ames. p. 1457. ROTALA INDICA (LYTHRACEAE) IN LOUISIANA.—During a _ recent visit to a rice field near Crowley to collect aquatic plants for class use I noted an individual of a species that was quite unknown to me. Bringing e minous United States only from Butte County, California (H. L. ason, A Flora of the Marshes of California, p. 602. 1957). On a return visit to the Crowley site I found, after a thorough search, only four more individuals of R. indica; these were left in situ to provide a seed source that would, hopefully, help to keep the species a part of the field’s flora. All the plants were growing in clear water up to 6 inches deep. All were unbranched. They were rooted in soft mud among Chara and were associated with Ammannia coccinea, Bacopa repens, Dopatrium junceum, Eleocharis atro- purpurea, E. obtusa, Heteranthera limosa, Ludwigia decurrens, Rotala ramosior, and Sphenoclea zeylanica. Collection data are as follows: in rice field, Rice Experiment Station, 2.5 miles NE of Crowley, Acadia Parish, Thieret 36102 (LAF), 19 September 1971.—John W. Thieret, University of Southwestern Louisiana, Lafayette 70501. NEW COMBINATIONS AND A NEW SPECIES IN HELENIUM (COM- POSITAE—HELENIEAE).—Three new combinations and one new species are reported at this time since a forthcoming systematic study of Helenium (A Chemo-systematic and Cytotaxonomic Study of Helenium Sect. Tetrodus (Compositae), Ph.D. dissertation, The University of Texas at Austin, 1971) will probably be at least one year before appearing in print. Two of the new combinations and the new species belong to Sect. Tetrodus and the other new combination (H. apterum) belongs to Sect. Hecubaea. The present work was supported by an N. S. F. Traineeship presented through the University of Texas at Austin. HELENIUM chihuahuensis Bierner, sp. nov. A H. mexicano foliis irregu- lariter laciniatis vel saltem irregulariter lobatis, partibus supernis inter- nodiorum breviter alis decurrentibus, pubescentia pedunculorum ligularum- que sparsa. MEXICO: Chihuahua. Cima, H. LeSueur 987, 9 Jul 1936 (Holo- type US; isotypes MO, PH, UC). Fig. 1. Annual herbs with several stems or one stem originating at the base, SIDA 5(1): 45. 1972. Figure 1. Helenium chihuahuensis Bierner, sp. nov. 47 sparingly branched above, 2.7—7.8 dm tall. Stems glabrous to very sparsely pubescent. Peduncles 2—12 cm long, only slightly broadened at the top, sparsely pubescent. Leaves variable, but most irregularly laciniate or at glabrous to very sparsely pubescent, midrib prominent, the lateral veins ebscure, hardly decurrent at all along the stem. Heads few to several, globose to slightly ovoid, 8—14 mm high, 8—14 mm wide (excluding rays). Receptacle globose to ovoid, 2.5—$.0 mm high, 3.0—6.0 mm wide. Involucre of two series of lanceolate bracts, the outer exceeding the inner and united at the base, reflexed at anthesis, sparsely pubescent. Ray florets 14—15, ligules 4.5—13.0 mm long, 2.5—6.0 mm wide, yellow, glabrous to very sparsely pubescent. Disc corollas 5-lobed, cylindrical to cylindric-campanu- late, 1.6—2.7 mm long, 0.8—1.1 mm wide, yellow with purple lobes. Achenes 1.2—1.7 mm long, 0.5—0.7 mm wide, pubescent mainly along the 6 ribs. Pappus scales 6, 0.7—1.2 mm long (including the terminal awn which is ca 1/3 the length of the body), 0.3—0.5 mm wide Distribution: Central and northwestern Chihuahua, Mexico. From label data, apparently occurring in ditches and around ponds at elevations of 6,000—7,000 ft. June—September. HELENIUM ELEGANS DC. var. amphibolum (Gray) Bierner, comb. nov. Helenium amphibolum Gray, Proc. Am. Acad. 9:202. 1874. Heleniastrum amphibolum eee Ktze., Rev. Gen. Pl. 342. 1891. Helenium stenopterum Rydb., N. A. FI. "34: 123. 1915. This taxon may be distinsuictied from the closely related var. elegans by its irregularly toothed or lobed lower leaves, usually longer peduncles, larger heads and more southern distribution. HELENIUM MICROCEPHALUM DC. var. ooclinium (Gray) Bierner, comb. nov. Helenium ooclinium Gray, Proc. Am. Acad. 9: 202. 1874. Heleniastrum ooclinium (Gray) Ktze., Rev. Gen. Pl. 342. 1891. This taxon may be distinguished oni the closely related var. micro- cephalum by its longer peduncles, longer ligules, larger heads and more southern distribution HELENIUM apterum (Blake) Bierner, comb. nov. Hecubaea aptera Blake, Contrib. U. S. Nat. Herb. 22:649. 1924. This species has been transferred to Helenium Sect. Hecubaea because of its obvious relationship to H. scorzoneraefolium; however, it may be dis- tinguished from the latter by its leaves which are decurrent for only a few millimeters below the point of attachment to the stem, possession of pappus scales and sterile ray florets.—Mar ierner, Department o Botany, The University of Texas at Austin, Texas 78712; present addres Department of Botany, University of Tennessee, Pe 37916, SIDA 5(1): 47. 1972. 48 ADVENTIVE RECORDS OF ERIOCHLOA CONTRACTA (GRAMINEAE) IN THE UNITED STATES.—The main distribution of Eriochloa contracta Hitche. is Kansas and Missouri, south to Louisiana and Texas, and west to Arizona and Colorado. The species is occasionally adventive elsewhere. A new record for Indiana is as follows: E. E. Terrell and F. G. Meyer 4319, Aug. 28, 1970, abundant for about 50 yards along roadside of route 460 about 0.2 mile west of its junction with route 462, Harrison County, Indiana (distributed to IND, NA, US). As it was abundant at this southern Indiana site, it is possible that it may persist or spread. G. N. Jones (Flora of Illinois, 3rd ed., 1963) reported the species from two counties in southern Illinois. There are specimens in the U.S. National Herbarium which represent new states not listed in Hitchcock’s ‘‘Manual of Grasses of the United States’? (1951, rev. by Chase). These represent adventive occurrences and are as follows: ALABAMA. C. M. Wilson, July, 1947, ‘‘eastern central Ala- bama’’; MISSISSIPPI: W. C. Young, Sept. 1955, 40 acres or more in natural stand near Clarksdale [northwest]; H. J. Jacob 784, Aug 29, 1954, dry soil, Doddsville, Sunflower County [northwest]. In addition, there are three collections from three counties in central California which also represent adventive or introduced occurrences. Munz & Keck (A California Flora, 1968) noted it as occasional in the state. Hitchcock’s Manual cited it as being adventive in Virginia, apparently on the basis of the following specimen: S. F. Blake 11675A, Aug. 7, 1933, one plant only in center of unfinished new highway near Glencarlyn, Arlington County, Virginia. This single plant collection should not have been con- sidered an extension of range.—Edward E. Terrell, New Crops Research Branch, Crops Research Division, U.S.D.A., Beltsville, Maryland 20705. TWO NEW SPECIES OF CHAMAESARACHA (SOLANACEAE) FROM TEXAS.—During a revisionary study of Chamaesaracha, two heretofore un- described species were encountered. My revision of the genus will not appear for several months, and it seems advisable to report these taxa at this time. Also, both species have appeared in Correll and Johnston’s Manual of the Vascular Plants of Texas (1970: p. 1741), and this will AESARACHA edwardsiana Averett, sp. nov. Herba perennis. Folia subsessilia 2.7—7.0 em longa 0.7—1.5 cm lata lance-linearia vel rhombica integra vel plus minusve lobata glabra. Chromosomatum numerus n = 24. Type: TEXAS: Travis Co.: % mi FE of Pedernales River on Hwy 620, 28 June 1968, J. I. Averett 289 (Holotype, TEX; isotypes, GH, M 5 ) Primarily on limestone soil in the Edwards Plateau of Texas. The species is probably most closely related to C. coronopus, differing from the latter in having glabrous, entire leaves. It is also related to C. pallida, differing in the narrow, linear-lanceolate, glabrous leaves. SIDA 5(1): 48. 1972. 49 CHAMAESARACHA pallida Averett, sp. nov. Herba perennis. Folia sub- sessilia 2.0—3.5 (—4.0) cm longa 1.0—1.5 cm lata rhombica vel late lanceolata integra vel sinuato-repanda dense stellato-pubescentia. Chromosomatum numerus n = 36. Type: TEXAS: Presidio Co.: SW edge of the Cuesta del Burro Mts., 35 mi SW of Marfa on Hwy 2810, 24 July 1966, J. E. Averett 155 (Holotype, TEX; isotypes, GH, MO, SMU, US). Chamacsdracia pallida is a rare species of Trans-Pecos Texas and north- ern Mexico, mostly confined to limestone soils. The species has its closest affinity with C. edwardsiana. However, a dense stellate pubescence is typi- cally found in C. pallida. Some specimens of C. edwardsiana and C. corono- pus have considerable pubescence, but not so much as C. pallida.—John E. Averett, Department of Botany, The University of Texas, Austin, Texas, 78712; present address, Department of Biology, University of Missouri- St. Louis, 63121 and Missouri Botanical Garden. A NEW VARIETY OF CAREX BICKNELLIL FROM ARKANSAS.—Three collections of Carex bicknellii Britt., by Dr. Delzie Demaree from river terraces in Prairie and Lonoke Counties, Arkansas represent a superficially puzzling variation because of its resemblance, in its large, nearly nerveless perigynia, to C. brittoniana Bailey. The latter characteristic and the fact that the perigynia are only partially translucent, are also suggestive of the northeastern C. merritt-fernaldii Mack., in which, however, the perigynia are much smaller (4—5 .o—3.5 mm). This variant may be known as: CAREX BICKNELLII Britt. var. opaea F. J. Herm., var. nov. A varietate typica recedit perigyniis valde concavis (margine externo excepto) ventra- liter vix aut haud nervatis, inter achaenium et alam suberosis, margine interno viride, externo hyalino. ARKANSAS: Prairie Co.: Demaree 60141. Holotype: US Differing from the typical form in having the perigynia (5.5—7 mm long, 4—4.75 mm wide), strongly concave rather than ‘‘very flat,’’ opaque (ex- cept for the outer margin), faintly if all nerved ventrally, corky between the achene and wing, and the inner margin green, only the outer hyaline. KANSAS: PRAIRIE COUNTY: river terraces (never plowed), rice region, Hazen, May 10, 1969. Delzie Demaree a (HOLOTYPE: US), 57815 (NY), 55595 (G), and low, wet ground, Ulm, peed 9, 1937, Delzie Demaree 14915 (Hermann Herb.); LONOKE COUNTY: common on terraces in wet areas, rice region, Charlisle, Delzie Demaree pie .—Fred- erick J. Hermann, Forest Service Herbarium, Colorado State University, Fort Collins, Colorado, 80521 SIDA 5(1): 49. 1972. DOCUMENTED PLANT CHROMOSOME NUMBERS 1972:1 GARY H. MORTON Department of Botany, The University of Tennessee, Knoxville, 37916; present address: Florida Junior College, South Campus, Jacksonville, 32205 The following work was initiated as an introduction to this phase of bio- systematics. Iam grateful to the Society of Sigma Xi for a travel grant to collect most of the Solidago here reported and to Bob MacDonald and the U.T. Arboretum for providing space in which to grow these plants. ASTERACEAE (COMPOSITAE) ARTEMISIA cee ene L. *(Fig. 1) n=9. Tenn., Anderson Co.; G. Mor- ton 2888 (TENN, SMU). CACALIA ae IA L. *(Fig. 2) n=26. Tenn., Union Co.;G. Morton 2880T-5 (TENN) and G. Morton 2880T-6 (SMU). In both plants figures of =24 were seen. ee TINCTORIA Nutt. *(Fig. 3) n=12. Tenn., Anderson Co.; G. Morton 2889 (TENN) and G. Morton 2890 (SMU). a PERFOLIATUM L. *(Fig. 4) n=10. NJ, Ocean Co.; G. Mor- ton 2900 (TENN). cc ere ek OR L. *(Fig. 5) n=9. NJ, Hunterdon Co.; G. Morton 3 (TENN). oo: BRACHYPHYLLA Chapm. (Fig. 6) n=18. Fla., Holmes Co.; G. Morton FL-9 (TENN, SMU). This is the first reported tetraploid plant for this species. SOLIDAGO GATTINGERI Chapm. *(Fig. 7) n=9. Tenn., Rutherford Co.; G. Morton TN-37 (TENN). SOLIDAGO JUNCEA Ait. *(Fig. 8) n=9. Penn., Bucks Co.; G. Morton 2897 (TENN) and G. Morton 2896 (SMU). SOLIDAGO JUNCEA f. RAMOSA (Porter & Britt.) Fern. °(Fig. 9) n=9 Penn., Bucks Co.; G. Morton 2899 (TENN) and G. Morton 2898 (SMU). piacere ODORA Ait. *(Fig. 10) n=9. NJ, Ocean Co.; G. Morton 2901 (TENN). Caen PINETORUM Small *(Fig. 11) n=9. NC, Anson Co.; G. Morton NC-8 (TENN). ‘Contribution from the Botanical Laboratory, The University of Tennessee, N. Ser. 371. “Species for which a similar chromosome count has been published previously. "Species for which a chromosome count differing from this one has been published pre- viously. © Species for which no chromosome count has been published previously. SIDA 5(1): 50. 1972 DOCUMENTED PLANT CHROMOSOME NUMBERS 1972:2 TOD F. STUESSY Academic Faculty of Botany, The Ohio State University, Columbus, 43210 JOHN L. STROTHER Department of Botany, The University of California, Berkeley, 94720 ASTERACEAE (COMPOSITAE) CALYPTOCARPUS VIALIS Less. n=12. Mexico, Nuevo Leon, Stuessy «& Renold 1280 (TEX); Mexico, Tamaulipas, Stuessy & Renold 1271 (TEX); Texas, Travis Co., Stuessy 1297 (TEX). These three counts substantiate the previous report of n—12 for this species (Turner, Ellison and King, 1961, Am. J. Bot. 48:221). SYNEDRELLA NODIFLORA (L.) Gaertn. (Fig. 1) n = 20. Nicaragua, Granada, Stuessy 617 (TEX). This taxon has been reported previously as n — 19 (Gajapathy, 1962, Proc. Ind. Acad. Sci. 55:72), 2n = 36 (Mangenot and Mangenot, 1962, Rev. Cytol. Biol. Veg. 25:431), n = ca. 20 (Turner in King, 1965, Phytologia 11:218), 2n = 40 (Harvey, 1966, Taxon 15:163), n — 20 (Mehra and Remanandan, 1969, Taxon 18:4388), and n = 19 or 20 (Powell and King, 1969, Sida 3:320). The present count of n = 20 further documents the 20 level in what may be an aneuploid series of n = 18, 19 and 20. SIDA 5(1): 32, 1972. BOOK NOTICES KEYS TO THE VASCULAR PLANTS OF THE BLACK GAP WILDLIFE MANAGEMENT AREA, BREWSTER COUNTY, TEXAS. Wm. F. Mahler. Third edition, 1971. 109 pp. Paperback. SMU Book Store, Box 759, Dallas, Texas, 75222. $3.00 ($3.65 postpaid).—This paperback edition is the first edi- tion available for distribution. The first (1969, Ditto) and second (1970, Xerox) editions were produced in limited quantities for the Museum and Field In- vestigations Course sponsored by the Department of Biology, SMU and the allas Museum of Natural History. REEVES’ FLORA OF CENTRAL TEXAS. Robert G. Reeves. Second edi- tion. 1972. Paperback. Prestige Press, P.O. Box 2504, Fort Worth, Texas, 76125. $6.85 ($7.50 postpaid). SHINNERS’ SPRING FLORA OF THE DALLAS-FORT WORTH AREA, TEXAS. Lloyd H. Shinners. Second edition. Edited by Wm. F. Mahler. 1972. Paperback. Prestige Press, P.O. Box 2504, Fort Worth, Texas, 76125. $7.25 ($8.00 postpaid). SIDA 5(1): 53, 1972, LOWELL DAVID FLYR, 1937-1971 “OH GOD, WHAT A HOPELESS TEXAN AM I!” Letter from Oxford, England; 9 Jan 1965 David was a Texas botanist. He'd like that said first and underlined. Suckled on the dry teat of a panhandle town in Texas (‘‘Stratford, write Stratford’; his ghost is in me!), dying by his own hand at thirty three, we wept wondering why? For he was gifted; played Bach, picking it out on the piano precocious child (‘‘clearly genius’? his music teacher told me once, confiding in our aloneness, knowing I'd tell David later, and I did, laughing it over coffee; ‘‘sheit’? he said, drawn out long through grinning teeth, pleased, warmed to his listening, ‘I’m the greatest Turner; better watch your women.”’) That David, I used to think he’s smarter than us all, the way he carries his 1575 (highest-on-the-campus) GRE score in his hip pocket like it were nothing ’cept an old photograph out of focus someone took once, catching him bent over at the hips looking at Leucophyllums, Texas Sage (L. frutescens). He was a westerner born out of his age; ‘‘maybe I'd of made it years before ...’’, he remarked once, ‘‘just standing around wait- ing for time to catch up.” He’d make a little remark like that once a day or so and I’d be haunted for hours wondering what he’d meant. Big man; large boned; carrying fat sometimes up to maybe two fifty; and ater, a couple of months maybe, he’d show up sixty pounds off in an ivy league suit speaking impeccable English chuckling with that little, deep- throated dog whimper endemic to him, he’d say, ‘‘remember me Dr. Turner? V'm full of wonderful vacuums’’, referring to shock treatments for depres- sion he’d taken voluntarily because it’s good for the thesis, works out phylogenies, ‘‘didn’t you know my mind was programed Fortran, needed plugging in.” But it did; within two months after shock treatment he wrote his entire thesis, handing it to me intact, saying, ‘‘Here it is; you don’t have to read it; it’s fine.”’ I said, ‘‘ok, don’t get bitchy.” Glancing over it briefly, remark- ing of a sudden, ‘“‘I call that piece a wonder, now’, (remembering a line from Brownine’s Last Duchess: oh, David loved me for that; he was an intellectual first, maybe even only.) About his thesis: for six years David wandered in and about the Botany Building at the University of Texas at any hour irregularly, taking courses whenever he couldn't talk his way out of them; disappearing suddenly into the field for no reason ‘cept as how he wanted away, collecting whatever he couldn’t avoid, pressing mostly Leucophyllum at first, but switching by degrees to Brickellia because, as I put it, “it grows daffodil in desert rock and nobody really gives a damn but you.’’ And he gathered a lot of field data about his genus, keeping it all in his head, rarely, if ever, making notes, except in contempt; and he retrieved it all, six years of it, in two _ SIDA 5(1): 54. 1972 aH) months time starting from scratch he emptied his vacuum. I was astonished. But more than once! He astonished me on our first field trip. We were in Big Bend National Park (where he always took off to in moments of stress) and him not knowing beans about botany, ‘‘not even the name of a Com- positae?”’ I asked. “No, but I’m-a-willing to learn.’’ He talked that way informally (to me at least; I took it to be his desire for identity with dirt farmers and things of his childhood and his father sitting on a tractor and David right there walking barefoot understanding every- thing excepting why he’d died when he was four: after that memories, only memories and long furrows). Leaning back, wickedly arrogant, David would ask ‘‘what’s that?’’ at 30 miles an hour along the roadside, “Engelmannias, Lindheimeras, Berlandieras’’ I'd respond, “‘lotsa genera’. And I think, swear, he learned to recognize and name maybe 300 species on that one trip, bragging how he’d have my job some day except that he wanted Pro- fessor Shinners’ position at SMU. He was his real hero. It is appropriate then that David’s obituary be buried here in SIDA. Lloyd was the first to recognize David’s interest in botany; indeed, he instilled it. When Shinners suggested that he work under my direction for a Ph.D., Lloyd wrote to me separately stating, “If successful, the effort you put into the development of David as a taxonomist will be worth that effort you might put into at least ten other ordinary doctorates.’’ And whether this might have been so or not isn’t important. What is important to me is that J] am a better taxonomist, teacher, person; having known him I carry a deeper sense of joy, laughter and, what other word, tragedy. He stood the test well: deeply sensitive and proud of his heritage, I never saw him flinch, beg excuses, cover his eyes with shakey fingers. David wore his profound (psychotic) depression so lightly (to me it was always so) that most of us mistook it for cautious wisdom, or aloofness born of the prairie or whatever else he managed to make it through the day with (and I’m thinking now that when he first corrected my illiterate pronounciation of the word faéade he did so simply because that word was important to him personally). I never understood that side of David except possibly once, just once I perhaps came close. We were atop the University Tower looking over the city, ‘‘gaining perspective’ he said ‘‘sometimes when I’m depressed I come up here and gaze westward looking to the plains, seeking solace with my ancestors.”’ ‘But it doesn’t help.’ I said ‘‘David, it doesn’t help, I feel sick to my stomach, I’ve a barnacled anchor heavy on my shoulders, I can’t breathe, I have no perspectives, I’m depressed, for the first time in my life, David, I am deeply depressed.’ David looked at me with a kind of eye-sharing we’d not experienced before, and he said, quite slowly, as though I might only this once hear the message, ‘“‘Dr. Turner, that’s the way I feel all the time’. I cried, and I think he understood. Neither of us ever mentioned the mo- ment again. In Memory Leucophyllum flyrii Turner, sp. nov. HOLOTYPE (TEX, isotypes to be dis- tributed): ca 4.5 E of Laguna Seca, 29 Jul 1966, Flyr 1113. Species unica fohorum basi 38 mm. lato (aliae omnes maxime 2 mm). Shrubs up to 1.6 m tall, much and somewhat intricately alternately- branched, especially above; foliage dense and somewhat canopy-like; bark dark grey-brown, conspicuously rugose or ridged and furrowed on oldest branches, inconspicuously rugose above; petiole scars and leafless somewhat spiny lateral branchlets tending to persist on old branches; branches leaf- less, except for the terminal 6-10 em; densely pubescent when young, grad- ually becoming glabrous after several growing seasons; leaves alternate, densely crowded (internodes often less than 1 mm long), sessile, the leaf base often 1 mm or more across at attachment to stem; blade shape varying but most characteristically obovate-spatulate, broadest 1/5-1/4 of distance from apex and then narrowed very gradually with nearly straight sides to the base; distally: either rounded or somewhat angled from the widest por- tion to the obtuse or somewhat acute apex; mature blades (10-)12-19 mm long, 5-9 mm wide; midrib slightly raised on the lower surface except near the apex, not visible above; no lateral veins visible; blade rather thick and felt-like by the dense mostly closely appressed pubescence; pubescence of young stems, leaves, and pedicels very complex: of many erect, closely- crowded tufts, each consisting of a central axis with many radii (or each tuft, through suppression of the axis, merely a pile of these lateral hairs), in older leaves the tufts becoming obscured and the surface tending to ap- pear a tangle of very fine hairs; flowers alternate or sometimes subopposite in the leaf axils, borne on pedicels 2 mm long and conspicuously more pubescent than the calyx, which is divided nearly to the base and consists of 5 lance-oblong sepals, 4.5-6.0 mm long, with somewhat attenuate tips: corolla appearing funnelform when pressed but dorsiventrally compressed and appearing subcylindrical in lateral view, narrowly campanulate or funnelform in dorsal view, 5(-6)-lobed and somewhat 2-lipped, 3(-4) lobes below the middle sparingly pilose, the middle 1(-2) extended, the lateral somewhat reflexed, the upper two less separate from each other than from the others; whole corolla up to 25 mm long, the lobes ca 6-7 mm long, the tube within almost wholly beset with many very small and dark purple spots up to 1 mm across, but intermixed with these are some oblong, dark-colored nectaries, internally sparingly pilose on or near the anterior lobe; veins somewhat prominent on tube, about 3 per lobe; stamens 4 or sometimes 5, alternating with the corolla lobes, didynamous, the posterior pair longer, filaments ca 8-9 and 11 mm long as measured from base of corolla, adnate up to half their length to the corolla, glabrous; anthers up to 4 mm long; ovary somewhat compressed laterally, with small swellings (nectaries?) around base, short pilose (or hairs somewhat branched) near summit, locules unequal (anterior larger); style ca 11 mm long, sparingly pilose, o7 stigma bilamellate; capsule 4.5 mm high, woody but thin-walled (less than 0.2 mm in diameter). Distribution: one or two localities northeast of San Luis Potosi, S.L.P., Mexico. Leucophyllum flyrii is known from a small area a few miles east and northeast of the city of San Luis Potosi. F. W. Pennell first discovered it at San Pedro which lies in low hills about 12 miles east of San Luis Potosi. As is true of other desert mining areas, very little woody vegetation remains in the area of San Pedro. I was unable to find L. flyrii there. Rzedowski, who collected San Luis Potosi rather thoroughly, made two collections of this species east of Laguna Seca, which lies a few miles north of San Pedro. After one unsuccessful search, I obtained from Dr. Rzedowski (in litt.) a detailed locality and was able to find the shrubs in flower. The very restricted range of this species suggests two possibilities: that it is limited by some edaphic factor, or that its present range is but a frag- ment of a larger former range. The species occurs at the southeastern edge of the more or less continuous distribution of Leucophyllum over the desert plateau of northern Mexico. To the south and east, increasing rainfall, with or without increasing elevation, allows development of a vegetation that does not include Leucophyllum until an isolated semi-arid area in Queretaro and Hidalgo is reached. Specimens Examined: MEXICO. San Luis Potosi: San Pedro, Sierra Madre Oriental, 2150-2200 m, 29 Jul 1934, Pennell 17735 (BM, F, MICH, NY); 8 km NE of Laguna Seca, 2250 m, 30 Aug 1955, Rzedowski 6335 (TEX). Except for the specific name, I have culled this newly described species of Leucophyllum, intact as to wording, from Dr. Flyr’s unpublished doctoral thesis. To me it seems appropriate that one of the taxa in this genus bear his name. Commenting to me about Leucophyllum once, he said, ‘‘we are lovers’. I liked that attitude and that’s why I’ve wed them here. Born: Lowell David Flyr, Stratford, Texas 24 Nov 1937 Parents: Bonnie Burns Almon Flyr and Lewis Anthony Flyr (deceased) Education: Attended public schools in Cleburne and Denton, Texas Graduated, Denton High School, 1956 B.A.—Southern Methodist University, 1960 Ph.D.—University of Texas, 1970 Honors: 1960—Phi Beta Kappa at SMU 1960—Recipient of the Avella Winn Hay award, as outstanding graduating male student 1960—Recipient, Research Award, Dallas Association of Phi Beta Kappa 1970—Research Fellow, Harvard University Died: by his own hand, Woodlawn Hospital, Dallas, Texas 2 Nov 1971. Buried: Stratford, Texas 38 Publications: 1966 Chromosome numbers in the Compositae. X. North American Species. Amer. J. Bot. 53:24-33 (with B. L. Turner). 1967 (Compositae). Sida 3:252-256. New names and records in Brickellia 1970 A systematic study of the tribe Leucophylleae (Scrophulariaceae). Doctoral Dissertation, The University of Texas, Austin. 1971 Thelesperma shinnersii Flyr (Compositae), a new species from Coahuila, Mexico. Sida 4:276-277 Dyssodia tenuiloba (Compositae): New to Mississippi. Sida 5(2). In press. B. L. Turner Department of Botany University of Texas Austin, Texas 78712 SIDA SSsonnes VOLUME 5 NUMBER 2 MAY 1973 CONTENTS Sex and the angiosperms. John W. Thieret. 59 Taxonomy of Perityle section Laphamia (Compositae-Helenieae- Peritylinae). A. Michael Powell. 6] Acknowledgments 61 Chromosomal considerations 62 Chromatographic considerations 63 Hybridization considerations 64 Morphological considerations 65 Phylogenetic considerations 68 Species concept TE Taxonomy 78 Excluded species 127 References 127 NOTES: Isoetes louisianensis (Isoetaceae), A new species from Louisiana. 129.—Dyssodia Thymelaeaceae): New to Louisiana. 132—-A new species of Frankenia (Frankeniaceae) from gypseous soils of north central Mexico. 132—Comments on New Mexico flora. 135. NOTICES 136 SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. Mahler, SMU Herbarium, Dallas, Texas, 75275, U.S.A. Subscription price $8 (U.S.) per volume; parts issued twice per year. ASSOCIATE EDITOR John W. Thieret University of Southwestern Louisiana Lafayette, Louisiana © SIDA Contributions to Botany, Volume 5 Number 2, pages 59—136 Copyright 1973 by Wm. F. Mahler SEX AND THE ANGIOSPERMS JOHN W. THIERET University of Southwestern Louisiana Lafayette, Louisiana 70501 Discussed in this essay is the almost universal application of sexual termi- nology to sporophytic structures of angiosperms. Sex in plants was not well understood until the latter part of the 19th century. Prior to that time, the use of sexual terminology in botany was conjectural—and was not noted for unfailing accuracy. Its misuse then does not justify its misuse now. Sexuality in plants, as it applies to alternation of generations, is well understood today; sexual terminology should be restricted to plants and plant structures that are morphologically sexual. The use—rather, misuse—of sexual terminology for sporophytic structures of angiosperms has long been rife with botanists. As a starting point for dis- cussion, however, we might select the Linnaean ‘‘Sexual System,’ a product of fanciful analogy between vegetable and man!‘ and of an era antedating factual knowledge of sex in plants. This system is now of historical interest only except for its attribution of sex to sporophytes, a concept to which, even in the enlightened 1970’s, many botanists tenaciously cling. Among the lower embryophytes—the bryophytes and pteridophytes—the use of terms indicative of sex is confined to the gametophytic phase. Thus, for example, we speak of a gametophyte as being male, female, or bisexual (hermaphroditic); we speak of an antheridium as being a male structure, of an archegonium as being a female structure. Among these plants, we do not apply sexual terms to the sporophytic phase. Such restriction of sexual terminology to gametophytes is not the case, however, with the ane IOSpeTNS: Not only is the gametophytic phase de- scribed with sexual termino logy .g., the embryo sac is female—but, by an astonishing extension of Heeaae. so is the sporophytic phase. Among the sporophytic structures thus endowed with sex are flowers (‘‘male,’’ ‘‘fe- male,’ ‘“‘bisexual’’), stamens (‘‘male’’), carpels (‘‘female’’), and even plants (‘‘male’’ and ‘‘female’’). The use of sexual terminology for sporophytic structures is, on occasion, even more ludicrous: in an otherwise eminently respectable journal I recently noted the terms ‘‘male sepal’’ and ‘“ pedicel.’”” Sexual terminology for sporophytic structures is not only misleading, in- consistent, and inaccurate but also superfluous, referring to precisely the same concepts as do the following terms, which are, it seems to me, above We all know about the Swedish attitude toward sex. >To extend this absurdity even further, why not “male” or “female” vessel elements or stomates, And one ee wonder if, at base of a ae that bears “male pedicels,” one might find ‘‘male root SIDA 5 (2): 59—60. 1973. 60 reproach: staminate (‘‘male’’), carpellate (‘‘female’’), imperfect (“‘uni- sexual’), and perfect (‘‘bisexual’’). By extension of the same reasoning (or lack of it) that permits the use of sexual terminology for flowers, the sporophytes of Selaginella and Marsilea could be called ‘‘bisexual’’ and the megasporangia and microsporangia of Selaginella could be called, respectively, ‘“‘female’’ and ‘‘male.” The correct and consistent use of sexual terminology for angiosperms will lead to increased exactness in one minuscule part of our taxonomic vocabu- lary. Taxonomy may never be an exact science but this does not excuse taxonomists from striving diligently for terminological precision. TAXONOMY OF PERITYLE SECTION LAPHAMIA (COMPOSITAE-HELENIEAE-PERITYLINAE A. MICHAEL POWELL Department of Biology, Sul Ross State University, Alpine, Texas. Section Laphamia comprises an assemblage of rock-dwelling species rang- ing from western Texas to southern California, throughout the southwestern United States, and northern Mexico. Thirty taxa are recognized, twenty-one species and nine varieties. Laphamia was described originally as a genus by Gray (1852) who noted its resemblance to Perityle. Several other investigators have been concerned with the close relationship of traditional Laphamia and Perityle, and several interpretations regarding generic status have evolved from their morpho- logically oriented studies (Shinners, 1959). Most workers have agreed that the only semi-consistent difference between Laphamia and Perityle is the presence of well-developed pappus squamellae in the latter and an absence of pappus scales in the former. I have proposed elsewhere that Laphamia and Perityle are best treated as sections of a single genus, based upon the evaluation of information from cytological, chromatographical, and hybrid- ization techniques (Powell, 1968a). A third section, sect. Pappothrix, is also recognized for Perityle (Powell, 1969). The closest generic relatives of Perityle are believed to be Pericome, Eutetras, and Amauria of the subtribe Peritylinae. Presently the Perity- linae is being re-evaluated with reference to Rydberg’s (1914) original con- cept of the phylad, and few other small genera with more distant relation- ship to Perityle probably will be added to the subtribe (Powell and Turner, unpublished). In fact, Peritylinae properly belongs with another tribe of Compositae, either Senecioneae or Heliantheae. ACKNOWLEDGMENTS I am deeply grateful to B. L. Turner for contributing in many respects toward the completion of this work. He originally directed my attention to the genus and has since provided invaluable systematic counsel. I am also grateful to students Sam Sikes and Dicke Patterson who have contributed much more than perfunctory technical assistance in the studies of Perityle. I wish to acknowledge gentleman-explorer Pave Smith of Taft, California, for facilitating the collection of P. inyoensis. I am further indebted to the late Lloyd H. Shinners for providing the Latin diagnoses included herein; to John Averett and Tom Watson for collecting bud material of P. stansburii and P. megalocephala; and to Tom Todsen and Rich Spellenberg for pro- viding new information about P. staurophylla. ' Supported by NSF Grants GB-2400, GB-5046, and GB-7740. SIDA 5 (2): 61—128. 1973. 62 My special appreciation is extended to the patient curators of herbaria from which specimens were borrowed. CHROMOSOMAL CONSIDERATIONS Chromosome numbers are available for 19 species of sect. Laphamia (v—17). Counts for most taxa have been recorded elsewhere (Powell, 1968b; Powell and Sikes, 1970). Chromosome numbers for P. stansburii “ 17), P. gracilis (n=—34), P. megalocephala var. oligophylla (n— 34), and P. megalocephala var. intricata (n—19+-1) were not previously reported eee 1). It has not been possible to obtain accurate counts for P. villosa and P. cochuilensis. Perityle grandifolia and P. gentryi have not been counted. TABLE 1. Species of Perityle sect. Laphamia examined for chromosome number. “Species mn Number ~ Location and Voucier P. gracilis 34 NEVADA: Lincoln Co. 4% mi N of Caliente. Powell and Patterson 1626. P. gilensis var. ca. 17* ARIZONA: Gila Co. Salt River Canyon. salensis S. Sikes 428. P. megalocephala ca. 19" NIVADA: Nye Co. 23 mi W of Indian var. intricata Springs. Watson and Averett 250 P. megalocephala 34 CALIF: Inyo Co. Lost Burro Gap, var. oligophylla Cottonwood Mts. Powell and Patterson 1612. P. stansburii 17 UTAH: Beaver Co. 1:5 mi EF of Minersville, Averett and Watson 422. P. stansburii ca. 17° UTAH: Tooele Co. 1.6 mi NE of Low, Watson 332. 1, wound fragment or panes observed in some cells. * Actually recorded ; 19 Tae 1; “In most cells aber 17 Il + 2 7; in few cells also observed 17 Il, 16 IT + 2 1, 17 1 + 4/1, 18 Il. The base chromosome number of sect. Laphamia is established as x—17. All but three species are on the base of x=17. Perityle inyoensis has been counted as a ‘‘questionable’? n=—18, and P. tenella and P. congesta are n= 16 (Powell, 1968b). The var. intricata (Table 1) is reported above to be n —19-+-1, and may be a fourth exception to the x—17 base, but its meiotic configurations were observed to be abnormal (17-20 II and 2-7 1). Regular meiosis was observed to be characteristic of polyploid populations (Powell, 1968b), with but a few exceptions. Meiosis in three individual plants from one population of P. megalocephala var. oligophylla (n—34) showed 1-3 multivalents (rings or chains of 3-4) and 2-4 associated but asynaptic univalents. One to five lagging univalents were common at anaphase I. No exact chromosome number was discernable in one population (Averett 418) of P. stansburii («—17), but meiotic behavior resembled that of a triploid. Approximately 20 Il and 6-12 I were commonly observed. Perityle coahui- 63 lensis tentatively was determined to be n=27-34 II, with irregular multi- valent and univalent behavior preventing an accurate count. Triploid meiotic behavior also was observed in P. villosa (tentatively n=ca. 17 II and 17 1). Meiosis was observed in plants from one population, and in a single plant from the same collection which was grown in the greenhouse. The number of apparent univalents varied from 3-19, the latter often being difficult to distinguish from bivalents because of bivalent heteromorphism. It is assumed that the plants of P. villosa examined are recent triploids. No direct evidence is available on the precise origin of polyploid popula- tions in sect. Laphamia. Autoploid origin has been assumed as a working hypothesis in most cases because polyploids generally are morphological reflections of diploids. Also to be considered is the fact that meiosis is normal in a large majority of the known polyploid populations. Certainly no reliable morphological or cytological evidence of alloploid genome constitu- tion has been detected in any polyploid taxon of sect. Laphamia, with the possible exception of P. coahuilensis. This subject is under experimental study CHROMATOGRAPHIC CONSIDERATIONS Simple pattern data from 2-dimensional paper chromatograms have been utilized in studying relationships among species and species groups of sect. Laphamia. The current investigations have corroborated preliminary studies which showed that most species of sect. Laphamia produce species-specific patterns (Powell and Tsang, 1966), although closely related taxa character- istically produce patterns which are much alike. General techniques have followed those outlined by Alston and Turner (1963), except that pure methyl alcohol was employed as an extracting solvent. Observations were made under ultra-violent light, with and without the presence of ammonia vapor. The major components have been identified as flavonoids (Powell and Averett, unpublished). Specific identity of several compounds is under study. It is deemed beyond the scope of this paper to present detailed chromato- graphic data in the form of figures or tables, even where such evidence has been helpful in understanding putative relationships. Instead, where appro- priate, conclusions drawn from comparative pattern data are presented in discussions. It might be argued that the explanations of ‘‘Chromatographic Considerations’? should be omitted from this work since the specific data upon which they are based have not been included. However, use of the “pattern character’? as a monographic tool is relatively innovative, and it is important that the taxonomic basis of its application be understood. It is strongly believed that simple chromatographic pattern data are best treated as merely another taxonomic character, and appraised as such in toto along with all other sources of information. Chromatograms of leaf extracts have been developed for 19 of the 21 species recognized for sect. Laphamia. No recent collections of P. grandi- folia or P. gentryi have been available. For each species, chromatograms 64 were produced from one to several individual plants and/or populations, depending upon the availability of fresh material. Considerable precautions were taken to ensure consistency in technique, and thus pattern variability is believed to represent primarily that which is intrinsic for each taxon. For the most part, systematic relationships inferred from chromatographic information have corresponded with affinities judged from morphological, chromosomal, and other data. Pattern analyses of taxa comprising related- species groups have been most useful. For example, seven geographically proximal species of the southwestern United States have been assumed to represent a monophyletic group on the basis of their similar morphologies. Five of these species appear to be the most closely related and have chrom- osome numbers of x—17 (n=ca. 18 in P. inyoensis). A base number of x—16 for the two remaining species (P. congesta and P. tenella), however, suggests the possibility of other kinship. Chromatographic analysis of all seven species has revealed a chemical profile which sets them apart from the rest of the sect. Laphamia. Furthermore, the chromatographic pattern character of each species of this ‘southwestern alliance’ corresponds exactly to presumed interspecific relationships which have been arrived at through morphological studies. Other morphologically founded species groups, such as P. lemmoni, P. dissecta, P. castillonii—, P. lindheimeri, P. angusti- foia—, and P. bisetosa, P. warnockii, also have been found to exhibit chemical integrity. The total chemical data gathered to date, especially the crude pattern data, are not yet sufficient to allow sweeping conclusions as to infrasec- tional and intersectional phylogeny within Perityle. HYBRIDIZATION CONSIDERATIONS Natural hybridization does not appear to have been an important factor in the evolution of sect. Laphamia. Perhaps this is simply because none of the species of this section are known to occur sympatrically. Nevertheless, artificial interspecific hybrids between numerous species have been obtained readily in the greenhouse, and thus there seems to be a general absence of strong genetic barriers to interspecific hybridization within the section (Powell, unpublished). Artificial hybrids between several species belonging to different sections of Perityle also have been produced in the greenhouse, and in some cases with rather amazing case. Examples of experimental intersectional hybrids in- clude the following combinations: P. bisetosa X P. rupestris (sect. Pap- pothrix); P. castillonit X P. vaseyi (sect. Perityle); P. quinqueflora (sect. Pappothrix) X P. vaseyi. Numerous other successful intersectional combi- nations could be cited. A few species belonging to separate sections of Perityle are known to have sympatric distributions. In the Chisos Mts. of Texas, natural intersectional hybridization involving P. rupestris var. albi- flora (sect. Pappothrix) and P. parryi (sect. Perityle) has been detected, and this phenomenon now has been documented (Powell, 1970). 65 Surprisingly enough, artificial intergeneric crosses also have been success- ful. Examples include Eutetras palmeri X P. cernua (sect. Pappothrix), E. palmeri X P. jaliscana (sect. Perityle), HE. palmeri X P. parryi (sect. Peri- tyle), and Pericome caudata X Perityle saxicola. Evidence oriented toward utilization of hybridization data in evaluating systematic relationships among the species of sect. Laphamia still is being accumulated. Experimentally, at least, closely related species of this section are so interfertile that several generations may be required in order to judge degrees of genetic affinity on this basis. Information about the general interfertility of species in Perityle has given reason to suspect the possibility of hybrid origin for several taxa of sect. Laphamia which have nebulous systematic affinities. Somewhat intermediate morphology, or other classical manifestations of possible hybrid involvement, has been detected in P. staurophylla, P. coahuilensis, P. cochisensis, P. villosa, and a population of P. lemmoni. Since all species of this section are mutually isolated geographically, it would appear that the mere breakdown of spatial barriers would be sufficient to allow hybridization. However, in all of the above suspected cases of hybridization, evidence of recent popula- tional confluence between putative parents is lacking. As a matter of fact, evolution of the peculiar rock-bluff habitats theoretically represents a rather permanent deterrent to secondary sympatry, at least with regard to ‘‘recent”’ time connotations. As mentioned earlier, a few species belonging to different taxonomic sections are known to occur sympatrically, but natural hybridi- zation has been detected in only one instance. This suggests that repro- ductive barriers between taxa of separate sections do exist under natural conditions even though hybrids can be synthesized in culture. Markedly low fertility is known to be characteristic for most artificial F, intersectional hybrids (Powell, unpublished). Still, the P. rupestris X P. parryi intersec- tional hybrid points to what could have happened in the course of Perityle evolution. It is evident that the possibility of rather ‘‘ancient’’ hybridization within the genus should not be overlooked in future systematic studies of the Peritylinae. MORPHOLOGICAL CONSIDERATIONS Numerous morphological features are useful in circumscribing taxa and evaluating systematic relationships among species of sect. Laphamia. Most notably these include habit, pubescence, leaf shape and size, capitulescence, head size and number of florets, flower color, presence or absence of ray florets, and achene-pappus morphology. The habits of most species are influenced to a large extent by their rock- bluff habitats. Plants may be erect, pendulous, or closely appressed to rocks, depending at least partially upon the angle of bluff exposures. Bluffs which are nearly perpendicular usually support plants which are somewhat pen- dulous, unless the stems are very short as in P. bisetosa, while in nearly horizontal exposures plants tend to become semi-erect or spreading. Both 66 extremes in habit often are exhibited by a single species. The height (or length) of plants varies tremendously, from approximately 2 cm in P. bise- tosa to about 50-60 cm in several species such as P. lindheimeri. Wide in- fraspecific variation in plant height is not uncommon. Unquestionably, sun exposure has an affect on plant size when exposure is a variable factor in a particular habitat. As would be expected, plants exposed to full sun for several hours per day characteristically are more diminutive than those which grow under relatively shaded conditions, Older plants of some species like P. staurophylla and P. megalocephala tend to become profusely branched and spread to widths approaching one meter, while in other species such as P. warnockit, plants have not been observed to exceed a diameter of ap- proximately 10 cm. The strong suffruticose or suffrutescent perennial con- dition exhibited by all species is judged to be an ancestral trait in Perityle. The only annual taxa of the genus are members of sect. Perityle. Pubescence appears to be significant in only a few species. Taxa such as P. bisetosa, P. staurophylla, and P. coahuilensis are essentially glabrous, while P. villosa, P. inyoensis, P. dissecta, and P. gentryi are characterized by pilose-villous pubescence. Most species are variably short-pubescent and the indument does not appear to be important taxonomically. Leaf shape and size are extremely variable among the species of sect. Laphamia. Even infraspecific variation is considerable in some taxa, but leaf morphology generally is the most important taxonomic character it this section. The smallest leaves are found in P. bisetosa (ea. 0.5-1.0 em long) while P. grandifolia produces the largest leaves (8-11 em long). The latter species, along with a few others, have essentially ovate leaves with entire to serrate or shallow-lobed margins. Leaf variability in sect. Lapha- mia is expressed to its greatest extent in the degree of margin dissection. The specific epithet of P. dissecta is particularly descriptive of one extreme condition where margins are irregularly cleft and divided. Perityle saxicola has leaves which are bi- to tripinnate with long, narrow segments. There is a strong tendency toward tripartitioning of leaves in taxa of this section, as evidenced especially by P. staurophylla, P. gilensis, P. gracilis, and P. coa- huilensis. In fact, 17 of the 21 species display tripartite leaves to some ex- tent, from shallow to deeply lobed. The capitulescence of most species is of solitary heads on short or long peduncles, depending upon the taxon being considered. In several taxa, how- ever, as in P. angustifolia and P. megalocephala var. oligophylla, the heads are arranged in loose clusters of two to four. The capitulescences of P. lindheimeri, P. staurophylla, and P. megalocephala var. intricata are of rather tightly clustered small heads (corymbiform), and are unique in sect. Laphamia with regard to this character. The corymbiform condition prob- ably is primitive in P. lindheimeri and P. staurophylla, but the feature ap- pears to be derived in P. megalocephala var. intricata. As might be assumed, head size varies rather consistently in correlation with the number of florets per head. Smallest heads are found in the Texas _ 67 species, P. bisetosa, P. warnockii, and P. lindheimeri (ca. 9-12 florets) and the largest heads are produced in the Arizona species P. saxicola and P. gilensis (ca. 100-200 florets). Most species have about 25-70 florets per head. The species of sect. Laphamia have yellow flowers, with the exception of P. bisetosa and P. warnockii which have white flowers. In four of the yellow-flowered taxa, P. dissecta, P. castillonii, P. lemmoni, and P. coahut- lensis, the disc corollas tend to become purple tinged at maturity. Flower color has limited infrasectional taxonomic importance simply because so few species differ in this trait. However, it does appear that ray and disc floret coloration will contribute significantly toward evaluating intersectional and intergeneric phylogenies within the subtribe Peritylinae. Preliminary hybridization studies have borne out the latter contention (Powell, unpub- lished). Normally, radiate heads characterize only seven species of this section: P. grandifolia, P. gentryi, P. stansburii, P. lindhetmeri, P. saxicola, P. gilensis, and P. staurophylla. A few specimens from an atypical population of P. lemmoni have ray florets, but this species otherwise is entirely dis- coid. Also, a single ray floret was formed in one head and two in another head of a plant of P. coahuilensis which was grown in the greenhouse. All field collections of the latter taxon are discoid. The ligules are yellow in all species. Achene and pappus morphology are considered to be among the most im- portant taxonomic features in sect. Laphamia. Generally speaking, species of the section are characterized by an absence of prominent pappus squa- mellae and by short-pubescent margins of flattened achenes, although these features are not absolute with regard to intersectional delimitation (Powell, 1968a). Achene structure per se is somewhat less valuable at the interspecific level than is the more variable pappus structure. In sect. Laphamia (and sect. Perityle), the pappus may be comprised of two parts, bristles and squamellae. Bristles are produced characteristically in 15 species, while in five species bristles are absent (or only rarely found). In taxa which usually have bristles, the actual number on each achene may vary from 0-2 (3-6). In fact, population studies have shown that variability in bristle number is typical of most species. For example, P. castillonii is characterized by achenes with 1-2 bristles, but some plants produce achenes with 3 bristles, and some populations are epappose. In one population of P. staurophylla, achenes usually are bisetose, but many plants were observed to have achenes with 3-6 bristles. Bristle number does appear constant in a few species, however, such as the wide-ranging P. stansburii in which only unisetose achenes have been observed. Judging from all correlated evi- dence, and with some reliance on the axiom that plant parts are more easily lost than gained, the presence of bristle(s) is thought to be an ancestral trait in sect. Laphamia. Certainly, the epappose species seem to have been derived from pappose ancestors, with the possible exception of P. grandi- folia. Best examples of the latter contention are expressed by the epappose 68 species P. angustifolia and P. megalocephala, which clearly seem to have been derived from P. lindheimeri and P. stansburii respectively. A crown of rather inconspicuous, hyaline squamellae, usually supported by a thin, callous crown, occurs as part of the pappus in some species of sect. Laphamia. In sect. Perityle, achenes of nearly every species have relatively prominent crowns of squamellae. Laphamia and Perityle have been separated as genera primarily on the basis of this character (Shinners, 1959; Powell, 1968a). There is no certainty that the crowns of squamellae in each section are homologous structures, but it seems most logical that those in sect. Laphamia represent reduced or vestigial forms derived from the prominent scales of sect. Perityle species. Careful examination has not re- vealed any morphological difference, other than size, between the reduced squamellae of sect. Laphamia and the prominent crown in sect. Perityle. It should be emphasized that even vestigial pappus squamellae are not evi- dent in all species of sect. Laphamia. Also, the small scales may be absent in some populations, individual plants, or even in some heads of species which normally exhibit squamellae. A thin callous crown remains on the achenes of plants in which no squamellae are found, and possibly should be regarded as the ultimate in reduction of a pappus crown. PHYLOGENETIC CONSIDERATIONS No clear picture regarding the evolutionary history of sect. Laphamia can be presented at this time. Partly this is because intersectional relation- ships within Perityle still require further study, but also because there is some indication that sect. Laphamia, as treated here, actually is a poly- phyletic assemblage. It appears significant, however, to explore some of the most reasonable conclusions which can be drawn on the basis of present knowledge about the section. Currently, it seems logical to assume that sect. Laphamia, at least in part, had its origin from sect. Perityle. Certainly the close relationship of the two taxa is reflected in their overall morphological similarity. That sect. Laphamia was derived from sect. Perityle, instead of the other way around, is suggested by the several characters of sect. Laphamia which appear re- duced from pre-existing structures in species of sect. Perityle. The most consistent of these reduced characters are the vestigial (or absent) pappus squamellae and short-pubescent achene margins, but other such features are revealed in succeeding discussions. All but two species of sect. Laphamia have yellow flowers. Perityle bise- tosa and P. warnockii have white flowers. The predominance of yellow- flowered species, along with other information, supports an assumption that the section had yellow-flowered ancestry, although this apparently does not apply to P. bisetosa and P. warnockii which may have originated from a white-flowered member of sect. Pappothrix. Ray florets normally are pro- duced in only seven species of sect. Laphamia. The ligules are yellow in all seven species, again suggesting that the section, at least in part, had its 69 origin from a yellow-flowered ancestor. Since sect. Laphamia presumably arose from sect. Perityle, ray coloration in that section might be expected to provide an important phylogenetic clue. All but three of the 24 species currently proposed for sect. Perityle (Powell, unpublished) have radiate heads. The ligules of 13 species are white, and nine taxa have yellow ligules. Among the nine species with yellow ray florets, all but four taxa can be eliminated as likely ancestors on other grounds (such as annual habit), thus considerably narrowing the search for a phylogenetic antecedent of sect. Laphamia. The remaining four taxa are P. palmeri, P. leptoglossa, and P. cordifolia which are found near the foothills of the Sierra Madre Occidental in northwest Mexico, and P. parryi which is distributed along the Rio Grande of West Texas. There is no attempt to place undue emphasis on the presence or absence of ray florets and flower coloration in evaluating the systematics of sect. Laphamia. It is expected that further studies, particu- larly of sect. Perityle, will result in an accumulation of additional evidence upon which to evaluate the above phylogeny Species of sect. Laphamia are distributed entirely in various mountain or rock formations at altitudes ranging from ca. 8500 ft. to ca. 3000 ft. Plants grow only in crevices of bluffs or boulders, or almost any type of exposed rock formation of considerable mass and stability. Occasionally a few plants of some species (e.g., P. gilensis and P. megalocephala) have appeared to be growing in soil at the base of bluffs, but further examination always has shown them to be rooted in subterranean crevices. As stated earlier, as far as is known, all species of sect. Laphamia are allopatric in distribution. Listed in Table II are characters which I consider to be primitive and advanced in sect. Laphamia. In attempting to evaluate such features, refer- ence to similar studies in the Helenieae by Carlquist (1956), Ellison (1964), Ornduff (1966), and in the family Compositae by Cronquist (1955), have been most helpful. No effort has been made to concoct a hypothetical ‘‘primitive species’’ for the section because of the likelihood that the taxon is poly- phyletic. Suspected primitive species are brought out in subsequent dis- cussions The center of origin for Peritylinae probably was in the Sierra Madre Occidental of northwestern Mexico, and the evolution of its constituent taxa probably had some correlatien with the proliferation of the Madro-Tertiary Geoflora (Axelrod, 1958). The Mexican distribution of Eutetras and Peri- come, genera considered to be ancestral to Perityle, and the distribution of supposedly primitive species of Perityle itself, appear to be supportive of the above genesis (Powell, unpublished). Presently, the native species of Perityle are fairly widely distributed in western North America, from Jalisco, Mexico, to central Texas, north- western Utah, and Baja California, Mexico (Fig. 1). A South American disjunct of the genus, P. emoryi (Closia Remy), occurs in Chile. Raven (1963) cited P. emoryi as one example in contending that most of the dis- juncts between North and South America may have attained their distri- 70 TABLE II. Primitive and Advanced Characters in Sect. Laphamia Primitive Character Advanced Character 1. Plants relatively mesophytic............ Plants xerophytic 2. Leaves serrate or shallow-lobed....Leaves lobed, cleft, or divided 3. Herbage pubescent Herbage puberulent or glabrous 4. Phyllaries numerous Phyllaries few D). Ray florets present (yellow)............ Ray florets absent 6. Disk florets numerousS.... -......0.0..0...... Disk florets few 7. Achenes slightly rounded or angular in cross section Achenes strongly flattened 8. Achenes pubescent... Achenes glabrous 9. Pappus or bristles and vestigial |Pappus reduced, especially squamellae squamellae 10. Achenes pappose Achenes epappose 11. Chromosome number n—17 (18)....Chromosome number below n—17 or hig ee ree polyploidy bution during the late Pliocene or Pleistocene, Since P. emoryi is considered to be a derived species of Perityle, one could assume that evolutionary divergence of the genus might have been underway before or during the Pliocene. It also follows that the relatively widespread dispersal of Perityle might not have occurred until late Pliocene or Pleistocene, because as Sharp (1953) and Dressler (1954) have suggested, the most propitious time for migration of temperate plants was during the Pleistocene. Evidence pre- sented by DeJong (1965) in favor of Pleistocene dispersal and evolution of Astranthium (Astereae) parallels other reasons for suspecting similar evo- lution in sect. Laphamia. These reasons are that (1) nearly all of the species are endemics, (2) related species occupy adjacent ranges, and (3) ‘‘most species share genic material, as indicated by artificial crosses, suggesting recent evolution.” Judging from present day distribution, it is possible to assume that sect. Laphamia diverged early from its presumed ancestral stock in sect. Perityle (Fig. 1). The northern distribution limits of Perityle, in northwestern Utah, are occupied by species of sect. Laphamia. Species of sect. Perityle are not known to extend past central Arizona (e.g., P. ciliata), and these species of sect. Perityle are not the most closely related to sect. Laphamia. Further- more, most species of sect. Laphamia are distributed in drier mountain regions to the north and east of their supposed ancestral birthplace in the more mesic mountains of northwestern Mexico. The bulk of the species of sect. Perityle still reside in the mountains of Mexico. Because of the ap- parent “‘close’’ relationship between sect. Laphamia and sect. Perityle, early divergence of the two phylads is not easy to reconcile, unless sect. Laphamia experienced relatively rapid evolution. It does seem likely that rapid evolution of sect. Laphamia would have been possible under Pleisto- cene climatic conditions, especially if its genome manifested some special “Kr Fig. 1. Distribution of the genus Perityle: sect. Pappothrix (shaded); sect. Laphamia (dotted); sect. Perityle (crosshatched). tendency for adaptation to drier environments. To some extent the biological evolution of Perityle species must have been correlated with post annis geological evolution of mountain systems through- out the southwestern United States and northern Mexico. To attempt critical evaluation of species-mountain evolution at this time seems prohibitive be- cause of the tremendous area to be studied, but some general observations concerning the subject seem pertinent. Many species now comprising sect. Laphamia might have emerged from a wide-ranging ancestral taxon which had populational segments become geographically isolated by ‘‘ancient’? erosion, or the phylad might have originated from a gradual expansion of pioneer populations into available habitats. Quite naturally, the latter statement represents an ambivalent quandary which has become an overriding problem in understanding phy- logeny within Perityle. Regardless of the actual beginning of sect. Laphamia, the successful evolution of many species must have been associated posi- tively with the geological evolution of new habitats, i.e., the formation of exposed rock bluffs. Essentially the same conditions were explained by Raven (1964) to have had a general influence on the evolution of plant species in the desert southwest where the encroachment of relatively xeric environments upon relatively mesic environments (and subsequent edaphic factors) provided marginal populations an opportunity for ‘‘catastrophic selection.’”’ When considering two or more related species of sect. Laphamia, often it is difficult to determine which actually occupies the driest habitat, but as a general rule it appears that derived species are associated with the most xeric environment. In discussing catastrophic selection with regard to closely rated pairs of species in Clarkia (Onagraceae), Lewis (1962) found that the derivitive species invariably occupy more xeric habitats than their progenitors. This also seems to be the case in sect. Laphamia where closely related pairs of species are common. Although the restricting rock- bluff habitat of sect. Laphamia species necessitates a somewhat modified interpretation of catastrophic selection and mesic to xeric adaptation, Raven’s and Lewis’ concepts appear to be particularly relevant to under- standing evolution in the section. The restricted habitat of the species of sect. Laphamia is thought to be an extremely important consideration in understanding speciation in the phylad. Several groups of related species (and varieties) offer evidence that geo- graphic isolation is a primary “‘mechanism’’ for speciation in sect. Lapha- mia, Beaman (1957) pointed out the importance of this type of spatial isola- tion in the infrageneric evolution of Townsendia (Astereae). Among the taxa which appear to exemplify this phenomenon in sect. Laphkamia are the clearly related P. lindheimeri and P. angustifolia, which occur in west- central Texas. Perityle lindheimeri occupies rather protected, moist lime- stone canyons in heavily vegetated areas of the Edwards Plateau. Perityle angustifolia is found in drier, more eroded and exposed caprock areas along the western edge of the Edwards Plateau. Morphologically, it appears that P. angustifolia evolved from P. lindheimeri as the former became isolated and adapted to drier habitats. Perityle bisetosa and P. warnockii, also found in dry areas of western Texas, provide another example of speciation fol- 73 lowing geographic isolation. These species are clearly related morphologic- ally, and occupy disjunct but similar limestone habitats. Four varieties of P. bisetosa have been recognized. All of the infraspecific taxa are distinctly isolated in severely eroded, small-mountain areas. One needs merely to examine the localities of other Laphamian taxa to find additional instances where speciation apparently occurred subsequent to geographical isolation. It is quite possible that polyploidy has played an initial role in speciation of some Laphamian taxa. Several isolated species and infraspecific popula- tions are polyploids (Powell, 1968b; Chromosomal Considerations), but in each particular case it is difficult to determine if an increase in ploidy level was an initial or subsequent isolation barrier. In fact, polyploidy is such a widespread phenomenon in the whole genus, that, more than likely, both sequencies of events have occurred. It is evident from previous compilations of chromosome numbers that additional cytological studies will contribute significantly to future systematic investigations in Perityle. There are five related-species groups which can be recognized in sect. Laphamia. The largest of these supposedly natural assemblages of taxa is comprised of seven southwestern species, distributed from California to northwestern Utah and northwestern Arizona. For easy reference this group has been designated as the ‘‘southwestern alliance,’’ and it includes P. villosa, P. inyoensis, P. megalocephala, P. stansburii, P. gracilis, P. tenella, and P. congesta. The four other related-species groups are: P. lemmoni, P. dissecta, P. castilloniti—; P. saxicola, P. gilensis—; P. lindheimeri, P. angustifolia—; and P. bisetosa, P. warnockii. The proper infrasectional dis- position of five rather anomalous species, P. staurophylla, P. cochisensis, P. coahuilensis, P. grandifolia, and P. gentryi, remains to be elucidated. A tentative phylogenetic scheme for sect. Laphamia, based primarily on morphological, chromosomal, and chromatographical data, is presented in Fig. 2. This graphic method of presentation is adopted in preference to lengthy discussions which would have limited significance because of their conjectural nature. Some discussion, however, with regard to what are judged to be the most a aliernatives for the phylogeny of species and species groups in the section, would appear to be informative. The exact infrasectiocnal relationships of the related-species groups have remained obscure. As stated earlier, perhaps the complexity of these inter- relationships partly is a reflection of the polyphyleticism which is suspected to have been involved in the origin of species which are recognized to com- prise sect. Laphamia. For example, it seems likely that most members of the section owe their origin to sect. Perityle, but good evidence suggests that P. bisetosa-P. warnockii were derived from a white-flowered member of sect. Pappothrix. In sect. Laphamia, these species are anomalous with regard to their flower color. Furthermore, there are morphological and ecological indications of a connection between P. bisetosa and P. vitreo- montana of sect. Pappothrix aL 1969). onsidering all available evidence, the other related-species groups ap- 3) Oo, is] ay tS n $3 ON : \ a0 ™ t 4 . 4 ge D \ 4 Pt - PT ‘ ie ee “ ‘ Ao \ i - a ‘ ae \ ar x 3 ) 4 Tee eae Se 4 n a t un +2 7 cS a r : { ’ os! ' ‘ - i ’ ! Z ‘ ‘ ¢ 1 , et ! ’ = I , @ hi ’ ‘ ; 1 2 i ’ ' 4 rf ! ¥ / ' H ‘ 1 ! ‘ ! , , t 1 ’ 1 ! t i 1 ‘ \ 1 1 , \ 1 t ¢ I ‘ \ 1 , \ t / 4 i ’ \ a 1 / - | ’ Pid ‘ / “7 ' , ! / \ t ‘ \ f ze re) t / c t / v ! 1 a A 4 t é ) ? 4 i “4 ‘ u) ‘ - 4 oi f ae qa a3 a se j a pa nt 2 a : oa t id a - ' 2 o o - ’ ig. 2. Supposed evolutionary relationships among the species of Perityle sect. Laphamia, based upon all available data. The arrangement of species names approximates their geographic juxtapositioning. Solid lines indicate the most certain relationships and directions of evolution. Broken lines indi- ate probable affinities, but not necessarily phylogenetic sequence. The shorter (incomplete) broken lines depict possible or suspected relationships. 79 pear to have some real affinity with sect. Perityle, although, with the excep- tion of the southwestern alliance and P. sawxicola-P. gilensis, the evidence for this relationship is not at all satisfactory. The achene and pappus morpho- logy of P. lemmoni-P. dissecta-P. castillonit and P. lindheimeri-P. angusti- folia is typically Laphamioid, but their habits are sufficiently different from other Laphamias (and Perityle) that one must consider the possibility of other kinship. For example, the habit of the yellow-flowered P. castillonii bears a very striking resemblance to the yellow-flowered P. rupestris of sect. Pappothrix. If P. bisetosa-P. warnockit were derived from sect. Pap- pothrix, then it is logical that yellow-flowered taxa also could have emanated from sect. Pappothrix. The P. castillonii group exists under similar environ- mental conditions as do P. bisetosa-P. warnockii, and the geographic juxta- position of both groups to sect. Pappothrix is the same (Fig. 2). With the possible exception of P. staurophylla and P. coahuilensis, there are no species of Perityle which have apparent relationship with P. lindhei- meri-P. angustifolia, and it should be remembered that the infrasectional position of the former two species themselves is not understoo ince P. lindheimeri is distributed in relatively old and stable Gustaccous: limestone exposures at the extreme eastern periphery of the Perityle range (Fig. 2), it is conceivable that the taxon represents a very early offshoot from the ancestral stock of sect. Laphamia. By most standards in Perityle, however, P. lindheimeri would not be considered a primitive taxon largely because its plants have subcorymbiform capitulescences with small heads, few florets, and short rays. The latter characters are found in the genus Peri- come (P. caudata is discoid), but not in any of the species of Perityle which are being considered as likely precursors of sect. Laphamia. Thus the pos- sibility that Pericome has contributed to the seemingly polyphyletic nature of sect. Laphamia cannot be overlooked in the course of future studies. Among the related-species groups of sect. Laphamia, P. stansburii of the southwestern alliance and P. saxicola-P. gilensis compare most closely with the yellow-flowered species of sect. Perityle, at least in terms of general vegetative and floral morphology. The species groups themselves, however, do not appear to be closely related. It has been difficult to determine whether or not these species groups constitute a direct link between sect. Laphamia and sect. Perityle. Perhaps both of them do. The radiate heads of P. saxicola-P. gilensis are the largest of any species in the section, and in this character especially they approach the supposedly ancestral species of sect. Perityle. Unfortunately, other features including their complex-lobed to pinnatifid leaves fail to support any close affinity with the pertinent species of sect. Perityle. It is tempting to speculate that P. saxicola-P. gilensis rep- resent modified relects of ancestral Laphamian stock which migrated from the mountains of northwestern Mexico into northern Utah-Nevada and gave rise to the southwestern alliance Perityle stansburii actually is no more revealing with regard to the inter- sectional affinity of sect. Laphamia than are P. saxicola-P. gilensis. Never- 76 theless, the radiate heads, habit, and distribution of P. stansburii (Fig. 2) all suggest the possibility that it had early divergence as ancestral Laph- amian stock, probably from sect. Perityle. Further discussion of the southwestern alliance seems to be in order, since it comprises a sizeable number of taxa, as compared with the other related- species groups of the section. Like most species of Perityle, these taxa evi- dently have evolved through geographic displacement and subsequent genetic differentiation. When one examines all the species of the southwestern alliance, it appears that P. stansburii fills the most expectations of an ancestral taxon. Perityle stansburti is the only species of the alliance which has both radiate heads and pappus of a bristle. Also, the latter species is the most widely dis- tributed, and has a chromosome number of «17. Other floral and vegeta- tive features (including habit) of P. stansburii conform to an expected proto- type of the southwestern alliance. None of the other species of this alliance produce ray florets. Pappus bristles (1-2), however, are found in P. gracilis, P. tenella, and P. congesta, and are rarely found in P. megalocephala var. oligophylla (Howell 3906), P. villosa (Sikes 100), and P. megalocephala var. intricata (Langenheim 3746). Perityle gracilis is limited in distribution, tetraploid (based on counts from one population), and probably derived from P. stansburii. Perityle tenella and P. congesta also are limited in distribu- tion, have chromosome numbers of x16, and are considered to be derived taxa. The rare occurrence of pappus bristles in the two varieties of P. megalocephala and P. villosa possibly reflects the retention of ancestral genes in some relatively closed populations, but it is considered very un- likely that any of these taxa represent a primal position in the southwestern alliance. Judging from morphological differences, chromosome numbers, and dis- tribution, P. villosa, P. inyoensis, P. tenella, P. congesta, and the southern populations of P. megalocephala, appear to have had early divergence from P. stansburii. Morphologically, the populations of P. megalocephala var. megalocephala from southwestern Nevada appear most closely related to P. stansburii, and their geographic position also suggests the most recent separation. Geographically and morphologically, P. gracilis (n—34) also appears to have had recent derivation from P. stansburii («—17). After reviewing the above discussions, one might well be stimulated in an attempt to explain a basis for the lack of apparent affinity between related- species groups of sect. Laphamia. In other words, is the section actually polyphyletic, or is the hiatus between each related-species group a reflection of differential response to varied environmental influences on taxa which underwent early divergence from a common Laphamian ancestor? A concept of polyphyletic origin from sect. Perityle would mean that the loss or reduc- tion in at least two characters, pappus squamellae and prominent pubescence of achene margins, occurred independently in the development of several related-species groups. The loss of pappus parts concomitant with speciation 77 can be demonstrated in all three sections of Perityle, but only in presumably ancestral yellow-flowered members of sect. Perityle (eg. P. cordifolia) are there instances in which reduction in pappus squamellae and achene margin pubescence have occurred togeiher. I must hasten to add that my observa- tions should not lead one to dismiss the possibility of polyphyletic origin of sect. Laphamia species from sect. Perityle, especially because the tendency for correlated evolution of more than one character is not uncommon in the Helenieae (Ornduff, 1966). The development of Laphamioid achene and pappus characters from species of sect. Pappothrix is quite another matter. The Pappothrix pappus is typified by a crown of 20-30 bristles, but both pappus squamellae and long-pubescent achene margins are laces. in species of this group. In one species of sect. Pappothrix, P. vitreomontana, the pappus is reduced to 2(3-10) bristles (Powell, 1969). If this trend (omard reduction in pappus bristles were continued, then achenes almost indistinguishable from the Laphamioid type would result. The latter might have occurred in the evolu- tion of P. bisetosa and P. warnockii. Eventually it may prove desirable to place these white-flowered species with sect. Pappothrix. At this time, how- ever, it seems best to recognize a more conservative (and convenient) sect. Laphamia, at least until it can be determined if P. castillonti, and perhaps other species, had similar origin from sect. Pappothria. If sect. Laphamia is polyphyletic, then of course this treatment is an un- natural one, but it does bring some infrasectional systematic order to the complex phylad. Various avenues of experimentation currently are being explored, and it is hoped that a better understanding of intersectional and infrasectional phylogeny will emanate in the near future. Additional infor- mation regarding systematic relationships is discussed with the respective species in the taxonomic portion of this paper. SPECIES CONCEPT The species concept which has emerged from systematic studies in the subtribe Peritylinae closely parallels that which was expressed by Cronquist (1947) in his monograph of Erigeron. Essentially, in sect. Laphamia, species are recognized as taxa which exhibit consistent morphological distinctive- ness in several characters and are isolated from one another by geographic barriers to gene exchange. The latter criterion is a reflection of the afore mentioned conclusion that none of the species occur sympatrically. The apparent occurrence of speciation through geographic isolation has strongly influenced an attempt to arrive at a phylogenetically oriented species concept in this phylad. Experimental crosses with several species have shown that F, hybrids can be created between any interspecific com- bination, although differential IF, sterility is expressed among the taxa. Artificial intersectional fertility is also common in the genus Perityle. It is obvious, therefore, that any attempt to incorporate reproductive criteria into a species concept would be undesirable, at least until fertility data are 78 studied. Quite simply, in sect. Laphamia (and sect. Pappothrix; Powell, 1969), it would seem that initial geographic isolation of taxa precluded a ‘necessity’? for the establishment of genetic barriers to gene exchange, and that truly genic strictures of interspecific fertility still are being accumu- lated as concomitants of continuing independent evolution. Infraspecific categories have been designated as varicties. Formal recog- nition has been given to populational entities which have attained only partial, marginal, geographic separation from the main body of a given species, and which differ consistently from the rest of the species by one or a few characters, or to populational entities which appear completely iso- lated from the parent species but exhibit only slight morphological or chromosomal differen It must be pointed out that I can feel confident of having obtained ‘‘ade- quate’’ populational surveys in but a few species of sect. Laphamia. Judging from the number of undescribed taxa which have come from these studies already, one cannot depend too heavily upon previous collections. Distres- singly large areas within the distributional ranges of several species have not been examined at all. Shortcomings in collecting Perityle often are dictated by the extremely rough topography in many areas of the south- western United States and northern Mexico. Many times it is necessary to provide oneself with a 4-wheel drive vehicle, drive into country with poor roads, or no roads at all, and climb hundreds of feet to a promising bluff habitat. In short, it has been physically impracticable to visit many locali- ties in search for a better populational understanding of Perityle species. Consequently, it has been necessary to extrapolate in making some syste- matic judgments. I have attempted to maintain a conservative approach to problems where inadequate information would be likely to influence a spurious nomenclatural decision. The reader will notice numerous instances where ‘‘anomalous populations” or similarly ambiguous problems are re- ferred to further study. TAXONOMY PERITYLE sect. LAPHAMIA (A. Gray) Powell, Sida 3:277. 1968. Laphamia A Gray, Pl. Wright. 1:101. 1852. Monothrix Torr. in Stansbury’s Exp. 390. 1852. Leptopharynx Rydb., N. Amer. Fl. 34:21. ca (in part) Plants suffrutescent perennials, growing from crevices of solid rock ex- posures, from 2-60 cm high, glabrous to variously pubescent, typically glandular-pubescent; leaves opposite or alternate, usually alternate, ex- tremely variable in size and shape, 0.5-11.0 em long including the petiole, 0.3-6.0 cm wide, linear-lanceolate, ovate, palmate, tending to be tripartite in most species, variously serrate, lobed, cleft, or divided to pinnate, glab- rous to pubescent, usually minutely glandular-punctate, viscid, bitter tast- ing, sessile or petiolate; capitulescence of solitary heads, or heads loosely to tightly aggregated in clusters of 3 or more; peduncles very short so that 79 heads are partially subtended by cauline leaves, or long with heads dis- tinctly raised above leaves; involucres subcylindric to broadly campanulate, heads 0.5-1.1 cm long, 0.4-1.2 em wide; involucre of 2 equal or subequal series, in some species seemingly either once or thrice seriate, inner bracts essentially like the outer except slightly wider and more membranous; bracts linear, oblong, lanceolate, flattened or keeled; ray flowers present or absent, ligules yellow, oblong to suborbicular, 2.5-7.0 mm long, 1.2-3.8 mm wide; disc corollas four-lobed, white or yellow, 2.2-6.5 mm long, tubes minutely glandular-pubescent (glandular-pilose in P. warnockii), throats cylindric to narrowly campanulate, typically sparsely glandular-pubescent, lobes typically reflexed and acute; style branches 1.0-3.0 mm long, linear, flattened, tapering to a minutely pubescent tip; achenes black when ma- ture, 2.0-3.8 mm long, distinctly flattened radially, often obscurely rounded or angled on one or both surfaces, linear, linear-oblong to narrowly oblance- olate in outline, outer ones cften slightly curved and partially clasped by bract, with conspicuous or inconspicuous callous margins (rarely absent), typically short-pubescent on margins and surfaces, rarely glabrous; pappus absent or of 1 or 2 (rarely 3-6) bristles, the bristles arising from the achene margins, 0.5-4.6 mm long, naked or minutely antrorse-ciliate distally, pappus also of inconspicuous, vestigial, hyaline squamellae, or the squamellae absent, revealing just a callous crown; anthers 4, 1.5-3.0 mm long; base chromosome number, «=—17. Type: P. lindheimeri A. Gray. KEY TO THE SPECIES la. Flowers white; plants 2-8 (12) cm high; pappus absent or typically of 2 rather flattened bristles. 2a. Pappus absent; leaves scabrous-pubescent 1. P. warnockii. 2b. Pappus typically of 2 bristles; leaves ese gels glabrous, except in var. scalaris . . P. bisetosa. 1b. Flowers yellow; plant: ieaelly seedeaine YD cm iene pappus absent or of 1-3 (6) rounded bristles. 3a. Heads radiate. 4a. Plants, especially stems, pilose-villous; achenes epappose; Sonora, inaloa. 5a. Leaves 8-11 cm long, 4-6 cm wide; Sinaloa . 20. P. grandifolia. 5b. Leaves 1.5-2.5 em long, 0.8-1.8 cm wide; PA gk ke ee ee ee ee ee 621. PL gentryi. 4b. Plants essentially glabrous or short-pubescent; pappus of 1-2 (6) bristles, rarely none; Utah, Nevada, Arizona, New Mexico, Texas. 6a. Florets ca. 16-22 per head; capitulescence usually of tightly ag- gregated heads on short peduncles; leaves broadly ovate to ovate-lanceolate; Tex ae 15. P. lindheimeri. 6b. Florets exceeding ca. a per aaeae: @apitilescence of several, loosely aggregated heads, or few heads on long peduncles; leaves 80 subovate to variously tripartite, or lobed, divided, or pinnate. 7a. Leaves usually subcruciform but also merely lobed or pinnate; New Mexico . . 17. P. staurophylla. 7b. Leaves ovate to eaiy Andee or ena iss Utah, Nevada, Arizona. 8a. Florets less than 100; leaves subovate to suborbicular, serrate to shallow-lobed; Utah and NE Nevada . . 3. P. stansburiti. 8b. Florets exceeding 100; leaves deeply dissected or pinnate; Arizona. Ya. Leaves typically pinnately trifoliate with segments much broadened distally . 2 gilensis. 9b. Leaves tri- to bipinnate with iene: linear to flat-filiform segments .. 10. P. saxicola. 3b. Heads discoid, except dates eel ea in Pp. ee 10a. Plants relatively long-pubescent, villous, pilose, pilose-hirsute, oc- casionally sparse-pilose in P. castillonii. lla. Leaves deeply dissected and divided into numerous irregular lobes, merely serrate-lobed in a population of P. lemmoni. 12a. Pappus bristle(s) rather stout; bracts attenuate: disc corollas tubular; Presidio-Brewster Co., Texas, and adjacent Chihua- hua, Mexico — dissecta. 12b. Pappus Baas: ee aaicac bracts acute; dise corollas broad-tubular to subfunnelform; SE Arizona —~ — : P. lemmoni. llb. Leaves serrate, aliens en: or sntiné: 13a. Florets 16-30; disc corollas 2.2-3.5 mm long, turning purplish at maturity; 1-2 pappus bristles usually present; E. Chihauhua and W. Coahuila, Mexico . . . 14. PP. castillonizi. 13b. Florets 30-75; disc corollas 4-5 mm ines not turning purplish at maturity; pappus bristles usually absent (rarely 1-2 bristles in P. villosa); Inyo and Panamint Mts., California. lda. Leaf margins regularly serrate-lobed; leaves opposite or alternate .. oo ah ee ow ms & 6. P. inyoensis. 14b. Leaf margins satire, or with 1-3 lobes per margin, leaves alternate —_~ oo de et ke a as “We ee ok ke oR wie 3. orllosad. 10b. Plants relatively short-pubescent, hispidulous, hirtellous, short pilose-hirsute in P. tenella, or essentially glabrous. 15a. Plants essentially glabrous, occasionally puberulent to hispidu- lous in P. cochisensis. l6a. Achenes typically epappose; limestone rocks; SW Texas P. ancuslitolia: 16b. Achenes with pappus of L2 bigs penile. Mexico, Arizona, 17a. eg 4-12 em high; involucres campanulate; Chiricahua Arizona ‘, oo ee ae ee me ee P. cochisensis. 17b. ae 15-35 cm high; involucres nearly eet SW n 81 Coahuila, Mexico . . 18. P. coahuilensis. 15b. Plants pubescent, seeasionally nieanly sisivone in P. gracilis. 18a. Pappus absent, rarely of 1-2 bristles in var. intricata; stems leafy, especially toward the base in var. megalocephala, stems with a few, small leaves to nearly naked in var. oligophylla and var. intricata; SW Nevada and adjacent California os & S . P. megalocephala. 18b. Pappus of 1-2 (3) bristles; seis epioasy leafy; SE Nevada, SW Utah, NW Arizona. 19a. Leaves 3-cleft, tripalmate, to subcruciform; SW Nevada do. P. gracilis. 19b. Leaves broadly detiola- one re subeouaate and serrate, or ovate to ovate-elliptic and entire, serrate, or shallow-lobed. 20a. Leaves broadly deltoid-ovate to subcordate, serrate; SW Utah and adjacent NW Arizonia. : . 8. P. tenella. 20b. Leaves ovate, ovate-lanceolate, to ayale. elliptic, entire, serrate, or shallow-lobed; Grand Canyon area, Arizona 9. P. congesta. 1. PERITYLE WARNOCKII Powell, Sida 3:177. 1966. TYPE: TEXAS: Val Verde Co. NW part of county: in crevices and small pockets of solid Cretaceous limestone exposures, ca. 40-50 mi NE of Dryden, ca. 1 mi E cf Pecos see! er, 11 Oct 1964, Powell 1311 (Holo- type, SRSC! ee. GH! SMU ! Plants 2-10 cm high, scabrous-pubescent, aoncely leafy; leaves opposite below, alternate above, 0.8-1.5 cm long, 0.4-0.8 (1.0) cm wide, scabrous- pubescent, suborbicular, ovate, to ovate-lanceolate, margins conspicuously serrate, acute at the apex, cuneate at the base; petioles 2-5 (8) mm long; capitulescence of solitary heads borne on short peduncles (1-7 mm long); involucres campanulate, heads 7-10 mm long, 4-7 mm wide, often partially obscured by subtending leaves; receptacle flattened, 1.0-1.5 mm wide; in- volucral bracts 9-10, 5-7 mm long, 1.2-2.2 mm _ wide, obscurely keeled, hirtellous; ray flowers absent; disc flowers, 9-12, corollas bright white, 5-6 (6.5) mm long, the tube 1.0-1.3 mm long, glandular-pilose, the throat sub- funnelform, sparsely gladular-pilose, the lobes attenuate-acute, (1.0) 1.2-1.6 mm long; style branches white, ca. 2.0 mm long; achenes 2.2-2.5 (2.8) mm long, oblong to oblong-obconical, truncate at the base and apex, often rounded or obscurely angled on one or both surfaces, with conspicuous callous margins, the margins glabrous, minutely pubescent on achene sur- faces; pappus absent; chromcsome number, n—17. Known only from the type locality. Fig. 3. Spring-Fall. Perityle warnockit is clearly related to P. bisetosa as evidenced by its similar, and in some instances reduced, vegetative and floral morphology. The species is delimited primarily by its epappose achenes and scabrous- pubescent leaves which are typically suborbicular to ovate with conspicuous- ly serrated margins. ; ‘ \ gy C Kent o * ca 2 5 X Af owed Fig. 3. Distribution of P. bisetosa var. bisetosa (open square); P. bisetosa var. scalaris (open triangle); P. bisetosa var. appressa (closed squares); P. bisetosa var, spathulata (closed triangle); P. warnockii (closed star): P. saxicola (open circles); P. gilensis var. gilensis (closed circles); P. gilensis var. salensis (closed arrow); P. cochisensis (open star circled): P. grandifolia (open star); P. gentryi (open arrow). Both P. warnockii and P. bisetosa were derived independently from a common ancestor (perhaps white-flowered Pappothrix, e.g., P. vitreomon- tana). Neither taxon has a combination of morphological features which are suggestive of having been derived from the other. The epappose achenes of P. warnockit reveal the possibility of reduction from P. bisetosa, but it seems unlikely that the larger, pubescent leaves of P. warnockii could have come from P. bisetosa. 2. PERITYLE BISETOSA (Torr. ex A. Gray) Shinners KEY TO THE VARIETIES la. Leaves subsessile or short-petiolate (1-2 mm), usually opposite, essen- tially glabrous, margins entire to subserrate; Texas 2a. var. Veta a: lb. Leaves distinctly petiolate (2-7 iy: distinetiy alternates glabrous or ispidulous, margins distinctly serrate; Texas and Coahuila, Mexico. 2a. Leaves scabrous-hispidulous; Texas . . . . . 2b. var. scalaris. 2b. Leaves essentially glabrous; Texas, Mexico. 3a. Leaves ovate, 1.0-1.5 ecm long, 0.4-0.8 em wide, Texas and Coahuila, exico . . ..... 2c. var. appressa. 3b. Leaves oblanceolate-spathulate, 0.7-1.2 cm ‘Gone: 0.2-0.4 cm wide; oahuila, Mexico . .... . . . 2d. var. spathulata. 2a. PERITYLE BISETOSA (Torr. ex A. Gray) Shinners var. BISETOSA, Southwest. Nat. 4:204. 1959 Laphamia bisetosa Torr. ex A. Gray, Pl. Wright. 2: 106. 1853. TYPE: TEXAS: Brewster Co. crevices of rocks in a canyon of the Rio Grande below Presidio del Norte, Oct, 1852, C. C. Parry s.n. (Holotype, GH! Isotype, NY!). Plants 2-7 (10) cm high, stems minutely pubescent, densely leafy; leaves usually opposite, less often alternate, 0.5-1.0 cm long, 0.2-0.6 cm wide, es- sentially glabrous, glandular-punctate, coriaceous, ovate, ovate-lanceolate, to lanceolate, margins entire to subserrate, often subundulate, acute at the apex; subsessile or short petiolate (1-2 mm); capitulescence typically of solitary heads (rarely 2) on very short peduncles; involucres narrowly campanulate, heads 7-8 (10) mm long, 3.5-7.0 mm wide, subtended and mostly obscured by leaves; receptacle flattened, 1.0-1.5 mm wide; involucral bracts 8-10 (12). 4.5-5.5 mm long, 1.0-2.0 mm wide, keeled; ray flowers absent; disc flowers 11-17, corollas white, (3.5) 4.2-4.8 mm long, the tube 0.8-1.0 mm long, the throat tubular to subfunnelform 1.5-2.5 mm long, the lobes acute, 0.8-1.0 mm long; style branches (1.0) 1.5-2.0 mm long; achenes 1.8-2.2 (2.5) mm long, oblong to oblong-obconical, often rounded or obscurely angled on one or both surfaces, with conspicuous callous margins, the margins glabrous, minutely pubescent on achene surfaces; pappus of 2 flattened bristles, 1.5- 2.3 mm long; chromosome number, n=17. Very restricted in occurrence, crevices and pockets of limestone exposures. Fig. 3. Spring-Fall. Specimens examined: TEXAS: Brewster Co. 7.1 mi SW of US 90 on road to Cox Ranch, D. Flyr 229 (SRSC); low white hills, Cox Ranch, 7.1 mi SW of Hwy 90, near Long- fellow, A. M. Powell 1370 (SRSC); McRae Canyon, Cox Ranch, B. H. War- nock 11418 (SMU, SRSC); Cox Ranch, 65 mi SE of Alpine, Warnock 18643 (SRSC) The type locality given for var. bisetosa (canyon of the Rio Grande below 84 Presidio del Norte) is at Presidio, Texas (or Ojinaga, Mexico), but P. bise- tosa is not known to have been collected along the Rio Grande near Presidio. Also, the area in question does not appear to offer the typical habitat for the taxon. More likely the type locality could have been below Mt. Carmel (Sierra del Carmen) along the Rio Grande, the only region in which P. bisetosa has been discovered, except for 2 localities south and east of Sierra del Carmen in Mexico. Perityle bisetosa is indeed a remarkable species. Its reduced habit, white flowers and bisetose pappus of flattened bristles set it apart from all other taxa in sect. Laphamia. The species exhibits clearly what is believed to be a very important evolutionary trend in Perityle, i.e., geographic isolation serving as a primary impetus for speciation. The area in which P. bisetosa occurs is relatively dry, and has been severely eroded, leaving numerous limestone habitats exposed as ‘‘caprocks,”’ hills, mesas, pinnacles, and can- yons. Four varieties of P. bisetosa have been recognized. Each seems to be isolated completely from the others, and each apparently has evolved inde- pendently after becoming geographically separated. With reference to the evolutionary history of P. bisetosa, one might visualize that as erosion of the countryside progressed, small colonies became stranded and therefore subject to the evolutionary forces of small populations. There is some evidence that P. bisetosa evolved from a _ white-flowered member of sect. Pappothrix, such as P. vitreomontana. The typical Pap- pothrix pappus is comprised of 20-30 bristles, but that of P. vitreomontana is reduced to 2(3) main bristles with several vestigial nubs. Perityle vitre- omontana apparently evolved in response to increasing aridity at the peri- phery of the Pappothrix range after becoming geographically segregated (Powell, 1969). The morphology and ecology of P. bisetosa suggests that it continued the trend toward reduction in pappus bristles, which is manifested in P. vitreomontana, in still more arid regions located to the south of the latter species. Not only do the bisetosa pappus and gross morphology indi- cate a connection between P. vilreomontana and P. bisetosa, but in var. appressa and var. scalaris 3-5 bristles have been observed, with 1-3 of them reduced to vestigial nubs. Additionally, the occurrence of a few 3-4 angled achenes in var. appressa suggests that the distinctly flattened achenes typical of P. bisetosa were derived from the flattened but obscurely 4-angled achenes of Pappothrix species. 2b. PERITYLE BISETOSA var. SCALARIS Powell, Sida 3:177-178. 1967. TYPE: TEXAS: Brewster he in crevices of lower limestone bluffs, Black Gap Game Preserve, ca. 4 mi S of ee enreen Cave Hill, E side, along Stairstep Mt.. 2% Sept 1965, A. M. Powell and T. Watson 1394 (Holotype, SRSC! Isotype, TEX!). Plants 6-12 cm. high, in small, once or pendulous clumps; leaves opposite below, alternate above, 8-12 mm long, 5-8 mm wide, scabrous-hispidulous, broadly ovate to suborbicular, with 1-2 shallow lobes per margin or merely serrate, the lobes and serrations acute; petioles 2-4 mm long; floral features 85 mostly slightly larger than in var. bisetosa, but otherwise the same, except pappus in some heads of 3 (4) bristles of equal length or with 1-2 reduced; chromosome number, n=ca. 102. Single population known only from the type locality. Fig. 3. Spring-Fall. 2c. PERITYLE BISETOSA var. APPRESSA Powell, Sida 3:178-180. 1967. E: TEXAS: Brewster Co. east face of the limestone caprock of a small mountain peak on the N rim of San Francisco Creek Canyon do July 1966, J. Scudday, S. Sikes, and A. M. Powell 626 (Holotype, SRSC! Isotypes, GH! SMU! TEX! US!). Plants 4-8 cm high, 12-20 cm broad, growing in dense, leafy clumps closely appressed to rocks; leaves distinctly alternate (rarely opposite below), 1.0- 1.5 cm long, 0.4-0.8 em wide, glabrous, ovate, serrate or serrate-sinuate to shallow lobed, typically 2-3 (rarely 4) serrations or lobes per margin, the serrations obtuse to subacute; petioles 3-7 mm long; pappus typically of 2 bristles, rarely 3 (5), with at least 1-3 reduced; chromosome number, n=17. Restricted to caprock bluffs. Fig. 3. Spring-Fall. Specimen examined: MEXICO: Coahuila: Colorado Canyon, Serranias del Burro, D. Patterson and R. Quifiones 77 (SRSC). 2d. PERITYLE BISETOSA var. spathulata Powell, var. nov. Fig. 4. TYPE: MEXICO: Coahuila: limestone beers - ae dead-end Boquilla Canyon, de la Encantada 92.5 mi NW of Musquiz, 22 May 1968, A. M. well and D. arene 1588 ‘Holotype. SRSC! ieetpned SMU! en ‘ Plantae 4-10 cm altae caespitosae, brevipendulosae vel ad saxa appressae; folia distincte alterna, 0.7-1.2 cm longa, 0.2-0.4 cm lata, glabra, oblanceolato- spathulata, ad laminae dimidium apicale 3-5 serrato-lobata, petiolis tenuibus laminas subaequantibus vel paulum brevicribus; chromosomata n=17. Single population known only from the type locality. Fig. 3. Spring-Fall. w PERITYLE STANSBURII (Gray) Macbride, Contrib. Gray Herb. n:s. 56:39. 1918 Laphamia stansburii Gray, Pl. Wright. 1:101. 1852. TYPE: UTAH: Tooele . erevices of limestone rocks Stansbury’s Island, 26 June 1850, Stansbury s.n. (Holotype, NY! Isotypes, GH! NY! aerate ee DS!). Monothrix stansburyana Torr. in Stansbury’s Exp. 390. Monothrix stansburii (Gray) Rydb. N. Amer. FI, 34:21. ie Plants 25-45 cm high, often in dense clumps to 60 cm across, hirtellous; leaves mostly alternate, less often opposite, 0.9-1.6 (2.0) cm long, 0.5-1.0 (1.4) em wide, hirtellous, typically broadly subovate, but varying from sub- deltoid to sub-orbiculate, typically 2-5 (6) lobed or serrate, less often tri- lobed or subentire; petioles (2) 3-8 mm long; capitulescence of solitary heads or subcymose clusters; peduncles rather stout, usually 1-6 cm long; in- volucres campanulate, heads 7-8 mm long, 5-9 mm wide; receptacle convex to short-conical, rarely somewhat flattened, 1.5-2.2 mm wide, 0.3-1.0 mm high; involucral bracts 16-21, 5-6 mm long, 1.0-1.9 mm wide, lanceolate to BE TA OF Pia WY Fig. 4. Perityle bisetosa var. spathulata: photograph of holotype (SRSC), Powell and Patterson 1588, 87 broadly oblanceolate, strongly keeled; ray flowers 12-14, pistillate and fertile, tube 1.6-2.2 mm long, ligules yellow, 3.0-5.5 mm long, 1.2-1.9 mm _ wide, typically oblong, less often subelliptic, 3-lobulate at the apex; disc flowers 60-80, corollas yellow, 4-5 mm long, the tube 1.2-1.5 mm long, the throat tubular or subfunnelform, 2.4-2.9 mm long, the lobes acute, 0.4-0.6 mm long; style branches 2.0-2.6 mm long; achenes 2.3-3.4 mm long, typically narrowly obconical, less often suboblong to obconical-elliptic, always smaller at the base, usually somewhat rounded on both surfaces, rarely angled on one surface, with thin callous margins, short-pubescent on the margins especially, and both surfaces; pappus of a single, rather stout bristle, 2.5-4.0 mm long, and a very small crown of hyaline squamellae, the bristle often tending to terminate abruptly as a bluntish point; chromosome number, n=—17 Common in crevices of rock exposures in NW and W Utah and NE Nevada. Fig. 5. Spring-Fall. Representative specimens: NEVADA: Elko Co. Wendover, M. E. Jones s.n. (RM); Loomis Pasture, NW of Wells, C.O. Land s.n. (POM, UC). White Pine Co. W slope of Sacra- mento Pass, H. D. Ripley & R. C. Barneby 9273 (NY). UTAH: Beaver Co. Minerville, Ripley and Barneby 9249 (CAS, NY); Wah Wah Mts., 35 mi W of Milford, R. C. Rollins and T. S. Chambers 2463 (DS, GH, RM, UC). Grand Co. Moab, M. E. Jones s.n. (POM). Juab Co. Thomas Creek, Deep Creek Mts., W. P. Cottam 3234 (UT); 12 mi E of Troutcreek, B. Maguire and R. J. Becraft 2855 (US); 13 mi from Troutcreek, Maguire and Becraft 2856 (GH, RM, UC); 25 mi SW of Eureka, Maguire 20752 (GH, NY), Granite Creek, Deep Creek Range, Maguire and A. H. Holmgren 21918 (GH, NY, US). Millard Co. Warm Spring, Cottam 3267 (UT); Gandy, Cottam 8157 (UT); Garrison, Jones s.n. (POM); House Range, 56 mi W of Delta, Maguire and Becraft 2857 (GH, POM, RM, US); 1.8 mi S of Garrison, Maguire 20851 (NY, US); House Range, 47 mi W of Delta, Maguire 21056 (GH, NY, US); Pruess Lake, near Garrison, F. W. Peirson 7537 (CAS, RSA); E of Skull Rock Pass, 51 mi SW of Delta, A. M. Powell 1270 (SRSC); near Preuss Lake, I. Tide- strom 11161 (US); Skull Rock Pass, Confusion Range, S. L. Welsh and G. Moore 2230 (NY). Salt Lake Co. Garfield, A. O. Garrett 2794 (UT); Garfield, W. W. Jones s.n. (UC). Sanpete Co. Fairview, Jones s.n. (POM); Route 25, Soldier Fork Canyon, Wasatch Mts., F. W. Pennel, R. D. Johnson and R. L. Schaeffer 22753 (NY); W of Sterling, Tidestrom 1307 (RM, US). Sevier Co. Gooseberry ranger station, Fishlake Nat'l Forest, Wasach Mts., W. W. Eggleston 10378 (US). Tooele Co. Stansbury Island, kK. Brizzee 7803 (UT); Thomas Creek, Deep Creek Mts., Cottam 3247 (UT); Stansbury Isl., Cottam 9185 (RSA, UT); near Low, A. Eastwood and J. T. Howell 380 (CAS, GH); base of Camel Back Mts., SW part of county, S. Flowers 10 (UT); North Point, Stansbury Mts., Flowers 1294 (UT); W side of Great Salt Lake, Gar- rett 5834 (DS, RUM, UT); Stansbury Isl., Garrett 8474 (CAS, RSA, SMU, UC, US); Ophir Canyon, Garrett 8815 (UC, UT); W of Grantsville, Jones 25582 (CAS, DS, ND, POM); Stansbury Isl., Jones s.n. (CAS, DS, ND, NMC, NY, POM, RM, UC, US); Granite Mts., Jones s.n. (POM); U \ | a NX \ ~ t eee i Q ONE al a oe, Vv _ q Lip * Ne a \ \ \ Wyy SN i) Kos i, eo NN | Ww oO \ Fig. 5. Distribution of P. stansburii (closed squares); P. megalocephala var. megalocephala (closed stars); P. megalocephala var. oligophylla (open stars); P. megalocephala var. intricata (open stars circled); P. inyoensis (closed circle); P. villosa (open square); P. gracilis (open circles); P. tenella (open triangles); P. congesta (closed triangles). Buent Sta., Dutch Mt., Jones s.n. (POM); Wendover, Jones s.n. (POM); Stansbury Isl., F. E. Leonard s.n. (DS, GH, NY, UC); S shore of Great Salt Lake, 13.5 mi W of Grantsville, Maguire, R. Maguire, and B. C. Maguire 5652 (UC, GH); 43 mi W of Salt Lake City, Maguire 16731 (POM, UC); Route 40 NW of Grantsville, F. W. Pennell and R. L. Schaeffer 22852 (NY, US); NW end of Stansbury Island, Powell and Calvert 1269 (SRSC); Mt. Flux, Ripley and Barneby 5512 (CAS, NY); Stansbury Isl., G. Saccoman- no 7667 (UT); Stansbury Isl., S. Watson 587? (NY, US). Utah Co. Rock Canyon near Provo, Garrett 3329 (GH, US, UT); S of Black Rock, Salt Lake, Garrett 8239 (DS, UT); Rock Canyon, B. F. Harrison 8563 (UT): Provo, Jones 5555 (NY, RM, UC, US). 89 The relatively wide ranging P. stansburii is clearly related to P. megalo- cephala. In fact, when examining herbarium specimens, the two species are difficult to separate if well-developed heads have not been preserved. Norm- ally, P. stansburii is easily distinguished from P. megalocephala by its radiate heads and pappus of a single bristle. As discussed elsewhere, P. stansburii is regarded as ancestral in a group of seven species designated as the southwestern alliance (see Phylogenetic Considerations). Even though other phyletic lineages are possible for the southwestern alliance, P. stansburit appears to be the only taxon which possesses a combination of supposedly ancestral features from which the other species could have derived their phenotypic appearances. Briefly, P. stansburii is large in habit, has relatively broad leaves, radiate heads, pappus of a bristle, and a chremosome number of «=17, all presumably ancestral in this alliance. 4. PERITYLE MEGALOCEPHALA (Wats.) Macbride KEY TO THE VARIETIES Leaves relatively copious, 0.7-1.5 (1.7) cm long, 0.4-0.9 (1.2) em wide, ovate, lanceolate-ovate, subelliptic to suborbicular 4a. var. ne qaleceenae: — © lb. Leaves relatively meager, 0.4-1.7 cm jee 0.1-0.4 (0.6) em wide, lanceo- late or linear. 2a. Capitulescence of solitary heads or loose clusters of 2-3 heads; pedun- cles (1.5) 2.0-5.0 ecm long . . 4b. var. oligophylla. 2b. Capitulescence of 2-6 heads in tight iaeteee: peduncles rarely ex- ceedinglcm .. . . 4c. var. intricata. 4a. PERITYLE MEGALOCEPHALA (Wats.) Macbride, var. MEGALO- ontrib. Gray Herb. nus. 56: 918. Laphamia megalocephala Wats. Amer. Nat. 7:301. 1873. TYPE: NE- DA: 1872, Wheeler s.n. (Holotype, US! Isotype, GH!). Monothrix megacephala (Wats. ?) Rydb. N. Amer. FI. 34:20. 1914. Plants 30-55 cm high, often in dense, suffrutescent clumps, commonly 30-60 em across, less often 90 cm across, profusely branched at the base and above, densely hirtellous; leaves mostly alternate, less often opposite, 0.7- 1.5 (1.7) em long, 0.4-0.9 (1.2) em wide, becoming progressively smaller toward apex, hirtellous, ovate, lanceolate-ovate, subelliptic, to suborbicular, the margins typically entire to subentire rarely serrate to serrate-lobed, acute to obtuse or rounded at the apex; petioles 1-5 (6) mm long; capitules- cence of solitary heads or of a few, loosely aggregated heads on peduncles of variable length; peduncles usually 1.0-4.5 (8.0) cm long, typically bearing small leaves; involucres ie gente heads 6-9 (10) mm long, 5-6 (8) mm wide; receptacle convex and rounded to short conical, 1.5-2.0 mm _ wide, 0.5-1.0 mm high; involucral 0 14-19, 5-6 mm long, 1.3-1.9 mm _ wide, lanceolate to suboblanceolate, keeled or with a rather prominent midvein; 90 ray flowers absent; dise flowers 45-60; corollas yellow, 3.5-4.2 mm long, the tube 1.0-1.6 mm long, the throat subtubular to subfunnelform, 1.8-2.2 mm long, the lobes acute, 0.40.7 mm long; style branches 1.8-2.0 mm_ long; achenes 2.5-2.9 (3.0) mm long, narrowly obconical to suboblanceolate, round- ed or less often angled on one or both surfaces, with thin callous margins, pubescent on the margins and both surfaces; pappus absent, of a callous crown, or of vestigial, hyaline squamellae especially evident above the achene margins; chromosome number, n= 17. Relatively common in volcanic rocks of arid mountains, southwest-central Nevada and adjacent California. Fig. 5. Spring-Fall. Representative specimens: CALIFORNIA: Inyo Co. Devil’s Gate Waucoba Canyon, A. M. Alexander and L. Kellogg 2541 (DS, GH, UC); N fork of Silver Canyon, 8 mi E from Laws, White Mts., E. I. Applegate 6936 (DS); Silver Canyon, White Mts., kK. Brandegee s.n. (UC); Black Canyon, D. Cassell 273 (DS); Marble Canyon, V. Duran 2845 (CAS, DS, GH, NY, POM, RM, RSA, UC); Westgard Pass, KE. C. Jaeger s.n. (POM); Squaw Flat, Inyo Mts., NW of Waucoba Mts., M. Kerr s.n. (CAS, RSA); head of Marble Canyon, Inyo Mts., P. A. Munz and J. C. Roos 20169 (CAS, NY, RM, UC); hill behind Blue Bell Mine, Badger Flat, Inyo Mts., J. C. Roos and A. R. Roos 5938 (CAS, DS, NY, RSA, UC); Mazourka Canyon, Inyo Mts., Roos and Roos 5979 (DS, NY, RSA); the Narrows, Marble Canyon, Inyo Mts., Roos and Roos 6217 (CAS, NY, RSA); N fork of Silver Canyon, White Mts., C. B. Wolf 2587 (CAS, DS, RSA). NEVADA: Esmeralda Co. Goldfield, H. M. Hall 10537 (CAS, GH, UC); 3 mi below Pigeon Springs on the W side of Magruder Mt., B. Maguire and A. H. Holmgren 25661 (CAS, DS, GH, NY, POM, UC); N end of Clayton Valley, H. D. Ripley and R. C. Barneby 38657 (NY); Soda Springs Canyon, W. H. Shockley 568 (DS, ND, POM, UC, US). Mineral Co. 1-3 mi from mouth, up Cory Creek, Wassuk Range, W. A. Archer 6849 (DS, NY, UC); base of Wassuk Range, US Hwy 95 along Walker Lake, 13 mi N of Hawthorne, R. S. Ferris 13100 (CAS, DS, NY, RM, RSA, TEX, US); 10 mi N of Haw- thorne, P. Train 4049 (NY, UC). Nye Co. 16 mi FE of Warm Springs near Hwy 25, W of Nyala turn off, A. M. Powell 1274 (SRSC); Tonopah, W. H. Shockley 122 (DS, POM, RM, UC, US). Pancake Range, Nev., C. A. Purpus 6341 (NY, POM, RM, UC, US). Perityle megalocephala var. megalocephala appears to have close affini- — ties with P. stansburii and P. inyoensis. Several distinctive features of var. mee cele wenn are best determined by consulting the above description, but essentially this taxon can be delimited from P. stansburti which has radiate heads, pappus of a single bristle, and acute-lobed leaves and from P. iy yoensis which has a smaller habit, pilose-villous pubescence, and essentially = ' opposite leaves which are ovate to ovate-deltoid with serrate-lobed margins. Perityle megalocephala var. megalocephala is discoid, epappose, large and profusely branching in habit (30-55 cm high, 30-60 cm across) and typically has hirtellous, ovate to suborbicular leaves. 91 4b. PERITYLE MEGALOCEPHALA var. oligophylla Powell, var. nov. Fig. 6 TYPE: CALIF.: Inyo Co. Lost Burro Gap, Cottonwood Mts., Death Valley Nat'l Mon., 17 Sept 1957, R. S. Ferris and W. R. Ernst 13162 (Holotype, DS! Isotyp pes, DS! RM! RSA! TEX! US!). Plantae 15-35 cm altae, dense hirtellae, ramosissimae; caules tenues vel crassi, parce foliosi (vel quasi); folia alterna, 0.7-1.7 cm longa, 0.1-0.4 (0.6) cm lata, lanceolata ad linearia, petiolata; capitula solitaria vel 2-3 laxe disposita; pedunculi (1.5) 2.0-5.0 cm longi; flores sicut in var. megalo- cephala nisi capitulis paulum ANOEIOUS (3.5-7.5 mm latis) et raro pappo setae unicae; chromosomata n= Relatively common but very saiemeral in protected areas or northern exposures in mountains of the Death Valley region, California. Fig. 5. Spring- Fall, when it rains. Specimens examined: CALIFORNIA: Inyo Co. Death Valley Canyon, Panamint Mts., F. V. Coville and A. F. Gilman 89 (US); Death Valley Canyon, Coville aia Gilman 118 (US); right fork of Death Valley Canyon, Panamint Mts., M. F. Gilman 1853 (US); Goodwin Spring, branch of Wildrose Canyon, Gilman 2222 and 2223 (US); Wildrose Canyon, Panamint Mts., Gilman 2224 (POM); Tin Mt., Cot- tonwood Mts., Death Valley Nat’l] Mon., Gilman 2676 (POM); gorge below Rest Spring, Cottonwood Mts., Gilman 3896 (GH, POM); Leadfield Canyon, Death Valley region, Gilman 4229 (POM); Surprise Canyon, Panamint Mts., J. T. Howell 3906 (CAS, DS, GH, US); W entrance of Lost Burro Gap, Cot- tonwood Mts., A. M. Powell and D. Patterson 1612 (SRSC). Most collections here attributed to var. oligophylla have been recognized as var. intricata by several taxonomists (herbarium sheet annotations). The establishment of an additional variety of P. megalocephala is warranted in spite of the fact that much more populational information about the species needs to be gathered. Perityle megalocephala var. oligophylla exhibits a meager complement of lanceolate to linear leaves, a consistent feature by which it can be delimited from the var. megalocephala. The variety is further characterized by its smaller habit, and disjunct distribution (Fig. 5), although the varieties per- ceptibly intergrade in Marble Canyon, Inyo Mts., California. No consistent floral differences have been detected between var. oligophylla and var. megalocephala. A chromosome number of n—34 has been obtained for var. oligophylla from one population (Powell and Patterson 1612). Further study will be necessary before it can be determined if the variety is consistently tetraploid. The var. Giclee lets was found to be diploid (n=17), but thus far only one population has been examined chromosomally. The habit of some specimens of var. eile especially Gilman 1853, is suggestive of a relationship with P. inyoensis. It is suspected that var. oligophylla evolved its meager leaves in response to extreme arid conditions, although plants of var. megalocephala also inhabit DUDLEY HERBARIUM OF STANFORD ea Plants of California PF Laghamia intricate Srandg Rounded dense masees growing tn limestone eee : crevices. In Lost Burro ore he ee a alle of Cotts Mts., Swath ¥ HOLOTYPE $8 ay cece oo Tayo on i galoseph Alt, 392 as | SVR Raseta $ Ferra & MA-Erast 13162 LSTo = : Sept. 37, ; Fig. 6. Perityle megalocephala var. oligophylla: photograph of holotype (DS), Ferris and Ernst 13162 93 very dry areas. It should be noted also that collections of the few-leafed variety have been found at altitudes up to 8,350 feet (Coville and Gilman 118). The distribution of var. oligophylla is centered in the eastern Death Valley region and many localities at 4500-5000 feet receive sparse annual precipitation (e.g., Cottonwood Mts.). Plants of both varieties have been grown under uniform conditions in the greenhouse, and both have retained their typical growth habits. The infraspecific taxonomic dispositions of P. megalocephala are based almost entirely upon morphological evidence. Chromosome numbers have been obtained only from single populations of var. megalocephala (n=17), var. oligophylla (n=34), and var. intricata (n=19-+ romatographic sampling has been equally deficient, but phenolic patterns of the south- western species are so much alike that they are of limited value at the infraspecific level. However, the pattern obtained for var. intricata is quite different from those of the other varieties. Much additional populational study of P. megalocephala, particularly for chromosome numbers, will be necessary before the taxon is properly understood. The larger leafed var. lee acai is morphologically uniform over a fairly large area (Fig. 5). The leaves of a few collections are slightly re- duced in size (e.g., Applegate 6936 and Shockley 122), but these do not really approach the narrow foliage characteristic of var. oligophylla. In Marble Canyon, California, however, some plants from the same collection (e.g., Duran 2845) exhibit leaf morphology lke var. megalocephala and/or oligo- phylla. Other specimens from the southern California region north of Death Valley show narrow leaves like var. oligophylla (e.g., Alexander and Kellog 2541 and Roos and Roos 5979) and some plants have a pubescence approach- ing P. inyoensis. The Marble Canyon population particularly has been inter- preted as evidence that var. megalocephala and var. oligophylla intergrade. Elsewhere in their respective distributional areas, the two taxa appear to manifest genetic integrity. One population, here relegated to var. megalocephala, is suggestive of hybridization with P. inyoensis—or perhaps it is merely indicative of recent contiguity between P. megalocephala and P. inyoensis. This is evidenced by some plants from a collection (Roos and Roos 5938) from near Blue Bell Mine, Inyo Mts., California, which are intermediate in habit between P. megalocephala and P. inyoensis and have pubescence approaching that of the latter taxon. Other plants from the same collection have leaves which resemble those of var. oligophylla. Further speculation about the intermedi- ate variability is not advisable until more populational information can be obtained, but the close relationship of the taxa involved is apparent. The controversial collection (Howell 3906) from Surprise Canyon, Pana- mint Mts., California, seems to belong with var. oligophylla rather than with P. congesta as indicated by Ferris (1959). Some plants of this collection have a single bristle, and along with Langenheim 3746 of var. intricata, constitute the only specimens of P. megalocephala which are known to have 94 pappus bristles. 4 o) PERITYLE MEGALOCEPHALA var. intricata (Brandeg.) Powell comb. nov. Laphamia intricata Brandeg. Bot. Gaz. 27:450-451. 1899. TYPIc: NE- Vv : Sheep Mountain, on rocks, May-Oct, 1898. C. A. Purpus 6134 erate UC! Isolectotype, DS! US!) Pahrump ‘Valley May-Oct., A. Purpus 6051 age NY! UC! US! ae DS!). Pik ee intricata (Brandeg.) Rydb. N. Amer. FI. 34:2 Laphamia megacephala Wats. subsp. intricata co ‘Keck, Aliso 4: 101. 19% Plants 13-35 cm high, subcanescent, profusely branched, stems slender, sparsely leafed; leaves few, alternate, 4-8 mm long, linear, sessile or petio- late; capitulescence of 2-6 tightly aggregated heads on short peduncles rarely exceeding 1 cm; floral features essentially like var. oligophylla except heads smaller; heads 4.0-6.5 mm long, 3.5-5.5 mm wide; corollas with well- differentiated tubes and throats, the throats broad-tubular; achenes coarse- pubescent on margins and surfaces; pappus of short cilia-like squamellae, rarely of 1-2 bristles. Rare in dry mountains of S Nevada. Fig. 5. Spring-Fall, when it rains. Specimens examined:? NEVADA: Clark Co. Muddy Mts., J. Langenheim 3746 (UC); Nye Co. 23 mi W of Indian Springs, T. Watson and J. Averett 250: Checkpoint Pass, P. V. Wells s.n. (RSA); Checkpoint Pass, P. V. Wells s.n. (RSA) Perityle megalocephala var. intricata differs considerably from the rest of the species. In fact, future study may reveal the taxon warrants specific recognition. Infraspecifiec status has been retained for var. intricata because it exhibits apparent affinity with P. megalocephala var. oligophylla and because very little biosystematic information has been obtained for the taxon. Only five collections of typical var. intricata are known to me, including the types. One other collection with rather intermediate morphology, Watson and Averett 250, approaches var. oligophylla. This variety is similar to var. oligophylla in habit (excluding capitulescence), but its leaves are fewer and more reduced (linear to subfiliform) than those of var. oligophylla. Probably it is the paucity of foliage which has prompted previous workers to include var. oligophylla with var. intricata. The most conspicuous delimiting feature of var. intricata is its capitulescence of tightly aggregated heads on short peduncles. Typically the capitulescence of var. megalocephala and var. oligophylla is of solitary heads, although some plants of the latter varieties * After this work was accepted for saa J. T. Howell called my attention to an anomalous cae of Perifyle from Utah (Grand Co., seepage on cliff of Colorado River cany¢ I mi of Moab aden el sa ft, 6 Sept 1968, Howell and True 44841, CAS) eee ly, I ae ve assigned the collection to var. infricafa because of similarities in’ habit, achenes, and pappus. The Utah specimen differs from var. infricata, however, in its capitu- lescence of loosely clustered heads, involucral pea acts, and shape of upper leaves. Eventually the Utah population might prove worthy of separate varietal status, or even warrant specific status as suggested by A. Cronquist (on the How ell and ‘True label). 95 have a capitulescence of 2-3 heads in loose clusters on rather long peduncles. The var. intricata also differs somewhat from other populations of P. megal- ocephala by its smaller heads, shorter disc corollas with well differentiated tubes, achenes which are coarse-pubescent on the margins and surfaces, and a rather conspicuous crown of pappus squamellae (cilia). The latter feature is characteristic of the supposedly ancestral P. stansburii, as is the pappus of 1-2 bristles found only in the Langenheim 3746 collection of var. intricata. PERITYLE GRACILIS (M. E. Jones) Rydb. N. Amer. Fl. 34:19. 1914. Laphamia gracilis M. E. Jones, Proc. Calif. Acad. Sci. 5:703. 1895. TYPE: ZONA: Below Nagle’s Ranch, edge of Buckskin Mts., crevices of limestone rocks, 7000 ft, in very dry places, 15 Sept 1894, M. E. Jones 6050c (Holotype, POM; Isotype, US!). sl fastigiata Brandeg. Bot. Gaz. 27:451. 1899. TYPE: NEVADA: p Mts., 4-5000 ft, May-Oct, 1898, C. A. Purpus 6142 (Holotype, UC! 1 pes, US!). Monothrix fastigiata (Brandeg.) Rydb. N. Amer. Fl. 34:21. 1914. Perityle fastigiata (Brandeg.) ean Southwest. 4:204. 1959. Plants 10-17 cm high, densely puberulent to nearly glabrous; leaves alter- nate, 1.0-2.3 cm long, 0.5-1.5 cm wide, often extremely reduced at plant apexes, densely puberulent to nearly glabrous, typically 3-cleft, tri- ee subcruciform, or with 2-3 lobes per margin, the lobes or marginal seg lanceolate to oblong-lanceolate, acute, occasionally ane lobed ke the distal segments, usually attenuate at the base; petioles 2-5 (7) mm long; capitulescence of solitary heads or small clusters borne on slender or stout peduncles (usually 0.5-1.5 em long); involucres campanulate, heads 5-7 mm long, 4.0-5.5 mm wide; receptacle convex, flattened, 1.7-2.0 mm wide; in- volucral bracts (8) 14-17, 3.5-5.5 mm long, 0.7-1.2 mm wide, sublanceolate to linear-lanceolate, keeled; ray flowers absent; disc flowers 30-42, corollas yellow, (2.5) 2.8-3.6 mm long, the tube 0.8-1.1 mm long, the throat tubular to subtubular, (1.3) 1.6-2.0 mm long, the lobes acute, 0.4-0.6 mm long; style branches 1.2-1.8 mm long; achenes 1.8-2.5 (2.8) mm long, oblong-obconical, rounded or angled on one or both surfaces, with thin callous margins, short- pubescent on the margins and both surfaces; pappus typically of 1 rather stout bristle (rarely 2, 3, or none), 1-2 mm long, shorter than 1.0 mm, or vestigial, a callous crown with vestigial squamellae often apparent espe- cially above the margins; chromosome number, n= 34 Common in its restricted area of occurrence, especially along Meadow Valley Wash in SE Nevada. Fig. 5. Spring-Fall. Representative specimens: A: Lincoln Co. Railroad Canyon, 3 mi NE of Panaca, D. H. Galway 8263 (US, UT); Caliente, M. E. Jones s.n. (POM); Panaca, M. E. Jones s.n. S, DS, GH, POM, RM); Meadow Valley Wash along Hwy 93, 0.5 mi N of Caliente, A. M. Powell and B. Calvert 1272 (SRSC): 0.5 mi N of Caliente, Powell and D. Patterson 1626 (SRSC); 20 mi S of Caliente, S end of Rainbow Canyon on road to Elgin, Meadow Valley Wash, P. Train 2453 (DS, NY, ol 96 UC). Like several other species of sect. Laphamia, P. understood biologically. Herbarium specimens indicate considerable varia- tion in leaf and flowering head morphology. A single population of this species from near Caliente, Nevada, has been observed in the field (Powell and Calvert 1272), and its morphology does not correspond entirely with that of specimens from other localities, including the types. Plants from the ‘leaves, stouter stems, larger heads, and in- gracilis remains poorly Caliente population have large1 volucral bracts which are connate at the base, as compared to plants from other areas. An explanation of this morphological disparity might be as- sociated with the tetraploid chromosome number (n—34) which has been determined for plants from the Caliente population. Chromosomal informa- tion for plants from other areas is not available. Infraspecific polyploidy is frequent in Perityle species (Powell, 1968b), and it is not uncommon to find some gigas features associated with an increase in ploidy level. There is a need for further investigation in order to ascertain whether or not P. gracilis is also comprised of diploids. The interspecific relationship of P. gracilis is not clear, logical and chromosomal evidence suggests an affinity with P. Considering especially the southern most populations of P. stansburii, the two taxa are not very different in total morphology, except that the leaves but most morpho- i stans burit. of P. gracilis are typically tripartite and the heads are discoid. The leaves of some specimens of P. stansburii show a strong tendency toward being trilobed. There is some indication that P. gracilis is related to P. congesta, and thus a question arises about evolutionary sequence in the derivation of P. gracilis. The similar total morphology of P. congesta, especially the trilobed leaves, is suggestive of kinship. Additionally, the type collection of P. gracilis, which was near the original collection of P. congesta (Fig. 5), has enhanced the possibility of a relationship between the two taxa. A careful review of most evidence which has been compiled for both species has failed to verify any close affinity, although the location of the Jones collection of P. gracilis remains an intriguing anomaly. Chromosomal information appears to sub- stantiate the derivation of P. gracilis (n—34) from P. stansburii (n—17), rather than from P. congesta (n—16). It follows then that Perityle congesta (and P. tenella; n= 16, n-—32) could have evolved from diploid P. gracilis by reduction in chromosome number, and thus reflect derived instead of ancestral morphological similarities. It must be emphasized, however, that any reliance on chromosomal data is at best tenuous because so little data of this sort are available for the species. 6. PERITYLE INYOENSIS (Ferris) Powell, Sida 3:278. 1968. Laphamia inyoensis ange ea ib. Dudley Herb. 5:104. 1958. TYPE: ORNIA: Inyo Co. in rocks facing NE ac ross divide above Cerro Gordo mines, Cer rro Go ac Peale 8200 ft, 7 July 1942, Alexander and Kellogg 3056A (Holotype, DS! Isotype CAS! GH! LL! NY! POM! RM! UC!). 97 Plants 12-25 cm high, in dense, leafy clumps, pilose-villous; leaves op- posite or alternate, 0.8-1.8 (2.1) cm long, 0.6-1.2 (1.5) em wide, pilose- villous intermixed with short glandular hairs, ovate to broadly so, occasion- ally somewhat ovate-orbicular or ovate-deltoid, margins serrate-lobed, lobes and apex acute to subacute; petioles 0.5-2.0 (5.0) mm long; capitulescence of solitary heads or small clusters of 2-3; peduncles usually 0.8-4.0 cm long; involucres campanulate, heads 7.0-8.5 (9.0) mm long, 5-7 (8) mm wide; receptacle convex and rounded, 1.5-2.0 mm wide, ca. 1.0 mm high; involu- cral bracts 14-21, 5.5-6.5 mm long, 1.1-1.5 mm wide, sublanceolate to linear- lanceolate, keeled; ray flowers absent; disc flowers 35-60, corollas yellow, 4-5 mm long, the tube 1.4-1.6 mm long, the throat subtubular to subfunnel- form, 2.0-2.4 mm long, the lobes acute, 0.6-0.7 mm long; style branches 2.0- 2.3 mm long; achenes (2.5) 3.0-3.3 (3.5) mm long, narrowly obconical to subcblanceolate, rounded or angled on one surface, occasionally both, with thin callous margins, pubescent on the margins and both surfaces; pappus absent except for an inconspicuous callous crown, rarely with conspicuous, vestigial, squamellae, especially evident above the achene margins; chromo- some number, n=ca A rare endemic apmanenty restricted to higher altitudes, 5900-8500 ft, in S Inyo Mts. of California. Fig. 5. Spring-Fall. Specimens examined: CALIFORNIA: Inyo Co. N slope of Cerro Gordo Peak, E of divide Inyo Mts., A. M. Alexander and T. Kellogg 3056 (DS, GH, LL, NY, RM, US); 2 mi S of Cerro Gordo Spring, Inyo Mts., M. D. Decker 746 (CAS, DS, RSA); Tale Canyon, Southern Inyo Mts., FE. C. Jaeger s.n. (POM); 4% mi N of Cerro Gordo Mine, E of ridge near road, Inyo Mts., M. Kerr s.n. (CAS, RSA); NIE of Cerro Gordo Mines, A. M. Powell and B. Calvert 1275 (SRSC). Because of the several Laphamian taxa which occur in the mountains of southeastern California (Fig. 5), it is possible that the area was a major center of speciation. Perityle inyoensis appears to be restricted to relatively high altitudes (5900-8500 ft.) on or near Cerro Gordo Peak. The latter species is manifestly allied to P. megalocephala which is more widely distributed to the north and east at intermediate and lower altitudes. Perityle villosa, which is poorly distinguished from P. inyoensis, is known only from the type locality at 6500-7000 ft. in Hanaupah Canyon. In addition, a collection from Surprise Canyon (Howell 3906) appears anomalous among the other taxa. Ferris (1958) remarked about its similarity to P. congesta of Arizona, but in the present treatment it has been placed with P. megalocephala var. oligophylla. Perhaps much of the taxonomic uncertainty surrounding the California taxa could be removed through an extensive field survey in the areas concerned, but this will be extremely difficult because of the inacces- sibility of habitats and limited flowering period. Perityle inyoensis can be distinguished from P. megalocephala by its smaller habit, pilose-villous pubescence, essentially opposite leaves which are ovate to ovate-deltoid with serrate-lobed margins, and short petioles. — 98 Perityle inyoensis is poorly delimited from P. villosa, except that the latter species has leaves which are entire or with 1-3 short lobes per margin. 7. PERITYLE VILLOSA (Blake) Shinners, Southwest. Nat. 4:205. 1959. Laphamia villosa Blake, Proc. Biol. Soc. Washington 45: 142. 1932. TYPE: CALIFORNIA: Inyo Co. in shaded rock crevices, middle fork of Han- aupah Canyon, Panamint Mts., 2090 m, 22 Sept 1931, FF’. V. Coville and A. I’. Gilman 108 (Holotype, US! Isotype, US! Photograph, DS!). Plants 13-20 em high, villous; leaves alternate, 1.2-2.2 em long, 0.4-1.0 em wide, villous, ovate to ovate-cuneate and entire or ovate-cuneate and lobed, with 1-3 short, pointed lobes on each margin, acute or cuneate at the base, acute at the apex; petioles 3-6 mm long; capitulescence of solitary heads or small clusters of 2-3; peduncles 1-2 (2.5) em long; involucres campanulate, heads 7.5-9.5 mm long, 5-7 mm wide; receptacle flattened to slightly convex, 1.3-2.5 mm wide; involucral bracts 13-23, ca. 6.0 mm long, 1.0-1.5 mm wide, oblong-lanceolate to linear-lanceolate, inconspicuously keeled; ray flowers absent; disc flowers 30-75, corollas yellow, 4-5 mm long, the tube 1.0-1.7 mm long, the throat subfunnelform to tubular, 1.8-3.0 mm long, the lobes acute, 0.7-0.8 mm long; style branches 1.8-2.5 mm long; achenes 3.0-3.5 mm long, linear to oblong-obconical, typically rounded on both surfaces, with callous margins, short-pubescent on the margins and both surfaces; pappus absent, except for a conspicuous to inconspicuous callous crown, or in some heads of 1-2 bristles of equal or unequal length, 1-2 mm long; chromosome number, n= 10-22 IT plus 3-19 I. Known only from the type locality. Fig. 5. Spring-Fall. Representative specimens: CALIFORNIA: Inyo Co. Hanaupah Canyon, Panamint Mts., F. V. Coville and A. I’, Gilman 109 (US); middle fork of Hanaupah Canyon, Panamint Mts., Death Valley Nat’] Mon., S. Sikes 100 (SRSC). The complexity of P. villosa has been magnified through examination of a recent collection (Sikes 100). Leaves of the type specimens are essentially entire, while different plants from the Sikes collection have leaves which are either entire or short-lobed. In addition, plants with opposing leaf forms also exhibit independently several other differing features, thus suggesting the side-by-side occurrence of two morphotypes at the Hanaupah Canyon locality. The Sikes specimens with entire leaves conform to the type specimens except that in a few heads of some individuals, achenes with a pappus of 1-2 awns are found. The plants with lobed leaves also differ by producing larger heads, wider receptacles, more bracts, longer disc corollas, and longer, tubular throats. In fact, the ‘‘lobed’’ and ‘‘entire’’? forms are suffi- ciently different, that if they occurred in separate localities, they probably would be recognized as separate taxa. There is some evidence which indicates that the two morphotypes of P. villosa, and accompanying variability, has resulted from the occurrence of diploids, tetraploids, and triploidy, in the population. Meiosis observed in “greenhouse buds” from a lobed-type plant revealed extremely abnormal 99 configurations of n=10-22 II and 3-19 I. This cytological information sug- gests triploidy on the expected basis of n=17 II + 17 I, and, pending further study, at least provides a possible explanation for the complexity observed in P. villosa. io 2) PERITYLE TENELLA (Jones) Macbride, Contrib. Gray Herb. n.s. 56:39. 1918 Laphamia palmeri A. Gray, — Amer. Acad. 13:372-373. 1878. TYPE: NA: at Beaverdam, growing in crevices of rock in canyons, 1877, E. Palmer 199 ee GH! Isotypes, ND! NY! US! Laphamia palmeri var. tenella E. Jones, Proc. Calif. Acad. IL. 9: 708. 1895. TYPE: AH: See in crevices of sandstone rocks, 4000 ft., 16 May 1894, M. E. Jones 5249aa (Holotype, POM! Isotype, US!). Laphamia tenella M. E. Jones, Proc. Calif. Acad. II. 5:703. 1895. Monothrix palmeri (A. Gray) Rydb. N. Amer. FI. 34:21. 1914. Plants 15-25 cm high, densely to sparsely short pilose-hirsute, densely leafy; leaves alternate, 1.0-2-3 cm long, 0.8-1.8 cm wide, sparsely to densely short pilose-hirsute, typically broadly deltoid-ovate, less often broadly sub- ovate to subcordate, the margins regularly to irregularly serrate or dis- sected into acute, shallow lobes; petioles 2.0-8.0 mm long; capitulescence of solitary heads or small clusters; peduncles usually 0.4-2.5 em long; involucres campanulate, heads 6-7 mm long, 4-6 mm wide; receptacle flattened, 1.5-1.8 mm wide; involucral bracts ca. 13, 4.0-4.5 mm long, 1.1-1.3 mm _ wide, lanceolate-elliptic, inconspicuously keeled; ray flowers absent; disc flowers 23-35, corollas yellow, 3.0-4.0 mm long, the tube 1.2-1.6 mm long, the throat subtubular to narrowly funnelform, 1.2-1.6 mm long, the lobes acute, 0.6-0.7 mm long; style branches 1.0-1.8 mm long; achenes 2.5-3.0 mm long, very narrowly obconical, rounded or obscurely angled on one or both surfaces, with thin callous margins, short pubescent on the margins and both surfaces; pappus of a single bristle, 2.0-2.6 mm long; chromosome number, n=—16. Crevices of rock bluffs in protected canyons, usually at lower altitudes, extreme SW Utah and NW Arizona. Fig. 5. Spring-Fall. Representative specimens: ARIZONA: Mohave Co. Burnt Canyon, W. P. Cottam s.n. (UT); 4 mi E of Beaverdam, A. M. Powell and S. Sikes 1386 (SRSC); 5.7 mi S of Jacobs Well, E base of Virgin Mts., P. H. Raven and D. Verity 15483 (RSA, UC). NE- VADA: Clark Co. 2nd canyon § of Limestone Point, Virgin Mts., P. A. Munz 16709 (DS). UTAH: Washington Co. Great Arch Trail, E of tunnel, Zion Nat’l Park, S. Braem 852 (DS); Zion Nat'l Park, Cottam 3442 (UT): St. George, Cottam 7244 (UT); near observation point, Zion Nat'l Park, O. Degener and L. Peiler 16544 (NY); Zion Nat'l Park, A. Eastwood and J. 7 Howell 1161 (CAS, NY); Clear Creek, Zion Park, Eastwood and Howell 6330 (CAS); Hurricane, Eggleston 14863 (US); Zion Canyon, A. D. Garrett 2958 (NY); Zion Canyon, M. E. Jones s.n. bea Ww mi-4 mi up E Rim Trail, Zion Canyon, Zion Nat’l Park, Powell and Calvert 1267 (SRSC); Zion Canyon along E. Rim Trail, Powell and D. aia 1627 (SRSC); Clear Creek 100 Canyon, Zion Park, H. D. Ripley and R. C. Barneby 4930 (CAS, NY); E Rim Trail, Zion Canyon, Zion Nat’l Park, K. E. Weights 964-A-Z (UT); Narrows Trail, Zion Canyon, Weights s.n. (UT); Zion Nat’l Park, A. M. Woodbury 21 Although Laphamia palmeri A. Gray is an older name, that specific epithet was preempted when Laphamia was transferred to Perityle (Shinners, 1959; Powell, 1968a). According to M. E. Jones’ Journal of Botanical Exploration (Howell and Munz, 1965), his specimens of P. tenella (5249aa) were collected in Zion Canyon above Springdale, Utah. Perityle tenella is best distinguished from the closely related P. congesta by its broadly deltoid-ovate leaves with margins which are regularly to ir- regularly serrate or dissected into acute, normally further serrated lobes. No differences worthy of varictal status were detected between Utah and Arizona populations. There is a striking similarity in the habits of P. tenella (n—16) and P. castillonii (n=17) of Chihuahua and Coahuila, Mexico, but their extreme geographic separation would appear to negate any consideration that the two taxa are closely related. Besides their chromosome numbers, the two species differ in but a few technical features. ro) PERITYLE CONGESTA (M. E. Jones) Shinners, Southwest. Nat. 4:204. 1959 Laphamia congesta M. E. Jones, Proc. Calif. Acad. Sci. 5:703. 1895. Pk: ARIZONA: in clefts of rocks on the mesa below the Buckskin ete 7000 ft., 21 Sept 1894, M. EF. Jones 6063 (Holotype, POM! Isotypes, OM! RM! UC! US! Photograph DS! Patel toumeyi Rob. and Greenm.., Amer. Journ. Sei. 50:176. TYPE: ARIZONA: in the Grand Cany on, 12 July 1892, J. W. Toumey 645 (Holotype, US! Isotypes, DS! GH! Photographs DS!). Monothrix congesta (M. E. Jones) Rydb. N. Amer. FI. 34:20. 1914. Monothrix toumeyi (Rob. and Greenm.) Rydb. N. Amer. Fl. 34:20, 1914. Plants 10-30 cm high, densely clumped and leafy, hirtellous; leaves alter- nate, rarely opposite, 1.0-2.5 cm long, 0.4-0.8 em wide, hirtellous, typically ovate and 3-lobed, but often ovate-lanceolate to ovate-clliptic and 3-lobed rarely 4-5 lobed, serrate, or entire, rounded or obtuse at the apex; petioles 2.5-10 (14) mm long; capitulescence subcymose, of small clusters, or less often of solitary heads; peduncles usually 0.3-1.7 cm long; involucres cam- panulate, heads 5.0-6.0 (6.5) mm long, 3.5-4.5 (5.0) mm _ wide; receptacle convex, typically rather flattened, less often slightly rounded, 1-2 mm wide; involucral bracts 13-16, 3.5-4.5 mm long, 0.8-1.4 mm wide, oblong-lanceolate to sub-lanceloate, rather flattened to keeled; ray flowers absent; disc flowers 30-40, corollas yellow, 2.5-3.0 mm long, the tube 0.8-1.0 mm long, the throat tubular to subfunnelform, 1.2-1.5 mm long, the lobes acute, 0.3-0.5 mm long, style branches 1.2-1.5 mm long; achenes (2.0) 2.5-3.0 mm long, narrowly obconical, rounded or angled on one or both surfaces, with thin callous margins, very short pubescent on the margins, slightly pubescent on both surfaces; pappus of 1-2 subequal to very unequal bristles, the longest 2.0-2.7 101 mm long, and an obscure crown of hyaline squamellae; chromosome num- ber, n=16. Common in rock bluffs along the high rims of the Grand Canyon, Arizona, and vicinity. Fig. 5. Spring-Fall. Representative specimens: ARIZONA: Coconino Co. Grand Canyon, M. Armstrong 216 (NY); Hava- supal Canyon, E. U. Clover 5008 (LL); near the top of Hualpai Trail, Clover 7152 (LL); North Rim, Grand Canyon, R. EF. Collom s.n. (US); North Rim, Grand Canyon, Collom 1116 (US); Jumpup . Station, Kaibab Plateau, Kaibab Nat'l] Forest, R. A. Darrow 2982 (CAS); Hualpai Hilltop, Kaibab Nat’l Forest, Darrow 3130 (CAS); Havasu Canyon, C. F. Deaver 1487 (CAS); Bright Angel Trail, Grand Canyon Ariz., A. Eastwood 22 (US); Grand Can- yon, Eastwood 3606a (CAS, GH); Grand Canyon, Eastwood 3506 (CAS); Hermit Trail, Hastwood 5817 (CAS); Grand Canyon, Kaibab Trail to Roar- ing Springs, Hastwood and J. T. Howell 1013 (CAS, POM); Kaibab Trail to Roaring Springs in Grand Canyon Nat’] Park, Eastwood and Howell 1021 (CAS); North Rim of Grand Canyon, Eastwood and Howell 7038 (CAS); Kaibab Trail to Roaring Springs, Grand Canyon, Hastwood and Howell 7059 (CAS, GH, US): N slope of Willow Point, L. N. Goodding 279-48 (UC); Oak Canyon, Kaibab Plateau, Goodding 425-48 (UC); Jumpup Canyon, Kaibab Plateau, Goodding s.n. (CAS); trail into Grand Canyon, A. E. Hitchcock 84 (US); head of Hualvai Canyon, Grand Canyon Nat’! Park, Howell 26369 (CAS, RSA); Hualpai Canyon, Grand Canyon Nat’l Park, Howell 26381 (CAS NY, UC); Havasu Canyon, Grand Canyon Nat’l Park, Howell 26441 (CAS, RM, SMU); Bright Angel Canyon, Grand Canyon, F.. W. Hunnewell 10898 (GH); Grand Canyon, F. R. Irvine s.n. (DS); Peach Springs, Jones s.n. (POM); Bright Angel, Grand Canyon, Jones s.n. (DS, POM); Bright Angel Trail to Grand Canyon, J. B. Leiberg 5954 (US); Bright Angel Trail, Grand Canyon, G. E. Osterhout 6980 (RM); Cape Royal, Kaibab Nat’l Forest, F. W. Peirson 7419 (CAS, POM, RSA); head of Bright Angel Trail, Grand Canyon Nat’l Park, A. M. Powell 1264 (SRSC); Mather Point, South Rim of Grand Canyon, Powell and B. Calvert 1265 (SRSC); Bright Angel Point, North Rim, Grand Canyon Nat’l Park, Powell 1266 (SRSC); Grand Canyon, C. A. Purpus 83807 (UC, US); Grand Canyon, V. Rattan s.n. (UC); Kaibab Trail, Grand Canyon Nat’! Park, P. H. Raven 13118 (CAS, DS, NY); top of Mooney Falls, Havasu Canyon, M. Sutton 8 (CAS); Grand Canyon, J. Toumey 645 (DS, GH, US). Mohave Co. Tuweep, at canyon rim, W. P. Cottam 8613 (UT); outer gorge, Toroweap, Cottam 13319 (CAS, UT); ae Horse Canyon, Grand Canyon Nat’! Park, Cottam 13535a (SMU, UT); g Cove, Outer Gorge, Grand Canyon Nat’] Monu., Toroweap, Cottam aS (UT). Yavapai Co. Grand Canyon, EF. D. Wooton 389 (US). Perityle congesta is a slightly variable species which occurs mainly around the upper rims of the Grand Canyon of Arizona. Morphologically the taxon is poorly delimited from P. tenella which occurs just to the north- west in Arizona and Utah. Perhaps the only character which can be used 102 consistently to distinguish P. congesta from P. tenella is the usually smaller leaves which are typically ovate and 3-lobed, but often ovate-lanceolate to ovate-elliptic and 3-5 lobed, serrate, or entire. Chromatographic patterns obtained for the two species are not noticeably different. The basic chromosome number of «—16 for P. congesta and P. tenella set the taxa apart from all others in sect. Laphamia (Powell, 1968b). Their re- lationship to other Laphamian species is not clear, but most evidence sug- gests affinity with P. gracilis (n—34) of the ‘‘southwestern alliance.’ Geo- grapically, the ranges of P. congesta and P. tenella are close to P. gracilis. In this genus, because of previously explained evolutionary tendencies, adjacent species are usually related. There is no compelling morphological indication of relationship with P. gracilis, but the three taxa are similar in general habit and floral structure. The chromatographic patterns of P. congesta and P. tenella are most like that of P. gracilis and thus, this chem- ical character might be indicative of a true relationship which has remained rather obscure on other grounds. It is interesting to note that both P. congesta (Jones 6063) and P. gracilis (Jones 6050c) were collected on or near the Buckskin Mts. of Arizona, which are located near the western edge of the Kaibab Plateau south of Fredonia (Howell and Munz, 1965). Jones’ Arizona collection of P. gracilis is out-of- place with all other records for the species. There is some indication that P. congesta is allied with P. megalocephala var. oligophylla of southeastern California as suggested by Ferris (1958). One collection of P. megalocephala var. oligophylla from Surprise Canyon in the Panamint Mts. near Death Valley (Howell 2906) is rather anomalous among the California taxa of sect. Laphamia, but closely resembles P. congesta. Any close relationship of P. congesta to the California species appears tenuous because of the present geographical gap, but it is suggested that they have ancestral affinity through a common ancestor of the south- western alliance, PERITYLE SAXICOLA (Eastwood) Shinners, Southwest. Nat. 4:204-209. 1959. Laphamia saxicola Eastwood, Proc. Calif. Acad, Sci. IV, 20:159-160. TYPE: ARIZONA: Maricopa Co. near the Roosevelt Dam on the road to Fish Creek, Apache Trail, 22 May 1929, A. Eastwood 17401 (Holo- type, CAS! Isotypes, CAS! DS! GH! POM! UC! US!). Plants 20-40 em high, spreading in clumps to ca. 30 cm wide, leafy to the tops, essentially glabrous except the peduncles glandular-puberulent; leaves opposite below, inconspicuously alternate above, 3.0-8.5 em long, glabrous, typically tripinnate, less often bipinnate, the segments long, linear, to flat- filiform; petioles 1.0-3.0 cm long, scarcely wider than the leaf segments; capitulescence of solitary heads or 2-3 in loose clusters; peduncles 1-3 ¢m long; involucres broadly campanulate to hemispheric, heads 7.5-10 mm high, 10-14 mm wide, mostly extending above upper leaves; receptacle convex, slightly rounded, 2-3 mm wide; involucre appearing uniseriate, bracts ca, 10. — 103 25, (7.5) 5-7 mm long, ca. 0.9 mm wide, linear-lanceolate, keeled; flowers ca. 100-200; ray flowers ca. 12-17, pistillate and fertile, the tube 2-3 mm long, ligules yellow, 4-6 mm long, 2.0-3.2 mm wide, oblong to oblong-ovate, tri- lobed at the apex; dise ccrollas yellow (4.0) 5.0-6.5 mm long, the tube (1.5) 1.7-2.0 mm long, the throat tubular to narrowly funnelform, the lobes acute, 0.5-0.6 mm long; style branches ca. 1.8 mm long; achenes 2.8-3.2 mm long, oblong-obconical to narrowly obconical, with conspicuous callous margins, shert-pubescent on the margins especially and both surfaces; pappus of a single bristle, 4-5 mm long, rarely reduced to ca. 2.0 mm; chromosome number, n=—17. Locally frequent but otherwise rare, growing in igneous rocks near the type locality, and Tonto National Monument. Fig. 3. Spring-Fall. Representative specimens: ARIZONA: Gila Co. E of entrance to Tonto Nat’! Monument, A. M. Powell 1262 (SRSC). Maricopa Co. rocks near Roosevelt Dam, A. Eastwood 8695 (CAS); Roosevelt Dam, R. H. Peebles 9420 (DS, US), % mi below Roosevelt Dam, S. Sikes 97 (SRSC The endemic P. saxicola (n=17) is clearly related to P. gilensis (n—ca. 17, 34-36). Perityle saxicola is easily distinguished by its thrice- to bi-pin- nately dissected leaves which have linear segments, but otherwise the two species are morphologically similar. 11. PERITYLE GILENSIS (Jones) Macbride KEY TO THE VARIETIES la. Petioles 0.9-18 ecm long ....... . . . Ila. var. gilensis. lb. Petioles 2.5-4.5(8.0) cm long . lib. var. salensis. lla. PERITYLE GILENSIS (Jones) ‘Macbude var. " GILENSIS, Contrib. Gray Herb. n.s. 56:39. 1918. ge eal gilensis Jones, Zoe 2:15. 1891. TYPE: ARIZONA: growing monet ocks, at Putnam’s Ranch, near the Gila River, 23 May 1890, Jon nes s.n. (Holotype, POM! Isotypes, UC! US!). noe eae aia Rydb., N. Amer. FI, 34:24. 1914. Laphamia eee nica Eastw., Proc. Calif. Acad. IV 20:159. 1931. TYPE: : Maricopa Co. Fish Creek, Apache Trail, 18 May 1919, A. Eastw a 8753 (Holotype, CAS!). Laphamia dura A. Nels., Amer. Bot. 23:266. 1936. TYPE: ARIZONA: aricopa Co. Salt River Valley, near Canyon Lake, 3 May 1925, A. Nelson 10323 (Holotype, RM! Isotype, DS! Photograph US!). Plants 22-40 cm high, often spreading to ca. 40 cm in dense clumps, essen- tially glabrous, appearing scarcely leaved except near the base; leaves essentially opposite, rarely alternate distally, basal leaves 1.4-3.0 cm long, 0.4-1.0 (2.0) cm wide, becoming progressively smaller upwards, essentially glabrous to sparsely puberulent, typically pinnately trifoliate with the seg- ments very broadened distally, often cruciform in appearance, less often merely trilobed to divided, terminal segment entire to lobed; petioles 0.9-1.8 cm long: capitulescence of solitary heads or groups of 2-4 on rather stout 104 peduncles; the peduncles usually 1.0-3.5 cm long; involucres broadly cam- panulate to hemispheric, heads 8-11 mm long, 9-12 mm wide, very conspicu- ous above the sparsely leafed stems; receptacle convex, rounded, or flat- tened, (2.0) 2.5-4.5 mm wide, 0.8-1.2 mm high; involucral bracts 25-30, 5-7 mm inns (.8-1.5 mm wide, sublanceolate to oblanceolate, attenuate, keeled; flowers ca. 100-200; ray flowers ca. 10, pistillate and fertile, the tube ca. 2.0 mm long, ligules yellow, 5.0-7.0 mm long, 2.0-3.8 mm wide, oblong to sub- ovate, tridentate at the apex; dise corollas yellow, (4.5) 5.0-6.5 mm long, the tube (1.0) 1.5-2.0 mm long, the throat subtubular to narrowly funnelform, 2.5-3.5 mm long, the lobes acute, 0.7-0.8 mm long; style branches (2.0) 2.5-3.0 mm long; achenes 2.6-3.4 mm long, narrowly obconical or oblanceo- late to narrowly obconical-elliptic, usually rounded on one surface and flat- tened on the other, with conspicuous callous margins, short-pubescent on the margins especially and both surfaces; pappus of a single bristle (rarely 2-3) 3.5-4.6 mm long, and a rather obscure crown of hyaline squamellae; chromosome number, n= 34-36. Common in southeast-central Arizona, in crevices of igneous bluffs and boulders particularly along canyon walls, sometimes found growing at the base of boulders, seemingly in soil, but probably always rooted in subter- ranean crevices. Fig. 3. Spring-Fall. Rerpresentative specimens: ARIZONA: Gila Co. San Carlos Indian Reservation, vicinity of Blue River Crossing, U.S. Hwy 60, B. Maguire 13070 (NY). Maricopa Co. Horse-Mesa Dam, along Apache Trail, A. Eastwood 17375 (CAS, GH, POM, US): Fish Creek Canyon, T. H. Kearney and R. H. Peebles 14490 (US); Fish Creek Hill, Kearney and Peebles 15133 (CAS, SMU); Fish Creek Canyon, FE. P. Killip 82723 (US); Apache Trail, near Fish Creek, A. Nelson and R. A. Nelson 1772 (NY, RM, UC); Fish Creek, Peebles, G. J. Harrison, and Kearney 5247 (US); near Roosevelt Dam, Fish Creek, Peebles and Harrison 5287 (US); crossing of Fish Creek and Hwy 88, Apache Trail, A. M. Powell and B. Calvert 1263 (SRSC); Apache Trail near Tortilla Flat, H. D. Ripley and R. C. Barneby 2830 (NY); top of Fish Creek Hill, Apache Trail, S. Sikes 107 (SRSC). Pinal Co. along Queen Creek, Thompson Arboretum, near Superior, BE. H. Graham 3721 (DS); Queen Creek Canyon, Kearney and Peebles 9224 (US); Devil’s Canyon, Peebles, Harrison, and Kearney 3183 (US); 4 mi NE of Superior, near Superior, near Hwy 60-70, Powell and B. McAfee 1317 (SRSC); Thompson Arboretum grounds, near Superior, J. Whitehead 2255 (RM). lib. PERITYLE GILENSIS var. saiensis* Powell, var. nov. Fig. 7. TYPE: ARIZONA: Gila Co. Salt River Canyon between Globe and Show Low, 18 Aug 1969, S. Sikes 428 (Holotype, SRSC! Isotypes, SMU! TEX!). * Laphamia gilensis subsp. longilobus Niles, Mem. N.Y. Bot. Gard. 21:51-54. 1970, is listed here as a synonym which appeared in print (Niles, 1970) after this work was ac- cepted for publication. 105 i om a i oe cnt $53 ae , photograph of holotype (SRSC) Perityle gilensis var. salensis: Fig. 7. S. Sikes 428. 106 Plantae 30-70 em longae, pendulae vel decumbentes; folia glabra, (3.0)3.5— 5.9(10.0) cm longa, superiora (raro basaliaque) ad filiformi-linearia ver- gentia vel apicem versus expansa (spathulata, lanceolata vel ovata), lobata vel integra, basalia et mediocria longipetiolata laminis latis vulgo trilobis ad subcruciformibus 1-2 cm longis 1.0-1.5 em latis; petiolis 2.5—4.5(8.0) cm longis; flores sicut in var. gilensi, sed radii 8-12 ligulis 0.7-1.0 em longis; chromosomata n=ca. 17. Restricted to the Salt River Canyon, between Globe and Show Low, Arizona. Fig, 3. Spring-Fall. Specimens examined: ARIZONA: Gila Co. Salt River Canyon, Globe to Show Low, R. A. Darrow (CAS); Salt River Canyon, Darrow (CAS). Few collections of P. gilensis var. salensis (n= ca. 17) have been examined, but its distinctiveness from var. gilensis (n—34-36) is clearly indicated by chromosome numbers, geographic separation (Fig. 3), and a few morpho- logical characters. Variety salensis is most easily recognized by its robust, trailing habit and long-petioled leaves which are larger than those of var. gilensis. Plants of var. salensis with distinctive filiform-linear leaves (see description) are perhaps nothing more than ecological variants, judging from the ontogeny of several greenhouse cultures. The distal leaves of such individuals may be filiform-linear, but as the plants grow older, basal leaves develop the typical blade-like (trilobed) appearance. Greenhouse cultures also indicate that both varicties of P. gilensis maintain their typical morpho- logies when grown under uniform conditions. Perityle gilensis var. gilensis has not been adequately sampled chromo- somally. Geographical and morphological evidence would seem to indicate that var. gilensis originated from var. salensis via migration down the Salt River and through the establishment of autopolyploidy. The probability of autoploid origin should be substantiated if, as suspected, diploid populations of var. gilensis are also found within its range. The once-pinnatifed tripartite leaves (with broad segments) exhibited by P. gilensis represent the most convenient feature for delimiting the taxon from P. saxicola. Distributional evidence (Fig. 3) and leaf morphology sug- gest that P. saxicola (n=17) evolved from P. gilensis var. salensis as prob- ably did var. gilensis. The infrasectional position of P. gilensis and P. savicola is still an open question. Their superficial morphology suggests the possibility that they were derived independently from a species of sect. Perityle somewhat like P. vaseyt. Perityle gilensis and P. saxicola do not exhibit any apparent mor- phological or chromatographical affinity to other radiate species of sect. Laphamia, with the possible exception of P. staurophylla. 12. PERITYLE LEMMONI (Gray) Macbride, Contrib. Gray Herb. n.s. 56:39. ety lemmoni Gray, Proc. Amer. Acad. 16:101. 1880. TYPE: ARI- ZON 1 Co. near Camp Lowell, Tucson, 1880, Lemmon s-n. 107 (Holotype, See Laphamia lemmoni var. pedata Gray, Proc. Amer. Acad. 16:101. 1880. Leptopharynx ee (Gray) Rydb. N. Amer. Fl. 34:24. 1914. Plants 6-17 (23) cm high, densely clumped and leafy, densely pilose or pilose-hirsute so that plants often appear cinereous; leaves typically opposite, seldom alternate above, 1.0-1.9 (2.5) cm leng, 0.7-1.5 em wide, densely pilose or pilose-hirsute, ovate and irregularly laciniate-serrate or lobed, to clearly thrice divided and palmate, the 3 main segments irregularly laciniate-dis- sected (merely serrate-lobed in one population); petioles 3-5 (8) mm long; capitulescence of solitary heads or small clusters on rather short, stout peduncles (1-4 mm long); involucres campanulate; heads 7-10 mm long, 5-9 mm wide, partially obscured by leaves; receptacle convex, rounded, rarely flattened, 1.0-2.0 mm wide, ca 0.5 mm high; involucral bracts 11-14, 4.0-6.4 (8) mm long, 1.0-1.7 mm wide, oblong-lanceolate to linear-lanceolate, rather flattened, keeled or with a rather prominent midrib; ray flowers absent; disc flowers 22-35, corollas yellow, often purple tinged at maturity, 3.8-4.7 (5.0) mm long, the tube 1.0-1.9 mm long, the throat broadly tubular to narrowly funnelform, 2.0-2.5 (2.9) mm long, 0.7-1.0 mm wide, the lobes acute, 0.4-0.6 mm long; style branches 1.8-1.9 mm long; achenes 2.5-3.2 (3.6) mm long, oblong-obconical to narrowly obconical, rounded or angled on one or both surfaces, with thin callous margins, short-pubescent on the margins and both surfaces; pappus of a single, very slender, delicate bristle (rarely 0-2), (0.1) 2.0-3.0 (4.0) mm long, and a vestigial crown of hyaline squamellae, or the squamellae absent, the bristles of various lengths even in same head; chromosome number, n=17 Common in crevices of granitic cliffs and boulders in SE Arizona. Fig. 8. Spring-Fall. Representative specimens: ARIZONA: Cochise Co. Douglas, rock crevices, L. N. Goodding 2292 (UC); Outlaw Canyon, Goodding 2371 (DS, RH, UC); Chiricahua Mts., J. G. Lemmon s.n. (UC, US). Graham Co. Frye Canyon, 4% mi above dam, Graham Mt., B. J. Maguire and R. R. Maguire 11733 (NY); ravine Canyon, NE Ceahan Mt., Maguire and Maguire 12067. Greenlee Co. Sandstone rocks Clifton, Frisco River, A. Davidson 341 (DS, NY, UC); Clifton, Davidson 341a (ND); above San Francisco River bank, S Clifton, Maguire, B. L. Piet and T. Mueller 11787 (GH, NY, UC, US). Pima Co. Sabino Creek, Sabino Canyon, L. Benson 9828 (POM); Soldier Trail Hwy., Santa Catalina s., Benson 11401 (POM); Santa Catalina Mts., W of Sabino, E. B. Bartram 427 (US); Sabino Canyon, Tucson, W. P. Cottam 10128 (UT); hills above Colossal Cave, A. Eastwood, s.n. (CAS); Colossal Cave near barbecue area, F. W. Gould and R. A. Darrow 2642 (LL, NY); Santa Catalina Mts., Gould 3440 (CAS, LL, NY, UC); Sabino Canyon, D. Griffiths 2575 (NY); near Tucson, J. G. Harrison 7184 (US); Santa Catalina Mts., Lemmon s.n. (DS, UC); Camp Lowell, W. F. Parish 90 (DS); near Hitchcock Rd. to Mt. Lem- mon, 1 mi below Molino Basin, A. M. Powell and B. Calvert 1260 (SRSC); = 108 Santa Catalina Mts., Windy Vista Point, Powell and Calvert 1261 (SRSC); Santa Catalina Mts., C. G. Pringle s.n. (DS, NY, US); Santa Rita Mts., Pringle s.n. (NY); Santa Catalina Mts., Pringle s.n. (NY); Goose-head rock, Mt. Lemmon, S. Sikes 96 (SRSC). NEW MEXICO: Hidalgo Co. limestone cliffs at NE end of Big Hatchet Mts., T. kK. Todsen 2001 (SRSC). This species has been merged with P. dissecta, or interpreted as subspe- cific with the latter taxon by several taxonomists including Everly (1947) and Niles (1970). However, several factors support the recognition of P. lemmoni as a distinct species, and in fact suggest a closer relationship with P. castillonii than with P. dissecta. Fig. 8. Distribution of P. lindheimeri var. lindheimeri (open circles); P. lindheimeri var. halimifolia (closed circles); P. angustifolia (open squares); P. dissecta (closed squares); P. castillonii (closed triangles); P. stauro- phylla (open stars); P. coahuilensis (black stars); P. lemmoni (open. tri- angles) 109 Unquestionably a similarity in gross leaf morphology has led to most opinions that P. lemmoni and P. dissecta are conspecific. The leaves of both taxa are indeed dissected, but there are differences in these structures. Essentially, in P. lemmoni, the leaf lobes and segments are usually broader than in P. dissecta and are densely pilose. As compared to P. dissecta, other distinguishing features of P. lemmoni include; shorter, broader, acute bracts which are pilose, broad-tubular to subfunnelform disc corollas, slender, delicate pappus bristles, and granite (or conglomerate) habitat. In view of these differences, and the distinct distribution of P. lemmoni (Fig. , 1 would suspect similarity to P. dissecta through convergence if chromato- graphic pattern data did not indicate a relationship to P. dissecta and P. castillonit. Perityle lemmoni differs from P. castilloniti most conspicuously in leaf morphology. The two taxa generally compare, however, in dense pubescence (including broader trichomes), shape of disc corollas, slender pappus bristles, and shape of involucral bracts. A closer relationship between P. lemmoni and P. castillonii is not sup- ported by chromatographic pattern data. The patterns of P. castillonii and P. dissecta are much more similar to each other than to P. lemmoni. In spite of the bulk of morphological evidence suggesting a closer affinity of P. lemmoni and P. castillonii, further sampling of intermediate populations, if they exist, will be necessary before arriving at a confident appraisal of true relationships among this related-species group. Two ‘‘atypical’’ collections which have been referred to P. lemmoni add to the populational complexity of the taxon. The atypical populations occur near the summit of Graham Mi. in Graham Co., Ariz. (e.g., Maguire and Maguire 11733), and near Clifton, Ariz. (e.g., Davidson 341). Specimens of the Graham Mt. collection show leaves which are ovate-deltoid in outline and are irregularly serrate-lobed to cleft or divided, but tend to be less lobed and dissected than typical P. lemmoni. These plants also have larger heads than other P. lemmoni, and produce a more conspicuous crown of pappus squamellae. Considering leaf morphology especially, it is possible to construe the Graham Mt. population as approaching the expected ancestral form of the species. According to label information on herbarium sheets, the Clifton population (Davidson 341a, 341) was proposed as a new species, Laphamia ambrosae- folia, by Greene. This name is not listed in the synonymy because I can find no record of it having been published. Plants from the Clifton population resemble P. dissecta in leaf, involucral bract, and pappus morphology (stout bristles) more than any cther specimens of P. lemmoni. Perhaps most in- teresting about the Clifton population is the occurrence of ray florets with well formed ligules up to 4.0 mm in some heads of the NY and UC speci- mens. Perityle lemmoni is otherwise entirely discoid. The presence of ray florets, and total morphology, suggests the possibility that P. lemmoni (at least the Clifton population) has hybrid origin, with the generally sympatric 110 P. coronopifolia (sect. Perityle) as one parent. In experimental crosses be- tween other radiate X discoid species, ray floret dominance has been ob- served through F, and F, generations. However, the presence of ray florets could, of course, simply depict an ancestral feature of P. lemmoni, but this in itself is an interesting factor for future consideration of evolutionary rela- tionships. None of the radiate species of sect. Laphamia appear at all closely related to P. lemmoni. Unfortunately, although attempts have been made, no recent collections from Graham Mt. and Clifton have been secured. 13. ev] PERITYLE DISSECTA (Torr. in Gray) Gray, Syn. Fl. N. Amer. 1:320. 1884. Cees dissecta Torr. in Gray, Pl. Wright. 2:81. 1853. TYPE: TEXAS: esidio Co, Presidio del Norte, August, 1852, J. M. Bigelow s.n. Y! Isotypes, GH! US! Leptopharynx dissecta (Torr. in Gray) Rydb., N. Amer. Fl, 34:24. 1914. Plants 7-20 cm high, hirsute-pilose, densely leafy; leaves opposite or alter- nate, usually alternate above, 0.9-2.5 em long, 0.4-1.5 ecm wide, pilose to hirsute-pilose, irregularly dissected, once, twice, or thrice pinnately divided, less often pinnately parted or cleft, the lobes irregular and rounded; petioles (3) 4-10 mm long; capitulescence of solitary heads or small clusters on rather stout peduncles 0).2-0.8 (1.7) cm long; involucres campanulate; heads 8-10 mm long, 4-5 mm wide, typically partially obscured by leaves; recep- tacle convex (rarely somewhat flattened), 0.8-1.8 em wide, 0.5 mm high; in- volucre obscurely 2-seriate, bracts 12-15, (5) 7-13 mm long, 0.5-0.9 mm wide, flattened, obscurely ribbed, linear-lanceolate, attenuate at the apex; ray flowers absent; disc flowers 20-30, corollas yellow, often purple tinged at maturity, (4.0) 5.0-5.5 mm long, the tube (1.0) 1.2-1.5 mm long, the throat typically distinctly narrowly tubular, less often barely subfunnelform, (2.0) m long, the lobes narrow, acute, 0.7-1.0 mm long; style branches 1.5-2.2 mm long; achenes (2.8) 3.0-3.8 mm long, oblong to narrowly obconical, rounded or obscurely angled usually on one surface, rarely both, with rather conspicuous callous margins, short-pubescent on the margins and both sur- faces; pappus typically of 1 (rarely 0-4) rather stout bristle, (1.0) 2.2-3.0 mm long, and a vestigial but conspicuous crown of hyaline squamellae; in tetraploids especially, the oP a appear to be reduced bristles; chro- mosome number, n=—17, 4. Infrequent on Geis walle of canyons. Fig. 8. Spring-Fall. Representative specimens: TEXAS: Brewster Co. Santa Helena Canyon, Big Bend Nat’l Park, lower part of Mesa de Anguila Escarpment, A. M. Powell 1244 (SRSC); lower walls, Santa Helena Canyon, Sikes and Patterson 414 (SRSC); Santa Helena Canyon, Big Bend Nat’! Park, B. H. Warnock 14196 (LL, SRSC, TEX). Pre- sidio Co. mts. near Ross Mine, 5 mi W of Shafter, L. C. Hinckley 3094 (LL, SRSC); canyon near old Ross Mine § side Chinati Mts., Hinckley 3407 (GH); canyon of dry creek, Livingston Ranch, ca. 6 mi SW of Shafter, 111 Powell and B. Calvert 1255 (SRSC); Livingston Ranch, ca. 6 mi SW of Shafter, Sikes and Patterson 413 (SRSC); on Santa Cruz Mt., near Presidio, Warnock T632 (TEX, US); Ross Mine area, Chinati Mts., Livingston Ranch, Warnock 18825 (SRSC); Ross Mine, Chinati Mts., Warnock 19133 (SRSC). MEXICO: Chihuahua. 4 km NW of Picachos station, W. P. Hewitt 178 (GH); Plomosas, Hewitt 373 (GH); La Campana Experimental Ranch, near En- cinillas, I. Knobloch 623 (SMU). In addition to its finely dissected leaves, P. dissecta is characterized by narrow, attenuate bracts, tubular disc corollas, sparse pilose to pilose- hirsute pubescence, stout main pappus bristle, rather prominent pappus squamellae, and limestone habitat. On the basis of most evidence, especially morphological, P. dissecta ap- pears related to P. lemmoni and P. castillonii. This contention is substan- tiated by chromatographic pattern data, although these characters indicate a closer relationship of P. dissecta and P. castilloniit. The precise affinity of the three species is not clear. Available distributional information indicates that the taxa are well separated, but much of the area in between known populations has not been investigated thoroughly. One collection from Chi- huahua, Mexico, Knobloch 623 (Fig. 8), is somewhat intermediate morpho- logically and geographically between all three species, and thus the possi- bility exists that either the taxa actually intergrade or that separation is not comp’ete. The Knobloch collection has epappose achenes like the north- western populations of P. castillonii, leaf shape and pubescence more like P. lemmoni, and involucral bracts more like P. dissecta. Admittedly, the Knobloch specimen narrows the gap between the three species in question. However, in view of the numerous differences between known populations, it seems best at present to recognize three distinct specific entities. Perityle dissecta, P. lemmoni, and P. castilloniti seem to represent a dis- tinct phylad within sect. Laphamia, but its exact origin remains obscure. Currently there are at least 3 plausible explanations for the origin of the group. 1) Perhaps most attractive is a possibility that the group was derived from a yellow-flowered member of sect. Pappothrix, such as P. rupestris. In one tetraploid population of P. dissecta (Sikes and Patterson 414), several vestigial bristles on somewhat angular achenes were observed. This possible reduction from a ‘‘Pappothrix pappus”’ is reminiscent of the situation in P. vitreomontana (sect. Pappothrix) and P. bisetosa. Also, in habit, P. castil- lonii is very similar to P. rupestris. 2) It is possible that the P. dissecta group represents taxa derived by ‘“‘reduction”’ from species of sect. Perityle. The occurrence of rather conspicuous pappus squamellae (typical of sect. Perityle species) in some populations of P. dissecta, P. castillonii, and P. lemmoni, as well as other vegetative and floral similarities to known sect. Perityle species, lends some credence to this hypothesis. 3) A more remote possibility is that the group arose through hybridization of sect. Pappothrix and sect. Perityle species. The latter possibility is based largely on an im- pression that the vegetative and floral morphology, e.g., that of P. castillonii, 112 could have been derived from such likely parents as P. rupestris and P. parryi (sect. Perityle). The natural hybrid P. parryi X P. rupestris var. albiflora and synthetic hybrids of similar parentage (Powell, 1970; unpub- lished data) exhibit recombination features which warrant consideration of the hybridization hypothesis. Three populations of P. dissecta have been sampled chromosomally, and in two of the colonies only tetraploid (n—34) plants were detected (Powell and Sikes, 1970). Tetraploids can not be distinguished from diploids morpho- logically except that some individuals have slightly larger vegetative and floral characteristics. 14. PERITYLE CASTILLONIT I. M. Johnston, Journ. Arnold Arb. 22:122- 124, 1941. TYPE: MEXICO: Coahuila. Canyon del Indio Felipe, Sierra Hechiceros, frequent in crevices of cliffs in deep canyon. 18 Sept 1940, Johnston and Muller 1359 (Holotype, GH!). Laphamia castillonii (Johnston) Everly, Contrib. Dudley Herb. 3:378. 1947. Plants 5-22 cm high, rather sparse-pilose to hirsute-pilose, densely leafy: leaves opposite below, becoming alternate above, 1-2.8 (3.5) em long, 0.8-2.2 cm wide, rather sparse-pilose to hirsute-pilose, deltoid-ovate to subcordate, irregularly serrate, lobed, or cleft, rarely parted, the major segments some- times further lobed, the lobes acute to rounded; petioles 4-9.5 (13) mm long; capitulescence of solitary heads or small clusters on rather stout peduncles (1-15 mm long); involucres narrowly campanulate or broad funnelform, heads 6-8 mm long, 4-6 mm wide, typically partly obscured by leaves; receptacle convex, 0.9-1.5 mm wide, 0.5 mm high; involucral bracts 10-11 (13-14), 3.3-4.8 mm long, 0.5-1.83 mm wide, somewhat flattened to subkeeled, oblong-lanceo- late, obtuse to acute at the apex: ray flowers absent; disc flowers 16-30, corollas yellow, purple tinged at maturity, 2.2-3.5 mm long, the tube 0.8-1.0 mm long, the throat narrowly or broadly tubular to subfunnelform, 1.0-1.8 mm long, 0.5 (1.2) mm wide, the lobes acute, 0.4-0.6 mm long; style branches (1.2) 1.8-2.0 mm long; achenes 1.8-3.0 mm long, oblong to very narrowly obconical, rounded or obscurely angled on one or both surfaces, with rather conspicuous callous margins, short-pubescent on the margins and both sur- faces; pappus absent, of an incomplete crown of extremely obscure, vestigial squamellae, or of 1-2 (3) slender bristles, 1.5-2.5 mm long, one or two often much shorter than the cthers, or merely a nub; chromosome number, n= 17. Common in crevices of igneous rock exposures, typically canyon walls and bluffs, in If Chihuahua and adjacent Coahuila, Mexico. Fig. 8. Spring-Fall. Representative specimens: EXICO: Chihuahua. crevices of lava cliffs near ‘“Virulento.’’? 16 mi S of Trincheras, I. M. Johnston and C. H. Muller 1430 (GH); in crevices and bluffs of mts. in Bolson Gigantes, ca. 50 mi S of Ojinaga, A. M. Powell, J. Scudday, and C. Surratt 1279 (SRSC); road to Barrillos, 9.2 mi SE of Ojinaga —Carmargo road, Sikes and Patterson 411 (SRSC); road to Barrillos, 5.8 mi SE of Ojinaga—Carmargo road, Sikes and Patterson 412 (SRSC). Coahuila. 113 Canon del Indio Felipe, Sierra Hechiceros, R. M. Stewart 10 (GH); Cafion del Indio Felipe, Stewart 525 (GH) Perityle castillonii differs most conspicuously from P. lemmoni and P. dis- secta by its deltoid-ovate to subcordate leaves which are irregularly serrate- lobed or cleft, but not deeply dissected. The species is further distinguished from P. dissecta by shorter, broader, and more acute bracts, shorter disc corollas, rather slender and delicate bristles (if present), and igneous rock habitat. Although P. castillonii is known only from 2 localities, the taxon shows the kind of variability which typifies most other species of sect. Laphamia. Specimens from the type locality exhibit 1-2 (3) delicate pappus bristles and much larger leaves than the ‘‘Virulento’’ plants which have epappose achenes and smaller leaves. Johnston (1941) explained this variability as ‘“‘evidently ecological,’ as it undoubtedly is to some degree in most species of Perityle. In addition, however, in many isloated populations where variability involves two or more characters, there must be some fixation of genetic differences. 15. PERITYLE LINDHEIMERI (Gray) Shinners. KEY TO THE VARIETIES la. Pappus bristle present; wide distribution . . 15a. var. lindheimevi. 1b. Pappus bristle absent, rarely present, but usually reduced; restricted distribution .. . . . . 1bb. var. halimifolia. 15a. PERITYLE LINDHEIMERI (Ge) Shinners var LINDHEIMERI Southwest. Nat. 4:204-209. 1959. ae lindheimeri Gray, Pl. Wright. 1:101. 1852. TYPE: TEXAS: mal Co. perpendicular ae on the banks of the Guadalupe River, ne Spring, near New Braunfels, May, 1850, Lindheimer 314 and 937 (Holotype, GH! Isotypes, GH! NY! UC! US! Photograph, DS ee He Rydb., N. Amer. Fl. 34:25, 1914. TYPE: TEXAS: San tar aa 3, 1852, Bigelow s.n., referred to L. rupestris by , (Ho , NY! Photograph, DS!). Beni ae (Rydb.) Shinners, Southwest. Nat. 4:204-209. 1959. Plants (10) 18-45 (60) cm high, stems sparsely pubescent, densely leafy; leaves opposite below, alternate above, broadly ovate to ovate-lanceolate, 2-5 cm long, 1.0-3.5 cm wide, glandular-punctate, essentially glabrous, serrate- lobed especially on anterior margins, apex and base acute (sometimes sub- attenuate); petioles 0.4-1.0 cm long; capitulescence of numerous heads loosely to densely aggregated at tips of slender, often dichotomously branch- ed peduncles; involucres narrowly campanulate, heads 6-7 mm long, 4-5 mm wide; receptacle flattened, ca. 2.0 mm wide; involucral bracts ca. 12, 3-4 mm long, ca. 2.0 mm wide, keeled; number of florets 16-22; ray flowers 3-5, pistillate and fertile, the tube 1.0-1.5 mm long, ligules yellow, 2.5-3.0 mm long, 1.5-2.0 mm wide, oblong to suborbicular, obscurely 3-lobed at apex; disc corollas yellow, 3.0-3.5 mm long, the tube 0.8-1.2 mm long, the throat narrowly campanulate, 1.0-1.5 mm long, the lobes acute, 0.5 mm long; style 114 branches ca. 1.5 mm long; achenes 2.0-2.8 mm long, linear-oblong to nar- rowly obeonical, often rounded or obscurely angled on one or both surfaces, with thin callous margins, minutely pubescent on margins and surfaces; pappus of a single slender bristle (rarely 2 or none), 0.5-1.8 mm long, anda vestigial, laciniate crown of squamellae; chromosome number, n= 17. Common in crevices of cretaceous limestone rock exposures, particularly beside streams and springs, Edwards Plateau, Texas. Fig. 8. Spring-Fall. Representative specimens: NEW MEXICO: Lincoln Co. at Alto, 7000 ft, G. L. Fisher 36184 (US). TEXAS: Bandera Co. head of Sabinal Canyon, 9 mi N of Vanderpool, R. C. Barneby 13542 (CAS); along Sabinal River, 6 mi N of Vanderpool, D. S. Cor- rell and D. C. Wasshausen 28046 (LL); Vanderpool, H. B. Parks 1045 (SMU); Vanderpool Canyen, Parks 3112 (SMU). Bexar Co. Classen Ranch N of San Antonio, R. D. Burr 287 (NY, TEX); San Antonio, H. D. Slater s.n. (US). Comal Co. 30 mi N of San Antonio, E. D. Schultz 145 (US). Edwards Co. 14144 mi SE of Rocksprings, V. L. Cory 13973 (TEX); 14% mi SIX of Rock- springs, Cory 20901 (GH); Polecat Creek, Cory 38947 (TEX); Rocksprings, G. O. Fisher s.n. (CAS); Frio Water Hole, R. T. Hill 42 (US); Northern Edwards Co., E. J. Palmer 10960 (A. CAS, DS, US); NIX exposures near Hwy 55, 13 mi S of Rock Springs, A. M. Powell and S. Sikes 1325 (SRSC); Hackberry, H. R. Reed 982 (TEX, US). Kendall Co. 5 mi S of Kendalia, M. C. Johnston, T. Melchert and Powell 6494 (LL, TEX); Spring Creek, near Boerne, Palmer 9827 (A, CAS, DS); Sabina Creek, near Boerne, Palmer 13650 (A); Edge Falls near Kendalia, Powell and B. Calvert 1235 (SRSC); Edge Falls, B. C. Tharp s.n. (TEX). Kerr Co. S fork of Guadalupe River, Barneby 13543 (CAS); Turtle Creek, W. L. Bray 185 (TEX, US); Lacey’s Ranch, Bray 229 (TEX, US); cliff above Turtle Creek, Correll and Correll 12793 (LL, SMU); near Kerrville, J. A. Drushel and S. G. Drushel 9883 (GH, NY); 13 mi SW of Kerrville, C. C. Johnson s.n. (DS); 5 mi W of Hunt, N fork of Guadalupe River, R. W. Kelting 49 (TEX); Lacey’s Ranch, Palmer 9986 (A, DS); above Guadalupe River, 54% mi SW of Hunt, G. L. Webster 4310 (SMU, SRSC, TEX); N of Hunt, E. Whitehouse s.n. (TEX); 12 mi W. of Hunt, on N Guadalupe River, Whitehouse 16325 (SMU, UC, US). Kimble Co. S fork of Llano River, 20 mi SW of Junction, R. McVaugh 8282 (DS, GH, LL, SMU, SRSC, TEX); 15 mi SW of Junction, J. M. Wilson 8815 (GH). Real Co. Echo Valley, J. Baird 57 (TEX); area of Mengas and Grapevine Canyons, Prade Ranch, Correll and FE. C. Ogden 25334 (LL); area of Mengas and Grapevine Canyons, Prade Ranch, Correll and Ogden 25335 (LL); 16 mi N of Leakey, Cory 8574 (LL); Prade Springs, west Frio River, Cory 12661 (TEX); Nueces Canyon, 7 mi S of Hwy 41, Cory 51873 (DS, SMU); Barks- dale, Palmer 13523 (A): 13 mi W of Leakey, Powell 1326 (SRSC); Horse Collar Bluff, 12 mi N of Leakey, B. L. Turner 3629 (TEX, SMU). Travis Co. Austin, FE. D. Schultz 388 (GH, US); N of Austin, Tharp 4714 (GH, TEX, US); Austin, Tharp s.n. (POM); W side of 3rd inlet to Lake Travis above Cow Creek, Tharp s.n. (TEX, SMU); Hamilton Pool, W of Austin, White- 115 house s.n. (NY, TEX, UC). Uvalde Co. Montell, Palmer 12986 (A, GH); Nueces River, Palmer 33694 (A, LL, NY); Reverchon 82 (GH); 1 mi N of Conean, H. D. oes R. C. Barneby 11092 (CAS, NY); Garner State Park, Turner 3852 (T Perityle ee ae is a widely distributed species of little variability, probably because of its rather uniform habitat and continuous distribution. The species is clearly related to P. angustifolia, and is easily distinguished from the latter by its ovate leaves, numerous, small, radiate heads, and pappus of a single bristle. According to label information, the type speci- mens with different collection numbers (314 and 937) are all from Lind- heimer’s original collecticn of the taxon. Specimens labeled Lindheimer 314 are deposited at GH and NY, while specimens labeled Lindheimer 937 are deposited at GH, NY, UC, and US. 15b. PERITYLE LINDHEIMERI var. HALIMIFOLIA (Gray) Powell, Sida Laphamia halimifolia A. Gray, Pl. Wright. 1:100. 1852. TYPE: TEXAS: Verde Co. crevices of rocks, on the summit of hills, near the ae Pedro River, July, 1849, Charles Wagn 301 (Holotype, GH! ypes, +H! NY! US! Photograph, Perityle halimifolia (Gray) Shinners, aan Nat. 4:204. 1959. Very similar to P. lindheimeri var. lindheimeri in vegetative and floral features; pappus absent, or of an obscure callous crown, or in some indi- viduals of a single bristle, 0.5-1.0 (1.5) mm long. Restricted and isolated in distribution, rare on cretaceous limestone ex- posures in SW Val Verde Co., Texas. Fig. 8. Apparently flowers only in April-May and ioe June. Specimens examined: Val ane Co. 8 2/3 mi S of Loma Alta, Cory 41691 (GH, TEX); 1234 mi S of Loma Alta, Cory 44461 (TEX); 8 miS of Loma Alta, Powell and Calvert 1234 (SRSC); 8 mi S of Loma Alta, Powell and Sikes 1322 (SRSC); Fawcett Lodge 20-30 mi up Devils’ River, B. H. Warnock 11175 (SRSC); be- tween Del Rio and Loma Alta, Warnock 11214 (LL, SRSC). After further study it may prove necessary to submerge this variety with P. lindheimeri. Even though the delimiting pappus character breaks down, varietal rank was designated because the population is localized at the drier western periphery of the var. lindheimeri range, and the plants generally are smaller and somewhat depauperate in habit. Currently, the essentially epap- pose var. halimifolia serves to link P. lindheimeri with the related species P. angustifolia. 16. PERITYLE ANGUSTIFOLIA (Gray) Shinners, Southwest. Nat. 4:204. 1959. Laphamia eee Gray, Pl. Wright. 1:100-101. 1852. TYPE: between Texas and El Paso, 1851, Charles Wright 1194 (Holotype, GH! Isotypes, GH! NY! US! oe ph, DS!). Laphamia angustifolia var. laciniata Torr., Bot. Mex. Bound. 82. 1859. 116 erties peer crevices of rocks along the Rio Grande, October, Bigelou (Holotype, NY! Isotypes, GH? US! ete oa !). hapten ea (Torr.) Rydb., N. Amer. FI, 34:25. 1914 Plants 8-25 cm high, stems minutely pubescent, densely leafy; leaves op- posite below, alternate above, 1.5-3.0 cm long (0.2) 0.3-1.0 (1.5) cm wide, coriaceous, glandular-punctate, essentially glabrous, linear, oblanceolate, lanceolate, rarely ovate, typically 3—5-lobed or cleft, the lobes acute to at- tenuate, sometimes subentire or laciniate, obtuse, acute, or attenuate at the apex, attenuate at the base; subsessile to petiolate, petioles when present 0.4-1.0 cm long; capitulescence typically of several heads loosely aggregated at tips of dichotomously branched peduncles; involucres campanulate to narrowly so, heads 5.5-7.0 (9.0) mm long, 3.5-5.0 (6.0) mm wide; receptacle flattened; involucral bracts 12-14, 4.0-4.5 mm long, 1-2 mm wide, keeled; ray flowers absent; disc flowers 20-33 (50), corollas yellow, 2.2-2.8 (4.5) mm long, the tube 1.0-1.2 (1.5) mm long, the throat narrowly campanulate, 1.0 (1.5) mm long, the lobes acute, 0.5 (1.0) mm long; style branches ca. 1.0 mm long; achenes 2.0-2.5 (2.8) mm long, narrowly obconical to linear-oblong, often rounded or obscurely angled on one or both surfaces, with thin callous margins, minutely pubescent on margins and surfaces; pappus typically absent, except for a minute, vestigiate crown of squamellae, rarely with 1-2 bristles, 1.5-2.0 mm long; chromosome number, n=17. Common in crevices of limestone, exposures (probably only Cretaceous), W Texas along and near the Rio Grande, and along the eroded W edge of the Edwards Plateau. Fig. 8. Spring-Fall. Representative specimens: TEXAS: Crocket Co. 32 mi NW of Ozona, V. L. Cory 44389 (TEX); below roadside park at top of Lancaster Hill along US 290, A. M. Powell and B. Calvert 1236 (SRSC). Pecos Co. SW of Sheffield, on and below rimrock of mesa, D. S. Correll and D. C. Wasshausen 27967 (LL); on a bluff above the Ft. Stockton-Sanderson road, 34-40 mi S of Ft. Stockton, L. C. Hinckley and Leon Hinckley 43 (SMU, SRSC); limestone cliffs on Owen's Ranch near Sheffield, J. Scudday 51 (SRSC); mesa 30 mi S of Ft. Stockton, B. C. Tharp s.n. (TEX); on bluffs 15 mi SE of Ft. Stockton, B. H. Warnock 20400 (SRSC): on bluffs 1 mi S of Sheffield, Warnock 20401 (SRSC); 1 mi S of Sheffield, Warnock 20402 (SRSC); bluffs 4 mi E of Brewster-Pecos Co. line, Warnock 20406 (SRSC). Terrell Co. limestone hill 8 mi W of Sanderson, Correll and M. C. Johnston 24588 (LL); limestone bluff 3 mi E of Sanderson, Johnston, Powell and T. Melchert 6478 (LL, TEX); Sanderson, C. R. Orcutt 728 (US): Mile Long Canyon at Langtry, J. Skiles 7 (SRSC); hills 1 mi E of Sanderson, Warnock 11887 (LL, SMU); 29 mi N of Dryden, Warnock and Johnston 17252 (SRSC); Blackstone Ranch in little Horse Head Canyon, 16 mi S of Sheffield, L. Webster 335 (TEX); Sanderson, EF. D. Wooten s.n. (US). Val Verde Co. Goldschmidt s.n. (TEX); mouth of Pecos River, V. Havard 33 (GH); road- side park, E side, Pecos River Canyon, Johnston, Powell, and Melchert 6486 (LL, TEX); limestone bluffs, Pecos River Bridge, Melchert and Powell 260 117 (TEX); G. C. Nealley 261 (DS, US); N of Hwy Bridge, W bluffs of Pecos River, D. Patterson 48 (SRSC); Pecos River Bridge below roadside park, Powell 1233 (SRSC); Pecos Canyon 12 mi W of Comstock, F. Shreve 3872 (GH); 45 mi N of Del Rio, Tharp s.n. (TEX); limestone crevices, mouth of Pecos River, Turner s.n. (SRSC); along Devil’s River at Highway Bridge, Warnock 11755 (LL, SMU, SRSC). Leaf shape and size is notably variable in this taxon. Plants exposed to the sun for most of the day typically exhibit narrower shorter, coriaceous leaves, while those of shade forms are usually much larger and thinner in texture. Plants growing in the shade also may grow larger than sun-exposed individuals, sometimes even trailing, and may form larger heads and florets. The taxon P. laciniata, recognized by Torrey and Rydberg, obviously is just a shade form, although one such larger plant growing in the shade recently has been found to be polyploid (n=ca. 51). Perityle angustifolia differs from the closely related P. lindheimeri by its reduced habit, leaf shape, larger and fewer heads per cluster, discoid heads, and epappose achenes. A single plant of one collection (Patterson 48) was observed to have 1-2 pappus bristles. The exomorphic features of P. angustt- folia, and the more xerophytic habitat, strongly suggest that the taxon evolved from P. lindheimeri by adaptation to drier conditions as it became geographically isolated by erosion of the Edwards Plateau caprock. 17. PERITYLE STAUROPHYLLA (Barneby) Shinners, Southwest. Nat. 4: 205 Laphamia foun senile Barneby, Leafl. West. Bot. 8:168. 1957. TYPE: NEW MEXICO: Otero Co. limestone ee in La Luz Canyon below High Rolls, western slope of Sacramento Mts., 2 Sept 1956, Barneby 12889 (Hclotype, CAS! ee GH! K, NY! RSA! UNM! US!). Plants 15-40 cm high, as rather small, singular plants or in dense clumps to 60 cm in diameter, scabridulous to nearly glabrous, densely leafy; leaves opposite or alternate but usually alternate, 1.7-3.8 (4.0) cm long, 1.0-2.5 cm wide, scabridulous to nearly glabrous, subcoriaceous, deeply divided into conspicuous narrow, flattened (rarely subterete) lobes, the lobes, sane the 2 lateral ones, often secondarily lobed, cleft or parted, lateral lobes at antrorse angles or perpendicular to central lobe, whole leaf subcruciform to cruciform (rarely bi-tripinnate) in appearance; capitulescence of several loosely (rarely densely) aggregated heads; involucres campanulate, heads 6.0-7.5 mm long, 4.5-6.0 mm wide, often partly obscured by upper leaves; receptacle convex, 1.5-2.0 mm wide, 0.5-0.8 mm high; involucre of 2 equal series (appearing uniseriate), bracts 13-16 (22), 3.5-4.0 mm long, 0.7-1.1 (1.8) mm wide, sublanceolate, keeled; ray flowers 4-8 (rarely absent), pistillate and fertile, the tube ca. 2.4 mm long, ligules yellow, 3.6-4.8 mm long, oblong, 3-toothed at the apex, an inner lcbelet of ligule sometimes present, often 2-3 vestigial anthers apparent; disc flowers 40-50, corollas yellow, 3.8-4.5 mm long, the tube 1.5-1.6 mm long, the throat subtubular to narrowly funnelform, 118 1.5-2.0 mm long, the lobes acute, 0.6-0.8 mm long; style branches 1.5-1.8 mm long; achenes 1.8-2.3 (2.7) mm long, oblong-obconical to obconical-elliptic, typically flattened, slightly rounded on both surfaces, rarely with a third angle on one surface nearer to one primary margin, with thin callous mar- gins, short-pubescent on the margins especially and both surfaces, pappus typically of 2-3 bristles, less often 4-6, and an obscure crown of minute hyaline squamellae which may appear as vestigial bristles or nubs, the squamellae sometimes seemingly absent, the bristles 1.8-2.5 mm long, third and fourth, if present, usually shorter than the main 2; chromosome number, n- : Locally common in limestone and granitic exposures in the SW Sacra- mento Mts. near the type locality, and in the San Andres Mts. Fig. 8. Spring-Fall. Specimens examined: NEW MEXICO: Dona Ana Co. San Andres Mts., FE slope of Goat Mt., just N of San Andres Pass, R. Spellenberg and T. Todsen 2641 (SRSC). Otero Co. Dog Canyon, Sacramento Mts., E. I’. Castetter 8124 (UNM); 1.5 mi W of High Rolls, along Hwy 82 toward Cloudcroft, Sacramento Mts., Powell 1276 (SRSC); 9 mi W of Cloudcroft, S. Sikes 67 (SRSC); mouth of Dog Canyon, 9 mi S of Alamogordo, Spellenberg and Todsen 2555 (SRSC). Sierra Co. San Andres Mts., Rhodes Canyon, 9 mi W of Rhodes Canyon Range Center, Spellenberg and Todsen 2537 (SRSC); San Andres Mts., near mouth of Cot- tonwood Cave, Spellenberg and Todsen 2547 (SRSC); San Andres Mts., 11 mi W of Rhodes Canyon Range Center, Spellenberg and Todsen 2637 (SRSC). Perityle staurophylla is an endemic species which is characterized by its variable pappus of 2-6 bristles, rather small heads with 4-8 ray florets, sub- corymbiform inflorescence, and variable leaves. The phylogenetic position of this species within sect. Laphamia is not at all clear, but several features of the taxon suggest possible relationships which are of theoretical interest with regard to the biosystematics of Perityle. The single most enigmatic feature of P. staurophylla revolves around the structure of its pappus. It should be remembered that the pappus is quite variable among the species of Perityle, and one must be careful in evaluating the taxonomic significance of this character. However, an understanding of the origin of the pappus in P. staurophylla would appear to be tantamount to understanding its phy- logeny. The variable pappus bristle structure (length and number) of P. stauro- phylla could be construed as a reduced Pappothrix condition (20-30 bristles) somewhat like that found in P. vitreomontana, sect. Pappothrix (Powell, 1969). In addition, some achenes of P. stawrophylla are 3-4 angled, and thus nearly intermediate between the more typical flattened achenes of P. staurophylla and the obscurely 4-angled fruits characteristic of Pappothrix species. However, as noted by Barneby (1957), the general appearance of P. staurophylla does not indicate close affinity with any of the species of sect. Pappothrix. The lack of close relationship with sect. Pappothrix is 119 further amplified by details of vegetative and floral morphology In leaf morphology, P. staurophylla resembles P. gilensis and P. saxicola (Arizona), and to a lesser extent, P. coahuilensis (Mexico). Leaves of the four taxa are not alike, but they are variably tripartite and subcruciform. Other characteristic features of these species do not suggest any close rela- tionship, however, and it is possible that the similar leaf shapes have been evolved independently. The tendency for trilobed leaves is apparent in many species of Perityle which appear unrelated to P. staurophylla. Since the pappus structure of this species is somewhat intermediate be- tween sect. Pappothrix and sect. Laphamia, several plausible explanations regarding its origin are amenable to consideration. Perityle staurophylla could have arisen from some ancestral species of sect. Pappothrix in much the same fashion as P. bisetosa is suspected to have been derived from sect. Pappothrix (Powell, 1969; present paper). No direct evidence is available to support the latter hypothesis. On the other hand, most morphological evi- dence, other than the pappus, suggests a closer relationship of P. stauro- phylla to species of sect. Laphamia than to species of sect. Pappothrix. Also, a relationship of P. staurophylla to sect. Laphamia is supported by chroma- tographic comparisons, where its pattern most closely resembles that of some southwestern species, especially P. gracilis. Certainly P. staurophylla does not appear to have any immediate affinity with P. gracilis, but its pre- sumed relationship with sect. Laphamia would seem to be strengthened by the crude chemical data. It is possible that P. stawrophylla resulted from interspecific hybridization with a species of Pappothrix as one parent. This could account for the leaf and pappus variability in P. stawrophylla, and for the restricted distribution of the taxon. The likelihood of intersectional hybridization has been en- hanced through greenhouse experimentation where numerous artificial F, combinations between all three sections of Perityle have been obtained (Powell, unpublished). A likely multibristled parental suspect would be P. quinqueflora (sect. Pappothrix) which is known to occur in the Guadalupe Mts. of New Mexico, not far south of the P. staurophylla habitat. Theo- retically, besides the pappus, P. quinqueflora could have contributed the following features to P. staurophylla: Yellow flowers, small heads, subcorym- biform inflorescence, subcoriaceous leaves, and glabrosity. If P. staurophylla did indeed have hybrid origin, another likely parent with close geographic distribution is not easy to find. The most attractive candidates appear to be P. lindheimeri, P. angustifolia, and P. coronopifolia (sect. Perityle). Peri- tyle cf. lindheimeri has been collected at Alto, New Mexico (Fisher 36184) although I have not been able to find it at that locality. Spontaneous green- house hybrids between P. lindheimeri X P. coronopifolia have been reported by Ripley (1957), but these do not resemble P. staurophylla. For various reasons none of the possible parental combinations mentioned above meet all the logical requirements for P. staurophylla ancestry. At any rate, further conjucture along this line of thought is not warranted. 120 18. PERITYLE coahuilensis Powell, sp. nov. Fig. 9. TYPE: MEXICO: Coahuila. deep narrow canyon, ca. 2 km SE of Puertecito, d of Sierra de la Madera, ca. lat. 27 15 N 21 Sept 1941, I. M. ie 9311 (Holotype, GH! Isotype, GH!) Plantae 15—35 cm altae, glabrae: caules albidi, glanduloso-tuberculati; folia vulgo alterna, raro opposita, 1.4—2.6 cm longa, 0.7—1.4 cm lata, glabra, glanduloso-punctata, subcoriacea, vulgo cruciformia ad palmatim triloba vel trifida segmentis vulgo lobulatis vel serratis (aliunde folia deltoidea vel ovata marginibus irregulariter serrato-lobulatis); petioli 0.6—1.7 cm longi; involucra anguste campanulata vel subcylindrica, capitula 0.8—1.0 cm longa, 4—6 mm. lata; flores radii vulgo desunt, disci 24—30 corollis flavis (Demum purpurascentibus), 4.0—5.5 mm longis faucibus tubulosis 2.0—2.5 mm longis; achaenia 3.4—3.6 mm longa, anguste obconica ad linearia marginibus (valde callosis) faciebusque brevipubescentibus; pappus seta unica valida (raro duabus); chromosomata n= 27-34. Known only from SW Coahuila, Mexico, growing in crevices of limestone bluffs in protected canyons. Fig. 8. Spring-Fall. Representative specimens: MEXICO: Coahuila. Cafion del Agua Grande, ca. 2 km W of Ejudo Las Delicias, S. Sikes and D. Patterson 399 (SRSC); Cafion del Agua Grande, Sierra del Sobaco, a few km W of Las Delicias, R. M. Stewart 2803 (GH). This rare and unusual species has commanded special attention with regard to systematic studies in sect. Laphamia. In the first place, it has not been possible to realize any certain interspecific relationship for P. coahui- lensis. The taxon is best delimited by its essentially glabrous, subcoriaceous, typically subcruciform leaves, discoid heads, and pappus of a single bristle. Its chromosome number (n==27-34) presumably is tetraploid on the base of x—17, although attempts to obtain an accurate count have not been successful. Considering leaf shape and texture, P. coahuwilensis resembles P. staurophylla (n=17), but several floral differences appear to negate any close relationship with the latter taxon. Superficially, in vegetative and floral habit (including disc corollas), P. coahuilensis compares with P. vaseyi (n—17) of sect. Perityle, but its achene morphology (including absence of pappus squamellae) definitely is Laphamoid in nature. Ontogenetic comparisons of P. coahuilensis with putative relatives have revealed initially surprising but rather convincing evidence that the taxon has affinity with P. angustifolia. Characteristically, P. angustifolia has much smaller heads than does P. coahuilensis, but shade forms of the former species are found to produce heads which equal those of P. coahuilensis in size and general morphological conformation. During the development of heads on plants of both species, just before anthesis, disc floret buds pro- trude uniformly from the involucral bracts in a manner which apparently is not matched exactly by any other species of the section. Further observa- tions also show that P. coahuilensis and P. angustifolia compare almost exactly in every aspect of habit except for very conspicuous differences in ABA MEMICOE costues Cenubsita $y TM. Bichnersss meng fl aa Fig. 9. Perityle coahuilensis: photograph of holotype (GH), I. M. Johnston 9311. 122 typical leaf shape (e.g., similarities in whitish basal stems, leaf glabrosity and texture, and leaf arrangement). Furthermore, it should be noted that leaves in a few plants of P. coahuilensis depart from the typical subcruci- form shape and approach P. angustifolia in being irregularly serrate-lobed. The above observations would appear to suggest the possibility that P. coahuilensis has arisen as a tetraploid offshoot from P. angustifolia. This possible lineage is not supported, however, by some important floral features of P. coahuilensis. The florets of P. angustifolia are epappose, have narrow- ly campanulate throats, and are much smaller than are those of P. coahui- lensis which exhibit a single bristle and have long, tubular throats. Indeed, such floral differences could have resulted from polyploidy and modification through speciation, but essentially only quantitative variation is character- istic of numerous other polyploid species in Perityle. Attention has been given to the possibility that P. coahuilensis is of inter- sectional hybrid origin since it exhibits some features which are character- istic of both sect. Laphamia and sect. Perityle. Among the species of sect. Perityle, P. coahuilensis bears closest resemblance to P. vaseyi, most notably in general habit, leaf shape, pappus bristle, and disc corolla char- acters. The distribution of P. vaseyi in Mexico is known to be proximal to that of P. coahuilensis. Several sect. Laphamia X sect. Perityle hybrids have been synthesized in the greenhouse (Powell, unpublished), thus supporting the possibility of natural intersectional hybridization. Besides P. angustifolia and P. vaseyi, there is some evidence that P. castillonii might have contributed to the genome of P. coahuilensis. The latter species and P. vaseyi are the only taxa which are known to occur close to the distributional range of P. coahuilensis. Interestingly, P. coahiu- lensis exhibits disc corollas which turn purplish at maturity, a relatively rare trait in yellow-flowered species of the genus Perityle. Among the species relevant to this problem, purplish corollas are found in P. castillonii, P. dissecta (related to P. castillonii) and P. aglossa (related to P. vaseyi). A chromatographic study of P. coahuilensis and possible relatives has not con- tributed any solid evidence with regard to its origin. However, one prom- inent yellow compound (as yet unidentified) of the P. coahuilensis pattern apparently occurs elsewhere only in P. castillonii and its ally, P. dissecta. Additional factors have given reason to suspect that P. coahuilensis is of hybrid origin. Unusual leaf variation (particularly in degree of lobing and dissection) has been detected in individual plants from a population near Las Delicias, Coahuila, Mexico. Population to population variation in leaf morphology actually is common among the species of Perityle, but plant to plant differences of the sort which have been found at Las Delicias are un- expected. Meiosis, observed only in greenhouse plants of P. coahuilensis, is irregular to such an extent that an accurate chromosome count could not be made. Although it is possible that the unnatural greenhouse environment (or autopolyploidy) could have some effect on meiosis, this cytological con- dition at least occasions a suspicion of hybrid involvement. Also, one green- 123 house plant of P. coahuilensis was observed to produce a single ligule (ray floret) in one head and 2 ligules in another head. The plant was grown from a woody caudex which was taken from the population near Las Delicias. The significance of this potential to sues a ray floret in a normally dis- coid plant cannot be interpreted at this time, but perhaps it is reasonable to anticipate that P. coahuilensis has a radiate ancestor in its pedigree. Additional conjecture with regard to the origin of P. coahuilensis would seem to have little value. After all, the species grows in one of the most inaccessible regions of North America, and is known from only 8 collections. Perhaps further botanical exploration of southwestern Coahuila, Mexico, and continued experimental studies will result in a better understanding of the taxon. 19. PERITYLE ecochisensis (Niles) Powell, comb. nov. Laphamia cochisensis Niles, Mem. N.Y. Bot. Gard. 21:47-49. 1970. TYPE: NA: Cochise Co. North-facing cliffs of Echo Canyon, elev. ca. 2000 m, Chiricahua Nat’! Mon., 5 Oct 1964, Niles 377 (Holotype, ARIZ). Plants 4-12 cm high, stems almost glabrous to sparsely or densely short- pubescent; leaves alternate, 1.5-3.5 cm long, 0.8-2.7 cm wide, subglabrous to hispidulous, subdeltoid, ovate, to deltoid-orbiculate in outline, irregularly lobed or cleft or subcruciform with 3-5 segments, segments serrate or lobed; petioles 0.7-1.5 cm long; capituiescence essentially of solitary heads on very short peduncles (ca. 0.5 em); involucres broadly campanulate, heads 0.8-1.0 em long, ca. 1.0 em wide; involucre of 2-3 equal series, bracts 5.0-6.5 mm long, 1.0-2.5 mm wide, lanceolate to lanceloate-ovate or ovate, acute-obtuse at the apex; ray flowers absent; disc flowers 30-45, corollas yellow, ca. 3.5 mm long, the throat funnelform to narrowly campanulate, ca. 2.0 mm long: achenes 2.5-3.0 mm long, linear to narrowly obconical, with thin callous margins, ciliate-pubescent on the margins, short-pubescent on the achene surfaces; pappus of one slender bristle, 2.0-2.5 mm long, or with the main bristle and a shorter one arising from the opposite margin, bristles sub- plumose distally, and pappus usually of an inconspicuous, partial, or com- plete crown of hyaline squamellae; chromosome number, ? Apparently restricted to granite rocks in canyons, Chcsua National Monument. Fig. 3. Spring-Fall. Representative specimens: ARIZONA: Cochise Co. Bonita Canyon, Chiricahua Nat’! Mon., O. M. Clark 8611 (UNM); crevices of granite rocks, near scenic hilltop, Chiricahua Nat'l Mon., A. M. Powell and D. Patterson 1628 (SMU, SRSC, TEX); N slopes of Reed Mt., 1 mi E of Southwestern Research Sta., Chiricahua Mts., T. Kk. Todsen 2007 (SRSC). The taxonomic position of this endemic species is uncertain. There is some evidence that the taxon is related to P. lemmoni which also grows in sand- stone rocks in southeastern Arizona. Perityle cochisensis and P. lemmoni are alike in certain aspects of floral structure, especially in achene shape, 124 corolla shape, and slender pappus bristles, but otherwise there is no clear morphological indication of relationship. For example, the lobed or cleft and almost glabrous leaves of P. cochisensis differ considerably from those of P. lemmoni which are laciniate-dissected and densely pilose. Also, the achene margins of P. cochisensis exhibit ciliation which approaches that of sect. Perityle species, and the pappus usually is comprised of 2 slender bristles, unlike P. lemmoni which has short-pubescent achene margins and usually one pappus bristle. s with a few other species which have nebulous infrasectional relation- ships, it seems appropriate to consider the possibility that P. cochisensis originated from intersectional hybridization. Both P. lemmoni and P. corono- pifolia (sect. Perityle) occur in the same general distributional area with P. cochisensis, although apparently not sympatrically, and the floral and vegetative morphology is somewhat like that which would be expected in hybrids between P. lemmoni and P. coronopifolia. Certainly the ciliated achene margins and 2 pappus bristles of P. cochisensis, as well as the vari- able leaf forms found among individual plants, are characters which could have been inherited from P. coronopifolia. It must be noted, however, that in several artificial intersectional crosses between radiate and discoid species (Powell, unpublished), F, hybrids always have developed ligules, and P cochisensis is discoid. 20. PERITYLE GRANDIFOLIA Brandeg. Zoe 5:224:225. 1905. TYPE: MEX- ICO: Sinaloa. Cerro Colorado, vicinity of Culiacan, 5 Nov 1904, T. S. Brandegee s.n. (Holotype, UC! Isotype, GH! US! Photograph NY!). Leptopharynx grandifolia Rydb., N. Amer. FI. 34:23. 1914. Laphamia grandifolia (Brandeg.) Everly, Contrib. Dudley Herb. 3:379. 1947. Plants 12-25 (50) cm high, stems pilose; leaves opposite or alternate, 8-11 cm long, 4-6 em wide, pilose, thin in texture, broadly ovate, apex acute, base truncate, margins serrate to doubly serrate or somewhat crenate; petioles 2.5-4.5 cm long; capitulescence subcymose, typically with 3-6 heads loosely aggregated on peduncles which are 3.5-4.5 cm long; involucres cam- panulate, heads 7-9 mm long, 7-8 mm wide; involucral bracts 5.0-6.5 mm long, 0.5-1.0 mm wide, linear, somewhat keeled, and with 2 conspicuous costae; ray flowers pistillate and fertile, the tube 2.0-2.3 mm long, ligules yellow, 5-6 mm long, 1-2 mm wide, oblong; dise corollas yellow, 3-4 mm long, the tube 1.0-1.5 mm long, the throat tubular, ca. 2.0 mm long, lobes 0.6 mm long, narrow and acute; style branches 1.5-2.0 mm long; achenes 2-3 mm long, linear-oblong and slightly curved, glabrous, callous margins absent, with a conspicuous, truncate, callous carpopodium, 0.1-0.2 mm long; pappus absent, but inconspicuous callous crown present; chromosome num- ber, unknown. Apparently very rare on limestone cliffs, at ca. 5000 ft. Fig. 3. Specimen examined: MEXICO: Sinaloa. Puerto a Tamiapa, H. S. Gentry 5850 (ARIZ). 125 Perityle grandifolia is easily set apart from other species of sect. Laphamia by its large leaves which are broadly ovate, 8-11 cm long, and 4-6 cm wide. The taxon does not exhibit obvious relationships with any other member of the section. Instead, P. grandifolia shows closest vegetative resemblance to the larger leafed forms of P. cordifolia (sect. Perityle), although its achene morphology is definitely Laphamioid. On the basis of most exomorphic characters (e.g., large, pilose leaves and radiate, yellow-flowered heads), P. grandifolia fills many expectations of an ancestral taxon in sect. Laphamia. The essentially epappose, emarginate, and glabrous achenes of P. grandifolia are considered to be derived features in the section, but an ancestral status can not be ruled out by these char- acters alone. In fact, in addition to the radiate heads and a few other fea- tures, this species can be linked with P. stansburii and the “southwestern alliance”? through an interesting technical character of the achenes. The rather prominent callous carpopodium of P. grandifolia has been observed to occur elsewhere in the section only in P. stansburii, P. megalocephala, and P. inyoensis, although in the latter taxa the carpopodium is a less prom- inent but obvious nipple-like structure. 21. PERITYLE gentryi A. M. Powell, sp. nov. Fig. 10. TYPE: MEXICO: Sonora. Saguaribo, Rio Mayo, transition pine country, 5500 ft., 2 Nov 1935, H. S. Gentry 2109 (Holotype, GH! Isotypes, ARIZ! UC! US!). Plantae 5—25 cm altae, prostratae vel pendentes, piloso-villosae; folia op- posita, pilosa, 1.5—2.5 cm longa, 0.8—1.8 cm lata, ovata ad ovato-deltoidea serrata vel duplo-serrata; involucra anguste campanulata, capitula 7—9 mm longa, 5—7 mm lata; flores radii 6—10, ligulis flavis, 4—5 mm longis, ca. 2.0 mm latis, oblongo-ovatis; disci flavae, 3—4 mm longae; achaenia 2—3 mm longa, oblonga ad angustissime obconoca, glabra nisi pilis brevibus ad margines faciesque adspersis, marginibus conspicue callosis; pappus nullus: chromosomata ignota. The single Gentry collection of this species was discovered among exsic- cata which has been lumped under P. leptoglossa and P. palmeri of sect. Perityle, probably because of vaguely similar radiate heads and a Perityle- like habit. Examination of the achenes, however, indicates that it is correctly placed with sect. Laphamia. It is worthy to note that P. gentryi further manifests the habit similarity of traditional Perityle and Laphamia, and serves to support congeneric recognition of the taxa (Powell, 1968a). Perityle gentryi is nct clearly allied to any of the Laphamian species. In some respects it resembles a miniature P. grandifolia. Both taxa have yellow-flowered radiate heads, similar leaves (although those of P. gentryi are much smaller), pilose pubescence, and epappose achenes. These species differ considerably in technical aspects of the achenes, however, and this characteristic in Perityle usually is a reliable indicator of taxonomic affinity. Based upon fruit morphology, P. gentryi would appear closer to the yellow- flowered discoid taxa, P. villosa or P. lemmoni. 126 Fig. 10. Perityle gentryi: photograph of holotype (GH), H. S. Gentry 2109. 127 EXCLUDED SPECIES Laphamia peninsularis Greene, Bull. nae Acad. Sci. in. I. pp. 8-9. 1884. TYP XICO: Scammon’s agoon, on the peninsula of Lower Cali- fornia, collector unknown. (Holotype, CAS? Fragment of Type US!).= Amauria rotundifolia Benth. ee scopulorum Jones, Contrib. West. Bot. 12:48. 1908. TYPE: MEX- : AHUA: Colonia Juarez, 6000 ft., Sept. 1908, M. E. Jones s.n. ee POM; fragment of Type, US!). — Perityle coronopifolia Gray. REFERENCES ALSTON, R. E. and B. L. TURNED: 1963. Biochemical systematics. Prentice-Hall, Inc Englewood C os li Jerse oe) AXELROD, D. Beaute of the Madro-Tertiary geoflora. Bot. Rev. 24:433-509. eo ee R, 5 ae A new species of Laphamia from New Mexico. Leafl. West. Bot. 8-1 Se AMANe I. HI. 1957, ba systematics and evolution of Townsendia (Compositae). Con- trib. Gray Herb. 183:1-151. CARLQUIST, S. 1956. o. the generic limits of Eriophyllum (Compositae) and related era, Madrofio 13:226-239. CRONOUI ST, A. 1947. Revision of the North American species of Erigeron, north of Mexico, Brittonia 6:121-300 —. 1955. Phylogeny andl taxonomy of the Compositae. Amer. Midl. Nat. 53:478- DE JONG, Ty. C, . A’ systematic ries - ia genus Asfranthinm (Compositae, As- ereae). Publ. ae Mik State Univ. Biol. 2:429-528 DRESSLER. R. L. 1954. Some floristic ne Geewsan Mexico and the United States. 36: 6. oe W. L. 1964. A systematic study of the genus Bahia (Compositae). Rhodora -86; 177-215; 281-311. E or RLY. M. L. 1947. A cra carey of the genus Perifyle and related genera. Contrib. Dudley are oe ford Univ. 3:377-396. FERRIS, R. 195 Taxonomic | notes on western plants. Contrib. Dudley Herb., Stanford niy. a. GRAY, A. 1852. Plantae wrig 3(5):9 7. — atianae Texano-Neo-Mexicanae: Parte I: Smithsonian Contrib. 3(5):99-101. gar eee J. T. and P. A. MUNZ (eds.). 1965. Botanical exploration ye adi E. Jones, LY ie An autobiographical account. Leatl. West a 10:189-2 JOHNSTON. I: is 1941. New phanerogams from Mexico, IV. J. fans De 22:110-124. EWIS, H. 196 Peni e aa selection as a factor in een on. Evolution 16:257-271. et S, W. E. 197 . Taxonomic investigations in the genera tear and Laphamia (Com- positae). Mem. NY. Bot. Gard. 21:1-82. ea eae R. 1966. A ag eae survey of the goldfield genus Lasthenia. Univ. Calif ub. in Bot., ti Calif. Press. 40:1-92 Sue LL, A. M. 1968a, Additional discssion pertaining to the congeneric status of Perityle and ee (Compositae). Sid 270-278 - : 1968b. Chromosome se ies in Panis and related genera (Peritylanae-Com- ae “Amer. J. B 55:820-828. See Taxonomy A Benth. (Compositae-Peritylanae). I. Section Pappo- thrix A. “Gray Whedors 71 3 - —_ 970. Natural ee a hybridization in Perityle (Compositae). Brittonia S. Sikes. 1970. a numbers of some Chihuahuan Desert Com- oe se Southwes ce 15:175- nd H. S. ies 166, Prctininary chtomatogtaphic studies of Laphamia and ated § peers of ee ositae. Southwest. 11:190-195, RAVE N, P. H. 196 )_Aophitropial itera in the floras of North and South America. Quart. Rey Biol, eee eee 64. ae rop ne ‘ehetin and Soran endemism. Evolution 18:336-338. RIPLEY, as mee Westerners in the east. Leafl. West. Bot. 8:118, RYDBERG, P. A. 1914. Carduaceae-Helenieae-Peritylanae. N. Amer. Fl. 34:11-27. 128 SHARP, A. J. 1953. Notes on the flora of Mexico; world eae . woody dicotyle- donous families sacl the origin of the modern vegetation. J. Ecol. 380. SHINNERS, L. H. 1959, a of Laphamia transferred to Perityle eae’ Helenieac). Southwest. Nat. 4:204 Manuscript received for publication ca October 1970 (Editor’s note). NOTES ISOETES LOUISIANENSIS (ISOETACEAE), A NEW SPECIES FROM LOUISIANA.—In April 1972, one of us (GL) collected megasporangiate specimens of a quillwort, Isoetes, in Thigpen Creek near Enon, Washington Parish, Louisiana. In September we gathered microsporangiate specimens. These collections represent an undescribed species Thigpen Creek, like many streams in eastern Tousiand: is cool and clear, has a silty-sandy bottom, and flows through a pine-hardwood forest. Besides Isoetes, aquatics in the stream were Orontium aquaticum, Potamogeton pusillus, and Sparganium americanum. Isoetes was found under a highway bridge and also about 4% mile upstream. While some plants were stranded, others were in the middle of the stream and would be stranded only if the stream dried up—an unlikely event. Isoetes plants, with surrounding soil, were moved in pots to the USL greenhouse where, submersed in an aquar- ium, they have thrived for 3 months, Enon is only 18 miles from the Louisiana-Mississippi border, suggesting that the new species may occur in the latter state, where Isoetes is not yet known. Further exploration will probably result in finding additional Lou- isiana sites for the species. The only Isoetes previously collected in Lou- isiana is I. melanopoda Gay et Dur., considered rare in the state The megaspores of I. louisianensis are like those of I. engelmannii var. caroliniana in that they possess “‘high reticulate ridges, much crisped cut with an irregular margin, producing [a] somewhat spiny effect’ (N. Pfeiffer, Ann. Missouri Bot. Gard. 9:207. 1922). The microspores and - velum are also similar to these of this variety. Sharply separating the new species from I. engelmannii var. caroliniana is its brown-spotted sporangial wall. The number of peripheral strands is variable. A few leaves had no strands at all; others, one to three, weakly to strongly developed; and some, as many as 28, evenly distributed. ISOETES louisianensis Thierct, sp. nov. Statura, velum, et sporae I. engel- mannii var. caroliniana similis, praecipue differt sporangiis maculatis. Cormi 2-lobati. Folia numerosa, plerumque 15-40 cm longa, plus minusve plana adaxialiter, rotundata abaxialiter; stomata praesentia; ligulae tri- angulares usque ovatae, 2.0-3.0 mm Jongae; fila peripheralia 0-28. Sporangia oblongo-elliptica, maculis brunneis conspersa, 6.5-8.0 mm longa, 3.0-4.0 mm lata, velo 1/3-%4 obsita. Megasporae albae, irregulariter cristato-reticulatae, 500-625 mm diametro; microsporae brunneolae, dense spinulosae, 25-35 mm ongae. HOLOTYPE (GH): Louisiana: Washington Parish. In and along Thigpen Creek 1 mile E of Enon, 21 Apr 1972, Garrie Landry 200. Isotype at MICH. PARATYPE (GH, MICH): Louisiana: Washington Parish. In and along Thigpen Creek 1 mile E of Enon, 30 Sep 1972, John W. Thieret 33660. SEDAN (2 129. 1973. 130 We acknowledge the advice of Dr. Warren Wagner in the preparation of this paper.—Garrie Landry and John W. Thieret, Department of Biology, University of Southwestern Louisiana, Lafayette, Louisiana 70501. DYSSODIA TENUILOBA (COMPOSITAE): NEW TO MISSISSIPPI.—In his monograph of the genus Dyssodia (University of California Publications in Botany, Volume 48. 1969), Dr. John Strother states that Dyssodia tenuiloba (DC.) Rob. var. tenuiloba, native to southern Texas and northeastern Mex- ico, has become established in Florida, Cuba, the Bahamas, Africa, and Asia. I have recently collected the taxon near the beach on the Gulf of Mexico in southern Mississippi: Harrison Co.: U. S. 90, between Gulfport and Biloxi; occasional; rays and disk yellow-orange. David Flyr 1358, 22 April 1970 (Specimens deposited at GH, SMU, UC). It seems likely that the specics may have been introduced as recently as 1969, when the center of the extremely destructive hurricane Camille passed over this portion of the Gulf Coast, having come across the Gulf from Cuba. There was no other typical beach or dune vegetation to be seen on this stretch of the coastal highway. The chromosome number of the plant has been determined by Dr. Strother to be 2n— 24. The plant is thus a triploid on a base of x—8. This chromo- some number was not among those published in Strother’s monograph for this taxon.—David Flyr, Route 1, Stratford, Texas 79084. (See SIDA 5(1): 58—Editor) ADDITIONS TO THE LOUISIANA FLORA.—The following species may be added to the flora of Louisiana. Eleocharis elongata Chapm. St. Tammany Parish: mostly submersed, flowering culms emersed, in Bayou Lacombe, 6 miles N of Lacombe, John W. Thieret 32416 (LAF), 10 Oct 1970; same locality, Allen 2901 (LAF), 23 Sep 1972. The previous recorded United States range of this species was Georgia and Florida (Svenson, 1957). Corallorhiza odontorhiza (Willd.) Nutt. St. Helena Parish: rich woods, 5 miles NNW of Chipola, Allen 1691 (LAF, LSU), 8 Oct 1971. This orchid was to be expected in Poliemns as it is found in neighboring Mississippi and Arkansas (Correll, 1950; also in litt.). Louisiana’s orchid flora is now 37 species, of which 19 occur in St. Helena Parish. Geranium texanum (Trel.) Heller. St. Landry Parish: weedy area, 3.5 miles NW of Whiteville along road to Bunkie, Thieret 32999, 30 Apr 1971 (LAF). Rapides Parish: weedy area, Chambers, 8 miles 8 of Alexandria, Thieret 33064, 1 May 1971 (LAF). This species previously was considered endemic to Texas (Correll and Johnston, 1970). Euphorbia hypericifolia L. Lafayette Parish: weed on USL campus, La- fayette, Allen 2934 (LAF), 5 Oct 1972. The nomenclature used here is that of Correll and Johnston (1970). Wheeler (1941) applied the name FE. glomeri- SIDA 3 €2)%-130,. 1973. 131 fera (Millsp.) L. C. Wheeler to this species, which he ascribed to Florida and Texas in the United States.—Charles M. Allen, University of South- western Louisiana, Lafayette, Louisiana 70501. REFERENCES rei LL, e Pe 1950, Aili orchids of North America north of Mexico. Chronica Bo- Mas CORRELL, ma . and M, C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. SVENSON, H. K. 1957, Eleocharis. pp. 509-540 in N. Amer. Fl. 18(9). WHEELER, L. C. 1941. Euphorbia subgenus Eran in Canada and the United States exclusive of south Florida. Rhodora 43:97-154, 168-205, 223-286. ae FILIFORM CREEPING RHIZOMES IN SOLIDAGO JUNCEA (COM- POSITAE).—Fernald (Rhodora 38:212. 1936) suggested an easy way to sepa- rate Solidago juncea Ait. from S. missouriensis Nutt.: the absence of fili- form creeping rhizomes (‘‘filiform stolons’’) in S. juncea and their presence in S. missouriensis. One exception he noted was that S. juncea from the sandy southeastern section of Massachusetts did develop rhizomes similar to those of S. missouriensis. I have found other areas where S. juncea develops rhizomes and thus would like to warn against using this character to separate the taxa. Solidago juncea at its southeasternmost limit—in Tennessee, Alabama, and Georgia— consistently has rhizomes, probably helping it become and remain estab- lished there. On Sand Mt. (northeastern Alabama) in 1970, in a large popu- lation of S. juncea, less than 2% of the plants flowered. The rest persisted as rosettes and freely produced rhizomes. In southern Illinois in 1972, an- other population of rhizome-producing S. juncea was observed; most of the plants were flowering. Arthur Cronquist (personal communication) stated that many plants of northeastern S. juncea also exhibit rhizomes (both the filiform type and ones that are much stouter); these are deep-seated and usually escape collection. In southern S. juncea I have collected, the rhizomes are easily broken off, giving the impression that the specimen possesses none. Thus, a few plants of the series below appear to lack rhizomes. Nevertheless, most plants from each site have them. Voucher specimens are: SOLIDAGO JUNCEA. TENNESSEE: Bradley Co., Morton 4425. ALABAMA: Jackson Co., Morton 4415 (only basal rosette), 4473. GEORGIA: Whitfield Co., Morton 4424 (all above dup. det. A. Cronquist; GA, NY, TENN). ILLINOIS: Alexander Co., Morton 5119 (NY, SMU). When identifying S. juncea or S. missouriensis one should be careful in using Fernald’s key (Gray’s Manual of Botany, 8th ed., 1950) because rhizomes seem much more prevalent in S. juncea than he supposed. Cron- quist’s (Gleason and Cronquist, Manual of the vascular plants of northeastern United States and adjacent Canada, 1963) use of leaf shape and venation SIDA 5 (2): 131. 1973. 132 pattern better distinguishes the two: S. juncea has leaves scarcely or not at all triple-nerved and 2-7.5 em wide, while S. missouriensis has leaves more or less strongly triple-nerved, often less than 2 cm wide. One chromosome count, n=—9, was made for S. juncea, agreeing with other published counts for the species. The voucher is: ALABAMA: Jackson Co., Morton 3366 (TENN).—Gary H. Morton, New York Botanical Garden, Bronx, New York, 10458, DIRCA PALUSTRIS (THYMELAEACEAE): NEW TO LOUISIANA.— Nevling (Jour. Arnoid Arb, 43:432, 1962) did not include Louisiana in the range of Dirca palustris L. In personal correspondence (1970) he indicated he was not aware of D. palustris having previously been collected from Louisiana. In Caldwell Parish I found a large population of this species covering several acres and scattered throughout two branches of a large valley in the drainage of Winn Branch 8 miles NNW of Columbia. In some places the plants formed extensive thickets. Leatherwood is often viewed as a facultative calciphile. I have collected it on limestone cliffs in Sevier Co., Tenn. and Independence Co., Ark. In Cald- well Parish it occurs on sandy moist soil in forests of Fagus grandifolia, Liriodendron tulipifera, and various Quercus. It is most abundant on the floor of deep valleys but spreads up adjoining slopes. It flowers in late February and the fruits fall by late April. There is disagreement in literature about the fruit color. In Louisiana the mature fruits are greenish yellow, turning purple as they dry before or after falling. The stems of leatherwood are very flexible; descriptions in literature often give the impression that the wood is pliable. However, the wood itself is extremely brittle; it is the bark that is pliable. Voucher specimens (Thomas et al. 12828, 28 Feb 1969) are available for exchange and are on deposit at SMU and GH.—R. Dale Thomas, Northeast Louisiana University, Monroe, Louisiana 71201. A NEW SPECIES OF FRANKENIA (FRANKENIACEAE) FROM GYPSE- OUS SOIL OF NORTH CENTRAL MEXICO.—FRANKENIA | leverichii Turner, sp. nov. Plantae perennes suffruticosae usque ad 3 dm altae. Caules parce ramosi farinosi. Rami oppositi basi fragiles pallide brunnei alibi griseo-virides ca. 1 mm diametro. Internodia 7-10 mm longa. Folia opposita obovata-elliptica vel elliptica 4-55 mm longa 1.0-1.5 mm lata apiculata basi attenuata supra minute pubescentia subtus farinosa margine revoluto. Petioli ca. 1 mm longi alati ciliati alis trans nodo confluentibus. Flores solitarii axillares pedicellis 2-3 mm longis bracteis 4 ca. 3 mm longis 1 mm latis. Calycis tubus manifeste 5-costatus 2-3 mm longus griseo-virides sed costae basi purpurescentes. Calycis lobi 5 ca. 1 mm longi. Petala 5 unguiculata oblanceolata pallide rosea apice erosa calycem 1.5 mm superantes. Petali dimidium inferum paginae ventralis squama lineari ligulari. Stamina saepe SIDA 5 (2): 132. 1973. 133 6 (raro 12 verticillis hexandris primum visum univerticillata), Antherae 1 mm longae basifixae. Filamenta alata alis conspicuis membranaceis ca. 0.5 mm latis. Pistillum 6.5 mm altum. Styli rami 3 ca. 3 mm longi. Sparingly branched suffruticose perennial to 3 dm high; stems branching oppositely, farinose, light brown and brittle below, grayish green above, ca. 1 mm in diameter; internodes 7-10 mm long; leaves opposite, obovate-elliptic to elliptic, apiculate, attenuate at the base, 4-5 mm long, 1.0-1.5 mm wide, margins revolute, the upper surface minutely pubescent, the lower surface farinose; petiole short, winged, ca. 1 mm long, with ciliate margin united with that of the opposite leaf; flowers solitary in the leaf axils on pedicels 2-3 mm long, subtended by four leaflike bracts 2-3 mm long and 1-2 mm ide; calyx tube strongly 5-ribbed, 2-3 mm long with 5 acute lobes 1 mm long, purplish on the ribs at the base, otherwise grayish green; petals 5, clawed, pale pink, oblanceolate, erose at the apex, exceeding the calyx by 1.5 mm, with a linear ligular scale on the lower half of the ventral surface; stamens mostly 6, but occasionally 12, 6 long (5 mm) and 6 short (4 mm), in 2 whorls of 6 each but appearing as a single whorl; anthers 1 mm long, basifixed, filaments with conspicuous membranous wings ca. 0-5 mm wide; pistil 6.5 mm high; ovary unilocular with 3 basally attached anatropous ovules, style branches 3, about half the length of the gynoecium, terminated by well-defined stigmatic lines. HOLOTYPE (TEX): MEXICO: Nuevo Leon: 100 km NW of Monterrey. Perennial in gypseous soil. 3 Sept 1971. J. D. Bacon, W. R. Leverich, & B. L. Turner 1076. Isotypes to be distributed. The species is found primarily on relatively barren gypseous hills and ridges dominated by two other gypsophilic endemics, Coldenia cf. hispidis- sima (Torr.) Gray, and Haplopappus johnstonii Blake. It also occurs along alluvial washes with a number of saliniferous species belonging to Allen- rolfea, Sesuvium, Suaeda, and Atriplex. Frankenia leverichii is apparently most closely related to F. johnstonii Correll, a recently described, white-flowered, shrubby species from southern Texas. The number of known species of Frankenia from North America has doubled in the last 2 decades (from 4 to 8); four of the most recently described are gypsophilic endemics of north central Mexico and adjacent xas. The species is named for Mr. William Richard Leverich, ‘‘quasi-hippie”’ and part-time graduate student in plant systematics at the University of Texas, who shared with me the pleasure of discovering this rare endemic. Actually, when it was first found, I could not believe we had chanced upon yet another new species in the genus, so I remarked with my usual en- thusiastic abandon, ‘‘Richard, if this is new, I'll name it for you.’ The re- mark caught me off guard and I’m now honor-bound to deliver on the promise. I hope this act of commitment will encourage an otherwise too SIDA 5 (2): 133. 1973. 134 relaxed student to get on with his doctoral study of Isocoma. I am grateful to Dr. M. C. Johnston for the Latin description and to Mr. Fig. 1. Frankenia leverichii. A. Habit, X 14%; B. Branch, X 3: C. Petal, X 9; D. Pistil, X 9; E. Bud, X 3; F. Fruiting calyx, X 10. 135 Geza Knipfer for the excellent line drawings. Field support for this study was provided in part by NSF grant 29576X.—B. L. Turner, The University of Texas, Austin, Texas 78712. COMMENTS ON NEW MEXICO FLORA.—This is the first of an anticipated series of notes presenting new information on the flora of New Mexico. Specimens are deposited in the herbarium of New Mexico State University (NMC) and for Silene plankii, the University of Washington (WTU) and the New York Botanical Garden (NY), and Perityle lemmoni Sul Ross State University (SRSC). Discussion follows each entry. Silene plankit Hitchc. & Maguire. Dona Ana County: crevices at base of sheer granite cliffs in box canyon, Long Canyon, Organ Mountains, Todsen 700802 (2 August 1970), Todsen 700815 (15 August 1970), R & M Spellenberg & Todsen 2401 (25 September 1970). These are the first collections in New Mexico since Plank’s original one in 1895. Correll collected S. plankii in the Franklin Mountains, El Paso County, Texas about 1965. Heuchera sanguinea Engelm. Hidalgo County: base of rock face above Turkey Spring, Animas Mountains, Todsen & Johnson 25 September 1966. Previously reported only from southern Arizona in the US Polygala rimulicola Steyerm. Dona Ana County: appressed on NE lime- stone cliff faces in crevices, Black Mountain, southern San Andres Mountains at 5400 feet, Todsen 2258, 18 June 1972. Previously known only from the type locality in the Guadalupe Mountains of West Texas. Eryngium lemmoni Coulter & Rose. Hidalgo County: open damp meadow near head of Cloverdaie Creek, Todsen August 1972. Known only from SE Arizona. Laphamia cernua Greene. Dona Ana County: NW granite seep face at the bottom of canyon, 1 mi W of Organ Peak, Fillmore Canyon, Organ Moun- tains, Todsen 4 July 1965; NE monzonite cliff face, Findley Canyon, Organ Mountains, Todsen July 1970; base of overhanging granite cliff, 4% mi E of Fillmore Spring, Fillmore Canyon, Organ Mountains, Todsen 6 August 1970; under overhangs and on N and E cliff faces, Long Canyon, Organ Moun- tains, Todsen 15 August 1970. This narrow endemic had been collected three times previously, all from a single cliff face near Dripping Springs, Ice Canyon, Organ Mountains. These new locations assure that this species will not become extinct. Perityle lemmoni (Gray) McBride. Hidalgo County: limestone cliffs in canyon at NE end of Big Hatchet Mountains, Todsen 2001, 28 August 1971. Known only from southern Arizona. Senecio salignus DC. Hidalgo County: in sandy loam of creek bottom, Guadalupe Canyon, Johnson’s Lower Ranch, Todsen 9-11, 23 February 1969. Reported only from southern Arizona in the US.—Thomas Kk. Todsen, White Sands Missile Range, New Mexico 88002. SIDA 5 (2): 135. 1973. NOTICES XII INTERNATIONAL BOTANICAL CONGRESS: JUNE 23-30, 1975 The Closing Plenary Session of the XI International Botanical Congress held at Seattle, U.S.A., in 1969, accepted an invitation issued by the Academy of Sciences of the U.S.S.R. to convene the XII International Botanical Con- gress in the City of Leningrad in 1975. In 1971 the Organizing Committee was appointed: consisting of a chairman (A. L. Takhtajan); four vice- chairmen (A. A. Prokofiev, A. A. Theodorov, N. V. Tsitsin, A. A. Yatsenko- Khmelevsky); a secretary-general (O. V. Zalensky); a scientific secretary . S. Snigirevskaya); and a number of members at large. The XII Inter- national Botanical Congress is intended to facilitate interdisciplinary com- munication among botanists as well as an informal exchange of ideas. A number of sections are planned, including special ones to accommodate mycologists (also lichenologists), phycologists, and bryologists. The Congress will be divided between organized half-day symposia and half-day contributed paper sessions. In addition to the opening and closing plenary sessions, two evening lectures are being scheduled. All special interest groups wishing to apply for space and time during the Congress should do so by writing as soon as possible to the secretary-general, Dr. Oleg Zalensky, Komarov Botanical Institute of the Academy of Sciences of the U.S.S.R., 2, Prof. Popov Street, Leningrad 197022, U.S.S.R. The sessions of the Nomenclature Section will take place, as usual, im- mediately before the opening of the Congress— in this instance June 20-23. Four days are set aside in order to enable the Section to convene for six to eight sessions of two to four hours eac A meeting of the International cae of Botanic Gardens (President Acadmician N.V. Tsitsin) will be held in Moscow at the Main Botanical Garden of the Academy of Sciences of the U.S.S.R. on June 20. A tentative schedule of scientific field trips has been planned for the im- mediate pre-Congress and post-Congress periods. The principle purpose of these trips is to acquaint visiting botanists with as many interesting and unique features of the flora and vegetation of various regions of the U.S.S.R. as possible. Some specialized trips for phycologists, lichenologists, bryolo- gists, and palaeobotanists are also planned. The double postcards announcing the XII International Botanical Congress will be mailed during the last months of 1972. Those who wish to receive further information on the Congress should return their interest cards by March 1, 1973, so that they will be placed on the mailing list for the First Information Circular expected to be published June-July 1973.—A. Takhtajan, Chairman, Organizing Committee. ATIONAL CHECKLIST OF CULTIVATED ILEX, Part I (Con- tribution 3, U.S. National Arboretum). Available without cost from the Inter- national Checklist Committee for Cultivated Ilex, U. S. National Arboretum, Washington, D.C. 20002 or the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, D.C. 20402 SIDA Suzie’ VOLUME 5 NUMBER 3 DECEMBER 1973 CONTENTS Guide to the botanical contributions of Lloyd H. Shinners (1918-1971). Jerry M. Flook Publications New names and combinations Index to generic names Published tributes By any other name... Wm. F. Mahler Chloris truncata R. Br. (Gramineae), an Australian wool-waste adventive in South Carolina. Lloyd H. Shinners. Shinnersoseris gen. nov. (Compositae: Cichorieae). A. Spencer Tomb. Publication notes. 182 183 190 SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. Mahler, SMU Herbarium, Dallas, Texas, 75275, U.S.A. Subscriptions: Li- braries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.) per volume; numbers issued twice a year. ASSOCIATE EDITOR John W. Thieret Northern Kentucky State College Highland Heights, Kentucky 41076 © SIDA Contributions to Botany, Volume 5 Number 3, pages 137—190. Copyright 1973 by Wm. F. Mahler GUIDE TO THE BOTANICAL CONTRIBUTIONS OF LLOYD H. SHINNERS (1918-1971) ERRY M. FLOOK Southern Methodist University Herbarium, Dallas, Texas 75275 The career of Lloyd H. Shinners in taxonomic botany was, despite its rela- tive brevity, a highly productive one. In addition to his master’s thesis and his doctoral dissertation, he authored 276 articles and a 514-page flora of the north-central Texas area. These publications contain 558 new names and combinations. In an effort to make this volume of information more readily accessible, I have compiled a bibliography of Shinners’ publications, cata- logued his new names and combinations, indexed the generic names appear- ing in his writings, and cited published tributes to Shinners. It is hoped that this assemblage will in some measure facilitate literature research by students of the North American flora PUBLICATIONS The publications of Lloyd H. Shinners are arranged as nearly as possible in chronological order. Years given are those borne by publications but do not necessarily agree with actual dates of distribution of the publications. 1940a. Aristida basiramea and its relatives. Am. Midl. Nat. 23: 633-634. 1940b. Notes on Wisconsin grasses—I. Additions to the grass flora. Am. Midl. Nat. 14: 757-760. 1941a. Studies of Wisconsin Compositae. M.A. thesis, Univ. of Wisc. pp. i-ili, 1-51, maps. 1941b. Epilobium paniculatum var. subulatum in Wisconsin. Rhodora 43: 335. 1941c. Notes on ogee grasses—II. Muhlenbergia and Sporobolus. Am. Mid]. Nat. 3. 1941d. The genus pe in Wisconsin. Am. Midl. Nat. 26: 398-420. 1942. The status of Aster longulus Sheldon. Rhodora 44: 338-339. 1943a. A revision of the Liatris scariosa complex. Am. Midl. Nat. 29: 27-41. 1943b. The grasses of Wisconsin. Ph.D. dissertation, Univ. of Wisc. pp. 1-iii, 1-68, 181 maps. 1948c. Notes on Wisconsin grasses—III. Agrostis, Calamagrostis, Calamo- vilfa. Am. Midl. Nat. 29: 779-782. 1943d. The genus Aster in Nova Scotia. Rhodora 45: 344-351. 1944a. Aster acadiensis nom. nov. Rhodora 46: 31. 1944b. Notes on Wisconsin grasses—IV. Leptoloma and Panicum. Am. Midl. Nat. 32: 164-180. 1945. The genus Aster in West Virginia. Castanea 10: 61-74. 1946a. Revision of the genus Chaetopappus DC. Wrightia 1: 63-81. 1946b. Revision of the genus Leucelene Greene. Wrightia 1: 82-89. 1946c. The genus Dichaetophora A. Gray and its relationships. Wrightia SIDA 5(3): 137-179. 1973. 19468, 1947a. 1947b. 1948a. 1948b. 1948c. 1: 90-94. Revision of the genus Aphanostephus DC. Wrightia 1: 95-121, Revision of the genus Kuhnia L. Wrightia 1: 122-144 Foreword. A supplementary gazetteer of Texas, 1875-1895, by Bessie Teeple Geiser. Field & Lab. 14: 88-89 Two anomalous new species of Erigeron L. from Texas. Wrightia 1: 183-186 Revision of the genus Krigia Schreber. Wrightia 1: 187-206, The vetches and pea vines (Vicia and Lathyrus) of Texas. Field & Lab. 16: 18-29 A new species of Chrysopsis Nuttall from the driftless area of Wis- consin. Wrightia 1: 218-219 Geographic limits of some alien weeds in Texas. Tex. Geogr. Mag. 12(1): 16-25, pl. 1-2. Nomenclature of species of dandelion and goatsbeard (Taraxacum and Tragopogon) introduced into Texas. Field & Lab. 17: 13-19 Notes on Texas Compositae—I. Field & Lab. 17: 23-30. Notes on Texas Compositae—II. Field & Lab. 17: 53-64, Early plant collection returns to Texas. Field & Lab. 17: 66-68. New names of Texas Chamaesyces. Field & Lab. 17: 68- 70. Aster ontarionis the same as A. pantotrichus (A. missouriensis). Rhodora 51 Aster coerulescens the same as A. praealtus. Rhodora 51: 91-92. Physostegia Correllii (Lundell), comb. nov. Rhodora 51: 120-122. Transfer of species of Petalostemum to Dalea (Leguminosae). Field & Lab. 81- The genus a Vache Petalostemum) in North-central Texas. Field & Lab 8 Arenaria Drummondii Shinners, nom. nov. Field & Lab. 17: 89. Delphinium virescens Nutt. var. Wootoni (Rydb.) Shinners, comb. nov. Field & Lab. 17: 89. . Asclepias tuberosa L. var. interior (Woodson) Shinners, comb. nov. Field & Lab. 17: 89. The Texas species of Conyza (Compositae). Field & Lab. 17: 142-144. Nomenclature of Texas varieties of Descurainia pinnata (Cruciferae). Field & Lab. 17: 145. Transfer of Texas species of Houstonia to Hedyotis (Rubiaceae). Field & Lab. 17: 166-169. Notes on Texas Compositae—III. Field & Lab. 17: 170-176 Revision of the genus Egletes Cassini north of South. America. Lloydia 12: 239-247. Two additions to the genus Egletes Cassini from northern South America. Lloydia 12: 248-250. The Texas species of Thelesperma (Compositae). Field & Lab. 18: 17-24. 1950k. 19501. 1950m 1950n. 19500. 1950p. 1950q. 139 Notes on Texas Compositae—IV. Field & Lab. 18: 25-32. Notes on Texas Compositae—V. Field & Lab. 18: 32-42. Parietaria ae aaa Muhl. var. obtusa (Rydb.) Shinners, comb. nov. Field & Lab. 2 Corydalis ina var. australis (Chapm.) Shinners, comb. nov. Field & Lab. 18: 42 The species of Matelea (including Gonolobus) in North Central Texas (Asclepiadaceae). Field & Lab. i8: 73-78. The Texas species of Cacalia (Compositae). Field & Lab. 18: 79-83. Ranunculus carolinianus DC. var. villicaulis Shinners, var. nov. Field & Lab. 84, Addenda on Texas Thelesperma (Compositae). Field & Lab. 18: 98-99. Forestiera pubescens Nutt. var. glabrifolia Shinners, var. nov. Field & Lab. 18: 99-100. Hedyotis crassifolia Raf. var. micrantha Shinners, var. nov. Field 8: 100. The North Texas species of Plantago (Plantaginaceae). Field & Lab. 18: 113-119. . Centaurium calycosum (Buckley) Fernald var. breviflorum Shinners, 30. var. nov. Field & Lab Gerardia tenuifolia Vahl var. leucanthera (Raf.) Shinners, comb. nov. Field & Lab Brassica Kaber (DC.) Wheeler var. stricta (Celakovsky) Shinners, comb. nov. Rhodora 52: 127-128. Sidelights from zoology on botanical nomenclature. Rhodora_ 52: 156-160. Notes on Texas Compositae—VI. Field & Lab. 18: 156-159. The Texas species of Psoralea (Leguminosae). Field & Lab. 19: 14-25. Ceanothus herbaceus Raf. for C. ovatus: a correction of name. Field Lab. 19: 33-34, Two new varieties of Solidago from North Texas. Field & Lab. 19: 34-35. Pinaropappus roseus var. foliosus (Heller) Shinners, comb. nov. Field & Lab. 19: 4 Cynanchum barbigerum (Scheele) Shinners, comb. nov. Field & Lab. 19: 65 The North Texas species of Heterotheca, including Chrysopsis (Com- positae). Field & Lab. 19: 66-71. Notes on Texas Compositae—VII. Field & Lab. 19: 74-82. Rhus aromatica Ait. var. flabelliformis Shinners, var. nov. Field & ab. 19: 86- The North Texas species of Hymenocallis (Amaryllidaceae). Field & Lab. 19: 102. Allium Fraseri (M. Ownbey) Shinners, comb. nov. Field & Lab. 19: 104 140 1951k. 1951s. 1951t. 1951u. 1951v. 1951w. 1951x. 1952a. 1952c. 1952d. 1952e. 19538¢. The Texas species of Hvax (Compositae). Field & Lab. 19: 125-126. Hybanthus linearis (Torr.) Shinners, comb. nov. Field & Lab. 19: 126. . Amsonia repens Shinners, sp. nov. Field & Lab. 19: 126-127. Phlox Drummondii Hook. var. McAllisteri (Whitehouse) Shinners, comb. nov. Field & Lab. 19: | Notes on Texas Compositae—VIII. Field & Lab. 19: 133-136. The Texas species of Euthamia (Compositae). Field & Lab. 19: 137-138. Two new Texas species of Physostegia (Labiatae). Field & Lab. 19: 166-168. Yucca Freemanii, a new species from northeastern Texas. Field & Lab, 19: 168-171 Agave lata, a new species from North Texas and Oklahoma. Field & Lab. 19: 171-173 The North Texas species of Mirabilis (Nyctaginaceae). Field & Lab. 19: 173-182 Vitis mustangensis Buckley var. diversa (L. H. Bailey) Shinners, comb. nov. Fie Lab. 19: 182 pata urticifolia Michx. var. texana Shinners, var. nov. Field & : 183 Croton capitatus Michx. var. albinoides (A.M. Ferguson) Shinners, comb. nov. Field & Lab. 19 3. Croton glandulosus L. var. ner Shinners, var. nov. Field & Lab. . ¢] Addenda on Texas Chamaesyce (Euphorbiaceae), Field & Lab. 20: -26. Cakile lanceolata (Willd.) O. E. Schultz var. geniculata (B. L. Rob- inson) Shinners, comb. nov. Field & Lab. 20: : Cyperus globulosus Aubl. var. robustus (Boeck.) Shinners, comb. ov, Field & Lab. 20: 34. Eragrostis oxylepis Torr. var. Beyrichii (J. G. Smith) Shinners, comb. nov. Field & Lab. 20: 34 The Texas species of Palafoxia (Compositae). Field & Lab. 20: 92-102. Cynanchum Grayi Shinners, nom. nov. Field & Lab. 20: 110. Synopsis of . United States species of Lythrum (Lythraceae). Field & Lab. 9. Seen nats the varieties of Monarda punctata L. (Labiatac). Field & Lab. 21: 89-92. Salvia farinacea Benth. var. latifolia Shinners, var. nov. Field & Lab. 21: 92. Valerianella florifera Shinners, sp. nov. Field & Lab. 21: 93. Pyrrhopappus georgianus Shinners, sp. nov. Field & Lab. 21: 93-94. Sida ciliaris L. var. mexicana (Moricand) Shinners, comb. nov. Field 21: 94 F ilago arvensis in Michigan: a second North American record. Rho- 141 dora 55: 228. Prairie variety of Solidago giganiea. Rhodora 55: 322. The bluebonnets (Lupinus) of Texas. Field & Lab. 21: 149-153. Synopsis of the genus Brazoria (Labiatae). Field & Lab. 21: 153-154. Notes on Texas Compositae—IX. Field & Lab. 21: 155-162. Allium lavendulare J. M. Bates var. Fraseri (M. Ownbey) Shinners, comb, nov. Field & Lab. 21: 164 . Dalea laxiflora Pursh var. pumila Shinners, var. nov. Field & Lab. 21: 164 Ipomoea trichocarpa Ell. var. Torreyana (Gray) Shinners, comb. nov. Field & Lab. 21: 164-165. alea Helleri Shinners, nom. nov. Field & Lab. 21: 165. Ruelliia nudiflora (Engelm. and Gray) Urban var. hispidula Shin- ners, var. nov. Field & Lab. 21: 165. George L. Fisher (1868-1953). Field & Lab. 22: 24-26. Rubus duplaris, a new species from northeastern Texas. Field & -30. Carex caroliniana Schwein. var. cuspidata (Dewey) Shinners, comb. nov. Field & Lab. 30 Cyperus virens Michx. var. arenicola (Steudel) Shinners, comb. nov. Field & Lab. 30. Trifolium gionieraian L. in Texas. Leafl. West. Bot. 7: 132. No tes on North Texas grasses. Rhodora 56: 25-38 George L. Fisher (1867-1953). Taxon 3: 938. Asclepias asperula (Decaisne) Woodson var. decumbens (Nutt.) Shinners, comb. nov. Field & Lab. 22: Eriogonum vespinum (Polygonaceae) a new species from Texas. Fielc Lab. 22: 68-6 Southwest desert plants. (Review. Trees and Shrubs of the South- western Deserts, by Benson and Darrow.) Southwest Review 39: 280-281. Botany in the Graduate Record Examination. (Letter to the Editor.) AIBS Bull. 5(2): 12-13. Botany in the Graduate Record Examination. Field & Lab. 23: 12-14. The Texas species of Potentilla (Rosaceae). Field & Lab. 23: 19-20. Heteranthera Liebmannii (Buchenau) Shinners, comb. nov. Field Lab. 23: 21. Notes on Compositae—X. Field & Lab. 23: 34-36. Crepis pulchra (Compositae) in northeastern Texas. Field & Lab. 23: 44 Non-validity of Nuttallian names in Fraser’s Catalogue. Rhodora 7: 290-293. Tower of ivy. Mustang (Southern Methodist Univ. ) Nov. 1955: 15-17. Technical name for the Polyantha rose. Baileya Francois Crépin on botanizing. Asa Gray Bull. II. 3: 65. 76. 142 1956a. 1956e. 1956f. 1956h. 19561. 1956). 1956k. 19561. 1956m. 1956n. 19560. 1956p. 1956q. 1956r. 1956s. 1956t. 1956u. 1956v. 1956w. 1956x. 1956aa. 1956bb. Authorship and nomenclature of bur clovers (Medicago) found wild in the United States. Rhodora 58: 1-13 Illegitimacy of Persoon’s species of Koeleria (Gramineae). Rhodora 58: 93-96. eee and pseudo-science, Taxon 5: 43-46. Review. The Ontario Field Biologist. Southwest. Nat. 1: 42-43. sen Vascular Plants of Ulinois, by Jones and Fuller. Southwest. Nat. 1: 44-47 Physostegia serotina (Labiatae), a new species from coastal Louisi- ana and Texas. Field & Lab. 24: 17-19 Zanthoxylon parvum (Rutaceae), a new species from Trans-Pecos Texas. Field & Lab. 24: 19-20 Yucca louisianensis Trel. var. paniculata (McKelvey) Shinners, comb, nov. Field & Lab. 24: 37 Quercus eae ie var. microcarpa (Torr.) Shinners, comb. nov. Field & Lab. Tragia Smallii eee sp. nov. Field & Lab. 24: 37. Euthamia pulverulenta Greene (Compositae) in southwestern Louisi- ana. Field & Lab. 24: 38. Chamaesyce perennans Shinners, sp. nov. Field & Lab. 24: 38. Argythamnia mercurialina (Nutt.) Muell.-Arg. var. pilosissima (Benth.) Shinners, comb. nov. Field & Lab. 24: Alchemilla microcarpa Boissier & Reuter (esacene) in Texas. Field & Lab. 24: 38-39. Yellow-flowered Oxalis (Oxalidaceae) of eastern Texas and Louisi- ana. Field & Lab. 24: 39-40. Technical names for the cultivated Capsicum peppers. Baileya 4: Elodea correct without being conserved. Rhodora 58: 162. Cenchrus parviceps (Gramineae), a new species from southern Texas. Field & Lab. 24: 73-74 Andropogon Ischaemum L. var. songaricus Ruprecht: technical name for King Ranch bluestem. Field & Lab. 24: 101-103. The Texas species of Limoniun (Plumbaginaceae). Field & Lab. 24: o- Ds Phylogeny in five dimensions. Given at Mo. Bot. Gard. 3rd Annual ymposium on Phylogeny and Taxonomy, St. Louis, 27 Oct. 1956. 4 pp. (mimeographed). Nuttall not the author of Fraser’s Catalogue. Rhodora 58: 281-289. Medicago polymorpha var. vulgaris. Rhodora 58: 310. Illegitimacy of the name Sorghum vulgare Persoon. Baileya 4: 141-142 Forestiera autumnalis Buckley (Oleaceae) in eastern Texas and western Louisiana. Southwest. Nat. 1: Hypochoeris glabra L. (Compositae) in Apieanees and Louisiana. 1956cc. 1956dd. 1956ee. 1956ff. 1957a. 1957b. 1957c¢. 1957d. 1957e. 1957f. 19572. 1957h. 19571. 1957). 1957k. 19571. 1957m. 1957n. Southwest Nat. 1: 88. Wind a factor in floral ecology and morphology. Southwest. Nat. 1: Review. Review of the Nearctic Viburnum, by W. L. McAtee. South- west. Nat. 1: 90-91. Review. Career Opportunities in Biology, by R. B. Stephens. South- west Nat. 1: 93-95. Review. Flowers and Their Histories, by Alice Coats. Southwest. Nat. 1: 142-144, Wahlenbergia marginata oe A. DC. (Campanulaceae) in Lou- isiana. Southwest. Nat. 2: Allium acetabulum (Raf.) oe comb. nov. Field & Lab. 25: 1-32 Linum imbricatum (Raf.) Shinners, comb. nov. Field & Lab. 25: 32. Cayaponia quinqueloba (Raf.) Shinners, comb. nov. Field & Lab. : 32 Three new names for North Texas Astragalus (Leguminosae). Field & Lab. 25: 33. Against conservation of Xanthoxylem and Pherosphaera. Taxon 6: 135-137 Sisyrinchium Bermudianum L. instead of S. angustifolium Miller. Rhodora_ 59: Translation and commentary. Salt cedars (Tamarix, Tamaricaceac) of the Soviet Union, by S. G. Gorschkova. Southwest. Nat. 2: 48-73. Searcity of wild bees and wasps uacnoptes ‘a, Aculeata) in North Central Texas in 1957. Southwest. Nat. Anthophorid behavior of a honeybee, ee neice (Hymenoptera, Apoidea). Southwest, Nat. 2: 131-132 Francois Crépin on botanizing. Field & Lab. 25: 65-78. Dianthera americana L. var. subcoriacea (Fernald) Shinners, comb. ov. Field & Lab. 25: 79-80 A modest plea in favor of divers taxons. Rhodora 59: 235-237. Polygonum bicorne Raf. instead of P. longistylum Small. Rhodora 59: 265-267 Synopsis of the genus Mustoma (Gentianaceae). Southwest. Nat. 2: Spring Flora of the Dallas-Fort Worth Area, Texas. Published by author, Dallas, Texas. 514 pp. New varietal names in Argemone (Papaveraceac). Southwest. Nat. 3: 213-2 Carduus nutans L. (Compositae), a European thistle in northwestern Louisiana. Southwest. Nat. 3: 220 Carthamus lanatus L, var. baeticus (Boissier & Reuter) P. Coutinho (Compositae): another introduced thistle in Central Texas. South- west. Nat. 3: 220. 144 1958e. 1959a. 1959b. 1959e. 1959d. 1959e. 1960. 1961a. 1961b. 1961¢. 1961d. 1961le. 1961f. 1962a. 1962b. 1962¢. 1962d. 1962e. 1962f. 1962¢. 1962h. 1962. Arabidopsis thaliana (.) Heynhold (Cruciferae) introduced in north- eastern Texas. Southwest. Nat. 3: 213. Hymenopappus filifolius var. alpestris (Maguire) Shinners comb. nov. Rhodora 61: 155. Typification of the genus Forestiera (Oleaceac), Rhodora 61: 293-294 Species of Laphamia transferred to Perityle (Compositae-Heleniecae). Southwest. Nat. 4: 204-206 Liatris tenuis n. sp. (Compositae), another endemic in southeastern Texas. Southwest. Nat. 4: 207-205 Species of Berendtia Gray (Scrophulariaceaec: not Goeppert, fossil Myrsinaceac) transferred to Berendtiella, Southwest. Nat. 4: 208-209. Ranunculus trilobus (Ranunculaceae) in southern Louisiana: new to the United States. Southwest. Nat Phlox bifida var. induta (Polemoniaceae), a new endemic in North Central Texas. Southwest. Nat. 6: 50-51. Ipomoea Shumardiana (Convolvulaceae), a neglected Oklahoma- North Texas endemic. Southwest. Nat. 6: Tragia nepetaefolia var. leptophylla instead of var. ramosa (Euphor- biaceae). Southwest. Nat. 6: 101. Nomenclature of Bignoniaceae of the southern United States. Cas- tanea 26: 109-118 Rhododendron Coryi (Ericaceae), a new repent species from south- eastern Texas. Castanea 26: 156-157. Viola Rafinesquii: nomenclature and native status. Rhodora 63: 327-335. Ilegitimacy of the names Iresine celosia L., I. celosioides L., and 1. paniculata (L.) Kuntze (Amaranthaceae), Taxon 11: 141-142 Legitimacy of the name Ipomoea trichocarpa Elliott (Convolvula- ceae). Taxon 11: Hyptis radiata (Gehintacs an illegitimate name. Rhodora 64: 184-186. Synopsis of United States Bonamia, including Breweria and Stylisma (Convolvulaceae), Castanea 27: 65-77 Three new Gerardia names under Agalinis, nom. conservandum (Scrophulariaceaec). Castanea 27: 87-88 Rhododendron nudiflorum and R. roseum (Ericaceae): illegitimate names. Castanea 27: Mazus reptans (Scrophulariaceae) in Maryland. Rhodora 64: 229-231. Salpichroa origanijolia instead of S. rhomboidea (Solanaceae), Leafl. est. Bot Evolution of the Gray’s and Small’s manual ranges. Sida 1: 1-31. Annual Sisyrinchiums (Iridaceae) in the United States. Sida 1: 32-42. New names in Arenaria (Caryophyllaceae). Sida 1: 49-52. Drosera (Droseraceae) in the southeastern United States: an interim report. Sida 1: 53-59. Key to southeastern glabrous-styled Tephrosia (Leguminosae), Sida 1962x. 1962y. 19622. 1962aa. 1962bb. 1962cc. 1962dd 1962ee. 1963a 145 1: 60-62. Calamintha (Labiatae) in the southern United States. Sida 1: 69-75. Synopsis of Collinsonia (Labiatae). Sida 1: Synopsis of Corradina (Labiatae). Sida 1: 84-88. Synopsis of Dicerandra (Labiatae). Sida 1: 89-91. Vegetative aa to the woody Labiatae of the southeastern coastal plain. Sida 92-93. Micromeria ee and its allies (Labiatae). Sida 1: 94-97. Weed transport in St. Augustine grass sod in Texas. Sida 1: 98-99. Eichhornia azurea (Pontederiaceae) in the Texas Coastal Bend: new to the United States. Sida 1: 99. Aneilema (Commelinaceae) in the United States. Sida 1: 100-101. Siphonychia transferred to Paronychia (Caryophyllaceae). Sida 1: 101-108 Stellaria Corei Shinners, nom. nov. (Caryophyllaceae). Sida 1: 103-104. Ranunculus trachycarpus (Ranunculaceae) in south central Louisi- ana: new to North America. Sida 1: 104-105. Warea il instead of W. amplexifolia of Small (Cruciferae). Sida 1: ee “grandnelta instead of W. macrantha (Malvaceae): no- menclatural corrections. Sida 1: 106-107. Gentiana DeLoachii (W. P. Lemmon) Shinners, comb. nov. (Gen- tianaceae). Sida 1: 107. Agastache breviflora (Gray) Epling var. Havardii (Gray) Shinners, comb. nov. (Labiatae), Sida 1: 107 . Scutellaria laevis (Labiatae), another endemic in Trans-Pecos Texas. ida 1: 107-108 Solanum Godfreyi Shinners, nom. nov. (Solanaceae). Sida 1: 108. Southeastern records of Stachys affinis (S. Sieboldii) and S. floridana (Labiatae). Castanea 28: 44-46 Gilia and Ipomopsis (Polemoniaceae) in Texas. Sida 1: 171-179. Leptoglossis and Nierembergia (Solanaceae) in Texas. Sida 1: 180- The varieties of Teucrium canadense (Labiatae). Sida 1: 182-183. Amaranthus microphyllus Shinners, sp. nov. (Amaranthaceac). Sida Scumellonia Thieretii (Labiatae), a new species from coastal Louisi- ana. Sida 1 1-252 Micranthemum glomeratum (Chapman) Shinners, comb. nov. (Scro- phulariaceae). Sida 1 Texas Evax transferred to Iilago (Compositae). Sida 1: 252-253. Verbesina Walteri Shinners, nom. nov. (Compositae). Sida 1: 253. Review. Flora of Henry County, Illinois, an Annotated List of the Vascular Plants, by R. J. Dobbs. Sida 1: 254-256. How to study the Florida flora. Sida 1: 257-261. 146 1964h. 19641. 1964). 1964k. 19641. 1964m. 1964n. 19640. 1964p. 1964q. 1964r. 1964s. 1965a. 1965b. 1965c. 1965d. 1965e. 1965f. 1965¢. 1965h. 1966a. 1966b. 1966c. 1966d. 1966e. Tigridia purpurea (Herbert) Shinners, comb. nov. (Iridaceae). Sida 95. Machaeranthera pinnatifida (Hooker) Shinners, comb. nov. (Com- positae). Sida 1: 295. Review. Roadside Flowers of Texas, by Wills and Irwin. Sida 1: 296-298. Application of the name Kuhnia eupatorioides var. angustifolia (Com- positae). Rhodora 66: 160. Calylophus (Oenothera in part: Onagraceae) in Texas. Sida 1: 337- 345, Texas Asclepiadaceae other than Asclepias. Sida 1: 358-367. New names and records for Texas Compositac. Sida 1: 373-379. Digitaria Ischaemum (Gramineae) in Mississippi and Texas. Sida 380-381 Cayratia japonica (Vitaceae) in southeastern Louisiana: new in the United States. Sida 1: 384 Three new varietal names in Sphaeralcea (Malvaceae). Sida 1: 384- 385. New varietal names for New World Ludwigia (Onagraceae). Sida 1: 385-386. Two Youngias (‘‘Crepis japonica’: Compositae) introduced in the southeastern United States. Sida 1: 386-388. Correct nomenclature of two Mexican narcotic plants. Taxon 14: 103-105. Ipomoea lacunosa (Convolvulaceae). Leafl. West. Bot. 10: 162. Ipomoea amnicola (Convolvulaceae), a South American waif in Mis- sourl. Rhodora 67: 200 Untypification for Ipomoea nil (..) Roth. Taxon 14: 231-234. Holosteum umbellatum (Caryophyllaceae) in the United States: pop- ulation explosion and fractionated suicide. Sida 2: 119-128. Cynanchum angustifolium Persoon instead of C. palustre (Asclepi- adaceae). Sida 2: Ipomoea & multifida (Rafinesque) Shinners, comb. nov. (Convolvu- laceae). Sida 2: 265. Bees and flowers. Given at Seventh Bee Pollination Conference, Texas A&M Univ., College Station, Tex., 29 Oct 1965. Verbena pulchella Sweet var. gracilior (Troneoso) Shinners, comb. nov. Sida 2: 266. Texas Polianthes, including Manfreda (Agave subgenus Manfreda) and Runyonia (Agavaceac). Sida 2: 333-338. Astranthium integrifolium var. triflorum (Rafinesque) Shinners, comb. nov. (Compositae). Sida 2: 348. Thelesperma filifoliwm var, intermedium (Rydberg) Shinners, comb. nov. (Compositae). Sida 2: 348. Cerastium glutinosum Fries (Caryophyllaceac) in Mississippi: new to 147 North America. Sida 2: 392-393. 1966f.. Hypochoeris microcephala var. albiflora (Compositae) in southeastern Texas: new to North America. Sida 2: 393-394 1967a. Brunnichia ovata (Walter) Shinners, comb. nov. (Polygonaceac). Sida 3: 1967b. Species of Biker (Tiniaria, Polygonum in part) transferred to Reynoutria (Polygonaceae). Sida 3: 117-118 1967c. Stray notes on Texas cane (Plantaginaceae). Sida 3: 120-122. 1967d. Tessaria sericea (Nuttall) Shinners, comb. nov. (Compositae). Sida 3: 122, 1967e. Polianthes Rosei Shinners, nom. nov. (Agavaceae). Sida 3: 183. 1967f. Bupleurum lancifolium (B. subovatum) (Umbelliferae) in Texas. Sida 3: 185 1968a. Calystegia sepium var. fraterniflora (Mackenzie & Bush) Shinners, comb. nov. (Convolvulaceac). Sida 8: 282 and Rogers MeVaugh. 1968b. Proposal to add Kuhnia Linnaeus (1763) as a nomen rejiciendum versus 8823, Brickellia Eliott, nom. cons. Taxon 17: 732-734. 1969a. Local floras: past, present, and future. Taxon 18: 167-176. Petrorhagia prolifera (Dianthus prolifer, Tunica prolifera) (Caryo- phyllaceae) in Arkansas and Texas. Sida 3: 345-346. 1969¢c. cae jejuna (Johnston & Warnock) Shinners, comb. nov. (Euphorbiaceae). Sida 3: 347. 1969d. Stylisma patens var. angustifolia (Nash) Shinners, comb, nov. (Con- volvulaceae). Sida 3: 347 1969e. Cryptantha mcr var. setosa (M. E. Jones) Shinners, comb. nov. Boraginaceae). Sida 3: 347. 1969f. Heterotheca aspera (Shuttleworth) Shinners, comb, nov. (Composi- tae). Sida 3: 34 19692. Croptilon divaricatum var. oe (E. B. Smith) Shinners, comb. nov. (Compositae) ida 1969h. Heleastrum Chapmanii (T. & G.) Shinners, comb. nov. (Compositae). Sida 3: 348. 197la. Bigelowia nudata var. australis (L. C. Anderson) Shinners, comb. nov. (Compositae). Sida 4: 4 1971b. Kuhnia L. transferred to Brickellia Ell. (Compositac) Sida 4: 274. 1971ce. Teucriuwm cubense Jacq. var. laevigatum (Vahl) Shinners, comb. ov. (Labiatae). Sida 4: 275. 1973. Chloris truncata R. Br. (Gramineae), an Australian wool-waste ad- ventive in South Carolina. Sida 5: 182 NEW NAMES AND COMBINATIONS The new names and combinations authored by Lloyd H. Shinners are ar- ranged alphabetically by genus, species, etc. Whenever possible, dates given are those of actual distribution of the publications. In cases in which issue 148 dates are not available the dates designated are those recorded in the Torrey Index. Type specimens are cited for the new taxa. Agalinis maritima var. grandifiora (Benth.) Shinners, comb. nov. Castanea 27: 88. 12. 962. Agalinis nuttalliit Shinners, nom. nov. Castanea 27: 88. 12 Jul 1962. Agalinis pseudaphylla (Penn.) Shinners, comb. nov. Castanea 27: 88. 12 Jul 1962. Agastache breviflora (Gray) Fpl. Sida 1: 107. 23 Nov 1962. Agave lata Shinners, sp. nov. Type: H. V. 171-173. 15 Oct 1951. Agrostis elliottiana Schult. f. molesia Shinners, f 14372 (SMU). Rhodora 56: 28. 17 Mar 195 Allium acetabulum (Raf.) Shinners, comb 15 Mar 1957 Allium acetabulum (Raf.) Shinners var. nov. Field & Lab. 25: 32. 15 Mar 1957. Allium fraseri (M. Ownbey) Shinners, 1 Jun 1951 Allium lavendulare Bates var ee ie sab. a1: on _ Feb 1 var. havardii (Gray) Shinners, comb. nov. Daly 61 (SMU). Field & Lab. 19: . nov. Type: L. H. Shinners i nov. Field & Lab. 25: 31-32. lavendulare (Bates) Shinners, comb. comb. nov. Field & Lab. 19: 104. . oe (M. Ownbey) Shinners, comb. nov. ? ryllus aa sp. nov. Type: U. T. Waterfall 6620 (SMU). ‘Sida ie “948. 249. “25 Jun 1964. Ambrosia caudata (Rydb.) Shinners, comb. nov. Field & Lab. 17: 174. 24 Oct 1949. Ambrosia grayi (A. Nels.) Shinners, comb. nov. Field & Lab. 17: 174. 24 Oct 1949. Ambrosia simulans Shinners, nom. nov. Field & Lab. 17: 173-174. 24 Oct 1949. Amsonia repens Shinners, sp. nov. Type: V. L. Cory 55089 (SMU). Field & Lab. 19: 126-127. 24 Jul 1951. Andropogon scoparius Mic var. virilis Shinners, var. nov. Type: L. H. Shinners 16009 (SMU: ee US). Rhodora 56: 36-37. 17 Mar 1954. Aphanostephus jaliscensis Shinners, sp. nov. Type: C. G. Pringle 9440 (GH isotype US). Wrightia 1: 115-116. 30 Apr 1946. Aphanostephus pachyrrhizus Shinners, sp. nov. Type: C. A. Purpus 5627 C; isotypes GH, MO, US). Wrightia 1: 117-118. 30 Apr 1946. Aphanostephus potosinus Shinners, sp. nov. Type: C. A. Purpus 5122 (UC; isotypes GH, MO, US). Wrightia 1: 108- no 30 Apr 1946. Aphanostephus skirrhobasis (DC.) Trel. f. incisifolius Shinners, f. nov. Type H. Shinners 8010 (SMU). Wrightia 1: 103-106. 30 Apr 1946. Aphanostephus skirrhobasis (DC.) Trel f. quasigigantiusculus Shinners, f. nov. Type L. H. Shinners 7104 (SMU). Wrightia 1: 103-106. 30 Apr 1946. ili skirrhobasis (DC.) Trel. var. thalassius Shinners, va ype: L. H. Shinners 7718 (SMU). Wrightia 1: 106-107. 30 Apr 1946. ~ nov. Arenaria anomala (Waldstein & Kitaibel) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962 Arenaria argaea (Boissier & Balansa) Shinners, comb. nov. Sida 1: 50. 23 Nov 1982. Arenaria drummondii Shinners, nom. nov. Field & Lab. 17: 89. 13 Jul 1949. Arenaria fassettii Shinners, nom. nov. Sida 1: 51. 23 Nov 1962. Arenaria fontinalis (Short & Peter) Shinners, comb. nov. Sida 1: 51. 23 Nov 1962 Arenaria godfreyi Shinners, nom. nov. Sida 1: 51. 23 Nov 1962. Arenaria iranica Shinners, nom. nov. Sida 1: 50. 23 Nov 1962. Arenaria jamesiana (Torrey) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962. Arenaria kotschyi (Boissier) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962. Arenaria lanuginosa (Michx.) Rohrbach var. cinerascens (Robins.) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962. Arenaria ludens Shinners, sp. nov. Type: B. H. Warnock 7419 (SMU). Sida 1: 51. 23 Nov 1962. Arenaria stephaniana (Willd.) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962. Arenaria stephaniana (Willd.) Shinners var. americana (Porter) Shinners, comb. nov. Sida 1: 50. 23 Nov 1962. Arenaria trigyna (Vill.) Shinners, comb. nov. Sida 1: 51. 23 Nov 1962. Argemone albiflora Hornem. var. texana (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 214. 1 June 1959. Argemone corymbosa Greene var. arenicola (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 214. 1 Jun 1959 Argemone grandiflora Sweet var. armata (G. Ownbey) Shinners, comb. nov. outhwest. Nat. 214. 1 Jun 1959. Argemone munita Durand & Hilgard var. argentea (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 218. 1 Jun 1959 Argemone munita Durand & Hilgard var. A (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 213. 1 Jun Argemone munita Durand & Hilgard var. een a (Rydb.) Shinners, comb. west. Nat. 3: 213. 1 Jun 1959. Argemone ochroleuca Sweet var. stenopetala (Prain) Shinners, comb. nov. Southwest. Nat. 3: 213. 1 Jun 1959. Argemone pleiacantha Greene var. ambigua (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 214. 1 Jun 1959 Argemone pleiacantha Greene var. pinnatisecta (G. Ownbey) Shinners, comb. nov. Southwest. Nat. 3: 214. 1 Jun 1959. Argemone squarrosa Greene var. glabrata (G. Ownbey) Shinners, comb. nov. Nat. 3: 213. 1 Jun 1959. Argythamnia humilis (Engelm. & Gray) Muell.-Arg. var. laevis (Gray) Shin- ners, comb. nov. Field & Lab. 24: 38. 15 Jun 1956. Argythamnia mercurialina (Nutt.) Mucell.-Arg. var. pilosissima (Benth.) Shinners, comb. nov. Field & Lab. 24: 38. 15 Jun 1956. Aristida basiramea Engelm. var. curlissii (Gray) Shinners, comb. nov. Am. Midl. Nat. 23: 633. 26 Jun 1949. Aristida dichotoma Michx. f. major Shinners, f. nov. Type: T. H. Kearney, s. n., 27 Sep 1896 (GH). Am. Midl. Nat. 23: 634. 26 Jun 1940. Aristida necopina Shinners, sp. nov. Type: V. H. Chase 5302 (SMU). Rhodora 06: 30 Artemisia ludoviciana Nutt. var. albula (Woot.) Shinners, comb. nov. Sida 1: 374. 30 Oct 1964 Artemisia ludoviciana Nutt. var. redolens (Gray) Shinners, comb. nov. Sida 374, 30 Oct 1964 Asclepias asperula (Dene.) Woods. var. decumbens (Nutt.) Shinners, comb. nov. Field & Lab. 22: 58. Apr 1954. Asclepias tuberosa I., var. interior (Woods.) Shinners, comb. nov. Field & Lab. 17: 89. 13 Jul 1949. Asclepias tuberosa L. var. rolfsii (Britt.) Shinners, comb. nov. Field & Lab. 17: 89. 13 Jul 1949 Aster acadiensis Shinners, nom. nov. Rhodora 46: 31. 12 Feb 1944. Aster cordifolius L. var. moratus (Shinners) Shinners, comb. nov. Castanea 10: 65-66. Oct 1945. Aster eulae Shinners, sp. nov. Type: L. H. Shinners 12027 (SMU). Field & Lab. 18: 35-38. 2 Jan 1950 Aster finkii Rybd. var. moratus Shinners, var. nov. Type: L. H. Shinners 3367 (WIS). Am. Midl. Nat. 26: 407-408. 30 Sep 1941. Aster furcatus f. elaciniatus (Benke) Shinners, stat. nov. Am. Midl. Nat. 26 405. 30 Sep 1941. Aster laevis L. f. latifolius (Porter) Shinners. stat. nov. Am. Midl. Nat. 26: 409. 30 Sep 1941 Aster laevis L. f. purpurascens (Farw.) Shinners, comb. nov. Am. Midl. Nat. 26: 409. 30 Sep 1941 Aster lateriflorus (L.) Britt. var. flagellaris Shinners. var. nov. Type: L. H. Shinners 9589 (SMU). Field & Lab. 21: 157-158. 15 Feb 1954. Aster lateriflorus (L.) Britt. var. indutus Shinners. var. nov. Type: E. White- house 17641 (SMU). Field & Lab. 21: 158. 15 Feb 1954 Aster macrophyllus f. eglandulosus Shinners, f. nov. Type: Monroe 11 (MIL). Am. Midl. Nat. 26: 405. 30 Sep 1941. Aster oblongifolius Nutt. var. angustatus Shinners, var. nov. Type: Fassett, Catenhusen, & Shinners 20764 (WIS). Am. Midl. Nat. 26: 418. 30 Sep 1941. Aster oblongifolius Nutt. var. orientis Shinners, var. nov. Type: W. V. U. Bot. Exped., 19 Aug 1929 (WVA). Castanea 10: 72. Oct 945. Aster oblongifolius Nutt. var. orientis Shinners f. roseus Shinners, f. nov. : Mr. & Mrs. H. A. Davis & E. L. Core, s. n. 24 Sep 1941 (WVA). Castanea 10: 72, Oct 1945. Aster oblongifolius Nutt f. pee (Benke) Shinners, comb. nov. Am. Midl. Nat. 26: 417-418. 30 S 941. Aster puniceus L. f. albiflorus ee Shinners, stat. nov. Am. Midl. Nat. 26: 414-415. 30 Sep 1941. — 151 Aster puniceus L. var. calvus Shinners, var. nov. Type: H. A. Allard 9035 WVA). Castanea 10: 71. Oct 1945. Aster puniceus L, - colbyi (Benke) Shinners, stat. nov. Am. Midl. Nat. 26: 414-415. 30 Sep 194 Aster puniceus L. 2 alabuetis Shinners, f. nov. Type: H. Cutler 315 (WIS). Am. Midl, Nat. 26: 414-415. 30 Sep 1941. Aster rolandit Shinners, sp. nov. Type: A. R. Prince & C. E. Atwood 1456 (WIS). Rhodora 45: 346-347. 11 Sep 1943. Aster sagittifolius Wedem. ex Willd. var. drummondii (Lindl.) Shinners, stat. nov. Am. Midl. Nat. 26: 406-407. Sep 1941. Aster sagittifolius Wedem. ex Willd. f. hirtellus (Lindl.) Shinners, stat. nov. Am. Midl. Nat. 26: 406. 30 Sep 1941 Aster scabricaulis Shinners, sp. nov. Type: L. H. Shinner 9504 (SMU). Field & Lab. 21: 156-157. 15 Feb 1954. Aster shortit Lindl. f. asper Shinners, f. nov. Type: Fassett & Shinners 20708 S). Am. Midl. Nat. 26: 408. 30 Sep 1941, Aster steeleorum Shinners, sp. nov. Type: E. S. & Mrs. Steele 324 (GH). Castanea 10: 67-68. Oct 1945, Aster subulatus Michx. var. australis (Gray) Shinners, comb. nov. Field & Lab. 21: 158-159. 15 Feb 1954 Aster subulatus Michx. var. cubensis (DC.) Shinners, comb. nov. Field & Lab. 21: 161-162. 15 Feb 1954 Aster subulatus Michx. var. ligulatus Shinners, var. nov. Type: L. H. Shin- ners 12057 (SMU). Field & Lab. 21: 159-160. 15 Feb 1954. Aster tenuipes (Wieg.) Shinners, stat. nov. Rhodora 45: 345. 11 Sep 1943. Aster texanus Burgess var. parviceps Pea var. nov. Type: E. White- house 20503 (SMU). Field & Lab. 21: 156. 15 Feb 1954. Aster virginiensis Shinners, sp. nov. oh pe: Ie. J. Grimes 4450 (GH). Castenea 10: 66-67. Oct 1945 Astragalus aeeiens (Small) Schulz var. pleianthus Shinners, var. nov. Type: L. H. Shinners 22900 (SMU). Field & Lab. 25: 33. 15 Mar 1957 Astragalus lambertii (Pursh) Spreng. var. abbreviatus (Greene) Shinners, comb. nov. Field & Lab. 25: 33. 15 Mar 1957. Astragalus lindheimeri Engelm. var. bellus Shinners, var. nov. Type: L. H. Shinners 18568 (SMU). Field & Lab. 25: 33. 15 Mar 1957 Astranthium integrifolium (Michx.) Nutt. var. robustum Shinners, var. nov. ia ie B. H. oe 5126 (SMU). Field & Lab. 18: 158. 11 Dec 1950. Astranthiun ifolium var. triflorum (Raf.) Shinners, comb. nov. Sida 2: 348. 20 ait 1966. Berendtiella coulteri (Gray) Shinners, comb. nov. Southwest. Nat. 4: 208. 1 Dee 1959 Berendtiella levigata (Robins. & Greenm.) Shinners, comb. nov. Southwest. Nat. 4: 209. 1 Dee 1959 Berendtiella rugosa (Benth.) Shinners, comb. nov. Southwest. Nat. 4: 209. ec 1959 _ ou bho Berendticlla spinulosa (Wats.) Shinners, comb. nov. Southwest. Nat. 4: 209. Dee 1959 Bigelowia nudata var. australis (L. C. Anderson) Shinners, comb. nov. Sida 22 Jul 1971 Bonamia patens (Desr.) Shinners, comb. nov. Castanea 27: 74-75, 12 Jul 1962. Bonamia patens (Desr.) Shinners var. augustifolia (Nash) Shinners, comb. nov. Castanea 27: 75. 12 Jul 1962. Brassica kaber (DC.) Wheeler var. stricta (Celak.) Shinners, comb. nov, Rhodora 52: 127-128. 10 May 1950. Brickellia adenolepis (Robins.) Shinners, comb. nov. Sida 4: 274, 22 Jul 1971. Brickellia chlorolepis (Woot. & Standl.) Shinners, comb. nov. Sida 4: 274. 22 Jul 1971 Brickellia eupatorioides (L.) Shinners, comb. nov. Sida 4: 274. 22 Jul 1971. Brickellia leptophylla (Scheele) _ var. mexicana (Shinners) Shinners, comb. nov. Sida 4: 274. 2 Jul 1 Brickellia aren ae (L.) oie var. ozarkana (Shinners) Shinners, comb. nov. Sida 4: . 22 Jul 1971 Brickellia ewpatorioides (l..) Shinners var. texana (Shinners) Shinners, comb, nov. Sida 4: 274. 22 Jul 1971. Brickellia leptophylia (Scheele) Shinners, comb. nov. Sida 4: 274. 22 Jul 1971. Brickellia leptophylla (Scheele) Shinners var. mexicana (Shinners) Shinners, comb, nov. Sida 4: 274. 22 Jul 1971. Brickellia mosieri (Small) Shinners, comb. nov. Sida 4: 274. 22 Jul 1971. Brickellia oreithales (Robins.) Shinners, comb. nov. Sida 4: 274, 22 Jul 1971. Brickellia schaffneri (Gray) Shinners, comb. nov. Sida 4: 22 Jul 1971. Brunnichia ovata (Walt.) Shinners, comb. nov. Sida 3: 115. 31 May 1967. Cacalia elliottti (Harper) Shinners, comb. nov. Field & Lab. 18: 80. 3 Apr 1950 bo ~J ee Cacalia lanceolata Nutt. var. virescens (Harper) Shinners, comb. nov. Field & Lab. 18: 80. 3 Apr 1950 Cacalia maxima (Harper) Shinners, comb. nov. Field & Lab. 18: 80. 3 Apr 1950 Cacalia plantaginea (Raf.) Shinners, comb. nov. Field & Lab. 18: 81-82. 3 Apr 1950 Cakile lanceolata (Willd.) Schulz var. geniculata (Robins.) Shinners, comb. nov. Field & Lab. 20: 33-34. 29 Feb 1952 Calamintha arkansana (Nutt.) Shinners, comb. nov. Sida 1: 72. 23 Nov 1962. Calamintha ashei (Weath.) Shinners, comb. nov. Sida 1: 73-74. 23 Nov 1962. Calamintha georgiana (Harper) Shinners, comb. nov. Sida 1: 74. 23 Nov 1962. Calylophus hartwegii (Benth.) Raven var. filifolius Gfastw.) Shinners, comb. nov. Sida 1: 345. 30 Oct 1964. Calylophus hartwegii (Benth.) Raven var. lavandulaefolius (T. & G.) Shin- ners, comb. nov. Sida 1: 345. 30 Oct 1964. Calylophus hartwegii (Benth.) Raven var. maccartii Shinners, var. nov. Type: R. E. Benavides 91 (SMU). Sida 1: 348-344. 30 Oct 1964. Calylophus hartwegii (Benth.) Raven var. pubescens (Gray) Shinners, comb. nov. Sida 1: 344-345. 30 Oct 1964. Calylophus hartwegii (Benth.) Raven var. towmeyi (Small) Shinners, comb. nov. Sida 1: 341. 30 Oct 1964. Calylophus serrulatus (Nutt.) Raven var. arizonicus Shinners, var. nov. Type: S. J. Preece, Jr. & B. L. Turner 2692 (SMU). Sida 1: 338. 30 Oct 964. Calylophus serrulatus (Nutt.) Raven var. spinulosus (Nutt., ined; ex T. & G ners, comb. nov. Sida 1: 339-341. 30 Oct 1964 Calystegia sepium var. fraterniflora (Mack. & Bush) shiners: comb. nov. Sida 3: 282. 21 Jun Capsicum annuum L. var. minus (Fing.) Shinners, comb. nov. Baileya 4: 82. 29 Jun Carex ouronniana Schwein. var. cuspidata (Dew.) Shinners, comb. nov. Field & Lab. 22: 30. Ja Castilleja purpurea (Nutt.) G. Don var. cifrina (Penn.) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 410. 3 Apr 1958. Castilleja purpurea (Nutt.) G. Don var. lindheimeri (Gray) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 410. 3 Apr 1958. Cayaponia quinqueloba (Raf.) Shinners, comb. nov. Field & Lab. 25: 32. 15 Mar 1957. Ceanothus herbaceus Raf. var. pubescens (T. & G.) Shinners Field & Lab. 19: 33-34. 18 Jan 1951 Cenchrus parviceps Shinners, sp. nov. Type: L. H. Shinners 16990 (SMU). Field & Lab. 24: 73-74. 25 Jul 1956. Centaurium calycosum (Buckl.) Fern. var. breviflorum Shinners, var. nov. Type: C. L. & A. A. Lundell 10022 (SMU). Field & Lab. 18: 130. 9 May 1950. Chaetopappa asteroides (Nutt.) DC. var. grandis Shinners, var. nov C.L. & A. A. Lundell 9982 (SMU). Wrightia 1: 76. 30 Apr 1946. Chaetopappa pcan (Gray & Engelm.) Shinners, comb. nov. Wrightia 1: 71-73. 30 Apr 19 , comb. nov. . Type: Chaetopappa néllivides (Gray) Shinners, comb. nov. Wrightia 1: 77-79. 30 Apr 1946. Chaetopappa bellioides (Gray) Shinners var. hirticaulis Shinners Type: H. H. Bartlett 10359 (SMU). Wrightia 1: 79. 30 Apr 1946. Chaetopappa effusa (Gray) Shinners, comb. nov. Wrightia 1: 68-69. 30 Apr 1946. , Var. nov. Chaetopappa keerlioides Shinners, sp. nov. Type: F. L. Wynd & C. H. Mueller 369 (MO; isotypes GH, UC, NY, US). Wrightia 1: 69. 30 Apr 1946. Chaetopappa pulchella Shinners, sp. nov. Type: I. M. Johnston 7151 (GH). Wrightia 1: 79-80. 30 Apr 1946. Chamaesyce carunculaia (Waterfall) Shinners, comb. nov. Field & Lab. 20: 24. 29 Feb 1952 154 Chamaesyce fendleri (T. & G.) Small var. chaetocalyx (Boiss.) Shinners, comb. nov. Field & Lab. 17: 70. 30 Mar 1949. Chamaesyce fendleri (T. & G.) Small var. triligulata (Wheeler) Shinners, comb. nov. Field & Lab. 17: 70. 30 Mar 1949 Chamaesyce golondrina (Wheeler) Shinners, comb. nov. Field & Lab. 17: 70. 30 Mar 1949 Chamaesyce jejuna (Johnston & Warnock) Shinners, comb. nov. Sida 3: 347. 6 Aug 1969. Chamaesyce missurica (Raf.) Shinners, comb. nov. Field & Lab. 17: 69-70. 30 Mar 1949. Chamaesyce missurica (Raf.) Shinners var. calcicola Shinners, var. nov. Type: L. H. Shinners 9367 (SMU). Field & Lab. 17: 69-70. 30 Mar 1949. Chamaesyce perennans Shinners, sp. nov. Type: B. L. Turner 3203, B. B. Tharp & B. H. Warnock 53-539 (SMU). Field & Lab. 24: 38. 15 Jun 1956. Chamaesyce polycarpa (Benth.) Millsp. var. simulans (L. C. Wheeler) Shinners, comb. nov. Field & Lab. 17: 70. 30 Mar 1949. Chamaesyce theriaca (lL. C. Wheeler) Shinners, comb. nov. Field & Lab. 17: 70. 30 Mar 1949. Chrysopsis wisconsinensis Shinners, sp. nov. Type: H. C. Greene, s. 7 Jul 1947 (SMU; isotypes GH, IU, MIL, ae Wrightia 1: 218-219. a Tah 1948. Chr seen viscidiflorus (Hook.) Nutt. var. ludens Shinners, var. nov. Typ . Shinners 9063 (SMU). Sida 1: 374-375. 30 Oct 1964. Cirsium terrae-nigrae Shinners, sp. nov. Type: L. H. Shinners 7486 (SMU). Fie'd & Lab. 17: 27-29. 18 Feb 1949 Cirsium texanun Buckl. var. stenolepis Shinners, var. nov. Type: L. H. Shinners 11192 (SMU). Field & Lab. 19: 81. 1 Jun 1951, Collinsonia L. subg. Micheliella (Briq.) Shinners, stat. nov. Sida 1: 76-77. 23 Nov 1962. Conradina brevifolia Shinners, sp. nov. Type: R. Garrett 41 (FLAS). Sida 1: 88. 23 Nov 1962. Conradina glabra Shinners, sp. nov. Type: F. H. Sargent 6219 (SMU). Sida 1: 85-86. 23 Nov 1962 Corydalis micrantha Engelm. & Gray var. australis (Chapm.) Shinners, comb. nov. Field & Lab. 18: 42. 2 Jan 1950 Corydalis micrantha Engelm. & Gray var. texensis (G. B. Ownbey) Shinners, omb. nov. Field & Lab. 18: 42. 2 Jan 1950. Croptilon eee hte var. graniticum (i. B. Smith) Shinners, comb. nov. Sida 3: 348. 6 Aug 1969. Croptilon . aricatum (Nutt.) Raf. var. hirtellum (Shinners) Shinners, comb. nov. Field & Lab. 19: 134. 24 Jul 1951 Croptilon divaricatum (Nutt.) Raf. var. hookerianum (T. & G.) Shinners, comb. nov. Field & Lab. 19: 134. 24 Jul 1951. Croton capitatus Michx. var. albinoides (Ferg.) Shinners, comb. nov. Field & Lab. 19: 183. 15 Oet 1951. 2) 155 Croton glandulosus L. var. hirsutus Shinners, var. nov. Type: V. L. Cory 54597 (SMU). Field & Lab. 19: 183. 15 Oct 1951. Cryptantha jamesii var. setosa (M. E. Jones) Shinners, comb. nov. Sida 3: 47. 6 Aug 1969. Cynanchum arizonicum (Gray) Shinners, comb. nov. Sida 1: 365. 30 Oct 196 Cynanchum barbigerum (Scheele) Shinners, comb. nov. Field & Lab. 19: 65. Cynanchum barbigerum (Scheele) Shinners var. breviflorwm Shinners, var. ov. Type: G. L. Webster 4340 (SMU). Sida 1: 360. 30 Oct 1964. Cynanchum blodgettii (Gray) Shinners, comb. nov. Sida 1: 365. 30 Oct 1964. Cynanchum grayi Shinners, nom. nov. Field & Lab. 20: 110. 8 Jul 1952. Cynanchum maccartit Shinners, nom. nov. Sida 1: 360. 30 Oct 1954. Cynanchum palmeri (Wats.) Shinners, comb. nov. Field & Lab. 19: 65. 1 Jun 1951. Cynanchum wigginsti Shinners, nom. nov. Sida 1: 365. 30 Oct 1964. Cyperus globulosus Aubl. var. robustus (Boeck.) Shinners comb. nov. Field & Lab. 20: 34. 29 Feb 1952 Cyperus virens Michx. var. arenicola (Steud.) Shinners, comb. nov. Field & Lab. 22: 30. Jan 1954. Dalea candida Willd. var. oligophylla (Torr.) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 409. 3 Apr 195 Dalea drummondiana Shinners, nom. nov. Field & Lab. 17: 83. 13 Jul 1949. Dalea emarginata (T. & G.) Shinners, comb nov. Field & Lab. 17: 84. 13 Jul 1949 Dalea glandulosa (Coult. & Fish.) Shinners, comb. nov. Field & Lab. 17: 83. 13 Jul 1949. Dalea grisea (T. & G.) Shinners, comb. nov. Field & Lab. 17: 84. 13 Jul 1949. Dalea helleri Shinners, nom. nov. Field & Lab. 21: 165. 15 Feb 1954 Dalea laxiflora Pursh var. pumila Shinners, var. nov. Type: E. Whitehouse 0515 (SMU). Field & Lab. 21: 164. 15 Feb 1954. Dalea multiflora (Nutt.) Shinners, comb. nov. Field & Lab. 17: 82. 13 Jul 1949 Dalea obovata (T. & G.) Shinners, comb. nov. Field & Lab. 17: 84. 18 Jul 1949. Dalea oligophylla (Torr.) Shinners, comb. nov. Field & Lab, 17: 82-83, 18 Jul 1949. Dalea phleoides (T. & G.) Shinners, comb. nov. Field & Lab. 17: 83. 13 Jul 1949. Dalea purpurea Vent. var. tenuis (Coult.) Shinners, comb. nov. Field & Lab. 21: 165. 15 Feb 1954. Dalea reverchoni (Wats.) Shinners, comb. nov. Field & Lab. 17: 84. 13 Jul 1949, Dalea sabinalis (Wats.) Shinners, comb. nov. Field & Lab. 17. 83. 13 Jul 1949. 156 Da —_— oOo = ea stanfieldii (Small) Shinners, comb. nov. Field & Lab. 17: 84-85. 13 Jul 49 Dalea tenuifolia (Gray) Shinners, comb, nov. Field & Lab. 17: 84. 13 Jul 1949. Dalea tenuis (Coult.) Shinners, comb. nov. Field & Lab. 17: 84. 13 Jul 1949. Delphinium virescens Nutt. var. wootoni (Rydb.) Shinners, comb. nov. Field & Lab. 17: 89. 13 Jul 1949. Descuraina pinnata var. glabra (Woot. & Standl.) Shinners, comb. nov. Field & Lab. 17: 145. 24 Oct 1949 Descurainia pinnata var. ochroleuca (Woot.) Shinners, comb. nov. Field & Lab. 17: 145. 24 Oct 1949. Descurainia pinnata var. osmiarum (Cockll.) Shinners, comb, nov. Field & Lab. 17: 145, 24 Oct 1949 Desmodium dichromum Shinners, sp. nov. Type: EE. Whitehouse 15866 (SMU). Spring Flora of the Dallas-Fort Worth Area, Texas, pp. 409-410. 3 Apr 1958. Dianthera americana L. var. subcoriacea (Fern.) Shinners, comb. nov. Field & Lab. 25: 79-80. 26 Sep 1957 Dicerandra frutescens Shinners, sp. nov. Type: F. H. Sargent 6600 (SMU). Sida 1: 89-90. 23 Nov 1962. Drosera leucantha Shinners, sp. nov. Type: A. Cronquist 5255 (SMU; isotypes NY, US). Sida 1: 57-58. 23 Nov 1962. Echinochloa crusgalli (L.) Beauv. var. macera (Wieg.) Shinners, comb. nov. Rhodora 56: 34. 17 Mar 1954. Echinochloa crusgalli (L.) Beauv. var. microstachya (Wieg.) Shinners, comb. nov. Rhodora 56: 34. 17 Mar 1954. Echinochloa crusgalli (l.) Beauv. ssp. muricata (Michx.) Shinners, comb. nov. Rhodora 56: 33-34. 17 Mar 1954 Echinochloa crusgalli (L.) Beauv. ssp. zelayensis (H. B. K.) Shinners, comb. nov. Rhodora 56: 33-34. 17 Mar 1954 Egletes commixta Shinners, sp. nov. Type: W. I. Broadway 4149 (GH: iso- type US). Lloydia 12: 243. 20 Mar 1950. Fgletes florida Shinners, sp. nov. Type: H. Pittier 6357 (US). Lloydia 12: 248. 20 Mar 1950. Egletes tebermanti Sch. Bip. var. eee Shinners, var. nov. Type: C. L. Lundell 1263 (GH; isotypes UC, F, , US). Lloydia 12: 246. 20 Mar 1950. Igletes repens Shinners, sp. nov. i. Cuatrecasas 3922 (US). Lloydia 12: 249. 20 Mar 1950. Iigletes viscosa f. bipinnatifida Shinners, f. nov. Type: A. H. Curtiss W. Ind. Pl. 697 (GH; isotypes F, US). Lloydia 12: 244-245. 20 Mar 1950. Egletes viscosa var. dissecta Shinners, var. nov. Type: J. G. Ortega 5156 (GH; isotype US). Lioydia 12: 245. 20 Mar 1950. Elymus canadensis L. var. villosus (Muhl.) Shinners, comb. nov. Rhodora 96: 28. 17 Mar 1954. Elymus trachycaulus (Link) Gould ex Shinners, comb. nov. Rhodora 56: 28. 17 Mar 1954. ts ee 157 Encelia scaposa Gray var. stenophylla Shinners, var. nov. Type: V. L. Cory 43870 (SMU). Sida 1: 375. 30 Oct 1964 Eragrostis oxylepis Torr. var. beyrichii (J. G. Smith) Shinners, comb. nov. Field & Lab. 20: 34. 29 Feb 1952 Ericameria laricifolia (Gray) Shinners, comb. nov. Field & Lab. 18: 27. 2 Jan 1950 Ericameria triantha (Blake) Shinners, comb. nov. Field & Lab. 19: 133. 24 Jul 1951. Erigeron geiseri Shinners, sp. nov. Type: L. H. Shinners 7205 (SMU). Wrightia 1: 183-184. 15 Dec 1947. Erigeron geiserit Shinners var. calcicola Shinners, var. nov. Type: L. H. Shinners 7329 (SMU). Wrightia 1: 184. 15 Dec 1947. Erigeron lobatus A. Nels. var. warnockii Shinners, var. nov, Type: Warnock 424 (TEX). Sida 1: 376. 30 Oct 1964. Erigeron mimegletes Shinners, sp. nov. Type: V. L. Cory 53650 (SMU). Wrightia 1: 184-186. 15 Dee 1947. Erigeron traversii Shinners, sp. nov. Type: Lundell & Lundell 11093 (SMU). Sida 1: 376. 30 Oct 1964. Eriogonum vespinum Shinners, sp. nov. Type: L. H. Shinners 18967 (SMU). Field & Lab. 22: 68-69. June 1954. Eustoma barkleyi Standl. ex Shinners, sp. nov. Type: A. Hernandez C., C. vell, & F. A. Barkley 16M548 (TEX). Southwest. Nat. 2: 39-40. 15 Oct 1957. Eustoma grandiflorum (Raf.) Shinners, comb. nov. Southwest. Nat. 2: 41-43. 15 Oct 1957 Eustoma grandiflorum (Raf.) Shinners f. bicolor (Standl.) Shinners, comb. nov. Southwest. Nat. 2: 41. 15 Oct 1957. Eustoma grandiflorum (Raf.) Shinners f. fisheri (Standl.) Shinners, comb. nov. Southwest. Nat. 2: 41. 15 Oct 1957 Eustoma grandiflorum (Raf.) Shinners f. flaviflorum (Cockll.) Shinners, Eustoma grandiflorum (Raf.) Shinners f. roseum (Standl.) Shinners, comb. nov. Southwest. Nat. 2: 41-42. 15 Oct 1957 Filago candida (T. & G.) Shinners, comb. nov. Sida 1: 259-253. 25 Jun 1964. Filago nuttallii Shinners, nom. nov. Sida 1: 253. 25 Jun 1964. Filago verna (Raf.) Shinners, comb. nov. Sida 1: 253. 25 Jun 1964. Filago verna (Raf.) Shinners var. drummondii (T. & G.) Shinners, comb. nov. Sida 1: 253. 25 Jun 1964. Forestiera pubescens Nutt. var. glabrifolia Shinners, var. nov. Type: L. H. Shinners 11331 (SMU). Field & Lab. 18: 99-100. 3 Apr 1950. Gentiana deloachti (W. P. Lemmon) Shinners, comb. nov. Sida 1: 107. 23 OV Gerardia tenuifota Vahl var. leucanthera (Raf.) Shinners, comb. nov. Field : 180. 9 May 1950. Pea tamuitolia Vahl var. polyphylla (Small) Shinners, comb. nov. Field 158 & Lab. 18: 130. 9 May 1950. Gilia ludens Shinners, sp. nov. Type: L. H. Shinners 19581 (SMU). Sida 1: 174-175. 12 Dee 1963. Gilia perennans Shinners, sp. nov. Type: D. S. Correll 13958 (SMU). Sida 1: 2 Dec 1963. Grindelia lanceolata Nutt. var. texana (Scheele) Shinners, comb. nov. Field & Lab. 19: 78. 1 Jun 1951. Hedyotis angulata Fosb. ex Shinners, nom. nov. Field & Lab. 17: 166-167. Hedyotis crassifolia Raf. var. micrantha Shinners, var. nov. Type: FE. White- house 20848 (SMU). Field & Lab. 18: 100. 3 Apr 1950. Hedyotis croftiae (Britt. & Rusby) Shinners, comb. nov. Field & Lab. 17: 167. 24 Oct 1949 Hedyotis greenmanii Fosb. ex Shinners, nom. nov. Field & Lab. 17: 167. 24 Oct 1949. Hedyotis nigricans (Lam.) Fosb. var. filifolia (Chapm.) Shinners, comb. nov. Field & Lab. 17: 168. 24 Oct 1949 Hedyotis nigricans (Lam.) Fosb. var. rigidiuscula (Gray) Shinners, comb. nov. Field & Lab. 17: 168. 24 Oct 1949. Hedyotis polypremoides (Gray) Shinners, comb. nov. Field & Lab. 17: 168. Hedyotis salina (Heller) Shinners, comb. nov. Field & Lab. 17: 169. 24 Oct 1949. Hedyotis subviscosa (Wright) Shinners, comb. nov. Field & Lab. 17: 169. 24 Oct 1949 Hedyotis taylorae Fosb. ex Shinners, nom. nov. Field & Lab. 17: 169. 24 Oct 1949 Heleastrum chapmanii (T. & G.) Shinners, comb. nov. Sida 3: 348. 6 Aug 1969. Heleastrum hemisphericum (Alex.) Shinners, comb. nov. Field & Lab. 17: 170-171. 24 Oct 1949. Heleastrum verutifolium (Alex.) Shinners, comb. nov. Field & Lab. 17: 171. 24 Oct 1949. Helianthus annuus L. var. tevanus (Heiser) Shinners, comb. nov. Sida 1: 377. 30 Oct 1964. Helianthus ludens Shinners, sp. nov. Type: Turner, Tharp, & Warnock 53-043 (SMU). Sida 1: 377-378. 30 Oct 1964. Heteranthera licbmannii (Buch.) Shinners, comb. nov. Field & Lab. 23: 21. 8 Jun 1955. Heterotheca aspera (Shuttlew.) Shinners, comb. nov. Sida 3: 348. 6 Aug 1969. Heterotheca breweri (Gray) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951 Heterotheca camporum (Greene) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951 Heterotheca canescens (DC.) Shinners, comb. nov. Field & Lab. 19: 68. 159 1 Jun 1951. Heterotheca echioides (Benth.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951. Heterotheca foliosa (Nutt.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951 Heterotheca fulcrata (Greene) Shinners, comb. nov. Field & Lab. 19: 71. Heterotheca gossypina (Michx.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun Heterotheca Graminiona (Michx.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun Heterotheca microcephala (Small) Shinners, comb. nov. Field & Lab. 19: 71. un 1951 Heterotheca nervosa (Willd.) Shinners, comb. nov. Field & Lab. 19: 68. 1 Jun 1951. Heterotheca oregona (Nutt.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951. Heterotheca pilosa (Nutt.) Shinners, comb. nov. Field & Lab. 19: 68. 1 Jun 1951 Heterotheca rutteri (Roth.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun ol, Heterotheca sessiliflora (Nutt.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun Heterotheca sienonha (Gray) Shinners, comb. nov. Field & Lab. 19: 68. 1 Jun Heterotheca en ee nie (Nutt.) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951 Heterotheca villosa (Pursh) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951 Heterotheca wisconsinensis (Shinners) Shinners, comb. nov. Field & Lab. 19: 71. 1 Jun 1951. Hybanthus linearis (Torr.) Shinners, comb. nov. Field & Lab. 19: 126. 24 Jul 1951 Hymenocallis eulae Shinners, sp. nov. Type: E. Whitehouse 16448 (SMU). Field & Lab. 19: 108. 1 Jun 1951. Hymenocallis liriosme (Raf.) Shinners, comb. nov. Field & Lab. 19: 102-103. 1 Hymenopappus filifolius var. alpestris (Maguire) Shinners, comb. nov. Rho- j Jul 1959 Hymenoxys glabra (Nutt.) Shinners, comb. nov. Field & Lab. 19: 80-81. 1 Jun 1951 ee scaposa (DC.) Parker var. villosa Shinners, var. nov. Type: y 53129 (SMU). Field & Lab. 19: 79-80. 1 Jun 1951. Hiplis: ala a (Raf.) Shinners, comb. nov. Rhodora 64: 184-186. 27 Jun 1962. Hyptis alata var. stenophylla Shinners, var. nov. Type: R. Kral 5657 (SMU; 160 isotype FSU). Rhodora 64: 185-186. 27 Jun 1963. Ipomoea & multifida (Raf.) Shinners, comb. nov. Sida 2: 265. 6 Apr 1966. Ipomoea shumardiana (Torr.) Shinners, comb. nov. Southwest. Nat. 6: Q-101. 1 Sep 1961. Ipomoea trichocarpa Ell. var. torreyana (Gray) Shinners, comb. nov. Fie & Lab. 21: 164-165, 15 Feb 1954 Ipomopsis aggregata (Pursh) V. Grant var. tevana (Greene) Shinners, comb. nov. Sida 1: 177. 12 Dee 1963. Ipomopsis wrightti (Gray) Shinners, comb. nov. Sida 1: 178. 12 Dee 1963. Isocoma megalantha Shinners, sp. nov. Type: J. C. Johnson 1638 (SMU), Field & Lab, 23: 34-36, 13 Jun 1955. Isocoma palmeri (Gray) Shinners, comb. nov. Field & Lab. 18: 27. 2 Jan 1950. Isopappus divaricatus (Nutt.) T. & G. var. hirtellus Shinners, var. nov. ‘ype: C. L. & A. A. Lundell 10813 (SMU). Field & Lab. 18: 157. 11 Dec — Qo | 1950. Isopappus divaricatus (Nutt.) T. & G. var. hookerianus (T. & G.) Shinners, comb. nov. Field & Lab. 18: 157. 11 Dee 1950. Iva angustifolia Nutt. var. latior Shinners, var. nov. Type: Lundell & TLun- dell 11947, Sida 1: 378. 30 Oct 1964 Koeleria gerardi (Vill.) Shinners, comb. nov. Rhodora 58: 95-96. 4 May 1956. Koeleria pectinata (Lam.) Shinners, comb. nov. Rhodora 58: 95. 4 May 1956. Krigia gracilis (DC.) Shinners, comb. nov. Wrightia 1: 205-206. 15 Dee 1947. Krigia occidentalis Nutt. f. mutica (T. & G.) Shinners, comb. nov. Wrightia 1: 200-202. 15 Dec 1947 Kuhnia aie ortoides L. var. ozarkana Shinners, var. nov. Type: E. J, Palm- er 33075 (MO; isotype GH). Wrightia 1: 136. 30 Apr 1946. rel inti L. var. tevana Shinners, var. nov. Type: L. H. Shinners 475 (S . Wrightia 1: 136-138. 30 Apr 1946 Kuhnia ieplopiiila Scheele var. mexicana Shinners, var. nov. Type: E. Palmer 935 (of the 1898 series) (GH; isotypes UC, MO). Wrightia 1: 128-129. 30 Apr 1946. Kuhnia microphylla Shinners, sp. nov. Type: Brother Nicolas 5932 (GH: iso- type ILL). Wrightia 1: 127. 30 Apr 1946. Lactuca hirsuta Muhl. var. albiflora (T. & G.) Shinners, comb. nov. Field & Lab. 23: 36. 13 Jun 1955. Liatris aspera Michx. var. salutans (Lunell) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 410. 3 Apr 1958. Liatris < deamii (Lunell) Shinners, comb. nov., stat. emend. Am. Midl. Nat. 29: 31-32. 6 Mar 1948 liatris x deamiti (Lunell) Shinners f. albina Shinners, f. nov. Type: N. C. Fassett 21207 (WIS). Am. Midl. Nat. 29: 31-32. 6 Mar 1943. Liatris glabrata Rydb. var. alabamensis (Alex.) Shinners, comb. nov. Field & Lab. 19: 76. 1 Jun 1951, Liatris < gladewitzii (Farw.) Farw. ex Shinners, comb. nov. Am. Midl. Nat. 29: 37. 6 Mar pare 161 Liatris ligulistylis (A. Nels.) K. Schum. f. lewcantha Shinners, f. nov. Type: J. Lunell, s. n., 9 Sep 1908 (MIN). Am. Midl. Nat. 29: 39-40. 6 Mar 1943. Liatris mucronata DC. f. tharpii Shinners, f. nov. Type: B. C. Tharp & B. H. Warnock 45-35 (SMU). Field & Lab. 23: 34. 13 Jun 1955. Liatris novae-angliae (Lunell) Shinners, stat. nov. Am. Midl. Nat. 29: 29. 6 Mar 1943 Liatris can aig (Lunell) Shinners f. albiflora Shinners, f. nov. Type: A. L. Page, s , 10 Sep 1909 (GH). Am. Midl. Nat. 29: 29-31. 6 Mar 1948. Liatris novae- ie (Lunell) Shinners var. nieuwwlandti (Lunell) Shinners, comb. nov. Am. Midl. Nat. 29: 31. 6 Mar 1948. Liatris novae-angliae (Lunell) Shinners var. niewwlandii (Lunell) Spears f, alba Shinners, f. nov. Type: Deam 43670 (IND). Am. Midl. Nat. 29: 6 Mar 1943, Liatris novae-angliae (Lunell) Shinners var. nieuwiandii (Lunell) Shinners f. trilisioides (Farw.) Shinners, stat. nov. Am. Midl. Nat. 29: 31. 6 Mar 1943. Liatris pycnostachya Michx. var. lasiophylla Shinners, var. nov. Type: V. L. Cory 549538 (SMU). Field & Lab. 19: 74-75. 1 Jun 1951 Liatris sphaeroidea Michx. f. asperifolia Shinners, f. nov. Type: N. C. Fas- sett 13469 (WIS). Am. Midl. Nat. 29: 36-37. 6 Mar 1943. Liatris sphaeroidea Michx. f. benkei (Macbr.) Shinners, comb. nov. Am. Midl. Nat. 29: 35-87. 6 Mar 1948. Liatris sphaeroidea Michx. var. salutans (Lunell) Shinners, comb. nov. Am. Midl. Nat. 29: 37. 6 Mar 1948. Liatris tenuis Shinners, sp. nov. Type: L. H. Shinners 26600 (SMU). South- west. Nat. 4: 207-208. 1 Dec 1959. Liatris * weaveri seas sp. nov. Type: ‘‘From seeds sent by Dr. Weaver to Dr. Gaiser, no. 16°’ (MO). Am. Midl. Nat. 29: 38. 6 Mar 1943. Limonium carolinianum (Walt.) Britt. var. compactum Shinners, var. nov. Type: V. L. Cory 51236 (SMU). Field & Lab. 24: 105-106. 7 Oct 1956. Linum imbricatum (Raf.) Shinners, comb. nov. Field & Lab. 25: 32. 15 Mar Linum rigidum Pursh var. filifolium Shinners ex McVaugh, var. nov. Type: . MeVaugh 7798 (SMU). Field & Lab. 17: 136-137. 24 Oct 1949. Ludwigia octovalvis (Jaeq.) Raven var. macropoda (Presl) Shinners, comb. nov. Sida 1: 385. 30 Oct 1964. Ludwigia octovalvis (Jacq.) Raven var. sessiliflora (Mich.) Shinners, comb. nov. Sida 1: 385. 30 Oct 1964. Ludwigia peploides (H. B. K.) Raven var. glabrescens (Kuntze) Shinners, comb. nov. Sida 1: 386. 30 Oct 1964. Ludwigia peploides (H. B. K.) Raven var. montevidensis (Spreng.) Shinners, comb. nov. Sida 1: 386. 30 Oct 1964. Lupinus perennis L. var. austrinus Shinners, var. nov. Type: Curtis School 21 (TEX). Field & Lab. 21: 152-1538. 15 Feb 1954. Lygodesmia bigelovit (Gray) Shinners, comb. nov. Field & Lab. 18: 31. Jan 1950 162 Lygodesmia pauciflora (Torr.) Shinners, comb. nov. Field & Lab. 18: 30. 2 Jan 1950 Lygodesmia tenuifolia (Torr.) Shinners, comb. nov. Field & Lab. 18: 31. 2 Jan 1950 Lygodesmia thurberi (Gray) Shinners, comb. nov. Field & Lab. 18: 32. 2 Jan 1950 Lygodesmia wrightii (Gray) Shinners, comb. nov. Field & Lab. 18: 31. 2 Machaeranthera annua (Rydb.) Shinners, comb. nov. Sida 1: 378. 30 Oct 1964. Machaeranthera arenaria (Benth.) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950. Machaeranthera aurea (Gray) Shinners, comb. nov. Field & Lab. 18: 41. 2 Jan 1950 Machaeranthera australis (Greene) Shinners, comb. nov. Field & Lab. 18: 42. 2 Jan 1950. Machaeranthera blephariphylla (Gray) Shinners, comb. noy. Field & Lab. 18: 38-40. 2 Jan 1950. Machaeranthera correllii Shinners, sp. nov. Type: L. H. Shinners 9059 (SMU). Field & Lab. 17: 56-57. 30 Mar 1949. Machaeranthera gracilis (Nutt.) Shinners, comb. nov. Field & Lab, 18: 41. 2 Ja Machaeranthera grindelioides (Nutt.) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950. dierent gymnocephala (DC.) Shinners, comb. nov. Field & Lab. 18: Rag 2 Jan Pie I t havardiit (Waterfall) Shinners, comb. nov. Field & Lab. 18: “40. M1. 2 os 1950. Machaeranthera juncea (Greene) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950 Machaeranthera laevis (Woot. & Standl.) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950. Machaeranthera phyllocephala (DC.) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950 Machaeranthera phyllocephala (DC.) Shinners var. annua (Rydb.) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950. Machaeranthera phyllocephala (DC.) Shinners var. megacephala (Nash) Shinners, comb. nov. Field & Lab. 18: 40. 2 Jan 1950. Machaeranthera pinnata (Nutt.) Shinners, comb. nov. Field & Lab. 18: 41-42. 2 Jan 1950 nT Machaeranthera pinnatifida (Hook.) Shinners, comb. nov. Sida 1: 295. 25 Jun 1964. Machaeranthera scabrella (Greene) Shinners, comb. nov. Field & Lab. 18: 42. 2 Jan 1950 2 Machaeranthera stenoloba (Greene) Shinners, comb. nov. Field & Lab. 18: 40. 2 2 Ja 163 Machaeranthera texensis (R. C. Jackson) Shinners, comb. nov. Sida 1: 378. 30 Oct 1964. Matelea albomarginata (Pittier) Shinners, comb. nov. Sida 1: 365. 30 Oct 6 Matelea aristolochiaefolia (Brandeg.) Shinners, comb. nov. Sida 1: 365. 30 Oct 1964. Matelea arizonica (Gray) Shinners, comb, nov. Sida 1: 366. 30 Oct 1964. Matelea calycosa (J. D. Smith) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964. Matelea chiapensis (Brandeg.) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964. Matelea cteniophora (Blake) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964. Matelea gonocarpa (Walt.) Shinners, comb. nov. Field & Lab. 18: 73. 3 Apr 1950. Matelea greenmanii Shinners, nom. nov. Sida 1: 366. 30 Oct 1964. Matelea johnstonii Shinners, nom. nov. Sida 1: 366. 30 Oct 1964. Matelea lasiostemma (Hemsl.) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964. Matelea oblongifolia (J. D. Smith) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964 Matelea sagittifolia (Gray) Woods. ex Shinners, comb. nov. Sida 1: 363-364. : Io. Matelea smithii Shinners, nom. nov. Sida 1: 366. 30 Oct 1954. Matelea stenantha (Standl.) Shinners, comb. nov. Sida 1: 366. 30 Oct 1964. Matelea woodsonii Shinners, sp. nov. Type: Lundell & Lundell 9926 (SMU). Sida 1: 364. 30 Oct 1964 Medicago polymorpha L. var. ciliaris (Ser.) Shinners, comb. nov. Rhodora 58: 9-12. 24 Jan 1956. Medicago polymorpha L. var. ciliaris (Ser.) Shinners f. apiculata (Willd., emend. Urban) Shinners, comb. nov. Rhodora 58: 11. 24 Jan 1956. Medicago polymorpha L. var. ciliaris (Ser.) Shinners f. tuberculata (Gren, & Godr.) Shinners, comb. nov. Rhodora 58: 11-12. 24 Jan 1956. Medicago polymorpha L. var. polygyra (Urban) Shinners, comb. nov. Rho- dora 58: 9. 24 Jan 1956 Medicago polymorpha L. var. tricycla (Gren. & Godr.) Shinners, comb. nov. do : 8. 24 Jan 1956. Medicago polymorpha L. var. vulgaris (Benth.) Shinners, comb. nov. Rho- dora 58: 310. 31 Oct 1956. Medicago polymorpha LL. var. vulgaris (Benth.) Shinners f. apiculata (Willd.) Shinners, comb. nov. Rhodora 58: 310. 31 Oct 1956. Medicago polymorpha L. var. vulgaris (Benth.) Shinners f. tuberculata (Godr.) Shinners, comb. nov. Rhodora 58: 310. 31 Oct Micranthemum glomeratum (Chapm.) Shinners, comb. nov. Sida 1: 202: 54. Micromeria bahamensis Shinners, sp. nov. Type: N. L. Britton & C. F. Millspaugh 2302 (GH: isotype NY). Sida 1: 96. 23 Nov 1962. Micromeria brownei (Sw.) Benth. var. ludens Shinners, var. nov. Bro. Alain 61387 (NY). Sida 1: 96. 23 Nov 1962. 164 Micromeria domingensis Shinners, sp. nov. Type: E. L. Ekman, s. n., 8 Sep 1929 (GH). Sida 1: 96. 23 Nov 1962 Mirabilis collina Shinners, sp. nov. Type: L. H. Shinners 10109 (SMU). Field & Lab. 19: 180. 15 Oct 1951 Mirabilis dumetorum Shinners, sp. nov. Type: L. H. Shinners 9536 (SMU). Field & Lab. 19: 179. 15 Oct 1951, Mirabilis eutricha Shinners, sp. nov. Type: lL. H. Shinners 11345 (SMU). Field & Lab. 19: 177. 15 Oet 1951. Mirabilis oe (Standl.) Shinners, comb. nov. Field & Lab. 19: 177-178. 15 Oct Mirabilis ae imeri (Standl.) Shinners, comb. nov. Field & Lab, 19: 175-176. 15 Oct 1951 Mirabilis serotina Shinners, sp. nov. Type: S. P. Gordon 74( SMU). Field & Lab. 19: 179. 15 Oct 1951. Monarda punctata L. var. arkansana (McCl, & Iepl.) Shinners, comb. nov. Field & Lab. 21: 90. 12 Jun 1953. Monarda punctata L. var. coryi (McCl. & Epl.) Cory ex Shinners, comb. nov. Field & Lab. 21: 90. 12 Jun 1953. Monarda punectata L. var. villicaulis (Penn.) Shinners, comb. nov. Field & Lab. 21: 90. 12 Jun 1953. Ovalis dillenti Jacq. var. radicans Shinners, var. nov. Type: L. H. Shinners 23872 (SMU). Field & Lab. 24: 39-40. 15 Jun 1956. Palafoxia callosa (Nutt.) T. & G. var. bella (Cory) Shinners, comb. nov. Field & Lab. 20: 94. 8 Jul 1952. Palafoxia cyanophylla Shinners, sp. nov. Type: L. H. Shinners 8792 (SMU). Field & Lab. 17: 25-27. 18 Feb 1949 Palafoxia hookeriana T, & G. var. minor Shinners, var. nov. Type: G. L. Fisher 50717 (SMU). Field & Lab. 20: 98. 8 Jul 1952. Palafoxia liebmannii (Sch. Bip.) Shinners, comb. nov. Field & Lab. 17: 25. 18 Feb 1949. Palafoxia pedata (Cav.) Shinners, comb. nov. Field & Lab. 17: 25. 18 Feb 1949 Palafoxia rosea (Bush) Cory var, ambigua Shinners, var. nov. Type: V. L. Cory 51241 (SMU). Field & Lab. 20: 95. 8 Palafoxia rosea (Bush) Cory var. papposa Shinners, var. nov. Type: H. B. Parks, s. n., 27 Aug 1934 (TAES; fragments, SMU). Field & Lab. 20: 95.. 8 Jul 1952, Palafoxia texana DC. var. i (Rydb.) Shinners, comb. nov. Field & Lab. 20: 97. 8 Jul 1 Palafoxia tripteris (DC.) ek comb. nov. Field & Lab. 17: 24-25. 18 Feb 1949 Palafoxia tripteris (DC.) Shinners var. brevis Shinners, var. nov. Type: V. L. Cory 40710 (SMU). Field & Lab, 20: 94. 8 Jul 1952. Panicum euchlamydeum Shinners, sp. nov. Type: L. H. Shinners & C. G. Shaw 4415 (WIS; isotypes GH, US, MIN, MII). Am. Midl. Nat. 32: 170-171. 165 27 Oct 1944. Pappopecus Shinners, sect. nov. Wrightia 1: 108. 30 Apr 1946. Pappophanus (Blake) Shinners, sect. nov. Wrightia 1: 101. 30 Apr 1946. Parietaria pennsylvanica Muhl. var. obtusa (Rydb.) Shinners, comb. nov. Field & Lab. 18: 42. 2 Jan 1950. Paronychia erecta (Chapm.) Shinners, comb. nov. Sida 1: 102. 23 Nov 1962. Paronychia patula Shinners, nom. nov. Sida 1: 102. 23 Nov Paspalum distichum L. var. indutum Shinners, var. nov. Type: L. H. Shin- ners 10564 (SMU). Rhodora 56: 31. 17 Mar 1954. Paspalum separatum Shinners, sp. nov. Type: L. H. Shinners 15566 (SMU). Rhodora 56: 32. 17 Mar 1954 Perityle angustifolia (Gray) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec 1959 Perityle bisetosa (Torrey) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec 1959 Perityle cernua (Greene) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec Perityle cengesta (M. E. Jones) Shinners, comb. nov. Southwest Nat. 4: 204. 1 Dee 1959 Perityle fastigiata (Brandeg.) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec 1959. Perityle halimifolia (Gray) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec 1959. Perityle intricata (Brandeg.) Shinners, comb. nov. Southwest. Nat. 4: 204. 1959. Perityle lindheimeri (Gray) Shinners, comb. nov. Southwest. Nat. 4: 204. 1 Dec. 1959. Perityle quinqueflora (Steyerm.) Shinners, comb. nov. Southwest Nat. 4: 205. ec. 19 oy Perityle rotundata (Rydb.) Shinners, comb. nov. Southwest. Nat. 4: 205. 1 Dec 1959 Perityle rupestris (Gray) Shinners, comb. nov. Southwest. Nat. 4: 205. 1 Dee 1959. Perityle saxicola (Eastw.) Shinners, comb. nov. Southwest. Nat. 4: 205. 1 Dec 1959. Perityle staurophylla (Barneby) Shinners, comb. nov. Southwest. Nat. 4: 205. 1 Dec 1959. Perityle villosa (Blake) Shinners, comb. nov. Southwest. Nat. 4: 205. 1 Dec 1959 Phlox bifida Beck var. induta Shinners, var. nov. Type: W. M. Thompson 1 (SMU; isotype TEX). Southwest. Nat. 6: 50-51. 10 Jun 1961. Phlox drummondii Hook. var. mcallisieri (Whiteh.) Shinners, comb. nov. Field & Lab. 19: 127. 24 Jul 1951. Phlox drummondii Hook. var. peregrina Shinners, var. nov. Type: V. E. Bono 6 (SMU). Field & Lab. 19: 127. 24 Jul 1951. 166 Physostegia correllii (Lundell) Shinners, comb. nov. Rhodora 51: 120-122. 8 Jun 1949 Physostegia edwardsiana Shinners, sp. nov. Type: C. L. & A. A. Lundell 13851 (SMU). Field & Lab. 19: 167-168. 15 Oct 1951. Physostegia praemorsa Shinners, sp. nov. Type: L. H. Shinners 11980 (SMU). Field & Lab. 19: 166-167. 15 Oct 1951 Physostegia serotina Shinners, sp. nov. Type: L. H. Shinners 22108 (SMU). Field & Lab. 24: 17. 15 Jun 1956. Pinaropappus roseus (Less.) Less. var. foliosus (Heller) Shinners, comb. nov. Field & Lab. 19: 48. 18 Jan 1951. Plantago patagonica Jacq. var. breviscapa (Shinners) Shinners, comb. nov. Sida 3: 121-122. 31 May 1967. Plantago patagonica Jacq. var. oblonga (Morris) Shinners, comb. nov. Sida 122. 31 May 1967. Plantago purshii R. & S. var. breviscapa Shinners, var. nov. Type: L. H. Shinners 11350 (SMU). Fie'd & Lab. 18: 118. 9 May 1950. Plantago purshii R. & S. var. spinulosa (Dene.) Shinners, comb. nov. Field & Lab. 18: 117-118. 9 May 1950. Polianthes brachystachys (Cav.) Shinners, comb. nov. Sida 2: 337. 20 Jul Polianthes brunnea (Wats.) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes debilis (Berger) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Poltanthes densiflora (Robins. & Fern.) Shinners, comb. nov. Sida 2: 336. 20 Jul 1966. Polianthes elongata (Rose) Shinners, comb. nov. Sida 2: 337. 20 Jul 1956. Polianthes guttata (Jacobi & Bouché) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes jaliscana (Rose) Shinners, comb. nov. Sida 2: 337. 20 Jul 1986. Polianthes lata (Shinners) Shinners, comb. nov. Sida 2: 335. 20 Jul 1936. Polianthes maculosa (Hook.) Shinners, comb, nov. Sida 2: 335-338. 20 Jul 1966. Polianthes oliverana (Rose) Shinners, comb. nov. Sida 2: 337. 20 Jul 196%. Polianthes planifolia (Wats.) Shinners, comb. nov. Sida 2: 337, 20 Jul 1966. Polianthes potosina (Robins. & Greenm.) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes revoluta (K1.) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes rosei Shinners, nom. nov. Sida 3: 183. 24 Nov 1967, Polianthes rubescens (Rose) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes runyonii Shinners, nom, nov. Sida 2: 335. 20 Jul 1966. Polianthes singuliflora (Wats.) Shinners, comb. nov. Sida 2: 337. 20 Jul 1966. Polianthes variegata (Jacobi) Shinners, comb. nov. Sida 2: 336. 20 Jul 1966. Polianthes virginica (1..) Shinners, comb. nov. Sida 2: 335. 20 Jul 1966. Polianthes virginica (l.) Shinners f. tigrina (Engelm.) Shinners, comb. nov. Sida 2: 335. 20 Jul 1966. Psilactis leptos Shinners, sp. nov. Type: L. H. Shinners 8918 (SMU). Field 167 & Lab. 17: 53-55. 30 Mar 1949. Psoralea digitata Nutt. ex T. & G. var. parvifolia Shinners, var. nov. Type: L. H. Shinners 11174 (SMU). Field & Lab. 19: 19-21. 18 Jan 1951. Psoralea latestipulata Shinners, sp. nov. Type: R. W. Pohl 4817 (SMU). Field & Lab. 19: 22-23. 18 Jan 1951. Psoralea scaposa (Gray) Macbr. var. breviscapa Shinners, var. nov. Type: L. H. Shinners 10190 (SMU). Field & Lab. 19: 24-25, 18 Jan 1951. Psoralea subulata Bush var. minor Shinners, var. nov. Type: L. H. Shinners 11024 (SMU). Field & Lab. 19: 23-24. 18 Jan 1951. Pyrrhopappus geiseri Shinners, sp. nov. Type: L. H. Shinners 8499 (SMU). Field & Lab. 19: 81-82. 1 Jun 1951. — georgianus Shinners, sp. nov. Type: A. Cronquist 4276 (SMU; isotypes GA, MO). Field & Lab. 21: 93-94, 12 Jun 1953 es ie. Shinn Buckl. var. microcarpa (Torr.) Shinners, comb. nov. Field & Lab. 24: 37. 15 Jun 1956 Ranunculus carolinianus DC. var. villicaulis Shinners, var. nov. Type: G. L. Fisher 48102 (SMU). Field & Lab. 18: 84. 3 Apr 1950. Reynoutria baldschuanica (Regel) Shinners, comb. nov. Sida 3: 117. 31 May 1967 Reynoutria cilinodis (Michx.) Shinners, comb. nov. Sida 3: 117. 31 May 1967. Reynoutria convolvulus (L.) Shinners, comb. nov. Sida 3: 117. 31 May 1967. Reynoutria scandens (L.) Shinners, comb. nov. Sida 3: 118. 31 May 1967. Reynoutria scandens (lL. )Shinners var. cristata (Engelm. & Gray) Shinners, comb. nov. Sida 8: 118. 31 May 1967. Reynoutria scandens (L.) Shinners var. dumetorum (L.) Shinners, comb. nov. Sida 3: 118. 30 May 1967. Rhododendron coryi Shinners, sp. nov. Type: V. L. Cory 57145 (SMU). Cas- tanea 26: 156-157. 15 Jan 1962. Rhododendron periclymenoides (Michx.) Shinners, comb. nov. Castanea 27: Rhus aromatica Ait. var. flabelliformis Shinners, var. nov. Type: V. L. Cory 54413 (SMU). Field & Lab. 19: 86. 1 Jun 1951. Rhus aromatica Ait. var. pilosissima (Engl.) Shinners, comb. nov. Field & Lab. 19: 86-87. 1 Jun 1951. Rosa ignota Shinners, sp. nov. Type: N. Stillwell, s. n., 21 Jun 1946 (SMU). pring Flora of the Dallas-Fort Worth Area, Texas, p. 409. 3 Apr 1958 Rosa < rehderi Shinners, nom. nov. Baileya 3: 171. 13 Dec 1955. Rubus duplaris Shinners, sp. nov. Type: L. H. Shinners 14465 (SMU). Field Lab. 22: 27-30. Jan 1954. Rudbeckia coryi Shinners, sp. nov. Type: V. L. Cory 52412 (SMU). Field & Lab. 17: 59-62. 30 Mar 1949 Ruellia nudiflora (Engelm. & Gray) Urban var. hispidula Shinners, var. nov. Type: V. L. Cory 55632 (SMU). Field & Lab. 21: 165. 15 Feb 1954. Salvia farinacea Benth. var. latifolia Shinners, var. nov. Type: Ii. Whitehouse 18117 (SMU). Field & Lab. 21: 92, 12 Jun 1953 168 Sarcostemma bilobum Hook. var. lindenianum (Dene.) Shinners, comb. nov. Sida 1: 365. 30 Oct 1964. Sarcostemma cynanchoides Dene. var. hartwegii (Vail) Shinners, comb. nov. Sida 1: 361-362. 30 Oct 1964. Scutellaria laevis Shinners, sp. nov. Type: D. S. Correll 13973 (SMU). Sida Scutellaria thieretti Shinners, sp. nov. Type: J. W. Thieret 16162 (SMU; iso- type USL). Sida 1: 251-252. 25 Jun 1964. Senecio warnockii Shinners, sp. nov. Type: Turner & Warnock 202 (SMU). Sida 1: 379. 30 Oct 1964. Sida ciliaris L. var. mexicana (Moric.) Shinners, comb. nov. Field & Lab. 21: Sisyrinchium pruinosum Bickn. emend. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 409. 3 Apr 1958 Solanum godfreyi Shinners, nom. nov. Sida 1: 108. 23 Nov 1962. Solidago gigantea Ait. var. pitcheri (Nutt.) Shinners, comb. nov. Rhodora D0: 322, 953 Solidago mollis Bartl. var. angustata Shinners, var. nov. Type: E. White- house 10912 (SMU). Field & Lab. 19: 34-35. 18 Jan 1951. Solidago rigida L. var, laevicaulis Shinners, var. nov. Type: E. Whitehouse 19269. Field & Lab. 19: 35. 18 Jan 1951. Spergula platensis (St. Hil. & A. Juss.) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 409. 3 Apr 1958. Sphaeralcea angustifolia var, oblongifolia (Gray) Shinners, comb. nov. Sida 384-385. 30 Oct 1964 oe digitata var. angustiloba (Gray) Shinners, comb. nov. Sida 1: 385. 30 Oct 1964. Sphaeralcea emoryi var. californica (Parish) Shinners, comb. nov. Sida 1: 384. 380 Oct 1964. Sporobolus asper (Michx.) Kunth var. canovirens (Nash) Shinners, comb. nov. Rhodora 56: 30. 17 Mar 1954. Sporobolus asper (Michx.) Kunth var. clandestinus (Biehler) Shinners, comb. nov. Rhodora 56: 30. 17 Mar 1954. Sporobolus asper (Michx.) Kunth var. macer (Trin.) Shinners, comb. nov. Rhodora 56: 29. 17 Mar 1954. Sporobolus vaginiflorus (Torr.) Wood var. neglectus (Nash) Shinners, comb. nov. Rhodora 56: 29. 17 Mar 1954. Sporobolus vaginifiorus (Torr.) Wood var. ozarkanus (Fern.) Shinners, comb. nov. Rhodora 56: 29. 17 Mar 1954. Stellaria corei Shinners, nom. nov. Sida 1: 103-104. 23 Nov 1962. Stylisma patens var. angustifolia (Nash) Shinners, comb. nov. Sida 3: 347. 6 Aug 1969 Tephrosia curtissii (Small) Shinners, comb. nov. Sida 1: 60. 23 Nov 1962. Tephrosia florida (Dietr.) C. Wood var. gracillima (Robins.) Shinners, comb. ov. Sida 1: 61. 23 Nov 1962. ~~ i) 169 Tephrosia seminole Shinners, sp. nov. Type: P. P. Sheehan, s. n., 12 Mar 1919 (NY). Sida 1: 60. 23 Nov 1962. Tessaria sericea (Nutt.) Shinners, comb. nov. Sida 3: 122. 31 May 1967. Tetraneuris scaposa (DC.) var. villosa (Shinners) Shinners, comb. nov. Spring Flora of the Dallas-Fort Worth Area, Texas, p. 410. 3 Apr 1958. Teucrium canadense L. var. boreale (Bickn.) Shinners, comb. nov. Sida 1: 183. 12 Dec 1963 Teucrium canadense L. var. nashii (Kearn.) Shinners, comb. nov. Sida 1: 183. 12 Dee 1963 Teucrium cubense Jacq. var. laevigatum (Vahl) Shinners, comb. nov. Sida 4: 275. 22 Jul 1971. Thelesperma filifoliwm (Hook.) Gray var. flavodiscum Shinners, var. nov. V. L. Cory 56160 (SMU). Field & Lab. 18: 98-99. 3 Apr 1950 Thelesperma filifolium var. intermedium (Rydb.) Shinners, comb. nov. Sida 2 1966 Thelesperma fraternum Shinners, sp. nov. Type: L. H. Shinners 7418 (SMU). Field & Lab. 18: 21-23. 2 Jan 1950. Thelesperma intermedium Rydb. var. ru leo Shinners, var. nov. Type: Shinners 7438 (SMU). Field & Lab. 19-20. 2 Jan 1950. Thelesperma megapotamicum _ Kuntze var. ambiguum (Gray) Shinners, comb. nov. Field & Lab. 18: 23, 2 Jan 1950. Trigridia purpurea (Herb.) Shinners, comb. nov. Sida 1: 295. 25 Jun 1964. Tragia nepetaefolia Cav. var. leptophylla (Torr.) Shinners, comb. nov. South- west. Nat. 6: 101. 1 Sep 1961 Tragia smallii Shinners, sp. nov. Type: R. McVaugh 8461 (SMU). Field & Lab. 24: 37. 15 Jun 1956. Tragia urticifolia Michx. var. tevana Shinners, var. nov. Type: L. H. Shin- ners 111383 (SMU). Field & Lab. 19: 183. 15 Oct 1951. Tridens flavus (..) Hitche. var. aristatus (Scribn. & Ball) Shinners, comb. nov. Rhodora 56: 27-28. 17 Mar 1954. Tridens flavus (L.) Hitche. var. chapmanii (Small) Shinners, comb. nov. Rhodora 56: 27. 17 Mar 1954 Tridens muticus (Torr.) Nash var. eiongatus (Buckl.) Shinners, comb. nov. h 56: 28. 17 Mar 1954 Valerianella florifera Shinhers. a nov. Type: E. Whitehouse 20889 (SMU). Field & Lab. 21: 93. 12 Jun 195 Verbena pulchella Sweet var. staweliata (Troncoso) Shinners, comb. nov. Sida 2: 266. 6 Apr 1966, Verbena pulchella Sweet var. gracilior (Troneoso) Shinners, comb. nov. Sida 2: 266. 6 Apr 1966 Verbena subincana (Troncoso) Shinners, comb. nov. Sida 2: 266. 6 Apr 1966. Verbesina walteri Shinners, nom. nov. Sida 1: 253. 25 Jun 1964. Vernonia marginata (Torr.) Raf. var. tenuifolia (Small) Shinners, comb. nov. Field & Lab. 18: 25-26. 2 Jan 1950. Vernonia vulturina Shinners, sp. nov. Type: J. Reverchon 404 (field no.) 170 (MO). Field & Lab. 18: 25. 2 Jan 1950. Viburnum cassinoides var. angustifolium (T. & G.) Shinners, comb. nov. Southwest. Nat. 1: 91. 22 Jan 1957 Vicia leavenworthii T. & G. var. occidentalis Shinners, var. nov. Field & Lab. 16: 22-23. 1 Mar 1948. ae ludoviciana Nutt. var. laxiflora Shinners, var. nov. Field & Lab. 16: 9-26. 1 Mar 1948. Vicia pinta Nutt. var. sou (T. & G.) Shinners, comb. nov. Field & Lab. 16: 23, 25. 1 Mar 1948 Vicia ing i ae Dictr. f. iependi (S. Wats.) Shinners, comb. nov. Field & Lab. 16: 21-22. 1 Mar 1948. Vitis mustangensis Buckl. var. diversa (Bailey) Shinners, comb. nov. Field & Lab. 19: 182. 15 Oct 1951 Warea auriculata Shinners, sp. nov. Type: M. L. Singeltary, s. n., 25 Sep 1937 (DUKE). Sida 1: 105-106. 23 Nov 1962. Wissaculy grandifolia Ii. G. Baker ex Rusby var. brevipedunculata (R. E. ries) Shinners, comb. nov. Sida 1: 106-107. 23 Nov 1962 Wissadula grandifolia Kk. G. Baker ex Rusby var. macrantha (R. E. Fries) Shinners, comb. nov. Sida 1: 106-107. 23 Nov 1962. Xanthocephalum amoenum Shinners, sp. nov. Type: Lindheimer 422 (SMU). Field & Lab. 19: 77. 1 Jun 1951. pea a amoenum Shinners var. intermediwm Shinners, var. nov. Type: V. L. Cory 50745 (SMU). Field & Lab. 19: 77-78. 1 Jun 1951. Xanthocephatum arizonicum (Gray) Shinners, comb. nov. Field & Lab. 18: 28. 2 Jan Xanthocephalum digynum (Blake) Shinners, comb. nov. Field & Lab. 18: 2 Jan 1950 Xanthocephalum discoideum (Gray) Shinners, comb. nov. Field & Lab. 18: 2 Jan ae dracunculoides (DC.) Shinners, comb. nov. Field & Lab. 1 Xanthocephalum glutinosum (Spreng.) Shinners, comb. nov. Field & Lab. Xanthocephalum grande (Blake) Shinners, comb. nov. Field & Lab. 18: 28. 2 Jan 1950 Xanthocephalum longipappum (Blake) Shinners, comb. nov. Field & Lab. 18: 29 50 Xanthocephalum microcephalum (DC.) Shinners, comb. nov. Field & Lab. 29. 2 Jan 1950. Xanthocephalum ramulosum (Greene) Shinners, comb. nov. Field & Lab. Jan : Xanthocephalum sarothrae (Pursh) Shinners, comb. nov. Field & Lab. 18: 29. 2 Jan peel be sphaerocephalum (Gray) Shinners, comb. nov. Field & Lab. 18: 29. 2 Jan 1950 171 Xanthocephalum sphaerocephalum ee Shinners var. eriocarpum (Gray) Shinners, comb. nov. Field & Lab. 18: 29. 2 Jan Xanthocephalum tenue (Greene) ce comb. nov. Field & Lab. 18: 29. 2 Jan 1950. Xanthocephalum texanum (DC.) Shinners, comb. nov. Field & Lab. 18: 28. Ximenesia nana (Gray) Shinners, comb. nov. Field & Lab. 18: 30. 2 Jan 1950. Yucca ee: Shinners, sp. nov. Type: H. A. Freeman, s. n., 6 Jul 1950 (SMU). Field & Lab. 19: 168-171. 15 Oct 1951. Yucca ee Trel. var. paniculata (McKelvey) Shinners, comb. nov. Field & Lab. 24: 37. 15 Jun 1956. Yucca necopina Shinners, sp. nov. Type: L. H. Shinners 20102 (SMU). Spring Flora of the Dallas-Fort Worth Area, Texas, pp. 408-409. 3 Apr 1958. Zanthoxylum parvum Shinners, sp. nov. Type: R. McVaugh 7890 (SMU). Field & Lab. 24: 19-20. 15 Jun 1956. INDEX TO GENERIC NAMES The generic names appearing in the publications of Lloyd H. Shinners are indexed alphabetically. The numbers (year) refer to the citations in the section on ‘‘Publications.”’ Abies 1956c Acerates 1950f, 1964m Achaetogeron 1946a, 1949r Achillea 1964n Achyropappus 19510 Achyrophorus 1951d Acinos 1962n Actinea 1951g Actinella 1951g Actinochloa 1954f Actinomeris 1951g, 1964e Adelia 1950j, 1959b Adenostyles 1950¢g Agapostemon 1957i, 1957] Agastache 1956v, 1962cc Agave 1951r, 1951s, 1966b, 1967e Aglitheis 1957b Agoseris 1964n Agropyron 1941c, 1954f Agrostis 1948c, 1954f, 1962t Aira Albizia 19571 Alchemilla 1956n Alhagi 1948c Alibertia 1966b Allionia 1951t Allium 1951j, 19521, 1955¢, 1956v, o/D Alopecurus 1956b, 1962t Alsine 1949k, 1962k, 1962x Alsinopsis 1949k, 1962k Amaranthus 1964a Amblyopappus 1951o Ambrosia 1949q Amellus 1950c, 1951f, 19641 Amorpha 1949i, 1955g, 1956v pike aac ue 1964m 5 1946a, 1950b, 1951f ee ae 1964m Amphitecna 1961d Amsonia 1951m Anacharis 1956q Anantherix 1954h, 1964m Andrena 19571 Andropogon 1941c, 1954f, 1956s drosaemum 1956q 172 Aneilema 1962v Anisostichus 1961d Antennaria 1946a Anthophora 1956cc, 1957) Aphanostephus 1946c, 1946d, 1947a, 949r, 1951c Aphora 1956m Apis 1957i, 1957) Aplopappus see Haplopappus Apogon 1947b Aptilon 1947b Arabidopsis 1958e Aragallus 1957e Arenaria 1949k, 1962k Arenbergia 19570 Argemone 1958b Argythamnia 1956m Aristida 1940a, 1940b, 1954f Arkezostis 1957d Arnoglossum 1950¢ Artemisia 1955g, 1964n Asclepias 1949m, 1950f, 1954h, 19541, 196 4m Asclepiodella 1964m Asclepiodora 1950f, 1954h, 1964m Ascyrum 1956q Astephanus 1964m Aster 1941d, 1942, 1943d, 1944a, 1945, 1946a, 1946b, 1946d, 1949c, ge 9510, 1951p, 1958k, 1956e, 1956ff, 1 1961f, 1969h Astragalus 1948c, 1949i, 1949j, 1951a, 19538n, 19541, 1955g, 1956v, 1957e Astranthium 1946a, 1946c, 1946d, 1949r, 1950q, 1966c Athanasia 1949q, 1964e Atropa 1962h Aureolaria 1962e Azalea 1962f Bahia 1949b, 19510, 1964n Baptisia 1951s Bartonia 1955g, 1956v Basistelma 1964m Bellis 1946a, 1949r Berendtia 1959e Berendtiella 1959e Berthelotia 1967d Bidens 1950a Bifora 1961f Bigelowia 1971la Bignonia 1961d Bilamista 19570 Bilderdykia 1967b Biventraria 1964m Boltonia 1946¢, 1956e Bombus 195 Bonamia ae 1964¢, 1969d Bothriochloa 1956s Bouchetia 19638¢ Bourdonia 1946a, 1949q Brachyactis 1946a Brachychaeta 1946a Brachyris 1950b Brassica 19500 Bravoa 1966b Brazoria 1953] Breweria 1962d, 1969d Breweriopsis 1962 Brickellia 1946e, 1949q, 1961f, 1964n, 1968b, 1971b Bromus 1940b, 1954f Browallia 1963c Brunnichia 1967a Bupleurum 1967f Cacalia 1950g Cactus 1955g, 1956v Cakile 1952b Calamagrostis 1943c Calamintha ss 1962p, 1962r Calamovilfa 1 Callisteris ] Calylophus 19641 Calymenia 1955¢g Calymmandra 1951k, 1964d Calyptocarpus 1950a, 1962t Calystegia 1968a Camassia 1956v, 1956ff Campsis He); 1961d 2g a. 1949b, 1958c, 1958d Carex 1954c, 1954f Carthamnus 1958d Ceanothus 1951b Cedronella 1962cc 1956r Centaurea 1951o, 1953g, 1958d, 1964n Centauridium 1950q Centaurium 1950m Centrocarpha 1949c Cephalobembix 19510 62q Cerastium te 1962t, 1966e Ceratina 1957i Ceropegia 1965f Chaetanthera 1946a Chaetopappa 1946a, 1946b, 1946c, 1 , 1949r, 1959c, 1964m Chamaesyce 1949e, 1949q, 1952a, 19561, 1969c Cheiranthus 19 Chilopsis 1956cc, “19614 Chlora 19570 Chloris 1973 Chondrosium 1954f Choretis 1951i Chrysanthemum 1946d, 1950a Chrysopsis 1946b, 1946c, 1948b, 1950q, 1951f, 1956v, 1969f Chrysothamnus 1950c, 1964n Chytra 1962e Cirsium 1949b, 1949q, 1951g, 1951s, 1954f, 1957i, 1964n Cissus 1964p Clinopodium 1962c, 1962n os ~l eo Cnicus 1949b Coleosanthus 1946e Colletes 19571 Collinsonia 19620 Collomia 1963b Commelina 1962v Conradina 1962p, 1962q, 1962r, 1964¢ Convolvulus 1957i, 1962b, 1962d, 1965d, 1968a Conyza 1946a, 1949n, 1949q, 1950d Coreopsis 1950a, 1950i, 1962t Corethrogyne 1949c Corydalis 1950e Cosmidium 1950a Cosmos 1950a Cotula 1949r, 1949s Cracca 1962m Crepis 1949b, 1952e, 1955f, 1956bb, 1964s, 1966f Crescentia 1961d bles 1946e, 1964k Cryptantha 1969e Cryptostegia 1964m Cryptotaenia 1962¢ bia 1947 Cynanchum 195le, 1952f, 1964m, 1965f Cynodon 1962t Cynthia 1947b Cyperus 1952e, 1954d Dahlia 1950a Dalea 1946e, 1949i, 1949j, 1950b, 1953m, 19530, 1954f, 1955g, 1956v Danthonia 1954f Dasystephana 1962bb Datura Delphinium 19491, 1951s Descurainia 19490 Diadasia 1956cc, 1957i Diallosteira 19620 174 Dianthera 1957] Ericameria 1950b, 19510 Dianthidium 19571 Erigeron 1946a, 1946c, 1946d, 1947a, Dianthus 1969b 1949n, 1949q, 1949r, 1951f, 1951o, Diaperia 1951k, 1964d 1953k, 1962v, 1964n Diatrema 1962b Eriochloa 1956e Dicerandra 1962q, 1962r Eriogonum 1954i, 1955¢, 1956v Diceratosmia 1957i Friophyllum 1949r Dichaetophora 1946a, 1946c, 1946d, Erythraea 19570 1949r Hschenbachia 1949n Dietaria 1950c Fucephalus 1946b Digitaria 1957a, 19640 Eupatorium 1946e, 1964n Diplacus 1959e Euphorbia 1949e, 1949q, 1952a, 1956], Diplopappus 1941d, 1946b, 1964i 1962t, 1969¢ Diplostelma 1946a Eustoma 19570 Distasis 1946a Eustylis 1964h Ditaxvis 1956m Kuthamia 1946a, 1950b, 1951p, 195$k Doellingeria 1941d, 1953k Evax 1951k, 1964d, 1964n Doxantha 1961d Evolvulus 1962d Dracocephalum 1949h Ixvolobus 1964m Drosera 19621 Eyselia 1949r Dupratzia 19570 Eysenhardtia 1949i Echinacea 1949c, 1964n Facelis 1949q Eichinocereus 1954f Fagara 1956¢, 1957f Echinochloa 1954f Festuca 1940b, 1956b Edisonia 1964m Filaginopsis 1951k, 1964d Figletes 1946c, 1946d, 1947a, 1949r, Filago 1951k, 1953g, 1964d, 1964n 1949s Fimbristemma 1964m Michhornia 1962u Fischeria 1964m Mlaeagnus 1955¢ Florestina 1949b, 1952e Elephantopus 1951¢ Forcipella 1962w Elodea 1956q Forestiera 1950j, 1956aa, 1959b Elodes 1956q Franseria 1949q Elymus 1954f Froelichia 1954i Emilia 1950¢ Funastrum 1950f, 1964m Enallagma 1961d Eincelia 1964n Gaertneria 1949q Enslenia 1950f, 1964m Gaillardia 1956v Enstoma 19570 Galinsoga 19538k Iphedra 1954f Galpinsia 19641 E'picion 1964m Garuleum 1949s Epilobium 1941b Geboscon 1957b Eragrostis 1940b, 1952d, 1954f, 1973 Gentiana 1949p, 19570, 1962bb EHremiastrum 1946a, 1951f Geoprumnon 1956e Eriastrum 1963b Gerardia 1950n, 1962e Gibbesia 1962w Gilia 1963b, 1968¢c Glandularia 1966a Glycyrrhiza 1955¢ Gnaphalium wa 1966f Criegeiies 1951¢, 1953g, 1956v Guara Cee 1946a, 1950b, 1951f, 1951p Gymnosperma 1950b Gymnostyles 1953k Halictus 19571 Haplopappus 1949q, 1950b, 1950c, 1950q d1f, 19510, 1956v, 1964n, 1969¢ Hedeoma 196 §2n Hedyotis 1949p, 1950k, 1953n, 1954i, 1961f, 1963b Hedypnois 1949a, 1950q Heleastrum 1949q, 1969h Helenium 1961f Heleochloa 1940b Helianthus 1950b, 1957i, 1964n Heliopsis 1951g Helleborus 1956ff Hemiachyris 1950b Hemianthus 1964c Hemiptilium 1950b Hemisia 1956ce, 1957] Herniaria 1962w Heteranthera 1955d Heterocallis 1956ff Heterosperma 1950a Heterotheca 1951f, 1956v, 1963a, 1969f Hibiscus 1956ff Himantostemma 1964m Hoffmannseggia 1946e Holcus 1956z Holosteum 1965c 175 Hordeum 1962t Hymenocallis 19511 Hymenopappus 1949q, 1959a Hymenoxis Hyoseris 1947b Hypericum 1956q Hypochoeris 1956bb, 1966f Fiyssopus 1955g Ilex 1962t Inula 1946b, 1950q, 1951f, 1969f Ionactis 1941d, 1946b Tonidium 19511 Ipomoea 1953n, 1961b, 1962b, 1965a, 1965b, 1965c, 1965d, 1965¢ Ipomopsis 1963b Tresine 1962a Ismene 19511 Isocoma 1950b, 1951p, 1955e Isopappus 1946c, 1950b, 1950q, 19510 Iva 1964n Jacaranda 1961d Justicia 1957] Keerlia 1946a, 1946d, 1949q, 1949r Keithia 1962p Kleinia 19502 Knifa 1956q Koeleria 1956b Krigia 1946a, 1947b, 1949q, 1956e, Krynitzkia 1969e Kuhnia 1946e, 1949q, 1953f, 1956e, 1964k, 1964n, 1968b, 1b Kuhnistera 1949i, 19530 Lachnostoma 1964m 176 Lacinaria 1948a, 1951¢ Mamillaria 19552, 1956v Lactuca 1949q, 1955e, 1955¢ Manfreda 1966b, 1967e Laphamia 1949q, 1959¢ Marshallia 1949q Lasioglossum 1956ce, 19571, 1957] Matelea 1950f, 1964m Lathyrus 1948a Matricaria 1949r Laxanon 1947b Mazus 1962¢ Leontodon 1947b, 1949a Medicago 1948c, 1956a, 1956w Lepachys 1949] Megachile 19571, 1957j Leptochloa 1940b, 1941¢e, 1954f Melinia 1964m Leptoglossis 1963¢ Melissa 1962n Leptoloma 1941c, 1944b Melissodes 1956cc, 1957i, 1957} Lespedeza 1941c, 1948c, 1956e Mellichampia 1964m Lessingia 1949¢c Melochia 1962t Leucelene 1946a, 1946b, 1946c, 1946d. Melosmon 1971c 1949q, 19510 Menispermum 1954f Leucopsidium 1946d Meriolix 19641 Liatris 1948a, 1946e, 1950c, 1951¢, Mesadenia 1950g 1955e, 1959d Metalepis 1964m Lilium 19552, 1956v Metastelma 1950f, 1951le, 1952f, Limnosciadium 1962t 1964m Limonium 1956t Micheliella 19620 Lianthus 1963b, 1963c Micranthemum 1964c Linum 1957¢ Micromeria 1962n, 1962s Liquidambar 1962¢ Microseris 1955e, 1964n Liquiritia 1955¢ Mikania 1950¢ Lisianthus 19570 Mimulus 1955e, 1959e Littorella 1967¢ Minuartia 1962k Loeselia 1963b Mirabilis 1951c, 1951t, 1954f, 1961f, Lolium 1969b 1963b Lophochloa 1956b Monarda 1958b, 1962c Ludwigia 1964r Monoptilon 1946a, 1951f Lupinus 195381 Monothrix 1959c Luthera 1947b Muhlenbergia 1940b, 1941c Lygodesmia 1949p, 1950b, 1953n, Myriactis 1949s 954i Lyonia 1964m Nabulus 1953k Lysianthus 19570 Narcissus 19511 Lythrum 1958a, 1956e, 1957i Nemostylis 1964h Nezera 1957¢c Macbridea 1962n Nierembergia 1963c Machaeranthera 1946c, 1949c, 1949q, Nomada 19571 1950c, 19641, 1964n Nothocalais 1964n Madia 1955e Nothoscordum 1957b Malva 1955g, 1956v Malvastrum 1953f Odontonychia 1962w Odontostephana 1950f, 1964m Oenothera 1953a, 1955g, 1956v, 19641 Oldenlandia ae Olearia Ophryosporus 1946e Oplismenus 1954f Orbexilum 1951a Oryzopsis 1940b Osmia 19571 Othake 1949b, 1952e calis 19560, 1957n Oxybaphus 1951t Oxypetalum 1964m Oxypterix 1964m Palafoxia 1949b, 1952e Pancratium 1951i Panicum 1940b, 1944b, 1954f q Paspalum 1954f, 1962t Pattalias 1952f, 1964m Paulownia 1961d Pediomelum 195la e Penstemon 1955g, 1956v Periploca 1964m Perityle 1959c Perizoma 1962h Persicaria 1957n, 1967b Petalostemum 1946e, 1949i, 1949), 19530 Petrorhagia 1969b Pharbitis 1962b, 1965a, 1965d Pherosphaera 1957f Pherotrichis 1964m Philibertella 1950f, 1964m Philibertia 1950f, 1964m Phlox 1951m, 1953n, 1956e, 1961a, 19638b, 19638c Phyla 19571 Physalis 1951s, 1962h Physostegia 1949h, 1951q, 1953), 956 Piloblephis 1962n, 1962r Pinaropappus 1951d Pinus 1956c Pityopsis 1951f Plantago 1950g, 19501, 1967c ee 19491, 1949s , 1967d Poa 1948c, 1954f, 1956b, 1962t Podostigma 1964m Polemonium 1963b Polianthes 1966b, 1967e Polistes 19571 Polyactidium 1946c Polyanthes see Polianthes Polygonum 1957n, 1967b Polypappus 1967d Polypteris 1949b, 1952e Porophyllum 1950g Potamogeton 1955d Potentilla 1955c Prenanthes 1950b, 1953k, 1964s Prionopsis 1949q, 1957i, 1964n Prochnyanthes 1966b Prunus 1956x Pseudobravoa 1966b Pseudotsuga 1956c Psilactis 1949c, 1964n Psoralea 1946e, 1949i, 195la, 1955¢ Psoralidium 195la Ptiloria 1950b Puccinellia 1940b Pycnanthemum 1962¢ Pycnothymus 1962n Pyrethrum 1949r 178 Pyrrhopappus 1951g, 1958e, 1953n, 1956bb, 19571 Pyrrocoma 1949q Quamoclit 1965a, 1965¢ Quamoclita 1965¢ Quercus 1956e, 19561, 1962t Rajania 1967a Ranunculus 1950h, 1960a, 1962t, 62y Ratibida 1949], 1955g¢, 1956v Rayania 1967a Reverchonia 1952a Reynoutria 1967b Rhinanthus 1962e Rhododendron 1961e, 1962f Rhus 195th Ridan 1964e Rivea 1965a Rosa 19551 Rosmarinus 1962p Rothrockia 1964m Roulinia 1950f, 1964m Rouliniella 1950f, 1964m Rubus 1952e, 1954b Rudbeckia 1949c, 1949}, 1949q, 1950b, 19552, 1956v Ruellia 1953p Runyonia 1966b Rygchium 19571 Salpichroa 1962h Salvia 1953¢c, 1956e Sarcostemma 1964m Satureja 1962n, 1962s Schizonoltus 1964m Scehkuhria 1949b, 19510, 1964n Schmaltzia 1951h Scilla 1956v Scorzonera 1947b Scutellaria 1962dd, 1968a, 1964b Seddera 1962d Selloa 1950b Senecio 1962t, 1964n Serinia 1946a, 1947b, 1949q Serophyton 1956m Serratula 1948a Seuleria 1964m Sibara 1962t Sida 1953f, 1973 Sideranthus 1946c, 1950c, 1955¢, Steglingia 1954f Silphium 1941c, 1949], 19510, 1954f Silybum 1958d Sinapis 19500 Stphonychia 1962w Sisyrinchium 1948¢e, 1951p, 1951s, 1957g, 1962], 1962t, 1966f Smilax 1962t Solandra 1961d Solanum 1962ece Solidago 1941c, 1941d, 1946a, 1946d, 949q, 1950b, 1950c, 1951c, 19510, 1951p, 1958h, 1953k, 1956k, 1964n Soliva 1953k Sophia 19490 Sophora 1955g, 1956v Sorghastrum 1954f Sorghum 1956z, 1962t Soxmantorum 1953n Sphaeralcea 1955g, 1956v, 1964q Sporobolus 1940b, 1941c, 1941d, 1954f i1f Stachys 19622, 1963a Stanleya 1962z Starkea 1950c, 19641 Stellaria 1949k, 1962k, 1962x Stenolobium 1961d Stenotaphrum 1962t Stephanomeria 1948c, 1950b Stevia 1949b Stiburus 1956b Stylisma 1962d, 1969d Swartzia 1961d Symphyopappus 1946e Synosma 1950g Tabebuia 1961d Tamarix 1948c, 1957h Tephrosia 1962m Tessaria 1967d Tetraneuris 1951¢ Thelesperma 1950a, 1950i, 1964n, 1966d Thuraria 1955g, 1956v Thurovia 1964¢ Thymbra 1962n Phymus 1962n, 1962s Townsendia 1950q Tradescantia 1962v Tragia 1951v, 1956j, 1961c Tragopogon 1947b, 1948c, 1949a Trichogonia 1946e Tridens 1954f Trifolium 1954e, 1962y Trisetum 1940b Triticum 1954f Troximon 1947b, 19552, 1964n Tunica 1969b Turbina 1965a Unamia 1941d Ungnadia 1954f 179 Uralepis 1954f Urananthus 19570 Valerianella 1953d Vaseyochloa 1964g Verbena 1966a, 1966f Verbesina 1950b, 1964e, 1964n Vitis 195lu, 1955g, 1956v Wahlenbergia 1957a Warea 1962z Wedelia 1951t Wedeliella 1951t Wissadula 1962aa Xanthisma 1950q Xanthium 1955e Xanthocephalum 1946d, 1949r, 1950b, 1951g, 195 Xanthoxylem see Zanthoxylem Xerobius 1949r Ximenesia 1950b Xylocopa 1956cc, 19571 Youngia 1962g, 1964s Yucca 1951r, 1955¢g, 1956h, 1956v Zanthoxylem 1956g, 1957f Zosterella 1955d PUBLISHED TRIBUTES oe DONOVAN S. 1971. Lloyd Herbert Shinners—a portrait. Brittonia -10: as. WILLIAM F. 1971. Lloyd Herbert Shinners 1918-1971. Sida 4: ROWELL, CHESTER M., JR. 1972. Lloyd Herbert Shinners, 1918-1971. In Memoriam. Texas J. Sci. 24: 266-271. VOSS, EDWARD G. 1971. News of Botanists. Michigan Botanist 10: 95. BY ANY OTHER NAME... Lloyd Shinners enjoyed classical music, mystery books, and poetry. He exchanged poetry with many of his friends and in one instance his poetic endeavors became intermingled with his journalistic pursuits. For his new journal, he selected a simple one-word title for (1) simplicity in reference citations, and (2), one that would be familiar to botanists all over the world. The generic name of SIDA (Malvaceae) was chosen as it fit these criteria. Upon receiving the first two numbers of the new journal, Professor Kwan penned some lines and a ‘‘retaliation’’ was immediately written. Reproduced copies of Shinners’ ‘‘retaliation’’ were inserted (loose-leaf) in the next issue of the journal. With Professor Ewan’s permission, the original is reproduced along with Lloyd Shinners’ response (Fig. Librarians, aware of the ‘‘retaliation’’ with the two lines referring to the abbreviation of SIDA, yet unaware of the generic name, have cataloged the journal in one library as an abbreviation. In another hbrary, however, the journal is cataloged as a one-word title. Inquiries are still_received from librarians seeking to determine the correct disposition. WFM Fig. 1. ~Holograph and typed response _of Joseph Ewan and Lloyd Shinners respectively. ' SIDA 5(3): 180-181, 1973. PG ek 3 From Subject: Remarks: AES TULANE UNIVERSITY Department of Botany Date 2¢ Peprenmherr IH uv ‘ss ce ae Be Sell L/tWe Loyd Hoeter Onde, ae ee pares “ < cathe, Ache, A Hi dhuk Ww ye Lily Aad \ cael et & conpel fret Jtasd tae ba Gove fr ava LL / Hoy See Zz, hius- Pro aj oe pts a er a tw linc. pin Prd erp Cong - Gs Aocln : fro Bet Preterruceatle ck tetil fod / HERBARIUM SOUTHERN METHODIST UNIVERSITY Daas 22, TEXAS LINES PENNED IN RETALIATION FOR J. EWAN'S DUBIOUS VERSES PENNED ON THE OCCASION OF HIS SUBSCRIBING TOS IDA ten, my children, gee sls shall hear r! Two issues were mailed - no less would do To hold its omnium-gatherum chatter On botany culture - unprintable matter] From Siberia to Bolivia plants now bear N. : e e en Please don't say that "It's. Gree me" You've two meanings to choo Shinners' Infamous Damnable Activities Shinners' Interesting Delightful Activities Is worse to come? Just wait and see! 26 Nov. 1962 se bu i you see: I8T 182 CHLORIS TRUNCATA R. BR. (GRAMINE Ads), AN AUSTRALIAN WOOL- WASTE ADVENTIVEID JN SOUTH CAROLINA.—The recent report of the African Wragrostis pliing Nees as a wool-waste introduction in South Caro- line makes it desirable to record another unusual alien, My thanks are due to Harry I. Ahles for showing me during April 1966 a large series of speci- mens from South Carolina localities where wool-waste had been discarded, and to Mr. Ahles and Dr. A. EF. Radford for a number of duplicates later sent to SMU in exchange. Among them were numerous species common in Texas, including at least one endemic to the state (Sida Lindheimer En- gelm. & Gray). An exotic-looking Chloris with elongate, digitate racemes and rather long-awned lemmas vas identified by matching with an Austra- lian specimen at SMU ofC. tray¢atasR. Br., received from the National Her- barium of New South Wales, Sydney: N.S.W., Narramine, R. Helms, XI. 1892 (Nat. Herb. N.S.W. No. 31321). The identification was later confirmed by plates and deseriptions in vols. 1 and\2 of Australian Grasses (Burbidge, 1966 and 1968). Like the African@ragrosis plana; this is apparently new to Nerth America. Three specimens are deposited at SMU. SOUTH CAROLINA. Berkeley Co.: waste ground around the Santee Wool Combing Mill, James- town on S. C. state route #45, Harry E. Ahles 25881 with John G. Haesloop, 20 May 1957. Florence Co.: waste ground around the Wellman Wool Combing Mill, north of Johnsonville on state route HALL Ahles 42889, 13 Jun 1958, Ahles 46959 with J. Haesloop, 21 Jul 1958.5 Lowa H_ Shinners: REFERENCES BURBIDGE, NANCY T. 1966, Eine Australian gra 1: 136, plate 54; 2: 144, plate 57. Angus & Robertson, Ltd., elbows ne. eee ription an plate repeated in volume 2.) 7 ORAK, JOSFF and FRANT IS K KUHN. 1966. E Bees Pflanzen in Areal der Wollspinnerei “Mosilana” in Brno (Brinn). Pralia 38: 327-332. (In Crech with German abstract. a - W. and R, I. LONARD. 1970. Eragrostis plana in South Carolina. Rhodora 72: HAY Ww a IDA os - FORGE Spire wor DRUCE. 1919. The Adventive Flora of Tweedside. Arbroa . Buncle & Co. 296 pp. LOUS. ; Y,. U3 cE. Mei ; census list of wool aliens found in Britain: 1946—1960. Proc. t. Soc. British Isles 4: 221-247. PRONST Ses 1949. Wolladventiyflora Mitteleuropas. Vogt-Schild AG, Solothurn. p) Be rosa: AU, ERNEST. 1953, Rhodora, Index to Volumes 1—50. p. 414: 9 titles with wool-waste or woolwaste (EDITOR’S NOTE: This is the last manuscript of Lloyd H. Shinners to be published.) SIDA 5(3): 182, 1973. SHINNERSOSERIS GEN. NOV. <&, SPENCTR TOM Department of Biological Sciences University of Illinois at Chicago Circle Chicago 60680 My recent work onCLygodesmtg revealed several taxa that, although they genus (Tomb, 1970a; 1970b; 1972). Lygodesmia rostrata (Gray) Gray is another species that does not belong in Lygodesmia. I have been reluctant to establish yet another small genus in the North American Cichorieae, since there are at present seven monosspie or ditypic genera in subtribe Stephano- meriinae ts ORES however, has a unique combination of characters and is difficu o place in an established genus. Furthermore, retention of this species in Lygodesmia would in my opinion make the genus polyphyletic. Therefore, the establishment of a new genus is required. The principal differences between the new genus, Ghinnersoseris (L. rostrata), and Lygodesmia are summarized in Table 1. TABLE I. PRINCIPAL DIFFERENCES BETWEEN SHINNERSOSERIS D LYGODESMIA Shinnersoseris _ ~- Lygodesmia Lower leaves opposite Lower leaves alternate Ligules 0.5-0.7 cm long, ca 1 mm Ligules 1.0-2.5 cm long, 0.4-1.2 em ide, erect, obscure wide, reflexed, showy Style branches short, ca 0.3 mm Style branches 0.6-4.0 mm_ long ng, white (most species have style branch- es 2.0-2.5 long), pink-lavender, purple Achenes 8-10-sulcate, dark brown, Achenes variously sculptured (ne- narrowed and scabrate below the ver 8-10-sulcate) pallid, ab- and summit, ab- and adaxial surfaces adaxial surfaces different (except similar in L. juncea) Pollen echinate, mean equatorial — echinolophate, mean diameter 44m, tetrads isobi- ial diameter 46-654.m, tetrade late ral nee dra Base chromosome number, 7—6 Base chromosome number, «=9 /CHROMOSOME STUDIES Stebbins, Jenkins, and Walters (1953) reported the first chromosome count of $Shinnersoseris rostrata) (Lygodesmia rostrata) as 2n—12. Additional SIDA 5(3): 183-189, 1973, 184 chromosome counts from 5 populations confirm the original count (Table I). The meiotic chromosome counts were made by using the procedures outlined by Turner and Johnston (1961). The karyotype of S. rostrata (determined from root tip squashes using a $0 min PDB pretreatment and aceto-orcein stain) consists of 6 chromosome pairs, all of almost the same total length. One pair is metacentric, three pairs are submetacentric, and the remaining two pairs are acrocentric (Fig. 3D). Satellites and secondary constrictions were not observed. TABLE II. CHROMOSOME COUNTS OF SHINNERSOSERIS ROSTRATA Locality & voucher | : n es NEBRASKA. Cherry Co.: Near Big Alkali Lake. Tomb 557. 6 a eee Cherry Co.: Neb. Spur 483, 9 mi SW of U.S. 83. Tomb 56 NE arg Cherry Co.: Entrance Valentine Wildlife Refuge. Tomb 561. ) OMING. Goshen Co.: 12 mi N of Lingle. Tomb 581. 6 12 WYOMING. Goshen Co.: 4 mi E of Bont Laramie. Tomb 583. 6 pany NOL OGY The pollen of Shinnersoseris r stratu is echinate, tricolporate (Fig. 1). Al- though ridges bearing spines aré clearly present, the highly ordered echi- nolophate pattern of Lygodesmia and most of the Cichoricae is never ap- proached in Shinne is. Sectioned exines of S. rostrata reveal a strati- fication pattern fiike Aer found in the Stephanomeriinae (Fig. 2). The elec- tron-dense extexine consists of short spines, a perforate tectum, columellae, and a thin foot layer. A cavus separates the columellar bases from the foot layer. Below the foot layer is the less electrondense endexine. A description of ite pollen of S. rostrata follows: equatorial diameter 43.7 pm (41.2-47.9 pm); polar diameter 38.2 pm (35.9-39.7 jem); spine length 1.7 pm (1.5-2.3 mn); grains echinate, spherical in polar view, almost spherical in equa- torial view; 9 lacunae (3 poral and 6 abporal); polar thickenings large. The pollen of S. rostrata is produced in isobilateral tetrads. Tetrahedral tetrads are found in all of the Stephanomeriinae examined to date. Shinners- oseris produces only 30-50 pollen grains per anther. This low pollen produc- tion and the nature of the flowers suggest that S. rostrata is self-fertile. ' Mean onne followed by the ranges in parentheses based on measurement of 100 grains from Tomb 557 Fig. 1. - Seanning electron micrograph of acetolized_S. _ rostrata pollen. Note abporal lacuna above germ pore. Fig. 2. Transmission electron micrograph of S. rostrata exine. B, Foot lay- er; Ca, Cavus; Fin, Endexine. Lines in each micrograph equal 1 fem. 185 186 TAXONOMIC TREATMENT Shinnerseseris Tomb, gen. nov. Herbae annuae. Folia fasciaria, integra, non rosulata, See inferioribus oppositis, foliis superioribus alternis et ee Capitula terminalia, floribus paucis. Involucra cylindracea, 7 ve acteis vere ipalibus ap- leases apicem versus, appendiculis carinoic ideis racteae exteriores ves, 2-seriales, catenin oe osculi ligulati, ligulis lavandu- ae quinquedent: atis, per ares ee rami stylorum breves. Cyp- selae 8 - 10-sulcatae, eae aceae, prope apices abrupte angustatae et scabrellae. ae -serialis, capillaceus, candidus C Shinnersoseris ros rata (G ray) Tomb, comb. nov. Qugod lesmia juncea var/ rostratasGray, Proc. Acad. Philad. 1863: 69. 1863. HOLOTYPE (GH!): Colorado; ‘On fae pins Rare.’ (between latitude 39° and 41°); Sep 1862; Elihu Hall «& . Harbour 347. Isotypes NY! From the aboye latitudinal data, im a pe was probably collected in the sandhill regiow of oe C Lygodesmia rostrat@ (Gray) Gray, Pree. Amer. Acad. 9: 217. 1874. Herbaceous taprooted annual, 1.0-8.5 dm tall (usually ca 3.0 dm tall); stems glabrous, green, sparingly branched and not striate in the basal third, becoming paniculately branched and striate in the upper two-thirds; lower leaves opposite for the first 5-8 nodes, not forming a basal rosette, linear, 6-10 cm long, 2-4 mm wide, entire, glabrous, obscurely 1-3 veined; upper leaves similar to lower except alternate, 6 cm or less in length, reduced to linear bracts below the heads; heads several to numerous, of 8-11 florets; involucre narrowly cylindrical, 1.2-1.5 cm long, 3-4 mm wide; the 7 or 8 principal involucral bracts narrowly linear, 1.6-1.9 cm long at maturity, gla- brous, the midribs obscure, the apices bearing a small keel-shaped appen- dage: outer involucral bracts lanceolate, in ca 2 series, forming a calycu- lum; ligules lavender with white apices, ca 6 mm long, 1 mm wide, erect at anthesis (i.e., not reflexed); corolla tube ca 8 mm long; anthers purple, ca 2 mm long, exserted ca 1.5 mm; style column purple, exserted ca 2 mm above the anthers; style branches white, ca 0.3 mm long; achenes subcy- lindric, 8-10 sulcate, 0.8-1.0 em long, abruptly narrowed and scabrate just below the summit; pappus 6-8 mm long, usually bright white (rarely sordid white), basally connate; sap cream colored; pollen grains echinate, tricol- porate, apertures lacunate, mean equatorial diameter 44 pm; chromosome number, n = 6 (Fig. 3D). DISTRIBUTION Sandy stream banks, sand dur mi and sand hills, growing with Muhlen- _bergim a a & Ss © a) fe) o Q S) REFERENCES STEBBINS, G. L. Jr., J. A. gira aoe M. S. WALTERS. 1953. Chromosomes and phylogeny in the Compositae, tribe Cichorieae. Univ. Calif. Publ. Bot. 26: 401-430 TOMB, A. S. 1970a. Novelties in shoes paso and Stephanomeria (Compositae: Cichorieae). 2: 1970b. A cytotaxonomic study of the aan ee (Compositae: Cicho- rieae). Ph.D. a ba The Univ. a Texas ates eee ment of the genus Prensnibel Rydb. (Compositae: Cicho- ~~ rieae). inaiea on TURNER, B. L. and M . JOHNSTON. 1961. Chromosome numbers in the Compositae. TI. Certain Mexican species. Brittonia 13: 64-69. PUBLICATION NOTES BOTANICAL SURVEY OF THE LAKE MONTICELLO AREA. Wm. F. Maher. 1973. 25 pp. SMU Contrib. in Anthrop. No. 9. $2.00, postpaid. Archae- ology Research Program, Department of Anthropology, SMU, Dallas, Texas, 75275. FLORA OF TAYLOR COUNTY, TEXAS. Wm. F. Mahler. 1973. 247 pp. Paperback. SMU Book Store, Box 759, Dallas, Texas, 75275. $7.25 postpaid. KEYS TO THE EMBRYOPHYTA OF TAYLOR COUNTY, XAS. Wm. F Mahler. 1966. 86 pp. H-SU Bookstore, Abilene, Texas. $3.00. Out of print; superseded by Flora of Taylor County, Texas. KEYS TO THE VASCULAR PLANTS OF THE BLACK GAP WILDLIFE MANAGEMENT AREA, BREWSTER COUNTY, TEXAS. Wm. F. Mahler. 1971. 109 pp. Paperback. SMU Book Store, Box 759, Dallas, Texas, 75275. $3.65 postpaid. REPRINTS. Incomplete sets of Lloyd H. Shinners’ reprints are still avail- able from the SMU Herbarium upon request. SHINNERS’ SPRING FLORA. Lloyd H. Shinners. Second Edition, 1972, . 014 pp. Paperback. Prestige Press, 4829A Gretna, Dallas, Texas, oir pan postpaid. SIDA, CONTRIBUTIONS TO BOTANY. Back volumes, 1, 2, and 3, are available at $6 (U.S.) per volume with volume 4 at $8 (U.S.). Current sub- scription price per volume is $8 (U.S.). Subscription rate per year is $6 (U.S.). Publication dates of the issues within each volume are printed on the back of the Title Page which is issued upon completion of each volume. Order from and make checks payable to : SIDA — Wm. F. hler. VIOLETS (VIOLA) OF CENTRAL AND EASTERN UNITED STATES: AN INTRODUCTORY SURVEY. Norman H. Russell. Sida 2 (1): 1-113. 1965. Separate. $1.75, postpaid. Order from and make checks payable to: SIDA — Wm. F. Mahler. WOODY VINES OF THE SOUTHEASTERN STATES. Wilbur H. Duncan. Sida 3 (1): 1-76. 1967. No longer available as a separate. SIDA toscraxy VOLUME 5 NUMBER 4 JULY 1974 CONTENTS A revision of Styrax (Styracaceae) in North America, Central America, and the Caribbean. Gene J. Gonsoulin. 19] Notes on the genus Najas (Najadaceae). Robert R. Haynes and W. Alan Wentz. 259 A new subfruticose Eriogonum (Polygonaceae) from northern Mexico. James L. Reveal. 265 Studies in American Rubiaceae 2. Ayuque, Balmea stormae, an endangered Mexican species. F. R. Fosberg. 268 Herbarium and field studies of Kentucky plants. Il. New state records and rarities. Edward T. Browne, Jr. 27 | Studies of the southeastern United States flora. IV. Oleaceae. James W. Hardin. 274 NOTES. Fatoua villosa (Moraceae) in Florida. 286.—Caperonia palustris (Euphorbiaceae) in Arkansas. 286.—Allium ampeloprasum (Liliaceae) and Trifolium vesiculosum (Leguminosae) in Oklahoma. 286.—Corallorhiza odontorhiza (Orchidaceae) in Louisiana. 287.—Hypo- choeris in Texas, 287. DOCUMENTED PLANT CHROMOSOME NUMBERS 1974: 1, 2. 290 REVIEWS 295 JOHN F. COOKE, JR., 1927-1970. 297 SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. Mahler, SMU Herbarium, Dallas, Texas, 75275, U.S.A. Subscriptions: Li- braries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.) per volume; numbers issued twice a year. ASSOCIATE EDITOR John W. Thieret Northern Kentucky State College Highland Heights, Kentucky © SIDA Contributions to Botany, Volume 5 Number 4, pages 191—300. Copyright 1974 by Wm. F. Mahler A REVISION OF STYRAX (STYRACACEAE) IN NORTH AMERICA, CENTRAL AMERICA, AND THE CARIBBEAN GENE J. GONSOULIN Department of Biology East Tennessee State University Johnson City, Tennessee 37601 The Styracaceae consists of about 12 genera, all small except Styrax, and about 150 species centering in eastern Asia, but with numerous species in the New World, none in Australia, one in the eastern Mediterranean re- gion, and about three in Africa. The characters distinguishing Styrax from other genera in the family are: stamens 10 or more; flowers perfect, in racemes, cymes, or panicles; ovary completely or almost completely su- perior; seeds not winged; filaments free or connate only in the lower part; and fruit drupaceous. Two of the genera, Alniphyllum and Pterostyrax (Chi- na and Japan), are considered most closely related to Styrax. Alniphyllum differs from Styrax in its winged seeds, filaments connate to the top into a tube, and capsular fruit, while Pterostyrax differs in its two-thirds to com- pletely inferior ovary (Hutchinson, 1959) Styrax, a genus of trees and shrubs, is cosmopolitan in its distribution and includes about 120 species, most of which are tropical and subtropical. It occurs in eastern Asia, New Guinea, the eastern Mediterranean region, the Caribbean, South America, Central America, and North America. Ecologic- ally, the species occupy habitats from high mountainous regions to swamps. Economically and horticulturally the genus is of substantial value. Many species are planted as ornamentals either singly on lawns or in hedges and fencerows. The white-flowered, pendulous racemes and spreading habit make these plants a noteworthy addition to any landscape. In addition to the orna- mental value, several species of the Mediterranean region are the source of the aromatic resins styrax and benzoin. This study is a reappraisal of Styrax in North America, Central America, and the Caribbean. The taxonomic status of many of the previously described species is doubtful. Some authors seem to have exploited the genus in order to name new species. This trend is reflected in Standley and Steyermark’s (1940) statement describing Styrax vulcanicola: The most closely related species is the recently ae Styrax mag- nus onde of the Volcan de Tacana, iapas. In t the shape and size of the leaves ee Styrax vulcanicola, but We pubescen nce con- sists of muc arger, relatively ax, soft, spreading hairs, making it fairly certain that two distinct species, as species of Styrax are divided, are represented. SIDA 5(4): 191—258, 1974. 192 These workers appear to have considered it justifiable to use frivolous characters to describe new species because other workers had done so. This trend was set by Perkins (1907), who monographed the genus and caused much confusion by finely and unnecessarily dividing the species complexes. Most species of the United States and Northern Mexico are quite distinct and easily delimited. However, difficulties arise in the southern Mexico and Central American species, primarily the Styrax argenteus complex. One of my main objectives is to clarify these matters. Although this revision is based primarily on morphology, attempts have been made to consider phylogenctic relationships. Approximately 3,500 speci- mens from 16 herbaria were examined. These were augmented by my per- sonal collection of many specimens from southeastern United States and southwestern Texas. Complete citations for all species are given, with the exception of S. amer- icana var. americana and var. pulverulenta and S. grandifolia. No more than five localities per state are given for S. americana var. americana and S. grandifolia. One locality per county is given for S. americana var. pulverul- enta. Specimens most widely distributed in herbaria were selected for cita- tion. A mimeographed list of collections examined may be obtained from me upon request. The following herbaria provided Styrax specimens—the abbreviations are those of Lanjouw and Stafleu (1959): DUKE, F, FSU, GA, GH, LAF, MO, NCU, NY, OKLA, PH, SMU, TEX, UC, US, and VDB. Acknowledgements are gratefully extended to the curators who provided specimens for study. I would like to express my sincere thanks to Dr. Robert Kral and Dr. Dean P. Whittier for encouragement and valuable suggestions. r. Delzie Demaree freely gave his time in providing collections of Styrax, and the late Dr. Lloyd H. Shinners provided valuable references and col- lections of several rare species. Publication costs were met by the Research Council, Kast Tennessee State University. GENERIC HISTORY “Styrax”’ is the ancient Greek name for a fragrant resin used as incense and as medicine and for the plant (Styrax officinalis) from which it was obtained. Accounts of Styrax were first given by Linnaeus (17385) in Systema Naturae. The genus was again treated by Linnaeus in Species Plantarum (1753) and Genera Plantarum (1754). The type species is Styrax officinalis. Cavanilles (1790) described the genus Strigilia from material of Styrax. The type species, Strigilia racemosa, was later transferred by de Candolle (1844) to Styrax as S. racemosa. Ruiz and Pavon (1794) applied the name Foveolaria ferruginea to a Peru- vian species that was later transferred to Styrax by Perkins (1907) under the name S. foveolaria. The generic name Cyrta was applied by Loureiro (1790) to an element of 193 Styrax that was later transferred to Styrax under the name S. agrestis (Lour.) G. Don (Don, 1838). The genus Tremanthes, described by Persoon (1805), included on publica- tion all species of Foveolaria Ruiz et Pavon as well as the type of Strigilia Perkins (1907) included Foveolaria ferrugineum, Tremanthes ferrugineum, and Strigilia racemosa in synonymy under Styrax foveolaria. If these rela- tionships are true, then the name S. foveolaria is incorrect. All four names represent the same taxon, so the name S. racemosa (Cav.) A. DC. has prior- ity since the specific epithet racemosa is the earliest given to this taxon. Rafinesque’s (1817) Florula Ludoviciana, significant as the first localized record of plants in southern Louisiana, was based on Robin’s Voyages dans Vinteriere de la Louisiana, de la Florida et dans les Isles de la Martinique. Robin’s Flore (a section of Voyages) contains descriptions of plants observed by him from Louisiana east to Pensacola, Florida. Many of the plants de- scribed are without adequate names. In trying to correct these errors, and in order to place such species into proper relationships, Rafinesque proposed new names for many of the species. Rafinesque made a number of mistakes since he had to work solely from written descriptions and illustrations and since he was not familiar with the flora of the region. In Florula Ludoviciana he named what he considered a new genus, Adnaria, and its only species, A. odorata. Robin had placed this shrub in the ‘‘Campanulacees”’ under Cam- panula, but Rafinesque recognized that it did not belong to that group and suggested that it was a member of the ‘‘Vaccinia.’’ Camp (1941) visited the area where Adnaria was said to have grown and found and identified the plant as Styrax americana. Camp said that the mistake in identification was caused primarily by Robin because Rafinesque was probably not familiar with S. americana in the field, but Robin listed and described S. americana in his Flore. The generic name Epigenia was proposed by Velloso (1825). Shortly there- after, Schott (1827), transferred all species of Epigenia to Styrax. Styrax benzoin was described by Dryander (1787). Hayne (1829) described a new genus, Benzoin, the type species being B. officinalis. Roxburg (1832), recognized Hayne’s error and listed B. officinalis as a synonym of Styrax benzoin. The most recent generic name associated with Styrax is Darlingtonia, published by Torrey in 1851. Torrey(1853) later stated, ‘“‘and a California plant, on imperfect specimens of which, I had recently indicated a genus under this name, proves to be only a species of Styrax.”’ Torrey’s combi- nation, Darlingtonia californica, was conserved in this publication and applied to a member of the Sarraceniaceae. The new styrax he named S. californica. During the later part of the 19th and the early 20th century numerous works treated Styrax from areas of Norh America, Mexico, and the Carib- bean. In addition to large treatments such as those by de Candolle (1844), Bentham and Hooker (1876), Gurke (1897), and Baillon (1892), which all 194 treated the Styracaceae, the most comprehensive and recent work is the monograph of Styrax by Perkins (1907). Perkins confused the taxonomy of the group by describing numerous new species many of which have proved not to be ‘‘good”’ taxa. MORPHOLOGY Life Cycle. The following general life cycle was formulated during my study from observations made on Styrax grandifolia and S. americana (southeastern United States). The winter buds unfold from late February through carly March. The in- ternodes elongate and the leaves expand and reach full size in late Maren or April. At the same time, the inflorescence develops. Before the winter buds unfold, the synsepalous calyx and the separate corolla lobes and stamens develop more or less simultancously. As the leaves reach full size, the corolla begins to project from the calyx. The aestivation of the corolla lobes may be imbricate or valvate depending on the subgenus. The gynoecium has developed by this time, and the lumen is expanded by a mass of tissue that later becomes the placentae. The flowers generally open about mid-April to mid-May. The period of flowering is normally 2-3 weeks for temperate species and somewhat longer for tropical and subtropical species. The flowers have a faint sweet odor and attract a variety of insects of which various bees seem the most active pol- linators. Although the stamens have introrse dehisence, they are reflexed considerably, this causing any visitor to be dusted with pollen. The flowers of most species remain open 3-5 days, after which the corolla lobes lose their turgidity and corolla and stamens are shed. Within a week after the fall of the corollas many ovaries are visibly swollen. Dissection of an ovary during the early stages of swelling reveals usually one but sometimes two developing ovules. The ovary or young fruit soon ex- pands so as to rupture the calyx and extend beyond it. The fruit ripens from July through September and eventually opens by one or more sutures. The seed coat is primarily a mass of stone cells forming a hard and brittle shell. The cotyledons are large and elliptic. There is a massive hypocotyl, but the plumule is scarcely detectable. The seeds often germinate in close proximity to the parent plant, resulting in dense stands. Habit. The storaxes are shrubs to medium-sized trees with a rather loose, spreading habit. The numerous, white, fragrant flowers, borne either singly or more often in few-flowered racemes, are followed by drupaceous fruits. Species of Styrax native in the United States are generally large shrubs, occasionally reaching tree size. Styrax grandifolia of the eastern United States often has a rather flabellate branching system and a rounded crown. The southwestern Texas and California species are medium-sized shrubs, rough in outline due to a somewhat irregular branching pattern. South through Mexico the species are progressively larger. Styrax argenteus of southern Mexico and Central America reaches a height of 30-85 ft., having 195 a rather freely branched crown and spreading habit. One of the largest species, S. glaber, which occurs in the Caribbean, reaches a height of 40 ft. but still has a somewhat spreading habit. The storaxes are generally understory shrubs or trees in the eastern United States but are often exposed shrubs in the southwestern and western United States. Those reaching tree size in Mexico and Central America often form a substantial part of the forest canopy or forest edge. Stem. The stems have smooth tight bark to rough flaky bark and may be glabrous to densely pubescent. Cork forms in the inner cortex, which includes a closed or slightly interrupted ring of sclerenchyma. Secondary phloem includes sclerenchymaious elements. The xylem forms a continuous cylinder traversed by narrow rays usually two to four (six) cells in width, markedly heterogenous. In the primary wood the vessels are narrow and scattered, with highly oblique, scalariform perforation plates. The fibers have bordered pits and are of medium length to moderately long. Parenchyma is diffuse and in regular uniseriate lines. The pith is solid. There are intercellular sec- retory cavities, containing balsamic resins, in the phloem, rays, and wood. Also, crystals of calcium oxalate are present in unlignified tissues (Metcalfe and Chalk, 1950). Mach leaf is supplied by a single vascular bundle, which is accompanied by a single gap in the stele (Sinnott, 1914). Pubescence. Pubescence in Styrax is generally in the form of stellate trichomes, these differing widely in size and structure. The arms of the tri- chomes vary from especially long and straight as in S. grandifolia (Fig. 4d) and S. argenteus var. hintonii (Fig. 11b), to very short and stout as in S. americana var. pulverulenta (Fig. 3d) and S. argenteus var. ramirezii (Fig. Four species possess lepidote trichomes: S. portoricensis (Fig. 15d), S. glaber (Fig. 16d), S. ochraceous (Fig. 14d), and S. conterminus (Fig. 12d). This type of trichome, very distinctive, is found predominantly on the lower leaf surface in these species. In S. obtusifolia of Cuba the lower leaf surface is covered with a light, fine tomentum mixed with scattered lepidote tri- chomes (Fig. 13d). Some relationship can be seen in the trichomes of S. platanifolia var. stel- lata (Fig. 5f), S. texana (Fig. 6g), and S. youngae (Fig. 6h). The leaf shape and trichomes of the last two species very closely resemble those of S. of- ficinalis var. fulvescens (Fig. 7f, 7g). However, S. officinalis var. fulvescens often has dimorphic pubescence—pale dense tomentum on the lower leaf sur- face and large dark brown or reddish-colored trichomes on the veins. These species, of rather arid regions, resemble one another in overall appearance. Styrax jaliscana of northern Mexico might be related to the above-men- tioned species. Although its leaves are more elliptic than suborbicular, the leaf texture and general appearance of the plant are very similar to S. of- ficinalis var. fulvescens. However, its pubescence is woolly, the trichomes (Fig. 8d) having long slender arms very much like those of S. grandifolia. 196 The leaves of S. youngae and S. jaliscana have uniseriate hairs on the upper surface. These hairs are fairly long and are angled toward the apices of the leaves. Styrax glabrescens var, pilosus bears a striking resemblence to S. grandi- folia in overall morphology. The pubescence is much heavier on the leaves of the latter species, often becoming woolly, but both species have the same type trichomes with long slender arms (Figs. 9d, 4d). In the S. argenteus complex there is variation in degree of pubescence: var. ramirezii (Fig. lle) has stout trichomes with short arms; var. hintonii (Fig. 11b) has trichomes with long slender arms that very much resemble those of S. grandifolia; while var. argenteus (Fig. lla) has trichomes of intermediate length. The degree of pubescence on leaves and on flowering and fruiting structures is used as a major taxonomic character in several species com- plexes. These include S. americana and its variety pulverulenta, S. platani- folia and its variety stellata, S. argenteus and its varieties ramirezii and hintonii, and S. glabrescens and its variety pilosus. Leaves. The leaves vary much in size, shape, and texture, ranging from broadly obovate to elliptic-oblong or semi-orbicular and from thin and fine to thick, coriaceous, and revolute. Many of these differences are employed taxonomically in this treatment. Leaf shape and geographic range together are good characters for separating several groups of species. Styrax grandi- folia, occurring in the eastern United States with S. americana var. pulveru- lenta—whose leaves may be also densely pubescent below—can be distin- euished from the latter species by its rather large, obovate leaves in mark- ed contrast to the smaller elliptic leaves of var. pulverulenta. The subor- bicular leaves of §S. plantanifolia, a species of southwestern Texas and northern Mexico, are distantly toothed toward the apices and cannot be con- fused with any other species in this study. Stomata are confined to the lower leaf surface. The vascular bundles of the smaller veins are accompanied by sclerenchyma. A_ transverse section through the distal end of the petiole reveals a median, cylindrical, vascular strand flattened towards the adaxial surface. Solitary and clustered crystals, probably of calcium oxalate, are reported to occur, especially around the vascular bundles (Metcalfe and Chalk, 1950). The epidermal pattern of the leaves of Styraxv is of both the regular- and the irregular-celled type. Although it is not likely that epidermal-cell form could be used as a specific taxonomic character, it is possible to see affini- ties between species said to be closely related. A study of epidermal cells reveals some possible relationships between the species of Styraa. In addition to the stellate pubescence, occasional scattered glands occur on the lower leaf surface. These glands, rather flattened and circular, are composed of from five to eight small cells surrounding a central depressed area; the function is unknown, A leaf-clearing technique, including the tannic acid-ferric chloride stain- 197 ing procedure of Foster (1934), was used for studies on the epidermal cells. Flowers. The inflorescenses are terminal or axillary, and have numerous white- or cream-colored, fragrant, often pendulous, solitary or racemose flowers. The pedicel of each flower is subtended basally by minute leaflike bracts. These bracts often also occur midway on the pedicel and on the calyx. The length of the bracts varies with the species. The calices of Styrax are campanulate, with the lobes reduced to teeth. The teeth vary from five to seven and range from inconspicuous as in S. argenteus to long acute-acuminate as in S. jaliscana. The calyx surfaces vary from glabrous and smooth, as in S. americana var. americana, to densely stellate pubescent, as in S. argenteus var. hintonii. The size of the teeth and the presence or absence and degree of pubescence may be used as taxonom- ic characters. The calices are persistent on the drupaceous fruits. The corolla is sympetalous, the tube being from 38-5 mm long. For this distance the stamens are completely adnate to the tube. The white- or cream- colored lobes number usually five, occasionally six. They are linear-oblong to elliptic-oblong or obovate and range, depending on the species, from 5 mm to 2.8 cm long. The lobes may remain straight, partially open, or widely flared, or the apices may be strongly reflexed. An unusual condition exists in S. ochraceus in that the lobes, up to 2.8 cm long and sympetalous basally, are partially fused for two-thirds their length with the apices reflexed. The effect is that of a long salverform corolla. Perkins (1907) divided Styraxz into two sections, Eustyrax and Foveolaria. Eustyrax includes those species having 16-24 ovules per ovary while Foveo- laria includes those having 3-5 ovules per ovary. Eustyrax was divided into two series, Valvatae and Imbricatae. Valvatae represents those species hav- ing valvate dehiscence of the corolla lobes; imbricatae, those with imbricate dehiscence. The species in Valvatae has lobes that are rather thick, often with a reddish tint on the inner surface. These include S. argenteus and its varicties, S. conterminus, S. portoricensis, S. obtusifolia, and S. glaber. The other species included in this study are in Imbricatae; their corolla lobes are rather thin and have the same color and texture on both surfaces. nectary was not detected at the base of the corolla: however, the flow- ers are often aromatic and attract a variety of insects. Copeland (1938) re- ports, of S. officinalis var. californica, that minute drops of liquid have been seen on the distal part of the ovary, but no taste of nectar was detected. The stamens are usually twice as many as the corolla lobes, monadelphous in a short tube basally, 4-6 mm long, and adnate to the sympetalous corolla tube but extending just beyond it. Styrax ochraceus, which characteristic- ally has long corolla lobes, has filaments fused nearly the entire length of the partially-fused corolla lobes. The anthers are basifixed, with linear introrse dehiscence. The filaments are flattened and often stellate pubescent. The pollen, yellow and tricolpate, varies little in size, shape, and texture among the species. The filiform style, exserted 1-3 mm past the stamens and about as long 198 as the corolla lobes, is usually glabrous but may be partially stellate-pube- scent. The stigma is inconspicuously 3-lobed. Fruit. The drupes, globose to oblong to broadly elliptic, are dry, loculicid- ally 3-valvate, and irregularly dehiscent or indehiscent. The one or rarely two seeds are subglobose to ellipsoidal with a basilateral hilum and a hard, smooth, or wrinkled testa. The cotyledons are large and elliptic, the hypo- cotyl is massive and the radicle and plumule are small. The calyx is persistent. In several species the style, or part of it, remains on the fruit as an apiculation. Fruit characteristics are not generally used as taxonomic criteria in Styrax. Styrax portoricensis, however, has fruits so large that it cannot be confused with any other species in this study. POLLINATING AGENTS The flowers of Styrax are considered to be entomophilous. The anther sacs have longitudinal, introrse dehiscence, which is usually associated with self- pollination. However, the anther sacs are strongly reflexed, so that any visi- tor becomes dusted with pollen. Also, the style is longer than the stamens, which suggests an outbreeding system. Some wind pollination may also occur. The pendant flowers are fragrant, and the rather wide, partially reflexed petals provide adequate landing sites for insects. Pollinators were observed on S. americana and on S. grandifolia. The following insects were most prevalent visitors: Hymenoptera Apis mellifera—Honey bee Bombus spp.—Bumble bees Anthophora spp.—Solitary bees Prosopis spp. Lepidoptera Danaus plexippus—Monarch butterfly Eurema nicippi—‘Sleepy orange”’ Diptera Fam. Blepharoceridae—The net-winged midges Fam. Drosophilidae Drosophila spp.—Fruit flies Thysanoptera Thrips tritiea—Thrips Of the Hymenoptera, Apis mellifera is the most frequent visitor, with sev- eral species of bumble bee next. The bumble bees have a more difficult time landing on the flowers than do smaller insects because the lax racemes do little to support their weight. Although the flowering period of S. americana and S. grandifolia lasts 2-3 weeks, insect visits are mostly during the first week. Those pollinators listed above were common on both species; how- ever, more Drosophila frequented S. americana. 199 CYTOLOGY Chromosome counts for several species of Styrax are found in the litera- ture. Counts for S. obassia and S. japonica of Japan were made by Mansard (1936). In addition, chromosome counts for S. officinalis var. californica were made by Copeland (1938), and for S. officinalis var. fulvescens by Raven et al (1965). These counts are listed in Table During my study I attempted to obtain chromosome counts for Styrax of eastern United States, including S. grandifolia, and S. americana var. ameri- cana and var. pulverulenta. The counts were based solely upon pollen mother cell smears of freshly gathered material and are listed in Table 1. The base number for Styraxv is n=8. Styrax grandifolia was found to have a haploid number of 16 (Fig. 1c) and is, therefore, a tetraploid. Both S. americana var. americana and var. pulverulenta were found to have a hap- Icid number of 8 (Fig. la, 1b), making them diploids. The relationship between the eastern species of Styrax is confusing. It seems that the morphological distinctness of the taxa is maintained by ternal rather than external isolating factors. Although the geographical ranges are sympatric, the ecological ranges rarely overlap. Styrax grandi- folia is more prevalent on rather dry, well-drained, sandy soils, commonly on slopes. It covers a wide range, most of which it shares with S. americana. Where the ecological ranges overlap there is no indication of hybridization between the species. There are at least two possible origins of S. grandifolia. Wood and Chan- nell (1960) noted that S. glabrescens, whose range includes northern Mexi- co, is a close relative of S. grandifolia. No chromosome counts are available for S. glabrescens, but the two species show very close morphological resem- blance. Styrax grandifolia also resembles S. officinalis and has been mis- identified as this species—as in Walter (1788). The resemblance to S. officin- alis is indeed striking on some specimens collected on high, well-drained, sandy soils. Styrax officinalis (a known diploid) is principally a Mediterran- ean species whole range also includes California. It may have ranged further east in North America in ages past. Styrax grandifolia could have arisen as an allo- or saa erage from the above species or possibly from S., americana. Further wor n the genomic characteristics is necessary to make any ae nArnrere The origin of S. americana is even more perplexing. Other than S. japoni- ca, there are no other species that, on morphological grounds, would have given rise to S. americana. Styrax japonica, a pentaploid whose range does not include North America, seems a most unlikely ancestor. A great deal more cytological work needs to be done in order to assemble conclusive evidence of the evolutionary relationships of the species of Styrax. As with many woody plants, it was rather difficult to determine the chro- mosome number, and it was only after many attempts that the counts for S. grandifolia and S. americana were obtained. There seems to be a very narrow time range within which meiosis occurs in Styraz. es 200 TABLE I. Chromosome Counts (2n) for Styraq. Species 2n Number Source of Count Styrax obassia 16 Manshard (1936) Styrax officinalis var. 16 Copeland (1938) californica Styrax officinalis var. 16 Raven et al. (1965) fulvescens Styrax americana var. 16 Gonsoulin (Louisiana: Washington Par- americana ish. Bogalusa. Gonsoulin 1032, 28 May 1969; flower buds collected 7 Mar 1970, 15 Mar 1972. NY, PH, SMU, US, VDB). Styrax americana var, 16 Gonsoulin (Alabama: Baldwin Co. Foley. pulverulenta Gonsoulin 1964, 20 Mar 1973. Herbar- ium of TIast Tennessee State Univer- Styrax grandifolia 32 Gonsoulin (Tennessee: Lewis Co. Gor- donburg. Gonsoulin 1022, 3 May 1969; flower buds collected 6 Apr 1969. NY, PH, SMU, US, VDB) Styrax japonica 40 Manshard (1936) SYSTEMATIC TREATMENT Styrax Linnaeus, Syst. Nat. ed. I. 1735; Gen. Pl. ed. I. 1483, 1737; Sp. Pl. 444. 1753; Gen. Pl. ed. 5. 208. 1754. Cyrta Lour., Fl. Cochinch. 278. 1790. Foveolaria Ruiz and Pavon, Fl. Peruy. Chil. Prodr. 57, t.9. 1794. Strigilia Cav., Monadelph. Class. Diss. 7:358, t. 201. 1790. Tremanthus Pers., Syn. Pl. 1:467. 1805. Adnaria Raf., Fl. Ludovic. 56. 1817. Epigenia Vell., Fl. Flum. 183. 1825. Benzoin Hayne, Getr. Darst. Beschr. Arzneyk. Gew., 11, t. 24. 1829. Darlingtonia Torr., Proc. Amer. Acad. Arts 4:191. 1851. Shrubs or small trees of warm or temperate areas of the Americas and Eurasia. Leaves alternate, estipulate, commonly deciduous, entire or serrate- serrulate, broadly ovate-obovate to elliptic-lanceolate, densely stellate-pube- scent to glabrous; apices mostly acute-acuminate; bases cuneate to round- ed. Flowers perfect, 5-merous, in pendulous racemes in the axils of the leaves and/or on terminal branches; pedicels glabrous to densely stellate pubescent; calices campanulate, truncate, gamopetalous, 5-—7-denticulate, glabrous to canescent or densely stellate-pubescent, fused basally, imbricate or valvate in the bud, eventually spreading or recurved. Stamens usually twice as many as the corolla lobes; filaments flattened, basally monadel- phous, inserted near the base of the corolla; anthers oblong, basifixed, with linear dehiscence. Ovary about half inferior to superior, 3-locular toward the 201 base, 1-locular above, ovules few to many in each locule; style simple, as long as or longer than stamens; stigma capitate, smooth or slightly lobed. Fruit globose to ovoid-oblong, dry, crustaceous, usually 1-seeded, irregularly dehiscent to indehiscent, basally surrounded by the persistent calyx. Seeds 1 or rarely 2, subglobose or elliptical with a basilateral hilum; testa hard, smooth or faintly wrinkled Type species: Styrax officinalis L.; common name: ‘‘Snow-bells.”’ There are about 120 species in the genus, most of them tropical and sub- tropical, in eastern Asia to New Guinea. (absent from the Philippines and Ceylon), the eastern Mediterranean region, South America, the West Indies, and Central and North America. Ten taxa occur in the United States, four of them varieties. Parts of the following key were modified from keys of previous workers. These include the Series Valvatae and Imbricatae recognized by Gurke (1897) and Perkins (1907). Key to North American, Central American, and Caribbean Styrax . Corolla lobes predominantly valvate in aestivation. ... Series 1. Valvatae . Corolla lobes predominantly imbricate in aestivation. eee Series 2. Imbricatae ht pe oS © Series 1. Valvatae Gurke la. Lower surfaces of the leaves covered either with long spreading stellate trichomes, or dense, matted or otherwise tomentose pubescence. . Lower surfaces of the leaves glabrous or covered with scattered to dense lepidote pubescence. a ae a a ed . Leaves 7-14 cm long, ablone -C iliptic. covered below with dense matted pubescence to dense long spreading stellate trichomes, apices short acuminate; mature flowers 13-17 mm long; central Mexico through Cen- tral America (Figs. 10 & 11). . . 9. Styrax argenteus 2b. Leaves 3-4 cm long, oblong to obov te, cover ea haaw with dense matted pubescence, apices rounded to acute; mature flowers 7-9 mm long; Cuba and Haiti (Fig. 13)... . 11. Styrax obtusifolia 3a. Leaves glabrous to covered wath seater jevidots pubescence below, the trichomes not commonly overlapping each other. . Leaves completely covered below with lepidote pubescence iupartine: a crusty appearance. Bock oe UO A ee Oe a. Oe. DD . Leaf blades obovate- elliptic: 2—5 cm long, apices rounded to acute; ma- ture flowers 2.5-3.4 cm long: drupes 8-19 mm long; Dominican Repvb- lic (Fig. 14). . 12. Styrax ochraceus . Leaf blades elliptic, 6— 1 cm iene apices cart acuminate; mature flow- ers 1.6-2 ecm long; drupes 2.5-3.5 em long: throughout Puerto Rico (Fig. 15)... eee eee «18. Styrax portoricensis — or bo jab) ve ao ee bad) = ao da. Mature flowers 12-15 mm long; drupes ob!ong-elliptic, slightly asymmet- rical, 12-15 mm long; Central America (Fig. 12). 10. Styrax conterminus db. Mature flowers 2—2.4 cm long; drupes ovoid to globose-ovoid, 2 cm long; throughout the British West Indies (Fig. 16)... . 14. Styrax glaber Series 2. Imbricatae Gurke la. Leaves orbicular to suborbicular or oblong-ovate; apices rounded to 2 acute ee fe ew se £ me Se se we & £ me & 1b Leaves narrowly to broadly elliptic or obovate, apices acute to long acuminate. 2a. Leaves sebenie? or — ing sparse eeiciea: coarse ste ae puhasecice, 2b. Leaves with fine, dense, silvery indumentum to woolly white or grayish 4 tomentum. oe «© « « 4 3a Leaves irres sateriy ped: or foomed ionand dig apices; penne: g pantous to finely puberulent, apices glandular and prominently denneae dwards Plateau, Texas, U.S.A. (Fig. 5). . 8. Siyrax ve ae 3b. Leaves not irregularly lobed or ee d buena the apices, lower sur- faces glabrous to fulvescent; calices puberulent to fulvescent, apices non-glandular; Mediterranean region and California, U.S.A. (Fig. 7). 6. Styrax officinalis 4a. Calices densely puberulent, ee apices aenauiak tipped; leaves bright silvery below with a very fine and dense indumentum; Edwards Plateau, Texas, U.S.A. (Fig. 6)... . 2. . . 4. Styrax texana 4b. Calices and lower leaf Surtees noe as aoe. da. Lower leaf surfaces covered with dense woolly nite ‘ a GS ee we om em & wee openal is var. Gumescens Ga. STYRAX OFFICINALIS Linnaeus var. CALIFORNICA (Torrey) Rehder, Mitt. Deutsch. Dendrol. Ges. 1915: 226. 1915. Styrax californica Torr., Smithsonian Contrib. 6(1V): 1-8, t. 12. 1853. Styrax officinalis L. var. californica (Torr.) Munz and Johnson, Bull. Torrey Bot. Club 51:297. 1924. Shrub to 3m, upright, branching, with irregular crown. Leaves glabrous to lightly pubescent below, blades broadly ovate to slightly orbicular, 4-6 cm long, margins entire, bases obtuse to rounded, apices acute fo obtuse; upper surfaces light green, smooth; lower surfaces paler, often with very fine scattered stellate trichomes; veins slightly impressed; petioles 5-8 mm long, slender, glabrous or very faintly pubescent. Flowers in terminal or axillary, 3-5 flowered racemes, borne on short leafy branches; ealices 5-7 mm long, faintly pubescent, shallowly toothed, teeth acute; corolla lobes 12-15 mm long, oblong to obovate, apices acute, surfaces with soft downy pubescence along the borders fading inward; stamens about 2 mm shorter than corolla lobes; filaments pubescent throughout their length; anthers 1/3 the length of the stamens; style barely exceeding the corolla lobes, pubescent on the proximal 1%; stigma barely lobed. Drupes maturing August—Septem- ber, globose, 8-10 mm in diameter, style persistent as an apiculation, seeds subglobose, shining, smooth, longitudinally grooved (Fig. 7). Styrax officinalis var. californica occurs generally in the foothills of the Sierra Nevadas along watercourses from 500 to 3,500 ft. elevation (Jepson, 1963). From Siskiyou County, its northernmost locality, it ranges southwest- ward to Lake and Alameda counties, eastward to Eldorado and Placer coun- ties and then to Fresno County, its southern limit (Fig. 18). Jepson (1963) gives the southernmost extension as San Bernardino County, but I have not seen material from this county that I would call var. californica. The known range of var. californica was, before my study, much more restricted than presently, extending only from Shasta County south to Amador and Cala- veras counties (Copeland, 1938). The flowers appear from April through mid-May. The colloquial name is ‘Snowdrop bush.”’ Holotype: U.S.A. CALIFORNIA. Sacramento Co.: Upper Sacramento. Fremont, 1845-47 (NY!). UNITED STATES. CALIFORNIA. Alameda Co.—Bracelin 1235 (F, NY), Bracelin 2586 (F, NY), Berkeley, Kelee (UNC). Amador Co.—Glenco, Bel- shaw 2379 (UNC). Butte Co.—Enterprise, Heller 1 1891 (F, GH, MO, N ~~ — 217 UNC). Calaveras Co.—Mokelumme R., Hanson 227 (GH, MO, NY). Colusa Co.—Clear Lake, Hoover 4988 (UNC); Leesville, Heller 13111 (F, GH, MO, NY, PH); Bear Valley, Bacigalupi 3187 (NY, UNC). El Dorado Co.—Natoma Ditch, Brandegee, 14 May 1907 (PH); American River, Smith 2540 (UNC). Fresno Co.—Miil Creek, ea 7686 (FSU). Glenn Co.—Neesville, Heller 11535 (F, GH, MO, NY, PH, UNC). Lake Co.—Leesville, Abrams 12577 (NY); Sulphur Banks, Eastwood pe (F, GH); Williams, Clausen 1027 (UNC); Clear Lake, Carter 353 (UNC); Borax Lake, Simontacchi 264 (UNC): Soda Bay, Neuns 165 (UNC), Schulthers, 9 Aug 1931 (UNC), Hastings, 28 Apr 1940 (UNC), Howell 15589 (GH); Bartlett Springs, Bleedlove 5180 (DUKE, SMU); Servier Lake, Bowmann, 11 May 1901 (GH). Placer Co.—Auburn, Carter 1485 (GH, NY, UNC); Auburn, Crum 2043 (UNC); Colfax, Hitchcock 6391 (NY); Forest Hill, Jump, 15 May 1939 (GH, UNC). Sacramento Co.—Upper Sac- ramento, Pringle 14757 (F, GH, MO); Sacramento R., Capt. Wilkes, 1838 (NY), Fremont, 1845 (NY), Hulse (NY); Folsom, Copeland 1614 (GH UNC), Nordstron 79 (UNC). Shasta Co.—Redding, Johannsen 149 (NY), Jo- hannsen 122 (UNC), Hitchcock 6486 (MO, UNC), McVaugh 6238 (UNC), Grant 7976 (UNC), Balls 20724 (NY); Kennett, Eastwood 1455 (GH), Smith 168 (GH); Lemoine, Fastwood 1804 (GH), Rose 34238 (F, NY, SMU); Pitt R., Mason 5876 (GH, UNC), Heller 13498 (F, MO, NY), Jones, 16 Jul 1931 (MO). Brown 273 (F, NY, UNC); Delta, Jepson 6177 (NY): Whiskey Town, Hovwell 29185 (NY), Rose 54029 (NY); Trinity, Balls 18665 (NY); Tall House, Baker, 30 Apr 1900 (UNC); Weaverville, Josephson 24 (UNC); Baird, Coombs, 9 Jul 1915 (GH). Siskiyou Co.— Pitt R., McMinn 2493 (UNC); McCloud, Sculphan 150 (NY). Tehama Co.— Paskenta, Crampton 7773 (UNC). Yuba Co.—Straw- berry Valley, Burks 38 (UNC); Smartville, Josephson 37 (UNC The type locality, ‘“‘Upper Sacramento,’’ means the sieiehborhood of the Sacramento and the Pitt rivers in what is now Shasta County. The nomenclature of this taxon is rather confusing. Its present rank is due to its strong resemblance to S. officinalis of southern Europe, from which it is chiefly distinguished by its few-flowered racemes, thickened pedi- cels (a character that does not hold up), and longer staminal tube. After Torrey (1853) described S. californica, it was reduced by Perkins (1907) to synonymy under S. ulate a) then resurrected by Rehder (1915) and, independently, by z and Johnston (1924) as S. officinalis var. californica. So the distribution = Z officinalis 7 the Mediterranean region and California. The movement of this entity through the land bridges that occurred across the Bering Sea is quite possible, as many species of animals and plants are known to have accomplished this. Several species of Styrax are represented in Asia. Some of them resemble S. officinalis. Among these are S. macran- thus Perk. of central Asia, S. japonicus Sieb. sige pace: of Japan, S. obassia Sieb. and Zucc. of China and Japan, and S. atissi s Champ. of China. The last species occupies the act ene: range of mode mentioned, Per- haps elements of S. officinalis moved northwestward during interglacial periods, eventually crossing the Bering Sea land bridge. As the climate cooled — 218 once again, the glaciers returned, destroying the species over much of the former range by forcing it south into terrain not suitable for it. Little is known of the fossil record of Styrax, especially in the region concerned. The California material differs from the Mediterranean material in the characters mentioned and in pubescence, being glabrate or lightly tomentose on the underside of the leaves instead of closely white tomentose. This pube- scence is dimorphic along the veins and midrib, with dark colored trichomes being scattered among and projecting above the pallid mass of the tomen- tum. Also, the style in Mediterranean material appears to be less com- pressed than in California material and is evidently less lobed (Munz and Johnson In that only differences between the European and the Californian exam- ples are minor, being neither constant nor significant, I agree with Munz and Johnston (1924) and Copeland (1938) and treat the Californian material as varicties of S. officinalis. For information concerning the embryology of S. officinalis var. californica the reader is referred to Copeland (1938). 6b. STYRAX OFFICINALIS Linnaeus var. FULVESCENS (Eastwood) Munz and Johnston, Bull. Torrey Bot. Club 51: 295-302. 1924. Styrax californica Torr. var. fulvescens Kastw., Bot. Gaz. 41: 283-296. 1906. Leaf blades broadly ovate-orbicular to subcordate, 4-6 cm long, margins entire, bases broadly ovate to subcordate, apices obtuse or abruptly acumin- ate, only slightly longer than broad; upper surfaces light green, smooth to roughened, glabrous or with a few scattered stellate trichomes; lower sur- faces much paler, pubescent to densely tomentose. Calices 5-7 mm long, shallowly toothed, teeth acute, usually white tomentose (Fig. 7). According to Munz and Johnston (1924), var. fulvescens ranges along the coastal slopes of California from the mountains near Santa Barbara to those near San Diego. The ranges of the two varieties (var. californica and var. fulvescens) are widely separated, with Santa Barbara County the north- ernmost extension. The known range of var. fulvescens is here extended as far north as Mendocino and Butte counties (Fig. 18). The flowers aafae pom mid-April through May. The colloquial name is “Hairy snowdrop Holotype: U.S.A. oinmonn. Santa Barbara Co.: Painted Cave Ranch, Santa Barbara Mountains. Wolf, 17 May 1904 (GH!). UNITED STATES. CALIFORNIA. Amador Co.—Zion, Hansen 840 (UNC). Butte Co.— Bidwell, May 1879 (GH). Calaveras Co.—Mokelumme Hill, Blais- dell (GH); Golden Gate Hill, Rosemary 225 (UNC); Calaveras R., Stanford 300 (GH, MO). El Dorado Co.—Salmon Falls, Robbins 1934 (GH, UNC). Fresno Co.—Trimmer, Bucksloe, 10 May 1953 (GH, UNC); Trimmer, Clark 46-14 (GH), Quibell 1927 (NY, UNC), Quibell 1565 (DUKE): Kings R., Craw- ford 25n (UNC). Lake Co.—Mt. Konocti, Blankinship, 1 May 1924 (MO); Lower Lake, Cuff 133 (UNC); Hough Springs, Abrams 6273 (NY, US). Meno- 219 docino Co.— Vasey, 1870 (GH). Orange Co.—Star Canyon, Hutt 373 (DUKE); Trabuco Canyon, Smith 5453 (F); Corona, Lewis 380 (UNC). Placer Co.— Folsom, Wolf 8609 (GH, MO, NY). Riverside Co.—Temecula, Stubblefield, 2 May 1938 (NY); nae, Spring 34 ee Sacramento Co.—Folsom, Cope land, 30 Apr 1939 (GH, F, MO, NY, C). San Bernardino Co.—Cajou Pass, Craig 922 (GH); San Bertiarding i Parish, 29 May 1888 (UNC), Parish, Apr 1887 (MO), Parish 307 (GH, MO, NY, PH, UNC), Parish 11389 (UNC); San Bernardino, Lemmon, May 1878 (F); Waterman Canyon, Jepson 0091 (VDB); San Savaine, Ewan 10982 (GH, MO); Mts., Engelmann 20 Sep 1880 (GH, MO); San Bernardino Mts., Clark 15916 (SMU); Big Bear Lake, McMinn 1560 (UNC). San Diego Co.—Rainbow, Jensen 141 (UNC); Fall Brook, Parry, May 1883 (MO); Mesa Grande, Gander 200 (UNC), Spencer 1147 (NY); Luis Rey R., Mason 3138 (GH, UNC); Henshaw Dam area, Munz 10353 (GH, UNC). San Luis Co.—Nipomo, Lee 635 (UNC). Santa Bar- bara Co.—Santa Inez Mts., Wolf 2302 (UNC); Painted Cave Ranch, East- wood 33 (F, GH, MO, NY, UNC), Wiggins 8508 (GH); Santa Barbara, Bran- degee, 1888 (F); Rattlesnake Canyon, Mason 456 (GH, UNC). Eastwood (1906) first described this variety as S. californica var. fulve- scens: “Differs from the typical Styrax californica in the broader, rounder leaves, cordate at base, the much denser stellate tomentum, and the gen- eral prevalence of rufous hairs especially on the calyx.’’ Munz and Johnston (1924) created a new combination, S. officinalis var. fulvescens, which I accept for reasons previously discussed. This variety ranges over a much greater area than originally thought. The ranges of var. californica and var. fulvescens do overlap considerably. Within this overlap are intermediate forms with various degrees of pube- scence. Thus, while the two varieties are generally separated geographically, there does exist an area, which includes El] Dorado, Amador, and Calaver- as counties, in which intermediates occur, 7. STYRAX JALISCANA Watson, Proc. Amer. Acad. Arts 26:144. 1891. Styrax Regi alis L. var. jaliscana (Wats.) Perkins, Pflanzenreich IV. 07. Shrub to 1 m, rather freely branched. Leaves subsessile, blades round- ovate or oblong-ovate, 4-10 cm long, the bases obtuse or somewhat cuneate, margins entire, apices abruptly short-acuminate; upper surfaces pale, dense- ly white tomentose and reticulately veined; petioles 3-4 mm long, densely white tomentose. Flowers in axillary, 1-5 flowered racemes, 8-12 mm long; pedicels 4-6 mm long, densely white tomentose; calices 5-7 mm long, dense- ly white tomentose, very evidently dentate, teeth 5-7, length 1-1.7 mm, acute; corolla lobes 12-14 mm long, oblong to elliptic-ovate, apices obtuse to acute, surface with soft downy pubescence; stamens 2-3 mm shorter than corolla lobes; filaments pubescent near the bases: style barely longer than the corolla lobes, essentially glabrous and slightly lobed. Drupes 8-12 mm in diameter, depressed-globose; seeds usually 3, smooth, with a ridge running from the hilum sideward (Fig. 8) 220 New branchlets densely white tomentose. Teeth evident on persistent calices. Styrax jaliscana occurs on rocky hillsides and rather dry slopes. Although only a few collections are available, they attest to the wide range of this species, which extends from northern Baja California through southern Mexico (Fig. 19). Holotype: MEXICO. JALISCO. Near Guadalajara. Pringle 2978, 10 Nov 1889 (GH!). MEXICO. BAJA CALIFORNIA. Sierra Laguna, Gentry 4420 (GH, UNC). GUERRERO. Galeana, Hinton 4583 (NY). JALISCO. Near Guadalajara, Pringle 2978 (GH), Pringle 4416 (F, MO, NY, PH, US, UNC), Pringle 11012 (F, MO, NY, US). SINALOA. Sierra Surotato, Gentry 6244 (MO, NY). Standley (1924) states that S. jaliscana may not be specifically distinct from S. officinalis. In gross morphology it does resemble the California va- rieties of the latter species, especially var. fulvescens, but, on closer exam- ination, differences of specific rank may be seen. The leaves of S. jaliscana have a slightly larger length/width ratio and are more acuminate, whereas those of S. officinalis tend to be more acute to very short-acuminate. Leaf pubescence in the latter species at its densest, in var. fulvesens, is mod- erate as compared to the dense woolly pubscence of S. jaliscana, which persists on young branchlets and racemes. The calices of both species differ not only in pubescence but also in presence of teeth, these being much more prominent in S. jaliscana. These two species may well be closely related but they have diverged enough to warrant separation. Styrax jaliscana is an extremely wide-ranging species. The collections from Sinaloa and Baja California have larger leaves, up to 10-12 cm long, than do the type and other collections from Jalisco. The species is probably relatively rare because only scattered collections are available. There are no species currently known from its range that, on morpholog- ical grounds, could have given rise to S. jaliscana. Therefore, I must con- clude, as Standley did, that its closest relative is S. officinalis. 8. STYRAX GLABRESCENS Bentham Key to Varieties Leaves essentially glabrous below (may have stellate trichomes in the vein axils); calices stellate pilose with rufous or yellowish hairs (Fig. 9). i. eae a a oe 8a. S. glabrescens var. glabrescens Leaves lightly to densely pilose below; calices minutely grayish tomentulose (Fig.9), 2. 2. 2... OB. SS. glabrescens var. pilosus 8a. STYRAX GLABRESCENS Bentham var. GLABRESCENS, Pl. Hartweg. 66. 1839. Styrax guatemalensis J. D. Smith, Bot. Gaz. 15: 27, 28. 1890. Small tree, 6-12 m high, spreading. Leaves essentially glabrous, ovate- 221 obovate to elliptic, the blades 8-14 cm long, 4-7 ecm wide, the bases cuneate- ly narrowed to nearly rounded, margins entire to distally toothed, apices short to long acuminate; upper surfaces dark green, glabrous; lower sur- faces paler, glabrous or with a few scattered stellate trichomes; surfaces reticulate-veiny; petioles 6-10 mm long, lightly stellate pubescent. Flowers in axillary, semi-drooping, 4-12 flowered racemes, 2-8 cm long, tomentose; calices 5-6 mm long, tomentose, with rufous hairs, shallowly toothed, teeth acute; corola lobes 12-16 mm long, ovate-elliptic, apices obtuse to acute, surfaces with soft down pubescence; stamens about 2 mm shorter than cor- olla lobes; filaments with scattered stellate trichomes; style essentially glab- rous; stigma slightly lobed. Drupes globose to subglobose, 8-12 mm in dia- meter, covered with dense matted tomentum; style not persistent; seeds us- ually 1, pale brown, with slightly roughened texture, with 2-3 grooves run- ning from the hilum sideward (Fig. 9). New twig growth covered with light stellate pubescence. Racemes covered with pubescence grading into dense tomentum toward the calices. Styrax glabrescens var. glabrescens occurs in mountainous regions along streams, on rocky slopes, and in heavy forest from southern Costa Rica through Honduras and Guatemala and into Mexico as far north as Hidalgo (Fig. 19). Holotype: MEXICO. VERA CRUZ. Near Jalapa. Linden, Jun 1838 (K; photograph of holotype examincd). COSTA RICA. ALAJUELA. La Palma de San Ramon, Brenes 6734 (NY), Brenes 3986 (F, NY), Brenes 13599 (NY), Brenes 6079 (F, GH, NY), Brenes 6734 (GH). CARTAGO. El Muneco, Standley 3350 (US), Standley 51137 (GH, US). GUATEMALA. ALTA VERAPAZ. Carcha, Standley 70316 (F); Sasis, Smith 1690 (GH, US). EL QUICHE. Chajul, Sharp 4684 (F), Sharp 4673 (F). HUEFHUETENANGO. Barillos, Holdridge 2338 (F), Steyermark 49738 (I). QUEZALTENANGO. Las Nubes, Standley 8397 (F); San Martin, Standley 67865 (I). SAN MARCOS. Tecutla, Steyermark 36807 (F); Finca, Steyermark 37378 (F). SUCHITEPEQUEZ. E\ Naranjo, Steyermark 46750 (F). HONDURAS. COMAYAGUA. oan Montecillos, Molina 7233 (F, US); El yous Yuneker 6175 (F, GH O). LA PAZ. Montana Verde, Ae 24404 (F). MEXICO. CHIAPAS. Siltepeg, Matuda 5102 (F); Boqueron, Matuda 5410 (F, MO); Letrero, Matuda 17816 (I), Matuda 4335 (GH, MO, NY, US); Ran- cho Phoenix, Purpus 10612a (F); Fenes, Purpus 10088 (NY); Monserrate, Purpus 10521 (GH, UNC). HIDALGO. Taeualipen: Moore 3231 (UNC), Martinez, Apr 1941 (GH); Honey Station, Pringle, 17 Jun 1908 (US); Trinidad, Pringle 13104 (MO, F, US) au 222 OAXACA. Montebello, Reko 41386 (US); Cuyamecaleco, Conzatti 3480 (US); La Sole de Vega, Carlson 2710 (F). PUEBLA. Huauchinango, Lundell 12631 (UNC, US); Villa Juarez, Rzedow- ski 17211b (TEX). VERACRUZ. Jalapa, Pringle 8129 (F, GH, MO, NY, UNC, US), McDaniels 942 (F); Huatusco, Purpus 1920 (UNC); Jalapa, Galeotti 2851 (US). Wood and Channell (1960) mention that S. glabrescens is a close relative of S. grandifolia. The latter species is a tetraploid, but no chromosome counts of the former have been made so no relationship along these lines can be established. They do, however, resemble each other in many morphological features. The leaves of both have the same texture, but those of S. glabre- scens lack the dense matted pubescence characteristic of S. grandifolia. Leaf shape differs somewhat, being broadly elliptic in the former and obo- vate in the latter species. The flowers are slightly larger in S. glabrescens and the petals have a slight purple tint contrasting to the creamy white petals of S. grandifolia. A good case for their close relationship exists but cytological work would be necessary for proof. Standley (1938) says of S. glabrescens, ‘“‘Frequent in forest of the Pacific slope, mostly at elevations of 500-1,500 meters: region south of Cartago; re- gion of San Ramon. Extending to Mexico. Easily recognized among Costa Rican species because of the broad, thin, almost glabrous leaves, green on the lower surface.” Styrax guatemalensis, described by Smith (1890), is treated here as a synonym of S. glabrescens. Smith states that it differs from the latter species by its ovate leaves, its one-half larger flowers, and its ovate petals. However, the leaves and flowers are identical to those of S. glabr 8b. STYRAX GLABRESCENS Bentham var. PILOSUS Senin Pflanzenreich IV. 241 (Heft 30): 72. 1907. Styrax pilosus (Perkins) Standl. Contr. U.S. Natl. Herb. 23:1129. 1924. Leaves ovate to obovate-elliptic, blades 6-12 cm long, 3-6 em broad; upper surfaces dark green, essentially glabrous; lower surfaces pale green, light to densely pilose; petioles 6-10 mm long, tomentose-pilose. Calices 4-5 mm long, stellate pilose with rufous hairs (Fig. 9). New twig growth and racemes are stellate-pilose with rufous hairs. Styrax glabrescens var. pilosus has a much more restricted range than var. glabrescens. It occurs in mountainous regions along streams, on rocky slopes, and in heavy forest in Chiapas, Oaxaco, Veracruz, and Hidalgo _ g. 19). Holotype: MEXICO. OAXACA. Chinantla and Rincon. Galeotti 2852, May MEXICO. CHIAPAS. Escuintla, Matuda 15995 (F), Matuda 4179 (GH, MO, NY, US); LaConcordia, Matuda 15922 (F); Monserrate, Purpus 9280 (UNC, US); Fenix, Purpus 10545 (GH, UNC): Jitotol, Breedlove 10188 (NY). HIDALGO, Zacualtipan, Martinez, May 1940 (GH). OAXACA. Galeotti 2852 (NY, US); Chinantla, Galeotti 3852 (F). 223 VERACRUZ, Chiconquiaco, Pompa 793 (GH). Only a few collections of this variety have been made since it was de- scribed by Perkins (1907). Standley (1924) raised this variety to specific rank. Since the only differences between the species and its variety are rela- tively minor ones I am herein following Perkins’ designation of this taxon as var. pilosus. 9. STYRAX ARGENTEUS Presl Key to Varicties Leaves covered beneath with a minute, very close, stellate tomentum (Fig. S. argenteus var. ramirezit ieagesa COV eiee below satin coarse ‘eueadie stellate hairs, the pubescence velutinous. Leaves less than half as wide as long; racemes and new branchlets cov- ered with very short stellate pubescence beset with lepidote trichomes (Figs. 10 & 11)... . . 9b. S. argenteus var. argenteus Leaves half or more as Gia. as roe racemes and new branchlets cov- ered with coarse, long, spreading velutinous pubescence, (Figs. 10 & 11). a 2 se a . 9c. S. argenteus var. hintonii . STYRAX ARGENTEUS Pres! var. ARG ENTEUS, Rel. Haenk. II. 60. 1836 co Press, Istin. WALTER, T. 1788. Flora Caroliniana. Londoni. WATSON, S. 1891. are to American botany. Proc. saunas Ra Arts 26: 124-163. WOOD, C. E. and R. B. CHANNELL. 1960. Styracaccae. pp. . In Vhe genera of the Ebenales in the Gone United States. J. Arnold Arbor. Aas our 236 1 Fig. 1. Early prophase I in pollen mother cells: a. Styrax americana var. americana; b. S americana var. pulverulenta; c. S. grandifolia. 237 Fig. 2. Styrax americana var. americana: a. habit sketch; b, c. upper & lower leaf epidermis (500); d, e. mature flower parts; f. mature fruit. 238 45mm b Fig. 3. Styrax americana var. pulverulenta: a. habit sketch; b, c. upper & lower leaf epidermis (500); d. trichomes on lower leaf surface (128); e. mature flower; f. mature fruit. Fig. 4. Styrax grandifolia: a. habit sketch; b, c. upper & lower leaf epider- mis (500); d. trichomes on lower leaf surface (128); e, f. mature flower parts; g. mature fruit. 240 Fig. 5. Fig. 6. Styrax platanifolia: a. habit sketch; b, ¢c. upper & lower leaf epidermis of var. platanifolia (500); d, e. upper & lower leaf epi- dermis of var. stellata (500); f. trichomes on lower leaf surface of var. stellata (128); g. mature flower; h. mature fruit. Styrax texana, S. youngae: a. leaf of S. texana; b. leaf of S. youngae; c, d. upper & lower leaf epidermis of S. texana (500); ec, f. upper & lower leaf epidermis of S. youngae (500); g. tri- chomes on lower leaf surface of S. fewana (128); h. trichomes on lower leaf surface of S. youngae (128); i. calyx & pistil of S. texana; j. calyx & pistil of S. youngae. 241 242 Fig. 7. Styrax officinalis: a. habit sketch of var. californica; b. c. upper & lower leaf epidermis of var. californica (500); d, e. upper & lower leaf epidermis of var. fulvescens (500%); f, g. trichomes on lower leaf surface (128), and leaf of var. fulvescens; h, mature flower; i. mature fruit. [i “gh: 243 Srmm | a i omr “e po) ~~ NY Se Fig. 8. Styrax jaliscana: a habit sketch; b, c. upper & lower leaf epider- mis (500); d. trichomes on lower leaf surface (128); e. mature flower; f. mature fruit. 244 55mm 14mm | Fig. 9. Styrax glabrescens: a. habit sketch; b, ec. upper and lower leaf epi- dermis (500); d. trichomes on lower leaf surface of var. pilosus (128); e. mature flower; f. mature fruit. 245 A tes a, S 2. e we FAN KN ; ( / ( Wu \\ { A, tp \ | ave C7 \ Sk rs C ITO. ee men rang {Sn ACG Pk Air ig Fig ie] 10. Styrax argenteus: a. leaf of var. argenteus; b. leaf of var. hintonii; c. leaf of var. ramirezii; d, e. upper & lower leaf epidermis of var. 246 Fig. 1. argenteus (500); f, g. upper & lower leaf epidermis of var. hin- tonit (500); h, i. upper & lower leaf epidermis of var. ramirezii (500 ). Styrax argenteus: a. trichomes on lower leaf surface of var. argen- teus (128); b. trichomes on lower leaf surface of var. hintonii (128 ); e. trichomes on lower leaf surface of var. ramirezit (128 x ); d. mature flower of var. argenteus; e. mature fruit of var. argenteus. 12mm \ 10mm 247 248 1Omm Fig. 12. Styrax conterminus: a. habit sketch; b, c. upper & lower leaf epi- i - d. trichomes on lower leaf surface (128%); e. ma- ture flower; f. mature fruit. | Vnnam 13f Fig. 13. Styrax obtusifolia: a. habit sketch; b, c. upper & lower leaf epider- mis (500); d. trichomes on lower leaf surface (128); e. mature flower; f. mature fruit. 6mm | 6mm amm jomm Hive | | | Poy tinal | Pep yh | 4e LAr | Fig. 14. Styrax ochraceus: a. habit sketch; b, c. upper & lower leaf epider- mis (500); d. trichomes on lower leaf surface (128); e. mature flower; f. mature fruit. 2951 Fig. 15. Styrax portoricensis: a. habit sketch; b, c. upper & lower leaf epi- 252 dermis (500); d. trichomes on lower leaf surface (128); e. ma- ture flower; f. mature fruit. Fig. 16. Styraxw glaber: a, habit sketch; b, c. upper & lower leaf epidermis (500 ); d. trichomes on lower leaf surface (128); e. mature flow- er; f. mature fruit. 293 6mm 17mm 166 S : 2 ep SS oe hectic ieee SALE Ses eo? 4 : ! ° e 7‘ \ a fyi 3 ° as Hdd ag ne e a) \, . e ° ” a ® “te: ovet ¥ “A rae) ae ie. Ses Ss Meee er Cee te ° od e\ Pee og ete . °, : l. e eee” oe oe _e : ah: . eo . eee - re wae 28 “3° Se, hos e , e ode eo y@ ° a oonds 3372 S.amenicana 2 i eee o 6%. - Pea ee women cabebe tt etetete, oe Mah cele Nec) OMe © ogre se 080s 06 9 ote of ee 2 %ee ; ®e. ee” be) e Ne 7 bs e eo. Sy : ¥ Var pulverulenta eee 24 oe es. MY : Pe o8| e : eee - ° en #i «asada? oie” ee eof 4 e Cia is ete eo at 8 e lad on _ y s aos ora : r ee © Se, . 0 aie tee oe vo oe Re : Sg rd Spore‘ e e oe e e ; be sie. e — . 7 a ° , 2% =so o oes? ot Paes e ? 7 , @ae® t °° ee. Se e % e e.°@ \ \ ee a ee : oaeet? ° oot. ; e oe? a MW oes ee ~ “o%ee Boece % 6 e °° ° >: Grand IToha “aoc ° mle eects e* : ® © f Siyrax americana var. americana, S. Fig. 17. Documented distribution of americana var. pulverulenta, and S. grandifolia. aie ve a Sy «var stellata \ oe “i \ ! . | | ee ge Ss youngae S.platanifolia — var platanifolia 7 ya 7 S. officinalis 5 var californica | ie 7 | ; =| aie a | ; toe | {ee res i _- ‘\ Ne es ; ex ae ; eae ) ‘ . % 4 \ als ; { \ cn 8 5 \ 1 @ . ee \ gets \ Le t i 7 ' Noetass ~y S. texana Fig. 18. Documented distribution of Styrax platanifolia var. platanifolia, S. platanifolia var, stellata, S. youngae, S. texana, S. officinalis var. californica, and S. officinalis var. fulvescens. 259 296 S.conterminus S. glabrescens var glabrescens Fig. 19. Documented distribution of Styrax jaliscana, S. conterminus, S. glabrescens var. glabrescens, and S. glabrescens var. pilosus. S.argenteus TR eye Van argenteus Co var hintonii Fig. 20. Documented distribution of Siyrax argenteus var. argenteus, S. ar- genteus var. ramirezii, and S. argenteus var. hintoniti. “a é S. glaber bad Fig. 21. Documented distribution of Styrax obtusifolia, S. ochraceus, S. por- toricensis, and S. glaber. aa a NOTES ON THE GENUS NAJAS (NAJADACEAE) ROBERT R. HAYNES and W. ALAN WENTZ Department of Botany, The Ohio State University, Columbus, 43210 and School of Natural Resources, The University of Michigan, Ann Arbor, 48104 Najas is a cosmopolitan genus of submersed aquatic plants with its great- est diversity in tropical and subtropical regions. Although floristic treat- ments have been prepared for certain areas, e.g., eastern United States (Clausen, 1936) and Panama (Wentz and Haynes, 1973), at no time since Rendle (1899) have all of the species, either worldwide or regional, been in- cluded in one revisionary treatment. Morphological variability within the group has been poorly understood. Because of the lack of a thorough re- visionary treatment and a poor understanding of the taxa, the genus is, we feel, in need of revision. Therefore at the present time, we are revising Najas in the Western Hemisphere. During the preparation of this treatment, we recognized two species (N. ancistrocarpa Magnus and N. wrightiana A. Br.) previously unreported from North America and a new variety of N. guada- lupensis (Spreng.) Magnus. These three taxa are discussed here. Najas ancistrocarpa was first collected in the United States by A. H. Cur- tiss (6865) in ‘“‘tidal creeks’? near Milton (Santa Rosa County), Florida on 3 Aug 1901. Fernald (1902) reported these specimens as N. conferta A. Br. (a synonym of N. wrightiana) and distributed them to many herbaria (GA, GH, MO, NY, UC, US, WIS) In 1947, R. F. Thorne collected N. ancistrocarpa from Open Pond (5537, GA) and Cane Water Pond (5538, GH, MO, UC) in Decatur County, Georgia. Thorne and W. C. Muenscher revisited the Milton, Florida location in 1948 and again collected the species (8612a, F, GA, GH, NY, US). All of the speci- mens collected by Thorne and Muenscher were identified as N. conferta. Most recently, N. ancistrocarpa has been collected in Lake Jackson, Leon County, Florida (Smith s.n., 1972, FSU, OS). This species should be sought in both fresh and brackish waters throughout the southeastern states. The vegetative organs of N. ancistrocarpa resemble those of N. minor All. and N. wrightiana by the presence of large teeth scattered alone the margin of quite narrow leaves. However, N. ancistrocarpa is easily distin- guished from the last two species by its fruits being so recurved they ap- pear crescent-shaped (ig. 1). We know of no other Najas with such curved fruits. Najas ancistrocarpa is native to Japan and, according to Miki (1935), has previously been known from only two Japanese islands, Honshiu and Shi- ~— ' Contribution from the Department of Botany, The Ohio State University (Paper No. 863). SIDA 5 (4): 259—264. 1974, 260 Fig. 1. Najas ancistrocarpa. A. Habit sketch (* 1); B. Fruit with involucre (X 9); C. Leaf (x 4%) koku. In addition to N. ancistrocarpa, other species in the aquatic flora of southeastern United States, e.g. Blywa aubertit and Ottelia alismoides (Thie- ret, Haynes, and Dike, 1969), are known to occur in isolated areas of North America and Eastern Asia. A second species, N. wrightiana, has recently been found in the United States. Clausen (1946) gives the range of this species as ‘‘Cuba, the coastal plain of Mexico north of the Isthmus of Tehuantepec, and the Yucatan Pe- ninsula.”’ We have seen specimens from the Dominican Republic, Brazil, 261 Venezuela, Guatemala, and British Honduras. The species is probably wide- spread throughout the West Indies and Latin America. Clausen (1946) recognized two subspecies based upon differences in leaf width and number of teeth along the leaf margin. We have examined nu- merous specimens from throughout the species’ range and do not think the segregation of infraspecific taxa is warranted. The first collections of N. wrightiana from the United States are from the Big Cypress Swamp in Collier County, Florida (Lakela 27796, ‘‘off Turner Read,” 24 Oct 1964, FSU, USF; Lakela 31526, Remuda Ranch Estates, 30 Jul 1968, USF). The Florida coliections of N. wrightiana apparently repre- sent the northernmost localities for the species. The collections we have seen are abundantly fruiting, and it is likely that the species will spread throughout southern Pe — Najas wrightiana is easily distinguished from other North American Najas by its large teeth scattered along the margin of quite narrow leaves, its short (1-1.5 mm) erect fruit with 5-angled areolae, and its bushy appearance due to axillary tufts of leaves and very short internodes (Fig. Variation in vegetative parts (plasticity) within individual taxa of aquatic plants has been recognized in Callitriche (Fassett, 1951), Halodule (Phillips, 1967), Nymphaea (Williams, 1970), Najas minor (Wentz and Stuckey, 1971), and Potamogeton (Haynes, 1973). These workers concluded that because of this plasticity, the taxonomy of the genera in question should rest primarily on reproductive characters. As a result of our own field work and the ex- amination of several thousand specimens we conclude that much of the vari- ability of N Oe ieee abets is vegetative plasticity and not of genetic origin. Clausen (1936 ), Ooststrcom (1939), and Wentz and Haynes (1973) discussed variability within N. aren and suggested that the complex should possibly be considered as several closely related species. It is possible that some taxonomists might consider this variation worthy of taxonomic recognition at the varietal level. We, however, regard varicties as morphogeographic subdivisions of a species that presumably reflect gene- tic differences (Kapadia, 1963). Thus far, in N. guadalupensis, we have found a definite correlation between a morphological type and its geographic dis- tribution only in a population in Florida. We have decided to consider the Florida population at the varietal level and not to give the other variants taxonomic rank. Clausen (1936) also mentioned that this population might be worthy of taxonomic recognition, The nomenclature of N. guadalupensis is difficult to explain. The taxon was named as Caulinia quadalupensis by Sprengel (1825). The original spe- cimen, subsequently deposited at Berlin, was destroyed during World War II (Gerloff, personal communication). We are uncertain which variety Spren- gel had before him. However, having seen specimens of only one variety from Guadaloupe—(the type locality), we consider that taxon as the typical variety. Several other varietal names of N. guadalupensis have been pro- posed. We have examined specimens of these and consider them to be the 262 » Fig. 2. Najas wrightiana. A. Habit sketch (x 34); B. Fruit with involucre 25); C. Leaf (x 2%) same taxon that occurs in Guadaloupe. Thus, the taxon limited to Florida has no varietal name. We therefore propose: NAJAS GUADALUPENSIS (Sprengel) Magnus var. floridana Haynes & Wentz, var. nov. Plantae glabrae, monoeciae. Caules 7-51 cm longi, 0.1-1.7 mm diametro, profuse ramosi, internodiis 0.1-9 cm longis. Folia 9-32.5 mm longa, divisa in laminas et auriculas. Laminae 0.5-2.1 mm latae, obtusae vel acuminatae apice, mucronatae; margines conspicuo serrulati, dentibus 18-42 unicellu- laribus, in uno latere; mucro dentibus lateralibus grandiore. Auriculae 1.2-2.5 mm latae, laminis latiores, rotundatae vel gradatim expansae prope basin, dentibus 5-8 in dimidio superiore quoque marginis, nullis prope basin; dentes similares sed illis laminarum grandiores. Flores solitari, staminati in axillis superis, carpellati in axillis interis; involucra ferruginea vel atro- 263 purpura, raro viridiba. Flores staminati, 1.5-2.4 mm longi; involucra abrupte angustata supra antheras in rostrum, 0.8-1.3 mm longa, apice lobis quatuor aculis; antherae 1-1.6 mm longae, 0.5-0.7 mm latae, theca una; filamenta emum 1-2.3 mm longa, basi dilatata. Flores carpelleti, 1-3.5 mm _ longi, fusiformes; involucra supra ovarium abrupte angustata in rostrum, 0.3—1.5 mm longa, findentia opposita carinae, apice lobis acutis tribus vel quatuor; styl usque ad 0.7 mm longi, lobis stigmaticis duobus, extendis supra rostrum involucrale. Fructus 1.6-2.2 mm longi, 0.3-0.8 mm lati, straminei vel per- pallidi-viridi, fusiformes; costa una, basaliter complanata, carinata; areolae inconspicuae, latiores quam longae, usque ad 0.1 mm longae, usque 0.2 mm latae, quatuor vel quinque angulatae, circa 20 serialibus longitudinalibus; semina fusiformia, stramineus vel lignicolor, extremis fuscat HOLOTYPE: UNITED STATES: FLORIDA: Dade Co.: cae in brack- ish water of ditch along St. Rt. 94 (east end) off U.S. Rt. 41 at edge of Ever- glades National Park, ca. 40 miles W of Miami, 5 Apr 1972, W.A. Wentz 670 (US!; isotypes, GH! MICH! OS!). The new variety (Fig. 3) can be distinguished from the typical varicty by its longer fruits (1.6-2.2 mm for var. floridana and less than 1.6 mm for var. guadalupensis). Also the leaves of the new variety at maturity are usually 2-3.5 cm long with 18-42 macroscopic teeth per side, while those of the typical variety are smaller and have about 100 minute teeth along each side. Najas guadalupensis var. floridana is widely distributed in rivers, streams, and ponds throughout Florida and Georgia. It is especially common in the Big Cypress Swamp and the Everglades region of southern Florida ACKNOWLEDGEMENTS We are grateful to Dr. Tod F. Stuessy for assistance with the Latin de- scription; to Dr. David J. Keil and Mr. Marvin L. Roberts for reading the manuscript; to the curators of herbaria from which specimens were bor- rowed; to Mrs. Jan S. Wentz for the illustrations; and to Mr. Howard F. Schnepp who made it possible to conduct field work in Florida during April 1972 REFERENCES eee - T. 1936. Studies in the genus Najas in the northern United States. Rhodora Oe Najas arguta in oo America and its relationship to N. wrightiana. Bulk pe Bot. Club 73: 363 FASSETT, N. C. 1951. Cle . the New World. Rhodora 53: 137-155, 161-182, 185-194, 209-222. ey eee M. L. 1902. Some little known plants from Florida and Georgia. Bot. Gaz. 33: mente R. R. 1973. A revision of North American aie ais subsection Pusilli Cour Nga eae Ph. D. Dissertation. The Ohio State University. 140 p. KAPADIA, Z. J. 1963. Varietas and ee a suggestion tow oe greater uniformity. oes 12: 257-259, MIKI, S. 1935. New water plants in Asia Orientalis II. Bot. Mag. (Tokyo) 49: 687-693. OOSTST ROOM S. J. van. 1939. Some notes on a collection of aquatic phanerogams from l Netherlands, fae Indies Islands, and from Venezuela and Colusa. Recueil ‘Trav Bot. Neéerl. 5-708 264 PHILLIPS, R. C. mae et species of the seagrass, Halodule, in Florida. Bull. Mar. Sci. Gul Caribbean 17: 672 a A. B. hae “ systematic revision of the genus Najas. Trans. Linn. Soc. London, 444, oe NG EL, C. 1825. Caulinia, p. 2 ting: THIE RE T, J. W., Re R. HAYNES, and D. H. DIKE. 1969. Blyva aubertii: (Aydrocharita- ceac) in i. New to North America. Sida 3: 343-344. WENTZ, W. A. and R. R. HAYNES, 1973. Family 3B. ae egy as _ oodson, R. Jr. and R. W. Schery. Flora of i ama. Ann. Missouri Bot. Gard. WENTZ, W. A. and R. L. STUCKEY. 1971. The changing yA oe Fr a7 genus Najas Najads reac) in Ohio. Ohio J. ae 71: 292-302. 70. Investigations in the white waterlilies (Nymphaea) of Michigan. ), In Linneaus, C. Systema vegetabilium, vol. 1. Got- a ” Fig. 3. Najas guadalupensis var. floridana. A. Habit sketch (x 34); B. Fruit without involucre (« 25): C. Leaf («x 2%); D. Leaf axil with fruit (x 9): Ie. Leaf apex (« 30); F. Leaf margin (x 30). A NEW SUBFRUTICOSE ERIOGONUM (POLYGONACEAE) FROM NORTHERN MEXICO JAMES L. REVEAL Department of Botany, University of Maryland, College Park, MD 20742 and National Museum of Natural History, Smithsonian Institution, Washington, DC 20560 Recent floristic studies by B.L. Turner of the University of Texas on vege- tation restricted to gypsophilous soils in southwestern United States and ad- jacent northern Mexico have resulted in the discovery of several interesting plants (Turner, 1972a, 1972b; Turner & Powell, 1972). Among the collections obtained by Turner from near San Roberto Junction in Nuevo Leon is a new Eriogonum (Polygonaceae) here described as: ERIOGONUM turneri Reveal, sp. nov. A EF. jamesii Benth. in DC. var. undulata (Benth. in DC.) Stekes ex Jones foliis lanceolatis 6-11(15) mm longis et 4-5(6) mm latis, inflorescentiis compactis capitatis vel biumbellatis, plantis subfruticosis et compactis differt. Plantae caespitosae et subfruticosae 2-10 em altae et 5-15(20) cm latae, apicibus lignosis et ramosissimis; folia lanceolata, laminis 6-11(15) mm longis et 4-5(6) mm latis, subtus albo-to- mentosis, supra glabris vel subglabris, marginibus crispatis, petiolis brevi- bus et 3-6 mm longis, floccosis; caules graciles, (1.5)2—5 cm longi, tomen- tosi; inflorescentiae capitatae vel biumbellatae, ad 4 cm longae et lataec, ramis tomentosis; bracteae 3-5, 0.7-4 mm longae; peduncli nulli; involucra solitaria, campanulata, 3-3.5 mm longa et 3.5-4 mm lata, sparse tomentosa, 5-6-lobatis, bracteolis brevibus, 0.8-1 mm longis et hirsutulosis, pedicellis 3-4 mm longis et glabris; flores stipitati, albi vel subrosei, 4.5-6 mm longi, puberuli, tepalis dissimilaribus, extimis tepalis oblongis, 3-3.5 mm longis et 0.9-1.2 mm latis, intimis tepalis spatulatis, 4-5 mm longis et 1.8-2.5 mm latis; stamina exserta, 3-4 mm longa, filamentis pilosis basi, antheris roseis vel purpureis, 0.5-0.8 mm longis, oblongis; achaenia infuscata, 4-4.5 mm Low cespitose subfruticose perennials 2-10 cm high with compact, woody caudex branches forming a mat 5-15(20) cm across, this arising from a stout woody taproot, the bark of the caudex branches and the taproot reddish brown, exfoliating in loose shaggy plates; leaves basal, in indistinct basal rosettes restricted to the base of the herbaceous flowering stems or at the tips of exposed caudex branches, the leaf blade lanceolate, 6-11(15) mm long, 4-5(6) mm wide, densely white-tomentose below, green and subgla- brous to glabrous above, the midvein remaining pubescent above in most, the margin enrolled and crispate, the apex acute, the base truncate to round- ed, the petiole short, 3-6 mm long, floccose to thinly tomentose, reddish SIDA 5 (4): 265—267. 1974. 266 under the tomentum, the petiole base triangular, 0.8-1 mm long, 0.5—9.8 mm wide, persistent and covering the upper part of the caudex branch, glabrous within, thinly pubescent without; flowering stems erect, slender, (1.5)2-5 cm long, thinly tomentose, often reddish under the tomentum; inflorescences capitate and consisting of a single terminal involucre with the head 1-1.5 cm across to biumbellate and up to 4 cm high and across, the branches thinly tomentose; bracts 3-5 at the base of the involucre and at cach node, usually with 1-3 semifoliaceous ones with one distinctly longer than the others, 2-4 mm long and similar to the basal leaves only more reduced, the others reduced and scale-like, 0.7-1.5 mm long, all connate at the base: peduncles lacking; involucres solitary, campanulate, 3-3.5 mm long, 3.5—4 mm wide, thinly tomentose without, glabrous within, the 5-6 rounded shal- low teeth less than 0.5 mm long, the bractlets small and peg-like, 0.8-1 mm long, densely hirsutulous, the pedicels numerous, 3-4 mm long, glabrous; flowers stipitate, pinkish-white with greenish to pinkish midribs and bases at anthesis, becoming pinkish in fruit, 4.5-6 mm long including the 1-1.3 mm long stipe, densely pubescent without especially along the midribs and stipe, glabrous within except for a few seattered minute glands, the tepals dissimilar, united about 14 the length of the flower, the outer whorl oblong, 33.5 mm long, 0.9-1.2 mm wide, becoming turned outward in fruit, the inner whorl spathulate, 4-5 mm long, 1.8-2.5 mm wide, remaining erect, the pu- bescence remaining rather dense from anthesis to fruit: stamens slightly exserted, 3-4 mm long, the filament thinly pilose basally, the anther dark red to purple, 0.5-0.8 mm long, oblong; achenes light brown, 4-4.5 mm long, the narrow base tapering to a long, sparsely pubescent, 3-angled beak. MEXICO: Nuevo Leon: Along Mexico Highway 57, 15.5 mi S of San Roberto Junction, 2.3 mi N of El Refugio, 0.5 mi W of highway on a small raised ridge of gypsum, with Isocoma gypsophila, Strotheria gypsophila, Frankenia margaritae, and Calylophus, 5700 ft elevation, 19 Sep 1972, Reveal & Hess 3147 (Holotype, US; 30 isotypes to be distributed from US). Additional specimens examined: MEXICO: Nuevo Leén: 16 mi S of San Roberto Junction, 0.5 mi W of highway, 24 Oct 1970, Turner & Crutchfield 6310 (TEX). The Turner Buckwheat, Eriogonum turneri, is a member of the subgenus Oligogonum Nutt, a complex of some 30 species typified by E. umbellatum Torr. (Reveal, 1969). The new species closely approaches I). jamesii Benth. in DC. var. undulatum (Benth. in DC) Stokes ex Jones, found in mountains and foothills of northern Mexico northward into western Texas and south- eastern Arizona. The new species may be distinguished by its short narrow leaves [5—11(15) mm long and 4—5(6) mm wide versus 1-2 em long and 0.5-1 em wide for var. undulatum]|; the short compact inflorescence (up to 4 cm long for E. turneri and up to 1.5 dm long for var. undulatum); and the generally smaller fruiting flowers (up to 6 mm long in the new species and up to 9 mm long in var. undulatum). Eriogonum jamesii occurs typically on sandy loam to 267 clayey soils but, so far as known, not on gypseous soils. Likewise, var. wn- dulatum in Mexico is found mainly in high mountains in well drained soils mostly above 7000 ft elevation. In most places, var. undulatum forms small to rather large mats (the latter especially in arroyos and on gradual rocky slopes), with its large leaves being a conspicuous part of the plant as is the tall (up to 5 dm) habit of the flowering stems and inflorescence. In E. turneri, the plants form very small compact mats, usually less than 2 dm across, on exposed valley bottoms at 5700 ft elevation. The plants, with their small in- conspicuous leaves, rarely exceed 1 dm in height. Both entities have cris- pate leaf-margins, unlike var. jamesii. It is a pleasure to name this species for Billie L. Turner, a Texan whose stature in plant taxonomy far exceeds that of this diminutive taxon. Field work was supported by NSF grant GB-22645, REFERENCES = age AL, J. L. 1969. The subgeneric concept in Eriogonum (Polygonaceae). Pages 229-249 2 J. E.G ae ed., Current topics in pan science. Academic Press, New York. TURNER, B. L. ee Two new species of Isocona (Comp Astereae) from north- central meee Sida 5: 23-2 = sarees 72b. oe (Coaipesiens. ‘Tageteac) a new monotypic genus from north- enna eens Am. J. Bot, 59: 180-182. = and A. M. POWELL. 1972. A new aypsophilic cet (Leguminosae) from north-c Seqteal Mexico and adjacent Texas. Phytologia 22: STUDIES IN AMERICAN RUBIACEAE 2. AYUQUE, BALMEA STORMAE, AN ENDANGERED MEXICAN SPECIES: F. R. FOSBERG Smithsonian Institution, Washington, DC 20560 It is difficult to establish the status of a continental tropical plant species as endangered, since the ranges of most of them are so inadequately known. The rubiaceous tree Balmea stormae, locally called ayuque, may conceivably be commoner than suspected in parts of Mexico as yet unstudied by botan- ists. However, one circumstance makes it likely that this species will soon become very rare in any place where it may be discovered. Due to its con- ical habit with pointed top when young, it is commonly cut and sold in mar- kets in the Uruadpan area as a Christmas tree. According to Miss Marian Storm, of Guadalajara, this use of Balmea arose when laws were enforced making it illegal to cut conifer saplings for this purpose. Miss Storm, noted writer and poet who brought ayuque to the attention of Professor Maximino Martinez and for whom he named it, has, in a series of letters between the years 1965 and 1971, described the gradual disappear- ance of this plant from the pedregales or lava beds near Uruapan where she discovered it in 1941. There is an amusing account of ayuque and her ad- ventures in bringing it to botanical notice in her charming book Enjoying Urudpan (1945; pp. 452-459) and a poem about it in her Poems of Sun and Snow (1955; p. 36) It seems worthwhile to offer an account of the present status and knowl- edge of Balmea stormae, with the hope that Mexico, with its growing inter- est in conserving its natural resources, may afford this fine potential orna- mental tree enough protection to enable it to survive and continue its role as a thing of beauty. Attempts are being made to bring it into cultivation. It might do well in warm temperate and subtropical rather dry regions. Balmea, a monotypic genus of the Cinchona tribe of the Rubiaceae, is en- demic to a few small areas in southern Mexico and Guatemala. It was first brought to the attention of the botanical public by Miss Storm although it had been collected as early as 1935 by George Hinton and had been given a manuscript name (as a Randia) on the basis of a 1939 Leavenworth and Hoogstraal collection sent to Paul C. Standley. BALMEA Martinez, Bull. Torrey Bot. Club 69: 438-441, 1942; An. Inst. Biol. 13(1): 36-41, 1942. This genus belongs to the group in Cinchonae characterized by the corolla contorted in bud, the circumscissile caducous calyx, and the elevated epi- ‘The first article of this series was published in SIDA 2: 386-389. 1966. SIDA 5 (4): 268—270. 1974. 269 gynous disk in fruit. In habit Balmea is a tree, pyramidal or conic when young, irregular when older, with broad, cordate leaves, slender, usually several-times trichotomous cymes, and seeds with a wing surrounding the body and tapering at both ends. Its closest relative, Cosmibuena, has elon- gate, often bifid wings at each end of the seed. Hillia, likewise close, has a small wing at one end of the seed and a small tuft of hair at the other. These seed differences are about the only tangible ones among the three genera, Otherwise Balmea is set off by a series of characters, any one of which would be regarded by itself as of scarcely specific importance. It differs essentially from Hillia only in its non-comose seeds (simply winged basally and apically, as in Cinchona, rather than with a conspicuous coma apically) and more superficially in its thinner much broader leaves with long sharply distinct petioles and in its pendent, usually much more com- plex inflorescences, those in Hillia being of 1 (often sessile) to 3 flowers. From Cosmibuena, Balmea differs in its pendent, more complex inflores- cence, its red rather than white flowers, its broad, cordate, much more veiny leaves, and its never bifid seed wing. In actuality the three genera could best be treated as one. I am deferring this reduction until all species of the three ean be placed side-by-side to be certain that no significant breaks among them exist. Martinez relates his genus also to Blepharidium Standl. This differs from the genera mentioned above in the aestivation of the four corolla lobes in bud, these being conspicuously overlapping but not contorted. The leaf vena- tion is more pinnate with many more veins on a side. It is in this general affinity but related also to Hintonia and several other genera. On both ends the seeds have wings that are broadly obtuse or rounded and erose or tri- dentate at the tips. BALMEA STORMAE Martinez, 1. c., figs. 1-6; 1. ¢., figs. 1-4. Small tree or shrub 4-10 m, entirely glabrous, bark thin, wocd fine grained, pale pinkish-cream; leaves suborbicular, somewhat broader distally, acu- minate at apex, cordate to subcordate at base, with 4-6 principal veins of unequal strength on a side, these subopposite to alternate, more prominent beneath, blades up to 9 « 14 cm, 2—3.5 cm long, apparently leafless at time of fruiting (McVaugh 22704); stipules early caducous; cymes terminal, pen- dent, once to usually twice (rarely 3 times) trichotomous, often with 1-2 extra flowers at basal ramification, peduncle 3-7 cm long, caducous subulate bracts at ramifications, branches 1-2 cm long, pedicels 1-3.5 cm long, elon- gating and becoming recurved in fruit; flowers becoming very fcaccant at night; calyx lobes 5, subequal or somewhat unequal, lanceolate, to 7-8 mm long, blunt, calyx caducous shortly after anthesis; corolla deep red to dark maroon, tubular, somewhat dilated distally, with 5 short recurved oval ob- tuse lobes, tube 20 to rarely 27 mm long, 5 mm wide at summit, on type col- lection notably marked with short white lines, lobes to 5 * 7 mm; anthers included, 5(-7), dorsifixed just below mouth of tube, broadly linear, panne at base; style 22-23 mm long, stigma somewhat enlarged, bifid, the blunt 270 lobes stigmatic on inside, disk (after corolla and other flower parts are shed), strongly elevated, conic, tipped with a short truncate style remnant, capsule oblong, 22-30 mm long, firm, dehiscing first septicidally, to the full length, then the disk loculicidally, the valves diverging somewhat, placentae lanceclate, fleshy, attached to septum, seeds reddish-tan, prominently winged, broadly lanceolate, (3)4-6 mm long, margins somewhat erose, one end narrowly acuminate, cellular reticulate, the other broader, acute to ob- tuse, cells arranged in a fan-like pattern, body of seed elliptic, 2 mm long. The specific epithet has been spelled several ways, but the simple fem- inine ending used above was the way it was published originally. The Eng- lish and Spanish versions of the original description were published almost simultaneously; I do not know which was first. The same ending was used in both publications. Specimens examined: MEXICO. Temascaltepee: Nanchititla, 19 Jun 1935, Hinton 7894 (US, 2 sheets, F-); Michoacan: Palo Verde, near Urudpan, Sep 1941, Storm (US); Palo Verde, Pedregales de Jicalan, near Urudpan, summer 1941, Storm (US); Palo Verde, Pedregales de Jicatlan et San Pedro, near Uruapan, Martinez 3400 (US); Palo Verde, near Urudpan, 20 (or 21) Sep 1941, Martinez 3400 es type, 2 sheets marked ‘‘tipo,’’ one selected and marked “‘lectotype’’ by F. R. Fosberg, 1972, because it is somewhat more complete; US, 2 sheets: i same locality, Martinez s. n. (F, 2 sheets without original labels, but marked ‘Isotype’? and, on Field ane label, “México, M. Martinez, Rec. Jan. 1944,’ probably duplicates of the type); Jicalan (as Jicatlan), near Uruapan, Martinez 3400 (MEXU, 4 sheets; sev- eral gatherings made on different days were all numbered 3400); La Laguna entrada al canon del Mal Pais, 15 Sep 1961, Alonso & Ramirez (MEXU), also 1961, Ramirez & Alonso (MIEXU); N.W. Aguililla, 6-7 km S of Aser- radero Dos Aguas, 2000 m, 3 Mar 1965, McVaugh 22704 (NCB) (fruiting twigs without leaves, both fruits and seeds longer than on type); 2 mi §S of Uruapan, 5577 ft, 29 Jul 1941, Leavenworth & Hoogstraal 1229 (F). GUATEMALA. Dept. Huehuetenango: NW of Cuilco, 2/3 way up Cerro Chiquihui above Carrizal, tree 25 ft tall, 1350-2300 m, 17 Aug 1942, Steyer- mark 50811 (US, F) (sterile but has the right leaves); Dept. Jalapa: Potrero Carrillo, 14 mi NE of Jalapa, 1500-1900 m, 11 Dee 1939 Steyermark 33056 (US, F) (Epiphyte with branches fleshy, leaves not or scarcely cordate on US sheet, cordate and normal for species on F sheet; stipules ovate); Dep Zacapa: San Lorenzo, Sierra de las Minas, shrub 10 ft tall, 1600 m, 24 Jan 1942, Steyermarl: 43177 (F). HERBARIUM AND FIELD STUDIES OF KENTUCKY PLANTS Il. NEW STATE RECORDS AND RARITIES EDWARD T. BROWNE, JR.’ Department of Biology, Memphis State University, Memphis, TN 38152 This paper records several species that are apparently new to Kentucky or else are extremely rare in the state. Specimens in the combined Gray Herbarium - Arnold Arboretum Herbarium, the New York Botanical Garden Herbarium, and the United States National Herbarium were examined. I ex- press appreciation to the curatorial staffs of these institutions. Collection numbers of most collections cited are those of the writer and his wife, here abbreviated EMB & ETB. Specimens are deposited in the Memphis State University Herbarium unless otherwise indicated, and duplicates are being distributed elsewhere. Cyperus rotundus L, This is a serious weed farther south. Although the species has been established in Kentucky for some time, this is apparently only the second report. Gunn (1968) listed C. rotundus for the area included in his eight-county survey but did not indicate a more specific locality. Ac- cording to Mr. D. C. Byers of Monticello, Kentucky, the species was intro- duced there with strawberry plants from Alabama in 1961, and it is now all over Wayne Co., carried about in soil on cultivators. Collection data: vege- table garden, 212 S$. Main, Monticello, Wayne Co., 20 May 1966, HMB & ETB 11833 (with John Warden) Wolffia papulifera C. H. Thompson. This small aquatic monocot is previ- ously unreported for the state; I found no Kentucky specimens in the her- baria examined. Gentry (1963), in an unpublished thesis, suspected that this and Lapsana communis L. (see below) were new state records. ae ees lifera periodically occurs in great numbers on the surface of pon tion data: duck ponds, Lexington Cemetery, Lexington, Fayette an. io Oct 1965, EMB & ETB 11590.3; N side of Lecompte’s Bottom in lake along Ken- tucky River, Henry Co., 20 Oct 1962, J. L. Gentry, Jr. 846. Chaenomeles speciosa (Sweet) Nakai (Chaenomeles lagenaria of auth., not (Loisel.) Koidz.) This, collected as an escape, has not been previously reported from Kentucky. I have seen no specimens from any of the sur- rounding states although Sharp (1960) reports Cydonia japonica Pers. as persistent in Tennessee, and this may be a synonym. Collection data: open mesophytic woods and clearings on river flood plain just N of jct. Ky. 986 at bridge over Little Sandy River, Rosedale, Carter Co., 23 May 1965, HMB = oO ~* Contribution No. 5, Kentucky Flora Project, Memphis State University. ppreciation is expr ae to the Faculty Research Committee, Memphis a University, for providing funds to enable me to visit the herbaria mentioned in this pape SIDA 5(4): 271—273. 1974. 272 & ETB 10173; 0.6 mi E of jet. Ky. 174 and US 60, between highway and C. & O. RR tracks, Rowan Co., 23 Apr 1966, EMB & ETB 11680, Arachis hypogaea L. This species is not grown commercially in Kentucky, and it is seldom cultivated in gardens. There are neither reports in the liter- ature nor records in the herbaria of its occurrence in the state (the Arachis specimens in US were out when I checked there). It is doubtful that A. hypo- gaea would survive winter conditions in Kentucky or much farther north. We collected A. hypogaea at a garbage dump, which we revisited ten years later to observe if the species had survived there. No trace of it was found. Considerable disturbance was evident—fire, flooding, clearing, and cultiva- tion. Any one of these could well account for its absence as well as that of many of the native species originally there. Collection data: garbage dump, Fk end of US 62 bridge over Green River, Ohio Co., 8 Jul 1962, EMB & ETB 5766 (KY), Erodium cicutarium (L.) L’Her. This species is apparently spread as an agricultural contaminant in fertilizers and/or feed: its habitat indicates dis- semination of this type. It might be expected to occur in Kentucky based on the range given in Gleason and Cronquist (1993): ‘ . . . throughout most of the U. S.” Unfortunately, folders 2-7 were out when the NY collections were checked, and so positive confirmation of FE. cicutarium as a species new to Kentucky must rest primarily on the literature. Collection data: off Cooper Drive along roadside by University of Kentucky Herbicide Building, Lexington, Fayette Co., 27 Apr 1965, Bruce W. Mook 70; in field, Agri- cultural Experiment Station Main Farm, University of Kentucky campus near Horticulture Herbicide House, Lexington, Fayette Co., 26 Apr 1965, Frank Melton s.n. (KY) Huonymus kiautschovica Loes. Common in cultivation in areas of dense population. Seeds are spread by birds, and the species is a frequent escape. Some may identify this as I. Fortunei (Turez.) Hand.-Mazz., a prostrate, evergreen species; FE. kiautschovica is tardily deciduous and upright unless growing against a tree or wall in which case it becomes attached by adven- titious roots (Rehder, 1940). Young BE. kiautschovica may be somewhat trail- ing and evergreen, but older plants become almost devoid of leaves by the end of winter. The species grows against almost every tree on the University of Kentucky campus. Collection data: plants established under shrubs and in fence rows from seed scattered by birds (no cultivated plants of this spe- cies are known to grow in the near vicinity), 1234 Kastle Rd., Lexington, Fayette Co., 19 Mar 1966, EMB & ETB 11634. Hedera helix L. This is a relatively infrequently cultivated ornamental in central Kentucky, where it is near its northern limit. There are no Kentucky specimens in the herbaria visited. Only Gunn (1968) and Duncan (1967) pre- viously listed this species as an escape in Kentucky. Duncan's report is based on the collection cited, Collection data: roadsides and C. & O. RR right-of-way 6 mi FE. of jet. Ky. 182 and US 60, Aden Springs, Carter Co., 23 Apr 1966, EMB & ETB 11713. 273 Chaenorrhinum minus (L.) Lange. Reported previously only by Gunn (1968). This species may be much more widespread and abundant than his and my reports indicate since it is usually small and easily overlooked. Col- lection data: 2.1 mi S of bridge over Hinkston Creek, roadside between high- way (US 68) and L. & N. RR tracks, Millersburg, Bourbon Co., 29 Jun 1966, EMB & ETB 12139. Artemisia ludoviciana Nutt. This species, an old, widely-grown ornamen- tal, may not be native to Kentucky. In both places where its varieties were collected, oe plants had obviously spread from cultivation—which may be expected to occur more frequently near homes now abandoned due to ad- verse economic conditions especially in the Appalachian region. Artemisia ludoviciana var. gnaphalodes is listed by Gunn (1968), but there are no other literature references or specimens in the herbaria. Collection data: var. ludoviciana: growing wild on roadside at jet. Ky. 94 and Ky. 307, Hickman , 29 Jun 1962, EMB & ETB 5636, !Arthur Cronquist; var. gnaphalodes (Nutt.) T. & G.: Ky. 174 roadside, 1.6 mi W of Rowan-Carter Co. line, be- tween highway and C. & O. RR tracks, Rowan Co., 15 May 1966, KMB «& ETB 11787. Lapsana communis L. This introduction is new to eae It might have been expected there, for Gleason and Cronquist (1963) sta “ . now found throughout our range’ and I saw collections from nae West Vir- ginia, and Virginia. Collection data: 1.3 mi W of jct. Ky. 389 and Ky. 202, hillside, edge of deciduous woods and grown-up field, Drennon Chapel, Henry Co., 3 Aug 1962, J. L. Gentry, Jr. 607. QD = REFERENCES DUNCAN, W. H. 1967. Woody vines of the southeastern states. 3: 1-76. GENTRY, J. L., JR. 1963. Vascular plants of Henry Co., ee Unpublished M. S. Thesis, Universes of Kentucky. near H. A., and A. CRONQUIST. 1963. Manual of vascu rand, Princeton, NJ. GUNN. Cc. R. 1968. a flora of Jefferson and seven adjacent counties, Kentucky. Ann, Soc. Nat. Hist. 1-322. ee A. 1940, i. ual of cultivated trees and shrubs. 2nd ed. Macmillan, New York. SHARP, A. J. 1960. A preliminary checklist of dicots in Tennessee. Knoxville. (Mimeo.) ar plants. D. Van No- STUDIES OF THE SOUTHEASTERN UNITED STATES FLORA. IV. OLEACEAE JAMES W. HARDIN Department of Botany, North Carolina State University, Raleigh, N. C. 27607 The Oleaceae is represented in our area by eight genera, four of which were introduced as ornamentals and have since become established to vary- ing degrees in our flora. In providing the treatment of the Oleaceae for the Southeastern Flora Pro- ject, a number of taxonomic decisions had to be made which demand some explanation prior to publication of the manual. The treatment of the Oleaceae for the “Generic Flora of the Southeastern United States’? (Wilson and Wood, 1959) should be consulted for a thorough description and discussion of the family, genera, and relationships. The distribution of species is based on the specimens seen at A, DUKE, FLAS, GH, NCU, NSC, SMU, and US. It is imperative for anyone working on plants of the southeast to see—as a bare minimum—the specimens of A, GH, NCU, SMU, and US. Key to Genera 1. Leaves pinnately compound, or reduced to one leaflet with an articulated petiolule; fruit a samara or 2-lobed berry. . Trees or large shrubs: flowers apetalous; fruit a samara; leaves usu- ally pinnate (Tribe Fraxineae) . I. FRAXINUS. 2. Shrubs; flowers with showy corollas: fr uit a 2- aubed Heer leaves pin- nate or reduced to one leaflet (Tribe Jasmineae) . . Il. JASMINUM. . Leaves simple or occasionally ternate in No. Ill; fruit a capsule, drupe, or drupe-like unlobed berry. 3. Flowers yellow, appearing before the leaves (Tribe Forsythieae) , Il. F ORSYTHLA. 3. Flowers white, lavendar to purplish, or apeleious: appearing before or with the leaves. a 4. Fruit a loculicidal capsule: flowers lavender to purplish or rarely white (Tribe Syringeae) . . . . . .) .) .) IV. SYRINGA. ! 1 from a manuscript and notes compiled for the Vascular Flora of the Southeastern United | es The area covered in this treatment is bounded by and includes Delaware, Maryland, West Virginia, Renmeky Arkansas, and Louisiana. The format follows Radford Any needed modifications or suggestions should be corrections can be made before the manual is in The first three parts of this numbered series with the same Mitchell Sci. Soc. 87 (1), 87 (2), and 88 (1). Approved as Paper No. 4004 of the Journal Series, NCSU Agric. Exp. Station. press. title are in a Journal Llisha SIDA 5 (4): 274—285. 1974. 275 4, Fruit a drupe or berry; flowers white or apetalous (Tribe Oleeae). . Corolla absent; flowers in scaly, short, lateral racemes or cymes; calyx minute or lacking . . V. FORESTIERA. 2. Corolla present; flowers in panicles: eal present. 6. Petals linear, united at base only; young inflorescence spreading or VI. CHIONANTHUS. 6. Hts ae biead aii ehoxe Bove a distinct funnelform tube; young inflorescence erect or merely spreading. . Inflorescence a few-flowered axillary panicle; leaves evergreen; drupe unilecular . . . . . . VII. OSMANTHUS. 7. Inflorescence a many- Mowened femme panicle; leaves decidu- ous or evergreen; drupe-like berry 2-locular VIIL. LIGUSTRUM. I. FRAXINUS L. Ash. This genus of some 65 species is represented in the southeastern U. S. by six species. Their relationships are discussed by Wilson and Wood (1959) and indicated in the ke The taxonomic treatment is a conservative one. The problematic complex- es are in Subsection Melioides where there is both ecotypic and ecophenic variation, as well as hybridization. All the complexes are in need of further biosystematic studies. 1. He deciduous or lacking; terminal bud scales foliose (Subsect. Fraxi- aes 4-angled; lateral leaflets petiolulate; flowers with small de- ‘iduous calyx Se Ge we oe oe eS .F. quadrangulata. 2. Branchlets hevele: ee leaflets dessiiee flowers without calyx . FF. nigra. 1. Calyx present and persistent in fruit; found pie ee at foliose (Subsect. Melioides Endl.). 3. Leaves papillose beneath; wing of samara terminal or only slightly de- current along upper third of body . . : 3. Ff. americana, 3. Leaves not papillose beneath; wing of samara decieen to near middle of body or beyond. 4. Body of samara flattened and winged to the base . 4. I’. caroliniana. 4. Body of samara terete or nearly so and winged to near the middle or slightly beyond. . Petiolule of lower leaflets winged nearly to base; samara wing usu- ally less than 7 mm. wide, and body less than 2 mm. wide = 2 5. F. pennsylvanica. 5. Petiolule of lower leaflets winglacs: samara wing more than 7 mm. wide, and body more than 2 mm. wide . . . . 6. F. profunda. Oo 1. F. quadrangulata Michx., Blue Ash. Dry or moist rich woods; var. prov., chiefly IP, Ala., Ark., Ga., Ky., Tenn., 276 W.Va. (Okla., Mo., Ill., Ind., Ohio). The distribution of Blue ash is spotty and in a number of physiographic provinces. See Little (1971, map 128-E) for details. 2. F’. nigra Marsh., Black Ash Wet woods, stream banks, bogs or swamps; pied. Del., Va., mts. Md., W.Va. (Il., Ind., Ohio, Penn., N.J.). Gleason and Cronquist (1963) record this species for Ky. No specimens have been seen to verify this record and neither Little (1971) nor Meijer (1971) include it for the state. A chromosome count is needed for this species. I’. americana L., White Ash. Rich woods, dry ae well-drained flat ne all ep SE (ALL). Incl. I’. biltmoreana Beadle, F’. catawbiensis Ashe acl. F. a, var. bilt- moreana (Beadle) J. Wright ex Fern—I*, R; incl. P. a. var, microcarpa Gray—I, Although all provinces are represented, White ash is absent from most of the lower woods of the Atlantic and Gulf Coastal Plain and Mississippi em- bayment. See Little (1971, map 126-]«) for details. This is one of the extremely polymorphic species, or species complexes, involving ecotypic (Wright, 1944a) and ecophenic variation, three ploidy evels (2n—46, 92, 148), plus hybridization (Miller, 1955). The Biltmore ash has frequently been distinguished as a species or variety by various authors. Although the extremes, involving a number of characteristics seem clear enough, there is neither a real morphological discontinuity between them, nor a clear ecological or geographical separation. In some areas, such as Mississippi, Biltmore ash occurs in acid soil while White ash is a calciphile, but this does not seem to hold up elsewhere. One interpretation (Miller, 1955) is that the Biltmore ash represents a hybrid between F’. americana and I’, pennsylvanica. This needs a thorough investigation, although sub- stantiated by the work of Santamour (1932). Variety microcarpa Gray seems to be a small-fruited form that appears randomly through the population. The type named by Ashe as F’. catawbiensis from the piedmont of Georgia and the Carolinas seems to be a pubescent form with some characteristics reminding one of either IF’. caroliniana on one hand or F. profunda on an- other. This needs critical study. Until the pattern and origin of the variation and presence of any real dis- continuities are known more exactly than now, I prefer to think of White ash as one polymorphic species without segregate species or varieties. See also Little (1952) for a discussion of varieties. 4. F. caroliniana Mill., Water Ash, Pop A., Carolina A. Low woods, river bottoms, cypress ae SWAMPS, Sa aanaaie chiefly ep., Ala., Ark., Fla., Ga., La., Miss Incl. F. el Nutt. _—s: incl. FE. c. var. saeaae (Curtis) Fern. & Schub., F. c. var. cubensis (Griseb.) Lingelsh.—F. AS oi by the synonymy above and the additional names given by aw — 277 Little (1952, 1953), this is an extremely variable species. In fact, Fernald (1950) considered it ‘‘our most variable species [of ash]’’ and recognized three varieties each with two forms. The variation involves the shape and size of samaras, and pubescence of petiole, rachis, and leaflet. As with White ash, the variation seems fairly random without any real morphological- ecological-geographical types that should be considered distinct at any level. o. F. pennsylvanica Marsh., Green Ash, Red A Moist or wet alluvial soils, river and stream margins, swamps; all prov. SE except penin. Fla. (ALL) Incl. F. darlingtonit Britt., FY. smallit Britt-—S; incl. F. p. var. subinte- gerrima (Vahl.) Fern.—F, A detailed distribution map is given by Little (1971, map 130-E). The variability in this species is seen in the pubescence, shape and tex- ture of the leaflets, and shape and size of the samaras. Three ecotypes have been distinguished by Wright (1944b). Again there appears to be no real dis- continuity between any of the forms. The glabrous variety subintegerrima is recognized by a number of authors but it grows with var. pennsylvanica and there is apparently no barricr to interbreeding. I view the variation in pubescence in this species as similar to that in White ash, thus for the sake of consistency, I am not recognizing varieties. e common names are somewhat confused. When the more common and widespread glabrous var. subintegerrima was recognized, it was called Green ash, and the pubescent var. pennsylvanica was called Red ash. When the two varieties are not distinguished, Green ash is the preferred common name and the one used by the U. S. Forest Service (Little, 1953). 6. F. profunda (Bush) Bush, Pumpkin Ash. Swamps and river bottoms; chiefly cp., SE except Del., Ky., W.Va., (Mo., Ill., Ind., Ohio). Incl. F. michauxii Britt.—S; F. tomentosa Michx. f.—R, The Pumpkin ash is possibly either an autopolyploid of F. pennsylvanica or an allopolyploid (hexaploid n=69) derivative of F. americana (n=46) « pennsylvanica (n=23). Wright (1944b, 1957, 1965) and Miller (1955) discuss this further. This needs a thorough investigation. The name of this species has been shifted back and forth between F. pro- funda and F. tomentosa for a number of years. According to Little (1952), F. tomentosa must be rejected as domenclaturaliv superfluous when pub- lished. This represents another nomenclatural change not too unlike Betula lutea to B. alleghaniensis (Hardin, 1971). Il. JASMINUM L. Jasmine. It is extremely frustrating to deal with the taxonomy and nomenclature of these groups that are only naturalized in the Southeast. There is no easy way to gain a real concept of the species and be certain of the correct nomen- clature. It is also very difficult to tell from herbarium specimens which spe- cies are really naturalized and not just escaped or persistent from cultiva- 278 tion. Collectors should note this distinction on the label. In this treatment of the Oleaceae, the Jasmines, Forsythias, and Ligus- trums have to receive a tentative treatment heavily dependent upon the available literature. Some of the nomenclature in Wilson and Wood (1959) and other authors apparently is incorrect according to Read (1962). Long and Lakela (1971) follow Read in naming the Florida plants, and I will do likewise here for those naturalized in the Southeast. 1. Corolla yellow. ye eG we ey ee cw le Sh eS yt. 1. oe white or sine fnest, 2. Leaves of 3-7 leaflets. 3. Leaflets 3; calyx very short compared with corolla ooo: OR : i avian nse. 3. Leaflets 5-7; calyx half or as long as corolla tube . . 4-9 J. officinale. 2. Leaves 1-foliolate (apparently simple). 4. Calyx glabrous or nearly so, the lobes less than 3 mm. long J. dichotomum. ~~] 4. Calyx pubescent, the lobes 4-12 mm. long. Calyx lobes densely pubescent .. . . . %& J. multifiorum. Calyx lobes hirsute on the edges or i1O8e: 6. Calyx lobes mostly 7-12; stems puberulent; leaves with tufts of pale hairs in axils of veins beneath . . . 6. J. sanvbac. 6. Calyx lobes 5-7; stems glabrous; leaves ne beneath . 7. J. nitidum. wai ai 1. J. mesnyi Hance, Primrose Jasmine. Icdees of woods, creck banks: cp. Fla., Ga. (Tex.). This species is a native of western China. It is not commonly grown and hence rarely found escaped or naturalized. 2. J. fluminense Vellozo, Azores Jasmine, Climbing J. Weedy in waste places and old fields; ep. s. Fla, Incorrectly called J. azoricum L. (Read, 1962), it is a native of Brazil. 3. J. officinale L., Poet’s Jasmine, Common J. Pinelands and thickets, waste places; ep. s. Fla. Inel. J. grandiflorum L.—S. This is a native of India and China and is the hardiest of the white-flowered Jasmines. The large flowered form represented by nearly all our materials, is known in the trade as var. grandiflorum (1.) Bailey (forma grandiflorum (L.) Kobuski). I prefer not to recognize these cultivars formally. . J. dichotomum Vahl, Gold Coast Jasmine. Vacant lots, roadsides, hammocks; cp. s. Fla. As implied by the common name, this is a native of Ghana and the west coast of Africa. 5. J. multiflorwum (Burm. F.) Andr., Furry Jasmine, Downy J. Waste places, 279 cp. s. Fla. at species has been found naturalized in Highlands Co. Fla. It is a native of India and southeastern Asia. 6. J. sambac (L.) Solander in Ait., Arabian Jasmine. Hammocks, woods, and thickets; ep. s. Fla. Naturalized from tropical Asia. 7. J. nitidum Skan Hammocks, disturbed sites; cp. s. Fla. Erroneously known as J. amplexicaule Wall. ex Don; or J. undulatum Ker.—S. Naturalized from the Admiralty Islands. II. FORSYTHIA Vahl, Forsythia, Golden Bells. Forsythia has not been considered heretofore as part of the Southeastern Flora. It was not included by either Small (1933) or Wilson and Wood (1959). Although it is a very commonly cultivated ornamental shrub, and may per- sist about old dwellings, it seldom becomes established or naturalized. One of the possible reasons for the lack of establishment and few seeds is its heterostylous nature. If the shrubs are self-incompatible, and both of the distylic forms are not close enough for cross-pollination, there would be no seeds produced. This hypothesis needs verification. As indicated under Jasminum, it is difficult to determine naturalization by information on herbarium sheets. There are a number of records, however, of local establishment of the two species included here. It appears that we have to include the Forsythias as part of the Flora. 1. Mature branches with hollow or irregularly excavated pith between the nodes; leaves oblong-ovate; branches generally arching or drooping ae . F. suspensa. 1. Mature branches with chambered pith; iaaves ahione mets branch- es usually erect . . . . . . el, . 2. F. viridissima. 1. F. suspensa (Thunb.) Vahl. Edge of woods, waste places; Va. . F. viridissima Lin Edge of woods, eee banks, road sides and waste places: Ark., Md., N.C., Va. (Il.). bo IV. SYRINGA VULGARIS L., Common Lilac. Waste places, fence rows, edge of woods; Md., Va. (Mo., N.J.). This native of southeastern Europe persists from old plantings, and spar- ingly escapes in the cooler parts of our area. The Persian Lilac (S. persica L.) is frequently cultivated and escapes rarely but has not become estab- lished in our area as far as I know. 280 V. FORESTIERA Poir., Forestiera. There is need for studies of ecophenic, ecotypic, and clinal variation and introgression as indicated by Wilson and Wood (1959). I am following John- ston (1957) here for the most part Shinners (1959) has determined F’. porulosa as the correct type of the genus. 1. Leaves evergreen, margins entire... . . IL. F. segregata. 1. Leaves deciduous, margins serrate or rc so. 2. Plants flowering before the leaves expand; fruits slender, length 2 or more times the width . . ......2.C~«~:~C~«SY 2. FF. acuminata. 2. Plants flowering after the leaves have expanded; ne broad, length less than twice the width . . ....... . . 3. F. ligustrina. 1. F. segregata (Jacq.) Krug & Urban, Florida Privet. Two varieties, that may merely represent the extremes of an ecocline, have been recognized by Johnston (1957) and Long and Lakela (1971). See the discussion of the Florida Scrub at the end of this paper. a. var. segregata Coastal dunes, hammocks, mangrove swamps, river bottoms, low pine- lands; cp., Ga., Fla. Incl. I’. glol ilans Small, F’. porulosa (Michx.) Poir.—s. . var, nie torwm (Small) - C. Johnsto High, dry pinelands; s. Fla I’. pinetorum Small—s. 2. F. acuminata (Michx.) Poir., Swamp Privet. Wet woods, swamps, river banks, hammocks; chiefly cp. Ala., Ark., Fla., Ga., Ky., La., Miss., S.C., Tenn. (Tex., Okla., Mo., Ill., Ind.) 3. FY. ligustrinad (Michx.) Poir. Rocky soils, sandy stream banks, dunes, calcareous glades and uplands, granite flat rocks, swampy woods, and hammocks: chiefly ep. and adjacent provinces, Ala., Fla., Ga., Ky., La., Tenn. (Tex.). F’. pubescens Nutt.—sensu S, not Nutt. om Cw VI. CHIONANTHUS L., Fringe-tree, Old-man’s-beard. Fringe-tree is variable in overall size, pubescence, leaf form, length of petals, stamen number, anther size, shape of anther tips, and fruit size. Some of the variants have been named as distinct species. For instance, Small (1924) recognized C. pygmaeus, a small shrub with small leathery leaves, blunt-tipped anthers, short petals, and large fruits, and an endemic of the sand-scrub in the lake region of south-central Florida. Li (1966) de- scribed C. henryae, a small shrub with small rather coriaceous leaves, pro- longed anther tips, and fairly short petals, and which he said occurred in northwestern Florida, southern Arkansas, southern Alabama, and west- central Georgia, In addition, four varieties have been named on the basis of leaf shape and pubescence. It appears that forms with small anthers and small petals are scattered throughout the area although more frequent in dry habitats southward; the anther tips vary from rounded or very blunt to very prolonged acuminate with the blunt form mostly in south Florida. This variation needs to be thor- oughly studied using fresh material and correlated with habitats and the sexuality of the flowers. At the present time, I prefer to recognize only two species. See the discussion of the Florida Scrub at the end of this paper. 1. Corolla lobes 1.5-3 cm. long; anther tips prolonged; drupe 1-1.5 cm. long 2. ew we LC. virginicus. 1. Corolla lobes about 1 cm. long; anther tips blunt or acute; drupe 2-2.5 Gi, Jone «= a-% & & 6 RA Soe te ee Se Se So e BOO OyoMCeUs: 1. C. virginicus, L. Dry sandy woods, rocky slopes, savannahs, pocosins, swamps, rich woods; all prov. SE (Tex., Okla., Mo., Ohio, Pa., N.J.). Incl. C. henryae Li; tentatively placed here on the basis of the long anther tips. 2. C. pygmaeus Small. Pine scrub; s. penin. Fla. VII. OSMANTHUS Lour., Devilwood, Wild Olive. Devilwood is variable in leaf size, texture, and shape; fruit size, shape, and color; and inflorescence size, pubescence, and compactness. Most of the variation in and between populations is found in peninsular Florida. There is need of further study from the standpoint of the degree of isolation between the variants. See the discussion of the Florida Scrub at the end of this paper. 1. O. americanus (L.) Gray. Two varieties are recognized following Green (1958). a. var. americanus. Hammocks, bays, low woods, swamps, maritime forests, rich woods; chiefly cp. Ala., Fla., Ga., La., Miss., N.C., 8.C., Va. (Tex.). Amarolea americana (L.) Small—S:; incl. O. floridana Chapm.—Ss. b. var. megacarpus (Small) P.S. Green. Pine scrub; penin. Fla. Amarolea megacarpa SmalH—Ss. VIII. LIGUSTRUM L. Ligustrum, Privet. The species of our area are all escapes from cultivation. They all could be more common and widespread than herbarium specimens indicate. Again, as with Jasminum and Forsythia, it is difficult to determine which are really naturalized in our area. Also, in this rather difficult genus, the taxonomy and nomenclature of our plants seem uncertain in a few cases. A revision of 28 th i 2 e genus is needed. Twigs pubescent. 2. Corolla tube equalling or shorter than the lobes. 3. Flowers sessile or subsessile 3. Flowers pedicellate. 4. Twigs densely pubescent: leaves 1. L. quihoui. pubescent on midrib beneath 2. L. sinense. 4. Twigs minutely puberulent; leaves glabrous 3. L. vulgare. 2. Corolla tube slightly or to 3 times longer than lobes. ). Calyx and pedicels pubescent. . 4. L. ¢ 5. Calyx and pedicels glabrous or li Shilsy mibescenl at base of calyx 6. Leaves 2-6 ecm. long; twigs conspicuously pubescent see do. L. amurense. 6. L. japonicum. »btusifolium. 6. Leaves 4-10 cm. long; twig sonantey uberiiedt . Twigs glabrous. . Corolla tube equalling or shorter than lobes. 8. Leaves persistent or tardily deciduous, 6-15 em. long . 7. L. lucidum. 8. Leaves deciduous, 3-6 cm. long . . . . 3d. L. vulgare. 7. Corolla tube slightly or to 3 times iengey ‘han icbes. 9. Leaves persistent and glossy, rounded or broadly cuneate at base; corolla tube slightly longer than lobes . .. . . 6. L. japonicum. Leaves deciduous or half evergreen, cuneate sip base: corolla tube 3 times longer than lobes 8. L. ovalifolium. 1. L. quihoui Carr., Wax-leafed Ligustrum, Rare in low woods, waste places; cp. N.C., Va. (Tex.). 2. L. sinense Lour., Chinese Privet. Low woods, fence rows, waste places; W.Va. (Tex., Okla.). This is probably the most frequently and widely escaped and naturalized of the privets. 3. L. vulgare L., Common Privet. echetss open woods, low woods; various proy. Ala., Ark., Ky., La., N.C., S.C., Tenn., W.Va. (Tex., Tl., Ind., Ohio, Penn., N.J.). 4. L. obtusifolium Sieb. & Zuce. Thickets, edge of woods, roadsides; rarely established in Ky., Md., N.C. (Penn.). d. L. amurense Carr., Amur Privet. Thickets, open woods; rare, cp. and pied. N.C. and Va. (Tex.). 6. L. japonicum Thumb., Japanese Privet. Low woods, infrequently established; ep. 7. L. lucidum Ait., Wax-leafed Privet. Low woods and thickets; ep. Ala., Ga., La., 8. L. ovalifolitum Hassk., California Prive Hammocks, roadsides, thickets; infrequent in ep. and pied., Fla., Ky., Md., N.C. (Tex., Mo., Penn.). various prov. SE except Del. and pied. Ala., N.C., S.C. (Tex.). Miss. (Tex., Penn.). 283 SPECIES OF THE FLORIDA SCRUB The distinctness of the ‘‘scrub” or ‘‘Florida scrub’’ has long been known and described (Nash, 1895; Harper, 1915, 1927; Mulvania, 1931; Kurz, 1942). These are areas of deep white to gray, fine sands on fairly recent or very ancient dunes, bars, or ridges. The vegetation is dominated by Pinus clausa and scrub oaks and a mosaic of shrub thickets and open white sand. It is an extreme habitat of low fertility, excessively drained deep sand, and high summer temperatures. Davis (1967) shows the distribution of these scrub forests along the north- western and eastern coasts, scattered small inland ‘‘islands’’ in the penin- sula, and large areas in Marion, Lake, Volusia, Polk, and Highland counties. The origin and distinctness of these habitats is important when consider- ing the Oleaceae of the Southeast for Chionanthus pygmaeus and Osmanthus americanus var. megacarpus occur in the scrub. Chionanthus pygmaeus is found in the scrub of Highlands, Hillsborough, and Polk counties. It seems to be fairly distinct as a small shrub with small- er leaves, smaller petals, blunt-tipped anthers, and larger drupes. Chionan- thus virginicus occurs in more mesic or wet habitats and the two seem both geographically and ecologically isolated. It is for this reason that I am recog- nizing two species. The problem here is that individual characters appear to vary throughout the range of Chionanthus as indicated earlier. However, only in the south Florida scrub is there the correlation of the features char- acterizing C. pygmaeus. The more recently described C. henryae is some- what similar to C. pygmaeus in morphology and habitat and, although dis- junct from C. pygmaeus, may be related or reflect similar ecotypic differ- entiation. This needs further study. Osmanthus americanus var. megacarpus is found in the scrub of Hernan- do, Highlands, Polk, Osceola, Sarasota, and Sumter counties. It is charac- terized by a smaller habit, often smaller leaves, and larger fruits. The typi- cal var. americanus is widespread and in hammocks, bays, low woods, and swamps. Although ecologically allopatric, the two are geographically sym- patric and probably not completely reproductively isolated. Intermediates do occur, and I prefer to use the varictal level for these extremes. Forestiera segregata var. pinetorum does not occur in the typical serub but is in the drier pinelands of Dade County. The extreme form is distinct with smaller leaves, puberulent stems, and smaller drupes. The typical var. segregata has larger leaves, glabrous stems, and larger fruits and occurs in more mesic habitats and is more widespread geographically. The two are geographically sympatric and probably not entirely isolated reproduc- tively. Intermediates do occur and I think the varietal level is appropriate here. It may seem inconsistent in this treatment of the Oleaceae to recognize these variants in the scrub or high pinelands on the one hand, and on the other not to formally recognize the tremendous variation in some of the ashes. My justification lies in the relative distinctiveness of the Florida scrub 284 vegetation. Until detailed biesystematic studies are made, I consider the differences in these plants to be more than mere ecotypes, for they seem relatively isolated from their counterparts in more mesic environments. In addition, they may have been isolated for a very long time since these habi- tats are considered ancient dunes or coastal bars or ridges. The adaptations seem rather obvious. The smaller habit, smaller leaves, smaller petals, and pubescence are all probably adaptations to the severe dry conditions of potentially high transpiration rates and insufficient soil water. The larger drupes of Chionanthus and Osmanthus may reflect a se- lective pressure at the time of seedling establishment. The larger seeds with more stored food may have a better opportunity for more vigorous growth and are thereby able to establish a more extensive root system and escape dessication following germination. Another possible explanation, and a rather intriguing one, is that the larger drupes reflect differential selection by the Florida gopher, or Gopher tortoise (Gopherus polypkemus). This tortoise is common in these dry sandy habitats and are known to eat fleshy fruits. One could imagine their selection for larger drupes, and then possibly a higher germination rate among the seeds that pass through their bodies and germinate in a nitrogen-rich environment. Regardless of the particular se- lection mechanisms, these appear to be good examples of adaptive radiation and parallel evolution in related genera. REFERENCES DAVIS, JOHN H. 1967, General map of natural vegetation of Florida. Fla. Agric. Exp. Sta., Gainesville. FERNALD, M. L. 1950. Gray’s manual of botany. 8th ed. New York: The Macmillan Co. GLEASON, H. A. A. CRONQUIST. 1963. Manual of vascular plants of northeastern NY. United States and adjacent Canada. D. Van Nostrand Co. Ine., GREEN, P. S. 195 3. A monographic revision of Osmanfhus in Asia and America. Notes | Ga Edinb. 22: 439-542 HARDIN, JAMES W. 1971. Sidi of = southeastern United States flora. I. Betulaceae. our. Elisha Mitchell Sci, Soe. 87: 39-4 HARPER, ROLAND M. 1915, Veretation types; natural resources of an area in central Florida. ee Geol. Surv., 7th Ann. Rept., pp. 142-143 27. Natural resources of southern Florida. Bla. Geol. Surv., 18th Ann. Rept., pp. ae : JOHNSTON, M. C. 1957. Synopsis of the United States species of Foresticra (Oleaceae). Southw. Nat. 2: 140-151. KURZ, Herman. 1942. Florida dunes scrub, vegetation and geology. Fla. Gel Surv. Bull. 2 LI, HUE LIN. 1966. — new species of Chionanthus. Morris Arb. Bull. 17: 63-6 ITT LE, Ee. JR. . Notes on een: (Ash) in the United States. i. Ww ash. Acac k list of native and naturalized trees of the United States (includin 41 ‘ Chee Alaska). Usb Lied Serv. Agric. Handb. ‘ Atlas of United iat trees. Vol. 1. Conifers and important hardwood Usb! e Fores Sen Miscl. Publ. No. ONC und OLGA LAKELA, oF 7 Bees of tropical Florida. Univ. of Miami Pres la. ; . 1971. Tree flora of Kentucky. Univ. of Kentucky, Lexington. MILLER, G. N. ie The Genus Fraxinus, pi ashes, in es h America, north of Mexic Cornell iv. Agric. Exp. Sta. 335: 1-64, ai, MULV ANIA, MAURICI 1931, Ecological survey NASH, GEORGE 1895, Notes on some Florida plants. of a Florida scrub. Ecology 12: 528-54 Bull. ‘Forrey Bot. Club 2 t. g S. S, oO. Q. ds 289 RADFORD, A. E., C. R. BELL, J. W. HARDIN, and R. L. WILBUR. 1967. Contributor’s guide for . ae ora ee the southeastern United States. Dept. of Botany, Univ. of North ata aes 1 Hill. READ, R. . Jasminum species in cultivation in Florida and their correct names. Fla. State ee ae Proc. ee 430-437. SANTAMOUR, F. S. JR. 1962. The relation between polyploidy and morphology in white and biltmore ashes. Bull. ane Bot. Club 89: 228-2 see ea ai H. 1959. Typification of the genus Forestiera (Oleaceae). Rhodora : 293-2 Shab J. K 1924 . Plant novelties from Florida. ae Torrey Bot. Club 51: 384. ay 1933 Manual of the southeastern flora. By the ea ieee Pa. WILSON ik a and C. E, WOOD, JR. 1959 ae genera of Oleaceae in the southeastern United Sates. Te Arn. Arb. 40: 369-384. WRIGHT, ee, Be 1944a, Genotypic variation in white ash. Jour. Forest. 42: 489-495. 44b. ae differentiation in red ash. Jour. Forest. 42: 591-597 ee ok New chromosome counts in Acer and Fraxinus. Morris Arb. Bull. 8: 33-34. 1965. Green Ash a axinus pennsylvanica Marsh.). In, Silvics of forest trees of the nived States. USDA Forest Serv. Agric. Handb. No. 271. 5 NOTES FATOUA VILLOSA (MORACEAE) IN FLORIDA.—Fatoua villosa (Thunb.) Nakai is a native of eastern Asia. Dr. John Thieret discovered it in Louisi- ana in 1964 (Sida 1: 248), the first time it was seen in America. I found the plant at Saint Leo College, Saint Leo, Pasco Co., Florida, growing in a thick patch as a pesty weed among a planting of zamias and bromeliads on our campus, Voucher specimens have been deposited in SMU and GH. I have some extra duplicates for distribution to individuals who request them.— Damian DuQuesnay, Saint Leo College, Saint Leo, FL 33574 CAPERONIA PALUSTRIS (EUPHORB IAC D AE) IN ARKANSAS.—Capero- nia palustris (L.) St.-Hil., widespread in warmer parts of America, ranges along the Gulf Coast from Florida to Texas. Correll & Johnston (Manual of the Vascular Plants of Texas, 1970) date its introduction into Texas at about 1920. In July 1971 a scrap of plant material collected by a representative of the Arkansas State Plant Board was sent to me for identification. The speci- men, from a rice field in east-central Arkansas, appeared to represent C. palustris. My student assistant, David Wheat, visited the rice field in Sep- tember 1971 and collected a number of specimens of undoubted C. palustris. According to the owner of the land, the plants at that time had successfully resisted all attempts at eradication with herbicides. Specimens collected by Wheat have been distributed through the Southern Appalachian Botanical Club. Collection data: small population in rice field 11 mi N of Stuttgart, Prairie Co., 11 Sep 1971, David Wheat s.n. Another Arkansas collection of the species was made in 1971 (UARK). Col- lection data: weed in soybean field near DeWitt, Arkansas Co., 6 Nov 1971, H. R. Durst, s.n., Extension Agronomist, determined by E. B. Smith.—G. E. Tucker, Arkansas Polytechnic College, Russellville, AR 72801. ALLIUM AMPELOPRASUM (LILIACEAE) AND TRIFOLIUM VESICU- LOSUM (LEGUMINOSAE) IN OKLAHOMA.—These species are new io the Oklahoma flora. Allium ampeloprasum 1.., found along a roadside, was asso- ciated with such native plants as Polytaenia nuttallii, Monarda fistulosa, Ca- calia plantaginea, Centaurea americana, Echinacea pallida, Engelmannia pinnatifida, and Silphiwm laciniatum. According to a local resident, the ‘“‘wild garlic” had been there for several years, each year becoming more abund- ant—in spite of the fact that he and others annually dug bulbs for culinary use. I counted 200 plants in a distance of 400 feet. They were producing many offsets. Collection data: roadside 6 mi NE of Madill, Marshall Co., 10 Jun 1973, Thieret 41147 (XNK). Trifolium vesiculosum Savi, reported in 1969 from Louisiana and Mississippi as new to the United Sates and in 1970 as new to Alabama, was seen along roadsides in several places in Johnston County. It was conspicuous because of its relatively large heads. Collection data: road- side 0.5 mi N of Milburn, Johnston Co., 20 June 1973, Thieret 41146 (KNK).— SIDA 5 (4): 286, 1! \S 74, 287 John W. Thieret, Northern Kentucky State College, Highland Heights, KY CORALLORHIZA ODONTORHIZA (ORCHIDACEAE) IN LOUISIANA— While working on a Flora of St. Helena Parish, Louisiana, I collected Cora lorhiza odontorhiza (Willd.) Nutt., a new state record. This orchid was to be expected in Louisiana as it had been recorded from neighboring Mississippi and Arkansas (D. S. Correll, Native Orchids of North America North of Mexico, 1950; also in litt.). Louisiana’s orchid flora is now 37 species, of which 19 occur in St. Helena Parish. Collection data: rich woods 5 mi NNW of Chipola, St. Helena Parish, 8 Oct 1971, Allen 1691 (LAF, LSU).—Charles M. Allen, University of Southwestern Louisiana, Lafayette, LA 70501. ~— 1 HYPOCHOERIS IN TEXAS.—Since the completion of the Manual of the Vascular Plants of Texas (Correll and Johnston, Texas Research Foundation, Renner. 1970) recent collections have extended the known range of Hypo- choeris microcephala var. albiflora and added two species of the genus, H. radicata L. and H. brasiliensis (Less.) Benth. & Hook., to the flora of the Hyco deh dees var. albiflora (Figure 1) was first collected in North Am a by L. H. Shinners (Sida 2: 393-394. 1966). This taxon has be- come an Ree dant eee weed in southeast Texas. In the last two years, it has replaced Pyrrhopappus as the most frequent Cichoriaceous taxon in some places along the freeway system and in vacant lots near and in Hous- ton, Texas. In the springs of 1969 and 1970, I collected Krigia and Pyrrho- appus in Harris County at several localities, where var. albiflora is now abundant. It must have been extremely rare or absent prior to 1970 in these localities. I have seen the following specimens of var. albiflora: LOUISIANA: Calcasieu Parish. Interstate 10 at Calcasicu River, 31 May 1969, Thieret 30814 (LAF). TEXAS: Jefferson Co. Port Arthur, 25 May 1959, Knight 128 (TEX 2). According to Mr. Leonard Knight, this taxon appeared in ditches and along roadsides in Port Arthur in the early 1960's (Pers. comm.); Hardin Co. just N. of Kountze, 21 May 1970, Correll & Correll 388138 (TIEX); Harris Co., SW Houston, Stella- link Rd. and Linkwood Rd., vacant lot. 27 Mar 1972, Tomb 724 (CHI, TEX): ca 5 mi N. of Houston city limits on Interstate 45, 7 April 1972, Tomb 743 (SMU, TEX). Orange Co. 6 mi N. of Orange, 17 May 1966, Shinners 31370 (SMU, TEX); Western edge of Vidor, 21 April 1970, Flyr 1354 (SMU). Hypochoeris microcephala var. apa ais a well adapted, rhizomatous, perennial weed that occurs in bo sand and heavy clay soils. Its small white heads and low pollen sean per anther suggest that it is self- fertile 288 289 The chromosome number of H. microcephala var. albiflora, n = 4II, is re- ported here for the first time (determined from Tomb 743 and 724). This number agrees with those reported for the other South American members of the genus. In 1970, the Correll’s made what appears to be the first collection of Hypo- choeris radicata L. in Texas: Newton Co., 3 mi S. of Burkeville, sandy soil, edge of woods, 19 May 1970, Correll & Correll 38775 (SMU, TEX). This Euro- pean species is widespread over the eastern half of North America, the Pa- cific Northwest, and in Central America. Another new record for Texas is Hypochoeris braziliensis: Anderson Co., SE edge of Palestine, growing in sandy soil, disturbed roadside, Tomb 745 (SMU, TEX). This species is also from South America and is closely related to H. microcephala. eee radicata and H. braziliensis are presently known from Texas from single collections and should be carefully watched for expansion and migration. a is too late to accurately retrace the initial introduction, local migrations and expansion of H. microcephala var. albiflora, yet it should be watched for continued expansion from its present North American dis- tribution. As Shinners (Sida 2: 119-128. 1965) said, we cannot continue to dis- regard weeds; they have much to tell us.—A. Spencer Tomb, Department of Biological Sciences, University of Illinois at Chicago Circle, 60680 Fig. 1. Hypochoeris microcephala var. albiflora. A. habit. B. palea from re- ceptacle. C. pappus bristle. D. achene. Correll and Correll 38813 ex- cept basal rosette and root from Flyr 1354. Drawings by Karen Baker. SIDA 5 (4): 289, 1974. DOCUMENTED PLANT CHROMOSOME NUMBERS 1974:1 CARL S. KEENER Department of Biology, The Pennsylvania State University, University Park, Pennsylvania, 16802 CHENOPODIACEAE® CHENOPODIUM ALBUM L. var. STEVENSII Aellen.* (Fig. 1.) 2n=54. WISC., Door Co.: H. M. Clark s.n. (WIS). CHENOPODIUM AMBROSIOIDES L.”* (Fig. 2.) 2n—32. CALIF., Ventura Co.: Pollard 23 October 1969 (CAS). CHENOPODIUM CALIFORNICUM S. a (Fig. 3.) 2n=18. CALIF., Los Angeles Co.: R. IF’. Thorne 37824 (RSA). CHENOPODIUM CARNOSULUM — var, PATAGONICUM (Phil.) Wahl? (Fig. 4.) 2n=18. CALIF., San Luis Obispo Co.: R. F. Hoover 9856 (PAC). CHISNOPODIUM GIGANTEUM Don.° (Fig. 5.) 2n=54. Original seed source of plants used in virus studies at The Pennsylvania State University ob- vane from Europe; voucher for the seed source used in this study in eer ODIuM GIGANTOSPERMUM Aellen.” (Fig. 6.) 2n=36. PA., Clear- fie H. A. Wahl 8288 with W. A. Niering (PAC). SHENOPODIUN GLAUCUM L. var. GLAUCUM? (Fig. 7.) 2n=18. WISC. Lafayette Co.: Palzkill 17 (WIS CHENOPODIUM HYBRIDUM Le » (Fig. 8.) 2n=18. SWEDEN: Malmo, Skane Province. Original seed source from Lund Botanical Museum (LD); voucher material in PAC, CHENOPODIUM MACROCALYCIUM Aellen.® (Fig. 9.) 2n=36. VA., North- ampton Co.: H. FE. Ahles 57612 (NCU CHENOPODIUM POLYSPERMUM IL. var. OBTUSIFOLIUM Gaudin." (Fig. 10.) 2n=18. SWEDEN: Malmo, Skane Province. Original seed source from Lund Botanical Museum (LD); voucher material in PAC, CHENOPODIUM PUMILIO R. Br.” (Fig. 11.) 2n=16. ARK., Clark Co.: Demaree 54131 (PAC). CHENOPODIUM RUBRUM L. var. RUBRUM.” (Fig. 12.) 2n=36. SWEDEN: Malmo, Skane Province. Original seed source from und Botanical Muse- um (LD); voucher material in PAC. * All counts were made from root tips of seedlings raised from seed taken from the in- dicated voucher specimens unless otherwise stipulated. "Taxon for which the same chromosome count has been published previously. ©Taxon for which a chromosome count differing from this one has been published previ- ously. “No chromosome number reported previously for this taxon. SIDA 5 (4): 290—291. 1974. w F oe Ne te e re oe oe ‘ > 3 oe 1 Se er . oe igs x rt a * c j x< K x SN + + <*> ¢ ~ 4 Ze 4 Lee ge / ’ a | th, ‘ O.OL mm o e o ~ & et Vrs Baty a ae % saan ¢ oe = SS % a ~= 3 * . oh Me ‘ on! waxy, @ 4 x az t 7 8 a \ a et ri >” an —~- oe? = fe «qt * . A“ ¢ = 10 de a uke 12 DOCUMENTED PLANT CHROMOSOME NUMBERS 1974:2 DANIEL J. CRAWFORD Department of Botany, The University of Wyoming, Laramie, 82070 ROBERT C. GARDNER Department of Botany, The Ohio State University, Columbus, 43210 This work was supported by a grant from the Division of Basic Research of the College of Arts and Sciences, University of Wyoming. In those instances where more than one population of a species were ex- amined, material from the collection which is cited first is the one from which the drawings were made. Only those determinations which are new or Which differ from previous reports are figured. All magnifications are ea, < 1250 RANUNCULACEAE ACONITUM COLUMBIANUM Nutt. ex T. & G. 2n = 16. WYO., Albany Co., Gardner 257 (RM); Carbon Co., Crawford 656 (RM). These determinations agree with previous reports (Wiens and Halleck, 1962, Bot. Notiser 115: 455; Kawano, 1965, Bot. Mag. Tokyo 78: 361'). ACONITUM COLUMBIANUM Nutt. ex T. & G. (Fig. 1) 2n = 18. WYO., Teton Co., Gardner 269 (RM). This determination represents the first re- his AQUILEGIA LARAMIENSIS A. ne (Fig. 2) 2n = 14. WYO., Albany Co., Gardner 223 (RM). This is the first report for the species, which is a rare endemic in the Laramie Range of southeastern Wyoming. CLEMATIS LIGUSTICIFOLIA Nutt. ex T. & G. 2n = 16. COLO., Larimer 0., Gardner 230 (RM) and Crawford 624 (RM). These reports agree with the two previous determinations for the species (Meurman and Therman, 1939, Cytologia 10: 1’; Kurita, 1956, Bot. Mag. Tokyo 69: 2391). DELPHINIUM BARBEYI (Huth) Huth (Fig. 3) 2n = 16. WYO., Carbon Co., Gardner 265 (RM). This is the first report for the species. DELPHINIUM GEYERI Green (Fig. 4) 2n = 16. COLO., Larimer Co., Craw- ford 623 (RM), and Gardner 231 (RM); WYO., Laramie Co., Gardner 239 (RM), Albany Co., Gardner 242 (RM). These appear to represent the first determinations for this common plains larkspur. DELPHINIUM NELSONII Greene 2n = 16. WYO., Albany Co., Gardner 225 (RM), and Gardner 229 (RM). These determinations agree with previous ‘These eae were taken ra pe 1969, Chromosome Numbers of Flowering Plants. Moscow: Acad. of Sciences the SR, V. L. Komaroy Botanical Institute, 926 pp. The original papers have not one seen. SIDA 5 (4): 292—294, 1974, 293 e @ 2 oe Ai? he? “. 1 = 2s - 3 NO ® Gait 9 1/6 5 g { 30) | | on 6 rN a | "% . Ce) Liove 8 reports for the species (Wiens and oe 1962, Hartman and Crawford, 7 Bot. Notiser 115: 455; 1971, Taxon 20: 158 DELPHINIUM OCCIDENTALE: (S. Wats.) S. ats, (Trig, Carbon Co., Crawford 641 (RM) and Gardner 263 (RM); Teton Co., 267 (RM). These are first reports for the species. DELPHINIUM RAMOSUM Rydb. (Fig. 6) 22 = 16. COLO., Las Animas Co., Crawford 677 (RM). This represents the first determination for this faxon: ANUNCULUS ACRIFORMIS A. Gray (Fig. 7) 2n = 28. WYO., Albany Co., Crawford 6389 (RM) and Gardner 240 (RM); COLO., Larimer Co., Gardner 5) 2n = 16. WYO., Gardner 294 236 (RM). These reports apparently represent the first for the species. RANUNCULUS ALISMAEFOLIUS Geyer ex Benth. var. MONTANUS S. Wats. 2n = 16. WYO., Carbon Co., Gardner 250 (RM); Albany Co., Craw- ford 6385 (RM). The present determinations are in agreement with the pre- vious report for the species (LOve and Kapoor, 1967, Taxon 16: 565). THALICTRUM SPARSIFLORUM Turcz. ex Fisch. & Mey. (Fig. 8) 2n = 14. COLO., Larimer Co., Gardner 233 (RM). The only previous report for this species gives a chromosome number of 2n = 42 (Kuhn, 1928, Jahrb. Wis- sensch, Bot. 68: 382') and contrasts sharply with the diploid determination presented here. REVIEWS THE PLANTS OF SOUTHERN NEW JERSEY. Stone, Witmer. Quarterman Publications, Inc., 5 South Union Street, Lawrence, Mass. 01850. $25.00. This is a reprint of Witmer Stone’s publication of some 63 years ago. There is a new foreword by Elizabeth M. Woodford. This tome has 914 pages and is well bound ee It is always a surprise when a ‘‘Manual” or ‘‘Flora’”’ is good reading but his one is! The chatty oe about many of the taxa are often de- lightful and betray the author’s intense personal interest in his work. The price is a bit high for the average botanist but well worth it. The usual problems of a reprint involving updating taxonomic concepts and the in- corporation of recent researches exist but do not seriously detract from its interest or usefulness.—Chester M. Rowell, Jr., Angelo State University, San Angelo, Texas 76901 THE EFFECTS OF ECOLOGICAL CHANGES ON BUCKEYE LAKE, OHIO WITH EMPHASIS ON LARGEMOUTH BASS AND AQUATIC VASCU- LAR PLANTS. John Bayless Judd and Stephen Taub. Biological Notes 6, Ohio Biological Survey, Ohio State University, Columbus, Ohio 43210. in $1.50 paperback. Biological Notes of the Chio Biological Survey are published at irregular intervals and include such works as Lichens of Ohio by Taylor (No. 3, 1967 and No. 4, 1968) and The Vascular Flora of Glen Helen, Clifton Gorge, and John Bryan State Park by Anliot (No. 5, 1973). The most current in this series deals with ecological changes in Buckeye Lake, Ohio. Buckeye Lake is a shallow, eutrophic lake with an area of approximately 1200 hectares. It was constructed in 1825 as part of the Ohio-Eric Canal and is thus a very old reservoir by today’s standards. This publication is divided into a discussion of historical changes, present and past plant associations, basic chemical limnology, and an extensive ex- amination of past and present fish populations including growth studies and parasitology. The botanist will find that the treatment of aquatic vascular plant ecology is particularly interesting. The authors found that species diversity has de- clined rapidly in the 20th century and important groups such as Elodea and Utricularia have been eliminated by various drainage and construction pro- jects. They in turn state that the decline in productivity of the Largemouth Bass is directly correlated to increasing cutrophication and lack of plant cover. Data presented concerning the physical limnology of Buckeye Lake was somewhat disappointing as relatively few sites were examined and _ para- meters studied were limited. Judd and Taub believe that increasing turbidity ~— SIDA 5(4): 295. 1974, 296 has played a severe limiting factor to the growth and stability of both aqua- tic plants and fish populations. Also, general statements concerning fish para- sites were based on too little data, but nevertheless, the fishery biologist, plant ecologist, and limnologist would find this work to be extremely valu- able. The Effects of Ecological Changes on Buckeye Lake presents a written and photographic documentation of eutrophication over an extended period of time. This type of work is all too rare in today’s literature.—John O. Me- com, Biology Department, Southern Methodist University, Dallas, Texas, 75275, THE VASCULAR FLORA OF GLEN HELEN, CLIFTON GORGE, AND JOHN BRYAN STATE PARK by Sture F. Anliot. vii + 162 pp., 13 figs. Biological Notes No. 5, Ohio Biological Survey, Ohio State University, Co- lumbus, Ohio 43210. 1973. $3.00 paperback. This annotated checklist follows the format which has become more or less standard for this type of work. A brief summary of the geography and geology of the study area in north-central Greene County, Ohio, is followed by descriptions of the vegetational communities encountered there. Photo- graphs of typical stands of these communities are included. The checklist reports 872 species, 25 varieties, and 13 forms, nearly half of which represent new reports for Greene County. Notations for each taxon in the checklist include common name, contemporary (only) synonyms, subjective estimate of abundance, habitat, growth form, flowering and fruiting period, and dis- tribution within the state. Origins of naturalized and adventive taxa and separate indexes to scientific and common names are included.—Jerry M. Flook, Herbarium, Southern Methodist University, Dallas, Texas 75275. Errata Graviora in SIDA 5(3) . 142, line 40. Insert: 1956x. Prunus americana var. lanata a synonym of P. nigra. Rhodora Ko] 1956y. Vicia grandiflora var. kitaibeliana in the Southeastern United States. Castanea 21: 151-152 p. 142, line 41. 1956x should read 1956z. p. 179, column 2, line 10. Should read Vicia 1948a, 1948c, 1956y. SIDA 5 (4): 296, 1974. JOHN F. COOKE, JR., 1927-1970 John F. Cooke, Jr. was Assistant Professor of Botany at Southern Univer- sity, Baton Rouge, Louisiana, when he died of an aortic aneurysm while driving his car in that city. Known as ‘“‘Jack’’ to his many friends at The Ohio State University and Cornell University, where he did graduate work, his career as a university professor and professional botanist was short-lived. Born 7 August 1927 in Woodbury, New Jersey, Jack attended Merion Gram- mar School in Merion, Pennsylvania, from 1933 to 1941, and Morse High School, Bath, Maine, from 1941 to 1945. Upon graduation from high school, Jack enlisted in the United States Navy, serving as a Reserve Seaman, 2nd Class. His tour of duty was for thirteen months, nine of which were aboard the U. S. S. DEVICE, a mine sweeper. Beginning in 1946, Jack enrolled as an undergraduate in the College of Arts and Sciences at Cornell University, Ithaca, New York, where he took various courses. Botany and Floriculture were his major interests, and he graduated in June 1950 with a B.A. in Botany. For 2 years following graduation, Jack was employed by the W. Atlee Burpee Company of Philadelphia. During summers, he worked at their Ford- hook Farms in Doylestown, Pennsylvania, doing planting, maintenance, flow- er trial rating, and pest control. In the winters he was in Philadelphia in- volved with seed ordering, the bulb department, and catalogue mailing; later he served in the purchasing department. From 1953 to 1956, he was employed as a garden superintendent by the Board of Education of Cleve- land, Ohio. While there he also was assistant supervisor of greenhouses for West Technical High School during 1953-1954. Horticulture as a science and as a practical phase of business was taught as a part of the science cur- riculum. Jack directed three to eight individuals. Later, 1954-1956, he became assistant to the instructor in charge of the school landscaping program. He became involved in the fall landscaping and spring planting program of the Cleveland Public Schools. In addition, he taught general gardening to adult night-school classes. From 1956 to 1960 employment was with S. C. Johnson, Inc., Cleveland, Ohio, S. S. Pennock, Co. (wholesale florists), Philadelphia, and with J. A. Pullen Greenhouses, Houma, Louisiana. In the last company he was a partner. Jack returned to Cornell University in 1960 and enrolled in the Graduate School for study in the Department of Floriculture and Ornamental Horti- e nts strated great ability in handling and instructing students, tsplaying strong initiative in presenting the subject matter. In the latter part of his term at Cornell University, Jack was given full charge of pode in the lab- ' Contribution at the Department of Botany (Paper No. 835) and the Herbarium, The Ohio State University. SIDA. 5 (4): 297—300. 1974. 298 oratory, of collecting plant materials used in classes and of maintaining the teaching herbarium. His research involved the cytology of and hybridiza- tion within and between Achimenes and Smithiantha. He produced the first tetraploid hybrid, «x Eucodonopsis, between these two genera, which was entered into trade with the cultivar name, ‘Diamond Lil.’ His M. S. Thesis, entitled, Interspecific hybridization in the genus Achimenes P. Br. (183 pp. + v, and several plates, diagrams, and tables), was completed under the direction of Professor Robert E. Lee. The M. S. degree was awarded in January 1963. Jack enrolled for graduate work in the Department of Botany and Plant Pathclogy at The Ohio State University in January 1963. He studied in the area of experimental plant taxonomy with emphasis on cytotaxonomy and cytogenetics under the guidance of Professors T. Richard Fisher and Elton I’. Paddock. He assisted and taught in the General Botany and Local Flora courses, and assisted in Plant Genetics, Basie Concepts in Botany, and Tax- cnomy of Vascular Plants. His background in horticulture and floriculture made him one of the most diverse students in the taxonomy program during the time Dr. Fisher was directing it. Jack frequently drew upon his knowl- edge of horticultural plants for examples in his teaching. The Ph. D. degree was awarded in the summer of 1968, with the completion of his dissertation, The chromatography and cytology of some cultivated taxa of the genus Hosta Tratt. (97 pp.), completed under the direction of Professor Fisher. Jack was elected to two honor societies, Pi Alpha Xi (Floriculture Hon- orary Fraternity) and Sigma Xi, and was a member of several socicties, the American Gloxinia and Gesneriad Society, American Society for Horticul- tural Science, American Society of Plant Taxonomists, Betanical Society of America, International Association for Plant Taxonomy, and the Ohio Acad- emy of Science. College teaching as a career began in the fall of 1968, when Jack became Assistant Professor of Botany at Southern University, Baton Rouge, Louisiana, where he taught courses in general botany and plant tax- onomy. His new career, just beginning, was short-lived. He died at the age of 42 on 12 January 1970. Jack never married. He is survived by his parents and one sister. Inter- ment was in Oak Grove Cemetery, Bath, Maine. At Morse aes Senoo! in Bath, Jack’s parents established in his memory The Dr. John e, Jr. scholarship to be presented annually to a graduating student iris to major in one or more of the life sciences, with a preference for botany. Jack had many interests other than those connected with plant science. He enjoyed the study of language, especially the derivation, history, and precise use of words. He was fun-loving; he enjoyed pleasant little jokes—those played on him as well as those he played on others. Jack could tell a good story, and he laughed easily. He liked cooking and was always on the look- out for new and exotic foods. The guests he entertained in his home were sure to enjoy the atmosphere of warm genial hospitality, the good food (prepared by Jack himself), and the sparkling conversation. All in all, he was a thoroughly delightful person, a highly respected colleague, and a ae? | Dr. John F. Cooke , Jr. (photograph taken in the fall of 1964) 300 staunch friend. His passing has left an abysmal void in the hearts of all of us who were privileged to be his associates. Publications: 1955 Gardening in the Cleveland Public Schools. Plant Life 11: 30-31. 1962 The chromosome number of Achimenes woodii Morton. Baileya 10: 53. The stigma of Achimenes glabrata. Baileya 11: 47. Plantain lilies bloom in shady areas. The Cleveland Plain Dealer Garden Planning Guide. 23 April. 11 1966 Trumpet best known of four a of lilies. The Cleveland Plain Dealer Gar- den Planning Guide. 15 April. 14, Hybridization within and between oa P. Br. and Smithiantha Kuntze (Gesneriaceae). Baileya 14: 92-101. (with Robert E. Lee). A new tetraploid x Eucodonopsis. ss ican Violet Magazine 20 (2): 45-56. 967 x Eucodonopsis ‘Diamond Lil.’ The es 17 (3): 25-26. Ronald L. Stuckey Department of Botany College of Biological Sciences The Ohio State University Columbus, Ohio 43210 SIDA suisse" VOLUME 5 NUMBER 5 DECEMBER 1974 CONTENTS A taxonomic re-evaluation of the genera Amphiachyris, Amphipappus, Greenella, Gutierrezia, Gymnosperma, Thurovia, and Xanthocephalum (Compositae). John Ruffin. ‘ 301 Further comments on Styrax L. Richard A. Howard. 334 Pollen morphology of Amphicarpaea (Leguminosae: Phaseoleae). Blanche W. Meeson. 338 Wood anatomy of Amphiachyris, Amphipappus, Thurovia, Gymnosperma, and the Xanthocephalum complex. John Ruffin. 34] NOTES. Bauhinia erythrocalyx (Leguminosae), new species from Mexico. 353.—Hydrilla verticillata eae new to Louisiana. 354.—Transcription error, 354 EULA WHITEHOUSE (1892-1974). INDEX TO VOLUME 5. Jerry M. Flook. SIDA, founded by Lloyd H. Shinners, is privately published and edited by Wm. F. Mahler, SMU Herbarium, Dallas, Texas, 75275, U.S.A. Subscrip- tions: Libraries—$6.00 (U.S.) per year; individuals—$8.00 (U.S.) per vol- ume; numbers issued twice a year. ASSOCIATE EDITOR John W. Thieret Northern Kentucky State College Highland Heights, Kentucky 6 SIDA Contributions to Botany Volume 5 Number 5, pages 301-361 Copyright 1975 by Wm. F. Mahler A TAXONOMIC RE-EVALUATION OF THE THUROVIA, AND XANTHOCEPHALUM (COMPOSITAE) JOHN RUFFIN Department of Biology, Atlanta University, Atlanta, Georgia 30314 The Compositae, one of the largest families of vascular plants, are one in which evolutionary modifications are complex and poorly understood. Generic boundaries within many of the groups are obscure and, therefore, controversial. Generic delimitation of Xanthocephalum, Gymnosperma, Gu- tierrezia, Amphiachyris, Amphipappus, and Greenella (Astereae, subtribe Solidagineae) is confused. As a consequence, I undertook the present inves- tigation, which has three objectives: (a) to reevaluate the taxonomic posi- tion of the genera by attempting to quantify and assess the gross morpho- logical variation among the various groups, (b) to conduct a comparative anatomical study to determine whether endomorphic characters support the relationships suggested by gross morphology and cytology, and (c) to determine phylogenetic or evolutionary relationships from anatomical data. A somewhat detailed historical account of most of these genera is given by Solbrig (1960). The present account adds some additional data. Xanthocephalum was the first genus of this group to be described—in 1807 by Willdenow, based on material collected and described by Humboldt (Sol- brig, 1961). Gray (1873), recognizing the close similarities between Xanthocephalum and Gutierrezia, placed both X. gymnospermoides and X, wrightii in Guti- errezia, emphasizing that the line separating the genera was not perfectly clear. Gray also held Gutierrezia to comprise Amphipappus as well as Am- phiachyris but later (Gray, 1884) felt that Amphipappus should be merged with Amphiachyris and kept separate from Gutierrezia. This transfer was justified because of the sterile disk florets and the quite setose pappus of both Amphipappus and Amphiachyris. Bentham and Hooker (1873) adopted Gray’s original treatment for the group. They merged Amphiachyris as well as Amphipappus with Gutierrezia and placed this group near Gymnosperma and Xanthocephalum. According to Nelson (1934), Amphipappus and Amphiachyris should re- main distinct genera, since Amphiachyris was based upon an annual herb, A. dracunculoides, ‘differing in many respects from Amphipappus.’”’ Porter (1943), agreeing with Nelson’s line of reasoning, considered Amphipappus distinct. SIDA 5(5): 301—333. 1974, 302 The morphological similarity of Greenella Gray to Gutierrezia and Xan- thocephalum is unquestionable, and both Gray (1884) and Shinners (1950) doubted the distinctness of that genus. According to Gray (1884), in the original description of the genus, “‘if the ray-flowers of this plant were yel- low instead of white, it would be referred to the genus es hae Nl and the habit is nearly that of X. linearifolium and X. sericocarpua More recently, further opposing views as to the taxonomic status i these genera have been recorded. According to Shinners (1950), the quite variable features of pappus, size of heads, and number of rays did not constitute dif- ferences sufficient to justify maintaining separate genera. To Shinners the fact that they were of one phylogenetic stock was unquestionable, and in view of the slight differences among them, Shinners felt they represented a single genus. Solbrig (1960) conducted a cytological and morphological study and concluded that certain morphological features in combination could be used to delimit the genera but that they often were insufficient when used alone. Chromosome number, however, proved to be a distinguish- ing character. This criterion, in Solbrig’s opinion, was sufficient to delimit the genera satisfactorily. Howe (1970) reported on megagametogenesis for Gutierrezia microcephala and G. texana and compared it with that of Amphiachyris dracunculoides. According to Howe, the differences in the female gametophytes supported the retention of Amphiachyris as a genus distinct from Gutierrezia, thus agreeing in part with Solbrig. While Solbrig’s cytological conclusion seems tenable, recent inconsisten- cies in chromosome numbers in the various genera, not reported in Solbrig’s work, raise questions as to the placement of the groups and warrant re- assessment of their status. Shinners (personal communication), Spencer Tomb (personal communica- tion), and Correll and Johnston (1970) suggested that the monotypic genus Thurovia Rose is also closely related to the Xanthocephalum complex. Rose (1895), in the original description, noted that, from the head structure and the general habit, Thurovia apparently belonged to the tribe Astereae and that the habit of Thurovia was like that of Gutierrezia or Greenella. Thus, my investigation also includes a study of Thurovia. MATERIALS AND METHODS Fresh plant material for anatomical studies was killed and fixed in either chromic-acetic-formalin or formalin-propionic-alcohol (CRAF and FPA, respectively; Sass, 1958), dehydrated through a graded tertiary butyl-alcohol series, infiltrated, embedded, and sectioned in paraffin. Dried material ju- diciously taken from herbarium sheets was restored by immersion in 3% aqueous sodium hydroxide and then was dehydrated and processed as de- scribed above Many of the data on gross morphology were obtained by measuring 15 characters on each of 10 plants per collection. To reduce error variation, 308 only plants in anthesis (showing pollen shedding) were chosen for measure- ments To standardize leaf measurements, the maximum length of the current year’s growth was first measured from the base of the year’s growth (or base of the plant, for annuals) to the tip of the longest branch. Measure- ments were made on the leaf nearest the midpoint of the current year’s growth. For involucral and floral characters one flower head was chosen from each plant, placed in 50% ethyl alcohol, and measured later. To stan- dardize this procedure and choose heads of the same relative maturity, the flower head nearest the terminal head on the branchlet nearest the midpoint of the current year’s growth was chosen. Whole leaves and flowers for venation studies were cleared and stained as described by Fuchs (1963). Table 1 shows the sources of the materials used in my morphological studies. OBSERVATIONS Involucre and receptacle. Comparative morphological and_ histological studies of the involucre and phyllaries (involucral bracts) show few major differences and are not useful in delimiting the genera. Greenella, Gutierrezia, and Xanthocephalum possess an involucre varying from narrowly turbinate to broadly campanulate. Amphiachyris has a some- what broadly campanulate involucre, and Gymnosperma, Amphipappus, and Thurovia possess elongate-turbinate involucres. Involucral bracts of the genera are glutinous, in two to many rows, im- bricated, and dark green at the tips. In all the groups the number of bracts appears to correlate well with the size of the involucre (Table 2). Histologically, in all species studied the outer phyllaries are four to seven cells thick. In Greenella, Gutierrezia, Gymnosperma, and Xanthocephalum (except X. wrightii and X. linearifolium) only one vascular trace enters each bract. Heads of X. wrightii have three traces in the outer bracts and one trace in each innermost bract. Xanthocephalum linearifolium shows three to five bundles in the outer bracts. In comparison, Amphipappus com- monly shows three bundles in its bracts, and Amphiachyris has as many as five traces in the outer bracts. Heads of Thurovia, like X. linearifolium, have three to five bundles in the outer bracts. One to five secretory canals occur in the outer bracts in all species stud- ied. These canals are, for the most part, conspicuous on the adaxial side of the vascular bundles. All of the species in each genus possess sclerenchyma in the outer bracts. The degree of sclerification is so pronounced that in most species the vas- cular bundles are usually enveloped in sclerenchyma, which is more abun- dant near the base of the bracts. Many inner bracts appear to be entirely sclerified except in the epidermal layers. In Amphipappus the central area of the larger bracts, six to seven cells thick, consists of a layer of spongy 304 TABLE 1. Sources of material used in morphological studies of Amphiachyris, Amphipappus, Greenella, Gutierrezia, Gymnosperma, Thurovia, and Xanthocephalum (Voucher specimens are in KSC unless otherwise indicated) Species Collection Location Amphiachyris amoenum var. amoenum (Shinners) Solbrig A. amoenum var. inter- medium (Shinners) Solbrig A. dracunculoides (DC.) utt. Amphipappus fremontii orr. & Gray Greenella arizonica Gray G. ramulosa Greene Gutierrezia californica DC. G. mandonti ssp. mando- Ruffin 7033 Ruffin 7034 Ruffin 7035 Ruffin 7030 Ruffin 7031 Ruffin 7032 Ruffin 7036 Ruffin 7037 Ruffin 7038 Ruffin 7039 Anderson 3005 Clokey 5962 Peebles et al. 3842 (US) Anthony 321 (MO) Krikorian s.n.* Venturi 8156 (US) Texas: Montague Co., 7 mi NE of Montague. Texas: Johnson Co., 13.7 mi SW of Cleburne. Texas: Lampasas Co., 3.4 mi NW of Lampasas. Texas: Milan Co., N side of Ben Arnold between hwy and railroad. Texas: Washington Co., 2% mi NE of Burton at ject of hwys 390 and 1948. Texas: Montague Co., 7 mi NE of Montague. Kansas: Geary Co., 9 mi W of Junction City along hwy 77. Kansas: Riley Co., 13% mi S of Manhattan along hwy 177. Kansas: Chase Co., 25 mi NE of FE] Dorado. Texas: Brewster Co., along hwy 118 S of Alpine. New Mexico: Chaves Co., 5 mi N of Roswell. Nevada: Clark Co., Glendale Junction, Rocky Hills, Larrea Belt. Arizona: Pima Co., Papago Reservation. Lower California: San Bartolome Bay area. California: Fresno Co., Parkfield Grade, NW of Coalinga. Argentina: Prov. Jujuy; nii (Sch. Bip.) Solbrig G. microcephala DC. G. sarothrae Pursh G. texana DC. Gymnosperma glutino- sum (Spreng.) Less. Thurovia triflora Rose Xanthocephalum centau- roides Wi X. gymnospermoides (Gray) Benth. & Hook. X. humile (HBK.) Sch. Bip. X. linearifolium (DC.) Ruffin 7005, 7006 Ruffin 7001 Ruffin 7002 Ruffin 7003 Ruffin 7004 Ruffin 7007 Anderson 3028 Ruffin 7008 Ruffin 7009 Ruffin 7010 Ruffin 7011 Ruffin 7012 Ruffin 7013 Tharp 280 (TEX) Palmer 501 (MO) Ruffin 7024, 7025, 7026 Pringle 7422 (US) Leavenworth 950 305 Dept. Rosario de Lerma. New Mexico: Grant Co., 15 mi N of Silver City off hwy 25, 6600 ft. (7005); 8800 ft. (7006). New Mexico: Grant Co., 6 mi N of Silver City along Signal Peak Road. Kansas: Logan Co., 20 mi SW of Oakley along hwy 83. Kansas: Scott Co., 10 mi E of Scott City. Kansas: Ness Co., 13 mi NE of Ness City along hwy 283. exas: Brewster Co., hwy 118 S of Alpine. Texas: Burleson Co., along hwy 290; S of Giddings. Texas: Washington Co., 1 mi NE of Burton along hwy 390. New Mexico: Grant Co., 5 mi N of Silver City on Cottage Sand Road. New Mexico: Hidalgo Co., 1 mi W of Cloverdale on hwy 77. along Texas: Brewster Co., 21 mi S of Marathon on road to Big Bend National Park. Texas: Brewster Co., along hwy 118 S of Alpine. Texas: Jackson Co., Coastal flat near mouth of Lavaca River. Mexico: City of Durango and Arizona: Cochise Co., Garden Canyon, Huachuca Mts., 6000 ft (7024); 5500 ft (7025); 6200 ft (7026). Mexico: Distrito Federal; northern puebla. Mexico: Serrania de Ajusco 306 Greenm. near Cima Station. X. sericocarpum Gray Palmer 143 (US) Mexico: city of Durango and vicinity. X. wrightti (Gray) Gray Ruffin 7020 New Mexico: Catron Co., 18 mi NE of Mogollon on hwy 78. Ruffin 7021 New Mexico: Grant Co., 11 mi NE of Silver City at bot- tom of Cherry Creek Canyon. Ruffin 7022, 7023 Arizona: Cochise Co., Garden Canyon, Hauchuca Mts., 6000 ft (7022); 5500 ft (7023). “Gutierrezia campornice has been described as being restricted to the area of San Francisco Bay, California (Solbrig, 1960). Material vollected by Krikorian places this species about 80 mi § of its eee known range TABLE 2. Averages (in mm) for floral features of Amphiachyris, Amphipappus, Greenella, Gutierrezia, Gymnosperma, Thurovia, and oe involucre flowers DOA mi 2 = 2 = = _s 8 € 3 a/o2 2 ¢¢ 2 8 5 5 5 o oo 2 Ss 2» | FF S&S F&F 8 os 6 5 Ss le G & = bs Species | Voucher | & EO s/2 3 & wa & 8B Amphiachyri is Ruffin 7033 3.8 8 Ey 40 16. 2 8.0 26.9 — 1.0 2.4 0.6 amoenum var, Ruffin 7034 4.0 3.6 16.3 7.9 266 — 0.9 2.6 0.6 amoenum Ruffin 70385 3.6 3.8 15.5 8.0 24.2 0.9 2.4 0.6 A, amoenum var. Ruffin 70380 3.6 3.3) 14.2 8.0 19.4 — 1.0 2.5 0.7 intermedium Ruffin 7031 3.8 3.38 14.7 8.0 20.6 — 0.9 2.6 0.7 Ruffin 70382 3.7 3.5 15.0 8.0 20.38 — 0.9 2.6 0.7 A. dracunculoides Ruffin 7086 2.7 2.3 14.7 8.0 28.5 — 1.0 2.4 0.5 Ruffin 7087) 2.8 2.5 15.0 8.0 28.1 — 1.0 2.5 0.6 Ruffin 7038 2.6 2.2 15.4 7.9 25.3 — 1.0 24 0.6 Ruffin 70389 2.9 2.5 14.3 8.0 26.2 — 10 2.4 0.6 Amphipappus Clokey 5962 4.4 2.0 8.0 1.6 4.2 1.0 2.6 3.6 0.7 fremontit Greenella arizonica Peebles et al, 3842 3.6 3.4 16.9 13.0 30.4 1.0 1.0 2.9 0.6 307 Gutierrezia californica Krikorian sn. 6.1 25 194 88 10.6 12 1.0 3.2 0.9 G. microcephala Ruffin 7005 3.1 10 45 12 13 — 10 2.3 0.5 Ruffin 7006 3.2 10 44 11 15 — 10 2.4 0.5 G. sarothrae Ruffin 7001 3.4 1.5 116 50 3.6 1.0 1.1 2.7 0.6 Ruffin 7002, 3.2 1.5 12.1 5.0 3.4 1.0 1.1 2.6 0.6 Ruffin 7003 3.8 1.4 11.2 4.9 3.2 10 11 2.6 0.7 Ruffin 7004 3.2 1.5 11.8 5.0 3.4 10 1.0 2.6 0.7 G. texana Ruffin 7007 3.1 3.3 17.2 10.4 17.2 1.0 08 1.9 0.8 Ruffin 7008 3.1 3.5 16.6 10.3 17.4 1.0 09 1.7 0.8 Ruffin 7009 3.3 3.2 17.9 11.8 17.9 1.0 0.9 1.8 0.8 Gymnosperma Ruffin 7010 3.8 16 10.1 71 44 10 13 3.3 0.8 glutinosum Ruffin 7011 3.6 16 11.0 7.3 48 1.0 1.3 3.3 0.8 Ruffin 7012, 3.9 1.7 10.6 7.4 49 1.0 1.2 3.2 0.3 Ruffin 7013 4.0 1.7 104 7.1 46 1.0 1.3 3.1 0.3 Thurovia triflora Tharp 280 3.5 2.1 7.2 0.0 3.0 1.0 0.0 24 0.8 Xanthocephalum Palmer 501 4.7 5.7 43.2 32.6 49.4 1.0 1.0 4.0 0.7 centauroides X. gymnospermoides Ruffin 7024 5.0 5.9 42.9 37.6 160.0 1.0 1.0 42 0.6 Ruffin 7025 4.9 5.5 41.8 34.3 157.8 0.9 1.0 4.1 0.6 Ruffin 7026 5.2 6.3 43.7 41.5 162.7 1.0 11 44 0.5 X. linearifolium Leavenworth 6.9 24.6 13.2 67.8 10 1.0 3.2 0.6 X. sericocarpum Palmer 143 5.7 6.7 33.2 32.4 120.6 1.0 0.9 3.5 0.3 X. wrightii Ruffin 7020 5.3 5.9 22.2 12.7 47.1 1.0 1.0 3.7 0.8 Ruffin 7021 4.7 5.6 244 12.4 45.6 10 1.0 2.9 0.8 Ruffin 7022 5.3 5.9 20.3 12.0 46.3 1.0 1.0 3.6 0.7 Ruffin 7023 5.4 6.0 23.4 11.9 47.4 1.0 1.1 3.7 0.6 mesopyhll, three to four rows of sclerenchyma adjacent to the abaxial epi- dermis, and two to three rows of parenchyma between the sclerenchyma and the adaxial epidermis. In all species studied, the inner bracts, in contrast to the outer, are rela- tively longer and narrower. Particularly noteworthy is the strong alignment of bracts in Greenella ramulosa (Fig. 15d), a condition not encountered in any other species. Uniseriate as well as multiseriate glandular trichomes may occur along the outer as well as the inner epidermis of most species of Gutierrezia, Greenella, Xanthocephalum, Thurovia, and Amphipappus. However, in G texana, G. sarothrae, X. gymnospermoides, and Amphipappus, trichomes about two to three cells high are especially abundant along both surfaces. In Gymnosperma and Amphiachyris, glandular trichomes occur almost ex- 308 clusively along the adaxial epidermis, The receptacles of most species studied appears flat or somewhat convex. However, in X. sericocarpum, the receptacle is more conical in form. Occurring on the receptacle of all species of Gutierrezia, Greenella, and Thurovia and of most species of Xanthocephalum are characteristic hook- like glandular trichomes (Fig. 14b-d); these are lacking only in X. gymno- spermoides, X. humile, and X. centauroides. They usually are four to eight cells long. The presence of these glands in the above genera appears to be a unifying feature; they are absent in Amphiachyris, Amphipappus, and Gymnosperma., Achene. In all of the genera both disc and ray achenes show little mor- phological variation, appearing turbinate and slightly flattened. A single vascular bundle from the receptacle extends into the base of the disc achene. According to Carlquist (1961), this condition is common in most Compositae. Bundle number in the groups is two to 10 in the ovary. Gutier- rezia taltalensis, Greenella ramulosa, and Gymnosperma glutinosum, with ten outer vascular bundles in the achene wall, are regarded as representing a more primitive condition. Xanthocephalum gymnospermoides, with two to three achene bundles, represents the advanced condition. Greenella ari- zonica is particularly interesting in terms of the secretory canal distribution on the wall of the achene. There are five bundles but 10 secretory canals. A secretory canal is adaxial to each bundle and one is between each two bundles. Amphipappus exhibits five to seven achene bundles with no secre- tory canals on the achene wall. In the other species the number of secretory canals correlates well with the number of bundles in the achene wall. In Gulierrezia microcephala (ig. 6) and all species of Amphiachyris (Figs. 9, 10, 11) the dise florets are aborted, but five bundles are present in the outer wall of the ray achene, Thurovia triflora shows about the same degree of specialization as the majority of Gutierrezia and Xanthocephalum species although commonly nine bundles were seen in the achene wall with a secre- tory canal adaxial to each bundle. Trichomes in all of the genera, when present on disc or ray achenes, are of the ‘“‘Zwillingshaar” type, a biseriate non-glandular trichome. In all spe- cies studied the tips of the two apical cells usually do not terminate at the same point. The length of these trichomes is generally 70-550 pm. The shortest of these occur in Gymnosperma, and the longest, in Thurovia. The only other variation noted was the papillate hairs of Thurovia and X. gym- nospermotdes; the trichomes appear to arise from little mounds on the achene wall (Fig. 14a). Dise and ray corollas. The ray corollas show significant differences in average number, width, and length among genera as well as within a single genus. The five-lobed disc corollas show minor differences in length and width but vary tremendously in terms of number per head. In all of the species studied the tendency is toward more disc florets than ray florets. Tables 2 and 3 show the counts and measurements of these. 309 TABLE 3. Averages (in mm) for floral features and leaves of Amphiachyris, Amphipappus, Greenella, Gutierrezia, Gymnosperma, Thurovia, and Species Xanthocephalum Voucher Amphiachyris amoenum var. amoenum A. amoenum var. intermedium A. dracunculoides Amphipappus fremontii Greenella arizonica Gutierrezia californica op) r. microcephala op) . sarothrae G. texana Gymnosperma glutinosum Ruffin 7033 Ruffin 7034 Ruffin 7035 7030 Ruffin 7031 Ruffin 7032 Ruffin 7036 Ruffin 7037 Ruffin 7038 Ruffin 7039 Clokey 5962 Peebles et al. 3842 . , length of ray Krikorian s.n, 3. Ruffin 7005 Ruffin 7006 Ruffin 7001 Ruffin 7002 Ruffin 7003 Ruffin 7004 Ruffin 7007 Ruffin 7008 Ruffin 7009 Ruffin 7010 Ruffin 7011 Ruffin 7012 Ruffin 7013 width of ray or to tO | aa 1 1.8 | length of style appendage (disc) length of style branch length of style branch (ray) — a) oOo oO _ jo length of pappus (disc) | | bo ew) 2.9 2.2 0.1% length of pappus (ray) 0.1% leaf width | ! | —) = HRY EYE YL DY | leaf length 310 Thurovia triflora Tharp 280 0.0 0.0 08 14 00 16 00 1.0 2.5 Xanthocephalum Palmer 501) 6.2 19 03 1.0 09 01 O1 2.4 46 centauroides X. gymnospermoides Ruffin 7024 7.4 1.5 0.2 1.0 0.9 0.5 0.1% 19.3 82 Ruffin 7025 7.5 14 03 1.0 0.9 0.4 0.1* 14.7 72 Ruffin 7026 74 1.7 0.2 1.0 0.9 0.5 0.1* 17.8 80 X. linearifolium Leavenworth 9 2.1 13 15 0.7 04 O.1* 3.1 44 X. sericocarpum Palmer 143° 7.4 14 0.6 1.0 0.8 1.2 0.1% 2.1 34 X. wrightii Ruffin 7020 7.8 2.8 1.2 1.5 0.7 0.1* 0.1% 3.1 42 Ruffin 7021 7.9 2.8 1.2 1. 0.7 0.1* 0.1% 2.4 30 Ruffin 7022 8.1 3.1 1.2 1.5 0.7 0.1" 0.1" 2.2 36 Ruffin 7023 8.0 3.0 1.2 1.5 0.6 0.1% 0.1% 2.1 29 * slightly less than 0.1. Xanthocephalum shows the greatest degree of variation. In terms of num- ber of ray florets present, it ranges from nine to 13 in X. wrightii to as many as 45 in X. gymnospermoides. Gutierrezia ranges from one to two ray florets in G. microcephala to as many as 14 in G. texana. Greenella commonly shows 13 ray florets, thus corresponding to the Fibonacci sequence. The number of ray florets in Gymnosperma glutinosum ranges from six to eight. The length and width of the ray corolla exceeds that of the disc corolla in all of the species studied except in Gymnosperma, where the average length of the ray corollas is shorter than that of the dise corolla. Gymnosperma also shows a reduction in the number of teeth on the corolla tip. Most of the species commonly show three lobes at the apex of the ray corolla; Gymnosperma, commonly two lobes. Amphipappus also shows a reduction in number of ray florets. Commonly one or two are present. The number of ray florets in Amphiachyris also corresponds to a Fibonacci sequence. All species of Amphiachyris studied had eight fertile ray florets. Ray florets are absent in Thurovia. Histologically, the ray corolla of most species varies little. The tissues are composed mostly of parenchyma. At their broadest point the rays are five to eight cells thick in transection. The cells of the lower epidermis in all species are papillate (Fig. 15c), some more pronouncedly so than others. Secretory canals in the ray florets are found only in X. wrightii, where they appear adaxial (Fig. 15c) to each vascular bundle. Uniseriate trichomes, usually of one to three cells, occur on the outer sur- face of the lobes of the ray and dise corollas. These trichomes are very infrequent but usually more abundant on the lobes of the disc corolla. Xan- thocephalum gymnospermoides has both uniseriate and biseriate trichomes present on the lobes of its disc and ray corollas, As in the ray florets, the inner surface of the lobes of the disc corolla is characterized by an abundance of papillate cells. The outer epidermal cells 311 of the disc floret in all species are highly cutinized. The disc lobes are from two to four cells thick. In each of the five lobes of the disc corolla in all the genera are two lateral vascular bundles. Very large secretory canals are associated laterally with each bundle. However they are less frequent and less conspicuous in the disc corollas of X. wrightii and X. sericocarpum and are absent in Amphipappus. e vascular anatomy of the disc floret in all the genera appears to be fairly advanced. Carlquist (1961) has given a summation of vascular evolu- tion in Compositae. Generally five vascular bundles occur in the base of the disc corolla. These bundles usually extend to the base of each lobe where the individual bundles fork, with a vein entering each adjacent lobe along the margin. The two veins in each of the lobes then meet and fuse at the very top of the lobe. In all species studied, the tube of the dise corolla is five to eight cells thick just above the nectaries, and no secretory canals are present. Occa- sional biseriate and/or uniseriate glandular trichomes may be present on the corolla tube. However, in X. gymnospermoides, X. humile, and X. cen- tauroides, the entire lower half of the corolla tube is covered with conspicu- ous glandular trichomes (Fig. 15a). This occurrence is of particular interest since these three species were the only members of the Xanthocephalum group studied that lacked the receptacular glands. Pappus. Pappus development shows degrees of variation among as well as within genera and thus is a somewhat questionable taxonomic character as far as absolutely delimiting the groups. Generally speaking, all of the genera except Amphiachyris and Amphipappus can be characterized as having a scaly or reduced pappus in the disc as well as the ray florets. The reduced condition appears slightly more pronounced in Gymnosperma (Fig. 8a, b). The pappus of both ray and disc florets in Amphipappus is composed of long well-developed setae that are slightly shorter in the ray florets (Fig. 7a, b). The latter condition also characterizes the florets of Amphiachyris, except the pappus of the ray floret commonly possesses a low scaly crown (Figs. 9, 10, 11). Anatomically the pappus of the various species shows very little cellular differentiation. In no instances were vascular traces seen in the pappus. The pappus in cross section appears variously shaped and is usually two to four cells thick at the widest. tyle. As described by Solbrig (1960), the styles of the ray florets are dis- tinctly different from those of the disc ones in all of the species considered. hese styles are characterized by two elongated branches with a border of papillae along both sides of the stigmatic branches. On the style branches of the disc florets the papillae are confined to the lower quarter on both sides or are absent, with collecting hairs covering the greater outside por- tion of the branches. Solbrig, in his assessment of this group, also pointed out that the style branches of the disc florets are shorter than those of the ray ones. However, actual measurements of the branches (Table 2) show 312 that the style branches are shorter in disc florets only in Amphipappus. and Amphiachyris with the difference being slightly greater in Amphipappus. Thurovia triflora shows the greatest degree of stylar variation (Fig. 12b). All the other taxa differ in minor respects. Style branches of Thurovia are characterized by short collecting hairs on the upper half of the branches. Just beneath the collecting hairs and some distance from the base of the branches are peculiar brush-like hairs. According to Rose (1895), this pe- culiarity led Hoffman in 1895 to conclude that perhaps Thurovia belonged with Heliantheae instead of Astereae. In Xanthocephalum, Gutierrezia, and Greenella the styles of the dise florets possess triangular or elongate tips with collecting hairs occupying various degrees of the outer surface of the branches. Node and leaf. Nodal anatomy proved to be a very conservative feature. In all species it is trilacunar with one trace per leaf gap (Fig. 15b). Leaves of Xanthocephalum are alternate, linear-lanceolate to lanceolate, simple, and entire or shallowly toothed. Leaves of X. gymnospermoides and X. centauroides show the greatest amount of serration or toothing, but en- tire leaves may occur as well. Xanthocephalum gymnospermoides has the broadest leaves of all Xanthocephalum studied. Leaves of Gutierrezia, Greenella, Amphiachyris, Thurovia, and Gymno- sperma are also alternate and linear-lanceolate. Those of Gymnosperma are much longer than those of most species studied. Leaves of Amphipappus are alternate, entire, and, in contrast to those of the other species, oval to elliptic or obovate. Table 3 gives averages for both length and width of leaves of certain species. Figure 13 shows some of the shape and venation patterns noted. Leaf anatomy of the genera shows striking similarities. A pronounced cuticle covers the upper and lower leaf epidermis of all species studied. At regular intervals on both surfaces of the epidermis are narrow cavities. Within each of these cavities is one to two multiseriate glands (Fig. 16a, b). This condition was found in all the species except Amphipappus. Solbrig (1966) found this condition to be uniform in the South American Gutierrezia. In addition to the multiseriate glands within the cavities, X. gymnosper- moides, Gutierrezia texana, G. sarothrae, G. microcephala, and Thurovia show infrequent uniseriate trichomes on the surface of the upper epidermis as well. In Amphipappus, uniseriate trichomes two to three cells high are frequent on the upper and lower surfaces. In all species studied the mesophyll contains abundant chloroplasts throughout the tissue except for the parenchymatous sheath surrounding the collateral vascular bundles. The leaves have prominent secretory canals associated adaxially with the vascular bundles. DISCUSSION AND CONCLUSIONS Features of the pappus were used chiefly by earlier botanists in delimiting these genera with less emphasis being placed on other morphological fea- 313 tures. The character of the pappus, when coupled with features from other parts of the plant, is still a very important diagnostic feature. Solbrig (1960) points out that to use pappus characters successfully one must consider the pappus of both ray and disc florets. In addition one must comprehend the variation present in different species as well as the magnitude of vari- ation within a given genus. Gymnosperma and Greenella are described by Solbrig (1960) as being similar in possessing, in both ray and disc florets, a reduced crown of minute scales instead of a well developed pappus. Yet in Greenella ramulosa both ray and disc florets have a well-developed pap- pus formed by lanceolate palae that resembles the characteristic pappus of Gutierrezia. The pappus of Xanthocephalum has been described as having only a low rim or crown in both ray and disc florets. Yet in X. sericocarpum, X. linearifolium, and X. gymnospermoides it is not uncommon to find well- developed pappi resembling to a great extent that in Gutierrezia. Certainly the nature of the pappus in Amphiachyris and Amphipappus is distinct enough to characterize these genera. Differences in structure and the fact that the ray floret in Amphiachyris does not show a well-developed pappus may serve as evidence for not including these two in the same genus. The style of Thurovia definitely distinguishes it from the other genera. Even though these show some degree of coherence in style structure, it is worth noting that the style branch of the ray florets is longer than that of the disc florets in Amphiachyris and Amphipappus and no stigmatic pa- pillae are present on the lower surface of the style branch. These characters can also be used to segregate the latter two genera from the others. Amphipappus appears to demonstrate the greatest degree of differences in characters. In addition to those characters already mentioned, the structure of the leaves in Amphipappus is distinct from that of the other genera. In addition, secretory canals, except in the phyllaries, appear to be lacking. Sectioned leaves of all the other genera are almost indistinguishable, and secretory canals are frequent in various plant parts. Certainly the occurrence of the hook-tipped receptacular glands in all spe- cies of Gutierrezia studied, in Greenella, and in Xanthocephalum adds uni- fying evidence to these previously distinct genera. In addition to the habit of Thurovia and other features already mentioned, the presence of these hook glands in Thurovia perhaps suggests the close relationship of Thurovia to the latter genera. A more thorough survey of the presence or absence of these glands in Astereae may prove valuable in determining the extent and usefulness of this character, Because such few chromosome numbers were known in certain of the genera, a clear cytological concept of the groups was not perceived by Sol- brig. At the time of Solbrig’s study, the base chromosome number was con- sidered to be four or a multiple of four in all Gutierrezia counted, five in Amphiachyris, seven in Gymnosperma, and nine in Amphipappus. Only one species of Xanthocephalum (X. gymnospermoides, n=6) and none of Green- 314 ella had been counted. In addition, Tomb (personal communication) has found n=5 for Thurovia triflora. Table 13 shows that counts now available for Greenella arizonica and for four more of the eight species of Xantho- cephalum indicate a base of n=4. These findings certainly eliminate the differences suggested by Solbrig (1960) among Xanthocephalum, Gutierrezia, and Greenella. TABLE 4. Previous meiotic chromosome counts in Amphiachyris, Amphipappus, Greenella, Gutierrezia, Gymnosperma, Thurovia, and Enea scecsacl "Species a Counts — Reference a Habit: Amphiachyris 5 Solbrig et al. | (1964), annual dracunculoides Solbrig (1960). Amphipappus n=J Raven et al. (1960), perennial fremontii Solbrig (1960). Greenella n=4 Solbrig et al. (1964). annual arizonica Gutierrezia n—8, 12, 16 Rudenberg & Solbrig (1963). perennial californica G. mandonii ssp. n=12 Solbrig (1966). perennial mandonii G. microcephala n—8, 16 Solbrig (1960), perennial Rudenberg & Solbrig (1963). G. sarothrae n=4, 8 Solbrig (1960), perennial Rudenberg & Solbrig (1963). n=8 Solbrig et al. (1964), Solbrig (1965). G. texana n=4 solbrig (1960), annual Turner & Ellison (1960), Rudenberg & Solbrig (1963), Solbrig et al. 1964). Gymnosperma n=7, 8 Solbrig (1960), perennial glutinosum Solbrig et al. (1964), Solbrig et al. (1969). Thurovia triflora n=5 Spencer Tomb annual (Unpublished count). Xanthocephalum n—6 Raven et al. (1960), annual gymnospermoides Solbrig (1960), Turner et al. (1962), Dejong & Longpre (1963). X. humile n—4 Turner et al. (1962). annual 315 X. linearifolium n=4 Turner (Solbrig, 1961), perennial Beaman & Turner (1962). XxX, sericocarpum n=4 Dejong & Longpre (1963). annual X. wrightii n=4 Solbrig (1961). annual Based on morphological and anatomical data and also recent cytological findings, I believe that the most reasonable concept for this complex would be to merge Xanthocephalum, Gutierrezia, and Greenella into one genus and to maintain Gymnosperma, Amphiachyris, Amphipappus, and Thurovia as distinct genera. Since Xanthocephalum is the earliest name in the group to be merged, it is the correct name for it. Even though Thurovia shares many morphological and anatomical fea- tures with members of the Xanthocephalum complex, its absence of ray flowers, its style, and its base chromosome number suggest that further bio- systematic studies are needed to assess its relationship with Xanthocepha- lum. However, morphological, anatomical, and cytological evidence clearly indicates that Greenella and Gutierrezia should not be maintained as dis- tinct and show enough unifying characters to be transferred into Xantho- cephalum. KEY TO THE GENERA A. Pappus of disc florets composed of well-developed setae basally united, almost equaling corolla. B. Involucre elongate-turbinate; ray florets 1-2; disc florets fewer than 10: pappus of both ray and disc florets composed of well-developed setae; perennial shrubs. . Amphipappus BB. Involucre broadly Giacni te ray raters more nae 9; disc florets more than 10; pappus of ray florets reduced to a low crown; annuals herbs (or possibly short-lived perennials) . . Amphiachyris AA. Pappus scaly or reduced to low crown in disc as well as in ray florets. C. Ray florets absent . . . . . 2... . OT hurovia D. Ray corollas shorter than or equal to the disc corollas Gymnosperma DD. Ray corollas longer than dise corollas . . . . Xanthocephalum GENERIC SYNONOMY AMPHIACHYRIS Nutt., Trans. Amer. Philos. Soc. 7:313. 1840. AMPHIPAPPUS Torr. & Gray, Boston J. Nat. Hist. 5:107. 1845. GYMNOSPERMA Less., Syn. Gen. Comp. 194. 1832. (nom. conserv.) 6. 1819. THUROVIA Rose, Contr. U. S. Nat. Herb. 3(5): 320. 1895. XANTHOCEPHALUM Willd., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 1:132. 1807 Brachyris Nutt., Gen. 2:163, 1818. 316 Brachyachyris Spreng., Syst. 3:574. 1825. Greenella Gray, Syn. Fl. N. Amer. 1:480. 1884. Gutierrezia Lag., Nov. Gen. et Spec. 30. 1816. Hemiachyris DC., Prodr. 5:313. 1836. Odontocarpha DC., Prodr. 5:71. 1836. ACKNOWLEDGEMENTS My appreciation is expressed to Dr. Loran C. Anderson for interesting me in this investigation and for giving helpful suggestions during the course of the study and preparation of the manuscript; to Drs. T. M. Barkley, L. C Hulbert, O. W. Bidwell, and C. W. Shenkel for reading the manuscript and offering suggestions and criticisms; to the late Dr. L. H. Shinners for sup- plying information on collection sites and for making valuable suggestions in work related to evolutionary relationships among the genera studied; and to Dr. Spencer Tomb for providing unpublished data and valuable advice. Financial support from the Kansas Agricultural Experiment Station and Grant RR-8006, General Research Support Branch, Division of Research Re- sources, National Institute of Health, is gratefully acknowledged. REFERENCES ANDERSON, L. C. 1963. Studies on Sai (Compositae). Anatomy, cytology, taxon- omy. Trans. Sere Acad. Sci. 66:632—68 BAILEY, IL W. 1953, The anatomical are to the study of genera. Chron. Bot. BEAMAN, J. H. and B. L. TURNER, 1962. Chromosome numbers in Mexican and Guate- malan Compositae. Rhodora 64:271—276. BENTHAM, G. and J. D. HOOKER. 1873. Genera plantarum 2:553. London. CAMP, W. H. 1940. The concept of the genus V. Our changing generic concepts. Bull. Torrey Bot. Club 67:386. CARLQUIST, S. 1961. Comparative plant anatomy. Holt, Rinehart, and Winston, New Tee CORRELL, D. S. and M. JOHNSTON, 1970. Manual of the vascular plants of Texas. Texas Research rah Renner, Texas. DEJONG, D. C. D. and FE. K, LONC GPRE. 1963. Chromosome studies in Mexican Com- positae. Rhodora 65:225—240. FUCHS, C. 1963. Fuchsin. staining i NaOH clearing for lignified elements of whole plants or plant organs. Stain Tech. 3 —144. aR oe . Notes on oo and characters of certain genera and species. Proc. Amer. Acad. Arts 8:632—63 ———.,, 1884, eae ee of North America. 1:480. New York. HOWE, T. D. 1970. The female gametophyte of Gutierrezia microcephala, Gutierrezia tex- ana, and Amphiachyris dracunculoides. Abstracts ofp: ape ers presented at the meeting of Botanical Society of America, Indiana University. Am J. Bot. 57:3 NELSON, A. or Rock Mountain Herbarium studies, tl. Amer, J. Bot. 21:573-—582 PORTER, C. L. 1943. The genus — a aie Gray. Amer. J, Bot. 30: 81- —483, RAVEN, P. H., . lr. SOLBRIG, D. S, and R. SNOW. 1960. Chromosome num- ber in Compositag. I. Astereae. rates . ene 7:124—-132. R ae J. N. 1895. Thurovia, a new genus of Compositae. Contr. U.S. Natl. Herb. 20—322. RUDE NBERG, L. and O. T. SOLBRIG. 1963. Chromosome eine and morphology in the genus Gutierrezia (Compositae). Phyton po entina) 20:199-204 ASS, J. E. 1958. Botanical microtechnique. 3rd ed. Iowa State Daven Press, Ames. oe RS, L. 1950. Notes on Texas Compositae IV. Field & Lab. 18:25—32. 317 SOLBRIG, O. T. 1960. The status of the genera Aanbiibab bus, Amphiachyris, Greenella, Gutierrezia, Gymnospernia and ae (Composita 62:43—S54. ———. 1961. Synopsis of the genus Xanthocephalum Cee. Rhodora 63:151-164. ———., 1965. The California species of ee fierrezia (Compositae). Madrofio 18:75-84. —.. 1966. The South American species of Gutierrezia. Conte Gray Herb. 197:3—42. SOLBRIG G, O. T., L. C. ANDERSON, D. W. KYHOS, P. H. RAVEN, and L. RUDEN- BERG. 1964. “Chicmecomne numbers in Compositae V. . Il. Amer. J. : SOL BRIG, O. ce L. C. ANDERSON, D. W. KYHOS, and P. H. RAVEN. 1969. Chromo- some numbers in Compositae VIL. Astereae III. Amer. J. Bot. 64:285-289. TURNE B. L. and W. L. ELLISON. 1960. Chromosome numbers in the Compositae. I. i ee counts for 25 species of Texas Compositae, including 6 new gen- eric — xas J. Sci. 12:146-151. TUR B. L., M. tn LL, and R. M. KING. 1962. Chromosome eee in the ee a Additional Mexican and Guatemalan species. Rhodora 64:251—271. 318 aoe SSW. le LAE ee Fig. 1. Flowering head of Gutierrezia texana X20: a. Ray flower X16. b. Dise flower X20. c. Style branch of ray flower X30. d. Style branch of disc flower X25. 319 Fig. 2. Flowering head of Greenella arizonica X16: a. Ray flower X10. b. Dise flower X10. c. Style branch of ray flower X16. d. Style branch of disc flower X10. 320 Fig. 3. Flowering head of Xanthocephalum wrightii X25: a. Ray flower X10. b. Disc flower. Note the presence of glabrous achene, X10. c. Style branch of ray flower X20. d. Style branch of disc flower X16. Fig. 4. Flowering head of Xanthocephalum gymnospermoides X12: a. Ray flower X12. b. Disc flower X10. c. Style branch of ray flower X16. d. Style branch of disc flower X16. Fig. 95. Flowering head of Gutierrezia sarothrae X20: a. Ray flower X20. b. Dise flower X20. c. Style branch of ray flower X25. d. Style branch of dise flower X25. Fig. 6. Flowering head of Gutierrezia microcephala X20: a. Ray flower X16. b. Disc flower X20. c. Style branch of ray flower X25. d. Style branch of dise flower X25. 324 ay flower X25. Fig. 7. Flowering head of Amphipappus fremontii X20: a. R c. Style branch of ray flower X16. d. Style branch of dise flower X25. b. Disc flower X16. 325 Flowering head of Gymnosperma glutinosum X16: Note ray flowers which do not surpass the tubular ones giving the head a discoid appearance: a. Ray flower X20. b. Disc flower X20. c. Style branch of ray flower X25. d. Style branch of disc flower X20. 326 Fig. 9. Flowering head of Amphiachyris dracunculoides X20: a. Ray flower X30. b. Disc flower X25. c. Style branch of ray flower X30. d. Style branch of dise flower X25. ae ee oe a m= _— a > XN a. Zz ~ pon 327 Fig. 10. Flowering head of Amphiachyris amoenum var. intermedium X16: a. Ray flower X16. b. Disc flower X20. c. Style branch of ray flower X20. d. Style branch of dise flower X25. 328 Fig. ll. Flowering head of Amphiachyris amoenum var. amoenum X16: ay flower isc X16. b. Dise flower X20. c. Style branch of ray flower X20. d. Style branch of dise flower X20. Fig. 12. Flowering head of Thurovia triflora X20: a. Disc flower X16. b. Style branch of disc flower X16. 330 = li, a Fig. 13. Leaf shape and venation. Only major veins are drawn: a. Xantho- cephalum linearifolium. b. X. gymnospermoides. c. X. wrightii. d. X. centauroides. e. X. sericocarpum. f. Gymnosperma glutino- sum. g. Greenella arizonica. h. Thurovia triflora. i. Amphipappus fremontii. j. Gutierrezia sarothrae. k. G. taltalensis. 1. Amphiachy- ris dracunculoides. m. A. amoenum var. intermedium. Fig. 14. Longitudinal sections: a. Xanthocephalum chee ala flower showing biseriate non-glandular ‘‘zwillingshaa trichomes. Note trichomes are situated on ‘‘mounts’’ aed on the wall of the achene. Also ncte nectary glands surrounding the style base, X89. b-d. Hook-like receptacular glands: b. Gutierrezia texana, X89. c, Greenella arizonica, X351. d. Xanthocephalum wrightti, X89. Fig. 15. we . a. Longitudinal section of Xanthocephalum humile flower showing abundant occurrence of glandular trichomes along lower half of corolla tube, X89. b. Cross-section of Gutierrezia texana stem showing trilacunar nodal structure, c. Xanthocephalum wrightii ray cross-section showing papillate epidermal ray cells and secretory canals adaxial to each vascular bundle, X89. d. Greenella ramulosa showing vascular bundle with conspicuous se- cretory canal. Note strong alignment of involucral bracts, X89. 333 Fig. 16. Leaf transections showing the presence of glands within cavities: a. Gutierrezia texana, X351. b. Gymnosperma glutinosum, X FURTHER COMMENTS ON STYRAX L. RICHARD A. HOWARD Arnold Arboretum, Jamaica Plain, Massachusetts 02130 In his recent monograph of Styrax (Sida 5: 191-258. 1974) Gonsoulin treated the species from the Lesser Antilles in such a way as to require comment and correction in relation to my own work on the flora of the islands. Curi- osity prompted the checking of one other taxon with the result that a new combination is proposed. I leave it to others to note these suggestions and to check the work further. Gonsoulin failed to explain his treatment of the gender of the name Styrax and was not consistent in the specific epithets used. In many cases the original spelling is altered without noting such changes. Styrax has been and can be treated as masculine, feminine, or neuter. However, Wood and Channell made a recommendation in a paper cited by Gonsoulin (Jour. Ar- nold Arb. 41: 1-35. 1960) on which he offered no comment. Wood and Chan- nell pointed out that Linnaeus consistently treated the name as neuter and that, under Recommendation 75A (1) of the International Code of Botanical Nomenclature, Linnaeus’ practice in this matter should be followed. The following neuter endings are preferred: Styrax americanum, S. grandifoli- um, S. argentum, S. conterminum, S. obtusifolium, S. ochraceum, and S. glabrum. It should be pointed out that general agreement on the gender to be used does not exist in even more recent practice. Entry 11/06408 in the Index Nominum Genericorum acknowledged the type species of Styrax to be “S. officinalis L. (officinale’).’’ Stearn (Bot. Latin 76. 1966) stated that the Pl. and Ferns, 8th ed., 1116. 1973) referred to ‘‘Styrax officinale.’ The rec- ommendation of the International Code of Botanical Nomenclature to follow “general usage’ can best be implemented by following the practice of the original author Linnaeus. In reference to the species in the Lesser Antilles, the editorial style adopt- by oe soulin is confusing and misleading. Swartz (Nov. Gen. Sp. PI. Prod. ei cee ublished S. es not @ glaber as used by Gonsoulin (op. — 1788, Cavanilles also published the same binomial as ‘‘Sty- Trax ee branes an ilegitimate substitute for S. americanum (En- cycl. Meth. 1: 82. 1783). Gonsoulin correctly listed the Cavanilles binomial in the synonymy of S. americanum but cited incorrectly the publication date as 1790. Although “‘Styrax glabrum Cav.”’ is illegitimate, a fact unnoted by Gonsoulin, the problem of two different uses of the same binomial in 1788 is easily settled by reference to Stafleu’s Taxonomic Literature (Reg. Veg. 52: 79, 462. 1967). The pertinent fascicle of Cavanilles’ Monodelphia classis dis- sertationes decem 6: page 340, plate 188, fig. 1 was published between Octo- SIDA 5(5): 334—337, 1974, 339 ber 12 and 25, 1788, while Swartz’ Prodromus was published in July of the same year. In the synonymy of S. glabrum Sw., Gonsoulin attributed the binomial S. occidentalis to Swartz ex Thunberg. This name also might have been indi- cated as illegitimate, as it was proposed in a thesis eae by L. J. Prytz, a student of Thunberg, when he noted the existence of S. glabrum Cav. and renamed S. glabrum Sw. as S. occidentalis Swartz. Why the name was at- tributed to Swartz is not clear. The citation of Swartz ex Thunberg is correct. To the synonymy of S. glabrum Sw. should be added Morisonia imrayl Griseb. (Fl. Brit. W. Ind. 19. 1859). Grisebach cited an Imray collection (K) from Dominica, and Urban noted this misidentification by Grisebach in a comment following the description of S. portoricensis (Bot. Jahrb. 15: 338. 1892). The designation of a holotype for S. glabrum Sw. also requires some com- ment, for Gonsoulin, possibly influenced by Urban’s citation of this record (op. cit.), has indicated it to be ‘‘Willdenow 8326.’ However, Willdenow 8326 is the number of an herbarium entry, actually two sheets, in the Will- the species, Swartz (Prodr. 74. 1788) stated ‘India occidentalis, Vincenti’’ with the classical symbol for a perennial or woody plant. Swartz later elab- orated on the description and stated (Fl. Ind. Occident. 2: 848. 1800) ‘‘Habi- tat in India occidentali; Ins. Sti Vincentii (Anderson).’’ Such a specimen from the island of St. Vincent, where Anderson was superintendent of the Bo- tanical Garden from 1785-1811, is in the herbarium of the British Museum (Natural History). This is the holotype. There are two sheets in the Willde- now herbarium as seen on microfiche (IDC 7740. 567. III. 1,2), the first a spe- cimen with flowers and the second a sterile specimen of but two leaves. Both specimens bear comparable labels mentioning the Swartz Prodromus page 74 and the habitat of ‘India occidentalis,’’ and both were annotated by Per- kins. The index to the Willdenow Herbarium on microfiche (IDC 7740. 27. III. 7) suggested with a question mark that the specimens might have come from the Vahl herbarium. The flowering specimen with its shortly acute to sub- truncate leaf apices appears to be a good match for a specimen collected by Ryan on Montserrat and deposited in the herbarium at Copenhagen, The questioned reference to Vahl’s herbarium may be accurate. Several errors of geography are also found in Gonsoulin’s citations. Bains- Jaunes, not ‘‘Bains-Tames,”’ is the classical collecting locality in Guade- loupe. ‘“‘Herbier de la Martinique’ is not a locality but a label heading. Duss collections cited under this title present other problems, for many specimens from different localities were distributed under the same num- ber. Duss himself (Fl. Phan. Ant. Franc. 389. 1897) noted that his collection 1728, a collection cited by Gonsoulin, was found in the ‘‘Bois de la Calebasse, de lAjoupa-Bouillon, de Fontaine-Chaude, de Case-Pilote.’’ In other cases a single Duss number may represent a mixed collection, or two or more 336 numbers may be given to a single specimen. Even a single location on a given herbarium label may vary from one herbarium to another and so can not be trusted. The Cowan collection from Tobago was made ‘‘Roxborough- Parlatuvier Road, Main Ridge: primary forest, 450-500 m. elev.’’—not at “Orxborough,”’ which is an unfortunate typographical error. Gonsoulin’s description of S. glabrum does not cover the full range of variation in the size and shape of the leaves of the species. He apparently followed the descriptions given by Urban (Bot. Jahrb. 15: 338. 1892) and Perkins (Pflanzenr, IV, 241: 63. 1907) but did not mention that both authors noted that their descriptions do not agree with that given by Swartz. Swartz described the leaves as lanceolate-oblong, acuminate, and subserrate to en- tire. The holotype has such characteristics, and I suspect this leaf shape is associated with vigorous shoots. Other specimens from the Lesser Antilles have shorter, elliptical to ovate leaves with apices acute to rounded or sub- truncate. A single specimen labeled Duss 3262/3707 (P) from Guadeloupe has fruit twice the size described by previous authors. Specimens have also been seen from the islands of St. Kitts (Howard 16588 [A], Wadsworth 495 [A]), Montserrat (Squire s.n. [A], Ryan s.n. [C]), and Dominica. The oc- currence of S. glabrum on Dominica was mapped by Gonsoulin but no speci- mens were cited. The Imray collection originally described as Morisonia documents such a record. It is possible that two species are represented in the Lesser Antilles but only a few fruiting specimens are available and no material of flower and fruit from the same individual plant or population is available. Styrax glabrum of the Lesser Antilles requires further study and careful examination in the field. Gonsoulin listed as a generic synonym of Styrax the name Darlingtonia Torrey, but he apparently failed to see an additional complication in this reference which affects the nomenclature he adopted. In 1851, Torrey (Proc. Amer. Assoc. Adv. Sci. 4: 190-193) described a new plant collected by Fre- mont in California for which he proposed the name Darlingtonia rediviva. In 1853, Torrey (Smithsonian Contr. Knowl. 6(4): 1-8, pl. 12) described the insectivorous genus Darlingtonia with one species, D. californica. The gen- eric name had been used earlier by A. P. de Candolle and its use by Torrey for the insectivorous plant has been conserved (3131). In the 1853 paper, Torrey stated: ‘“. . . a Californian plant, on imperfect specimens of which, I had recently indicated a genus under this name, proves to be only a spe- cies of Styrax.’’ Torrey then stated in a footnote: ‘‘Having recently obtained good flowering specimens of this plant, the following description of it is appended: -Styrax californicum ... .’ [Emphasis added.] The binomial Darlingtonia rediviva remained in limbo until L. C. Wheeler reviewed the matter (Bull. S. Calif. Acad. Sci. 44(3): 94. 1945) and made the combination Styrax rediviva (Torrey) L. C. Wheeler. This new combination was over- looked by Gonsoulin and is not considered in recent floras of California. If a variety is to be recognized (as Gonsoulin and Rehder recognized var. 337 californica), then the correct name, citation, and synonymy are the follow- ing: Styrax officinale L. var. rediviva (Torrey) Howard, comb. nov. Darlingtonia rediviva Torrey, Proc. Amer. Assoc. Adv. Sci. 4: 190-193. 1851. Styrax rediviva (Torrey) L. C. Wheeler, Bull. S. Calif. Acad. Sci. 44(3): 94, 1945. Styrax californicum Torrey, Smithsonian Contr. Knowl. 6(4): 1-8, pl. 12. Styrax officinalis var. californica (Torrey) Rehder, Mitt. Deutsch. Dendrol. Ges. 1915: 226. : POLLEN MORPHOLOGY OF AMPHICARPAEA (LEGUMINOSAE: PHASEOLEAE) BLANCHE W. MEESON Herbarium, Biology Department, Southern Methodist University Dallas, Texas 75275 This study concerns itself with the determination of the pollen character- istics of the members of the genus Amphicarpaea. Based on the external morphology of the members of the genus Amphicarpaea, Turner and Fearing (1964) regarded the genus as consisting of three valid species, A. africana (Hook.) Harms., A. bracteata (L.) Fern. and A. edgeworthii Benth. Each of these species exists on a different continent, A. africana from Africa, A. bracteata from North America, and A. edgeworthii from Asia. It was also pointed out that A. bracteata and A. edgeworthii are often quite difficult to distinguish based on external characteristics. MATERIALS AND METHODS The pollen samples were prepared using the standard acetolysis method devised by Erdtman (1952, 1960). The filtration of pollen was performed during the first washing after acetolysis. The pollen grains were mounted in glycerin and sealed with paraffin. RESULTS The pollen grains of Am a africana, A. bracteata and A. edge- worthii were found to exhibit istinotls different characteristics. All species had spherical grains, but differed with regard to texture of the sexine and colpi-ora characteristics. Pollen characteristics of specimens previously identified as A. comosa were also examined and found to have pollen charac- teristics identical to A. bracteata. The pollen characteristics for the three species of Amphicarpaea are as follows: A. africana (Fig. 1) Spherical, tricolporate, zonotrene ora, reticulated sexine, blending into smooth regions near the colpi, diameter ca. Ow. (Uganda: Taylor 2845, ISC; Tanzania: Richards 24654, NY) A. bracteata (Fig. 2) Spherical, tetraporate, zonotrene ora, smooth sexine, ora slightly raised or lipped, diameter ca. 28. (U.S.A.: Demaree 16228, SMU; Kral and Godfrey 3562, SMU) A. edgeworthii (Fig. 3) Spherical, triporate, zonotrene ora, smooth sexine, ora slightly raised or lipped, diameter ca. 28. SIDA 5(5): 338—340. 1974. 339 Figs. 1-3. 1. Pollen grain of Amphicarpaea africana. Note reticulation and colpora. X 1600. 2. Pollen grain of A. bracteata. Note number of pores. X 1600. 3. Pollen grain of A. edgeworthii. Note number of pores. X 1600. (Japan: Charette 1401, MO; Kirkino 24104-364, SMU) DISCUSSION AND CONCLUSION The pollen grain morphology supports the results obtained by Turner and Fearing (1964). There are only three distinct species of Amphicarpaea based on pollen morphology. In regard to the relationship of these three species, it could be hypothesized that A. africana possesses pollen most similar to 340 that of the precursor of the genus because it possesses a primitive tricolpo- rate grain with a reticulated sexine. The other species, A. bracteata and A. edgeworthii, appear to be modifications of this basic form by loss of the colpi and reticulated sexine and an increase or retention in the number of pores. This supports the statement made by Turner and Fearing (1964) that africana is ‘“‘a very distinct species being more distant from A. edge- worthi and A. bracteata than the latter taxa are from each other.” In fact, the external characteristics of A. bracteata and A. edgeworthii are so simi- lar that Turner and Fearing (1964) stated: ‘“‘The taxa are almost identical and even the technical characters listed in the key to the species often fail to place an occasional specimen. If it were not for their continental isolation, the taxa would probably have been treated at no more than varietal rank.” Although the pollen shows similar characteristics for these species, it also indicates that they are distinctly different. On the basis of pollen morphology, the species can be distinguished as follows: 1. Pollen tricolporate . 2...) . | . . A. africana 1. Pollen porate. 2. Pollen tetra-pentaporate . . . . . . . . . A. bracteata 2. Pollen triporate A. edgeworthii Using the pollen ahavacienieies in “conjunetion with the ewe morpholo- , the members of the genus Amphicarpaea can be definitively identified. ACKNOWLEDGEMENTS I would like to thank Dr. Wm. F. Mahler for his continuous guidance while preparing the specimens and the manuscript and Dr. Joan Nowicke for her suggestions and comments. REFERENCES ERDTMAN, G. 1952. Pollen morphology and plant taxonomy-angiosperms. Vol. 1. Alm- qvist ee Wiksell, Stockholm. — 60. The acetolysis method. A revised description. Svensk Botanisk Tidskrift. SP TURNER, B. L. and FEARING, O. S. 1964. A taxonomic ee of the genus Amphicar- paea (Leguminosae). The Southwestern Naturalist, 9 (4) :207 WOOD ANATOMY OF AMPHIACHYRIS, AMPHIPAPPUS, THUROVIA, GYMNOSPERMA, AND THE XANTHOCEPHALUM COMPLEX JOHN RUFFIN Department of Biology, Atlanta University, Atlanta, Georgia 30314 The problem of delimiting Amphipappus, Amphiachyris, Greenella, Gutt- errezia, Gymnosperma, Xanthocephalum, and Thurovia (Compositae) has been reviewed by Gray (1873), Bentham and Hooker (1873), Rose (1895), Nelson (1934), Porter (1943), Shinners (1950), Solbrig (1960), Correll and Johnston (1970), and Ruffin (1971). Unfortunately, the phylogenetic relation- ship of the genera is still an open question. This is the second paper in a multiphase investigation intended to assemble new evidence toward a more satisfactory taxonomic treatment of the genera. Comparative mor- phological and anatomical studies—of ray and disc florets, phyllaries, re- ceptacle, achene, pappus, style, and leaves—and recent cytological findings reported in the previous article (Ruffin, 1971) disclosed evidence favoring the merger of the previous distinct Xanthocephalum, Gutierrezia, and Green- ella into one coherent genus and maintaining Gymnosperma, Amphiachyris, Amphipappus, and Thurovia as distinct genera. Xanthocephalum, the earliest name in the group merged, is used throughout this report to include the pre- viously distinct genera. Although the details of wood anatomy presented here are by no means conclusive it is hoped that they will be of value in properly interpreting the limits of and evolutionary relationship among the genera. MATERIALS AND METHODS Most wood samples were sectioned between 15-20 microns on a sliding mi- crotome and were stained with safranin or with safranin and fast green with tannic acid followed by ferric chloride. When material allowed, macerations were made using Jeffrey’s 10% chromic acid-10% nitric acid solution for those wood features difficult to measure from sectioned material. For wood samples taken from herbarium sheets, 71 measurements were made from each collection except Xanthocephalum ramulosa and X. _ lineari- folium where only 51 were made due to inadequate longitudinal sections. Thurovia triflora lacked wood rays; thus other wood features totalled 51 measurements. Wood averages from field material represent 20 measure- ments for each character, with the exception of widest vessel diameter, taken from two individuals in each population. In all, 4,697 measurements were made. In order to measure characters such as average diameter of vessels, SIDA 5(5): 341—352. 1974. 342 average number of vessels per group, average length of vessel elements, etc., where there are many measurable elements per section, a random sam- ple of such elements was measured. A starting point for scanning a slide was chosen by using random coordinates on the mechanical stage. Moving the slide horizontally (from right to left), I scanned the microscope field between two major lines on an ocular micrometer, Each element measured was given an arbitrary reference point. Whenever these arbitrary points fell within the two lines on the ocular scale, those characters were measured or counted. OBSERVATIONS AND RESULTS Table 1 summarizes qualitative and quantitative features for the species studied. Tables 2 thru 9 present a summation of quantitative characters in each of the genera. Xanthocephalum ramulosum and X. linearifolium have been omitted from Table 1 because of reasons stated elsewhere. However, those features that could be measured are included in the averages of the various genera. In addition, features not included in the tables will also be discussed, Vessels. Although I did not attempt to determine the exact frequency of vessels seen in a transection, vessels appear to be very numerous in most species of Xanthocephalum (X. gymnospermoides, X. sarothrae, X. micro- cephalum, X. texanum, X. wrightit). The vessels are also distributed in radial rows in the majority of Xanthocephalum species studied. Xantho- cephalum sericocarpum shows some tangential pairs, a few solitary vessels, and a few short radial rows. In X. microcephalum and X. sarothrae, solitary vessels as well as long radial chains are present. A significant condition in the latter two species is a tendency toward ring porosity. In each species the ring-porous condition takes the form of more numerous and larger ves- sels in the late wood (Fig. 1). Also noteworthy is the presence of vascular tracheids in the latter two species. Extensive grouping of vessels also characterizes most Xanthocephalum species. In a number of instances as many as 12 vessels per group re seen. In those members of the genus showing the ring-porous condition, the early wood exhibit smaller numbers of vessels per group whereas the late wood show extreme grouping. Vessel grouping is, however, less extensive in X. centauroides, X. mandonii spp. mandoniti, and X. sericocarpum. Vessels in the latter species are in groups of two to eight, mostly two to five. In Amphiachyris, vessels appear numerous in all species. Distribution and amount of vessel grouping are diverse and not so extensive as in most spe- cies of Xanthocephalum. This genus is characterized by solitary vessels, short radial rows of two to six vessels, and some tangential pairs. Amphi- achyris amoenum var. amoenum and A. amoenum var. intermedium, in every population studied, tend toward a ring-porous condition. Bot ave been described as annuals, but a simple type of growth ring is represented in which growth appears to be accompanied by diminution in diameter of Table 1, Comparative features of the wood in Amphiachyris, Amphipappus, Thurovia, and Xanthocephalum Gymnosperma, E ne 2 og : E ze 7 fe. Ge §2 G8 ©. » & . “65: By. Ee S) 5 Ss £5 Sa 5 = gs AS AS 25 -— Te Ge eE0OU oC (tia OE OR Ot OE EZ §: ae BE co. of = Ge ee cee oF Species Age Collection 3 $ ae @ Be z= a ra iS ZE 5 SE 25 Amphiachyris amoenum var. amoenum 2? Ruftin 7033 62.5 34.6 165.7 11.5 397.1 + 1.3 + + 3.7 a? Ruffin 7034 57.5 37.0 167.6 11.6 425.5 + — 1.5 + + 4.0 2 Ruffin 7035 66.0 33.0 185.5 11.9 453.3 + — 1.4 + + 3.0 A. amoenum var. intermedium 2? Ruffin 7030 62.5 38.0 176.2 11.1 424.4 + — 1.6 + -f- 4.0 22 Ruftin 7031 67.5 Sot 185.3 10.8 413.4 + — 1.3 + + 3.8 2? Ruffin 7032 725 42.0 156.1 11.6 431.5 + — 1.1 + + 4.3 A. dracunculoides 1 Ruffin 7036 35 31.3 119.0 13.2 455.2 + — 985 + + 3.2 1 Ruffin 7037 97.9 39.5 136.0 11.5 450.7 + — 852 0 + + 2.8 1 Ruffin 7038 60 38.1 139.8 15.4 473.7 + _ 9430 + + 2.6 1 Ruffin 7039 57.5 33.9 135.0 13.7 511.7 + — 686 + + 2.6 Amphipappus fremontii 7 Clokey 5962 50 29.0 119.5 10.6 304.5 = 1.2 + + 4.0 Gymnosperma glutinosum 8 Ruffin 7010 62.5 3L.3 133.9 10.2 133.8 — — 1.0 + + 3,2 | Ruffin 7011 67.5 29.5 111.8 10.2 359.9 — — 888 + + 3.6 9 Ruffin 7012 67.0 34.0 110.9 ny Wes 375.6 = — 946 + + 3.4 Ebe Thurovia triflora Xanthocephalum centauroides X. gymnospermoides X. mandonii spp. mandonit X. microcephalum ca . sarothrae a . sericocarpum X. texanum * wrightii — oO — SS eS Be ee ee Oo Ruffin 7013 Tharp 280 Palmer 501 Ruffin 7024 Ruffin 7025 Ruffin 7026 Venturi 8156 Ruffin 7005 Ruffin 7006 Ruffin 7004 Palmer 143 Ruffin 7007 Ruffin 7008 Ruffin 7009 Ruffin 7020 Ruffin 7023 62.5 Table 1. (Cont.) 29.2 31.6 122.0 116.0 +++4 +ettt¢etttzt++e+ +t +++ + ++tttttt+++4¢+4+4++ pre 345 Table 2. Comparison of vessel element diameter in Xanthocephalum, Amphiachyris, Gymnosperma, Amphipappus, and Thurovia Genus cae of Number of Range Range of means Mean individuals zm. zm. em Xanthocephalum 10 37 10-75 18.5-42.7 31.7 Amphiachyris 3 20 10-70 34.8-38.5 36.8 Gymnosperma 1 4 8-60 31.0 Amphipappus 1 1 10-50 — 29.0 Thurovia 1 1 20-50 — 31.0 Table 3. Comparison of vessel element length in Xanthocephalum Amphiachyris, Gymnosperma, _Amphipappus, and Thurovia Genus oS of Sanne i Ri Me Range of means Mean ta individuals um em Xanthocephalum 10 37 40-330 72.0-199.4 118.5 Amphiachyris 3 20 45-300 132.4-172.5 159.2 Gymnosperma 1 4 50-210 — 119.6 Amphipappus 1 . 1 90-150 — 119.5 Thurovia 1 1 §0-165 — 116.0 vessels (Fig. 3). This condition was not present in any of the populations of A. dracunculoides. Vascular tracheids were not observed in any species of Amphiachyris. Vessels in eerie glutinosum are very numerous. Vascular tra- cheids also occur in the xylem. Degree of ring porosity in the wood is strong- ly portrayed. In terms 7 vessel distribution, no one pattern seem to pre- dominate. Numerous solitary vessels, tangential clusters, and short radial rows are all pronounced. Grouping of vessels is not so extensive as in Xan- thocephalum but does resemble the grouping in Amphiachyris. In Amphipappus, vessels are also numerous. Some vessels appear solitary but most show extensive grouping in radial and tangential chains (Fig. 4). The ring-porous condition is also well developed. Large vessels are seen in early wood while narrower vessels and vascular tracheids occur in late wood. Vessels in wood of Thurovia show a somewhat unusual pattern. Most large vessels occur in primary tissues of the xylem while the more numerous narrower vessels and vascular tracheids are in the later-formed xylem tis- sue. Because of the difficulty of distinguishing vascular tracheids from nar- row vessel elements, patterns of vessel distribution and vessel grouping in Thurovia were very difficult to estimate. Mean vessel length and diameter for each species, range, range of means, and the mean for all species studied in each of the genera are represented in Tables 1, 2, 3. Mean vessel diameter and mean vessel element length in 346 Table 4. Comparison of libriform fibers diameter in Xanthocephalum, Amphiachyris, Gymnosperma, Amphipappus, and Thurovia Genus Number of Number of Re it ~ Range of means Mean taxa individuals um. wm Xanthocephalum 10 37 ~ 10,0-15.8 9.0-13.5 10.9 Amphiachyris 3 20 9.1-18.2 11.1-13.4 12.0 Gymnosperma 1 4 6.8-13.6 — 10.6 Amphipappus 1 1 9.1-11.3 — 10.6 1 1 9.1-11.3 — 10.9 Thurovia Table 5. Comparison of libriform fiber length in Xanthocephalum, Amphiachyris, Gymnosperma, Amphipappus, and Thurovia _ Genus oe of Number of Range Range ee means ince ndividuals um = 10 37 210-556 ~-263.5-463.2 340.1 Amphiachyris 3 20 240-695 423.1-472.8 440.4 Gymnosperma 1 4 270-480 — 310.3 Amphipappus 1 1 250-340 — 304.5 Thurovia 1 1 250-340 — 292.0 the various genera are quite similar except in Amphiachyris. In Amphiachy- ris mean vessel diameter and mean vessel element length exceed the figures noted in the other genera. Vessel-element perforations, in all the genera, were simple. In any given sample the end walls of the vessels may vary from transverse to oblique. Intervascular pits are 3-5 jem in diameter for all of the species considered and are alternately arranged. Helical thickenings are found on the inner vessel wall of many of the spe- cies studied. Helical thickening, however, is not noticeably present on the vessel wall of any of the species of Amphiachyris. These thickenings are es- pecially conspicuous on walls of the narrower vessel elements and on walls of vascular tracheids in Xanthocephalum sarothrae, X. microcephalum, Gymnosperma, Thurovia, and Amphipappus. In all of the above species ex- cept Amphipappus these helical thickening are grooves connecting the aper- tures of pits. In Amphipappus, coarse bands were noted on each side of a ser- ies of pits forming a helix on the inner vessel wall. None of the elements of the xylem of Amphiachyris and Gymnosperma was storied. Except for the vascular tracheids present in Xanthocephalum sarothrae, X. microcephalum, Thurovia, and Amphipappus, none of the elements in the xylem of these genera appeared storied. Libriform fibers. Average dimensions of libriform fibers appear related to dimensions of vessel elements. This is indicated by the fact that shorter fibers occur in those species with shorter, narrower vessel elements. This correlating of average dimensions is clearly shown by Tables 2, 3, 4, 5. The Fig. 1. Cross-section of Gutierrezia microcephala wood showing ring-porous condition. Note solitary as well as long radial chains of vessels, X89. 2. Tangential section of Xanthocephalum wrightti wood show- ing short and widely spaced rays, X89. 3. Cross-section of Amphi- achyris amoenum wood where growth appears to be accompanied by diminution of vessels, X89. 4. Cross-section of Amphipappus fremontii wood showing wide rays, arrangement, and distribution of vessels, X89. 348 Table 6. Comparison of maximum ray width in Xanthocephalum, Amphiachyris, Gymnosperma, Amphipappus, and Thurovia | Gariue Number of Number of Range Range of Niset taxa individuals Means Xanthocephalum 8 35 1-14 cells 1.9-5.5 cells 3.5 cells Amphiachyris 3 20 1-6 cells 2.8-4.0 cells 3.5 cells Gymnosperma 1 4 1-6 cells — 3.5 cells Amphipappus 1 1 2-6 cells — 4.0 cells Thurovia 1 1 No rays — Table 7. Comparison of multiseriate ray height in Xanthocephalum, Amphiachyris, Gymnosperma, Amphipappus, and Thurovia ee Number of Number of Range Range of means Mean taxa individuals mm mm mm Xanthocephalum 8 35 ().23-3.0 1.1 -2.1 1.1 Amphiachyris 3 20 0).25-2.7 ().86-1.4 1.2 Gymnosperma 1 4 0).23-2.1 — 96 Amphipappus 1 1 0.37-2.8 —_ 1.2 Thurovia 1 1 No rays _ — longer vessel element in Amphiachyris parallels the dimensions given for fiber length in this genus. Also, the large vessel elements in Xanthocephalum gymnospermoides correlates with the long fibers present in this species. Amphiachyris is also distinct from the other genera in that fiber dimor- phism occurs in the wood of all three taxa. Extremely long, thick-walled fibers and shorter fibers occur within the same wood samples. As pointed out by Anderson (1963) ‘‘this condition may be indicative of a trend toward greater parenchymatization which in turn may be related to a more her- baceous habit.’ Even though slight differences exist in fiber diameter both among as well as within genera, these fibers are considered to be somewhat narrow for the Astereae. According to Carlquist (1960) the diameter of libri- form fibers of most Astereae is 20-35 pm. Fiber walls of all species studied are 2.3-5.1 pm thick. Rays. According to Carlquist (1962a) elimination or loss of uniseriate rays is common in Compositae. Nanthocephalum demonstrates this clearly. In only three species, X. wrightii, X. gymnospermoides, and X. texanum, were uniseriate rays abundan n no instances, however, were uniseriate rays as frequent as niuitiseriate ones. Uniseriate rays in the three species are four to six cells high. In all of the other species of Xanthocephalum studied, uniseriate rays were completely absent or rare and very limited in height (usually one cell high). Average ray width in Xanthocephalum is wide (two to four cells) to very wide (four to six cells). Xanthocephalum wrightii, where most rays are bi- Fig. 5. 349 Tangential section of Xanthocephalum gymnospermoides showing high and wide rays, X89. 6-8. Longitudinal sections showing pris- matic crystals in ray cells: 6. Gutierrezia microcephalum, 7. Gu- tierrezia sarothrae, 8. Xanthocephalum centauroides, X89. 390 Table 8. Comparison of vessel element length in annuals*—Xanthocephalum, Amphiachyris, and Thurovia Genus — of Number of Range Range of means Mean individuals um num. um Xanthocephalum 4 21 50-330 110.5-199.4 146.7 Amphiachyris 3 20 45-300 132.4-172.5 159.2 Thurovia 1 1 90-165 — * Amphiachyris amoenum var. amoenum and A. amoenum var. intermedium may be biennials Table 9. Comparison of vessel element length in perennials— Xanthocephalum, Amphipappus fremontii, and Gymnosperma Genus — of Number of Range Range of means Mean individuals um nm mm Xanthocephalum 6 16 40-175 72.0-128.0 99.7 Amphipappus fremontii 1 1 90-150 — 119.5 Gymnosperma 1 4 50-210 — 119.6 seriate, and X. teranum are characterized by the former condition while the remainder of the species in this genus are characterized by the latter pattern. Ray width in species of Xanthocephalum seems to correlate with ray height: species characterized by wide rays also have high rays. In con- trast, those species with narrower rays are limited in their vertical extent. Xanthocephalum gymnospermoides (Fig. 5) and X. sarothrae show the high- est rays in this genus while X. wrightii (Fig. 2) shows the least amount of ray height development. In terms of ray number and distribution, as seen in tangential view, all species studied except X. wrightii showed numerous closely-spaced rays. In X. wrightii the rays are less numerous and widely dispersed. Uniseriate rays were lacking in Amphiachyris amoenum var. amoenum and A. amoenum var. intermedium and for the most part scarce in A. cunculoides. When present, they are one to four cells high. Ray cells are one to four cells wide in A. dracunculoides and two to six cells wide in the other two taxa. Ray height is moderately high in all three taxa. Ray height is 0.4-2.7 mm in A. amoenum var. amoenum and A. amoenum var. inter- medium and 0.25-1.8 mm in A. dracunculoides. In wood of Gymnosperma glutinosum uniseriate rays one to two cells high were occasionally observed. e rays were one to six cells wide. Average ray height for the populations studied is 0.88—1.0 mm. In Amphipappus fremontii no uniseriate rays were observed in the wood. The multiseriate rays ranged from two to six cells wide and 0.37-2.8 mm high. Rays of the genera studied are heterocellular, having both erect and pro- 301 cumbent cells. However, both multiseriate and uniseriate rays are absent in wood of Thurovia. Carlquist (1966) lists the few genera of Compositae in which rayless woods have been found; Thurovia can now be added to the list. Barghoorn (1941) states clearly conditions which may promote ray- lessness. Resin deposits and crystals. Resin deposits are noticeably present in wood of all species studied except Xanthocephalum gymnospermoides, X. ramu- losum, Thurovia triflora, and Amphiachyris dracunculoides. Carlquist (1960) had previously described the conspicuousness of these resinous deposits in X. microcephalum (as Gutierrezia microcephala). In many instances nu- merous vessels are entirely filled with resinous contents. Carlquist (1960) also described the presence of prismatic crystals in the ray cells of X. microcephalum (Fig. 6). In my investigation, crystals were also observed in X. sarothrae (Fig. 7) and X. centauroides (Fig. 8). These crystals are much like those figured by Carlquist for X. microcephalum. Conspicuous crystals were not observed by me in any of the other species. DISCUSSION AND CONCLUSIONS With the wood data, it seems easier to show levels of specialization than to indicate relationships. However, there are some clear distinctions worthy of mention The genus Amphiachyris is distinctive because of its large vessels, the longer and wider libriform fibers, and the presence of fiber-dimorphism. Even when average vessel-element length of annuals is compared, Amphi- achyris maintains this distinctiveness. In contrast, Thurovia, in addition to its peculiar distribution of vessels, shows the shortest average vessel element length among the annuals (Tables 3, 8). The large vessel-element length among annual species of Xanthocephalum is due solely to the large xylem elements present in X. gymnospermoides. The fact that relatively long and wide vessel elements occur in X. gymnospermoides is very inter- esting. Perhaps these large elements of the xylem, the larger leaves, little branched inflorescence, and the large chromosome number (n=6) qualify this species as the most primitive member in the genus. However, when an- nuals and perennials are combined, even though some vessel elements and libriform fibers in Xanthocephalum are wider and longer than those in Gymnosperma, Amphipappus, and Thurovia, there are little differences in the mean for these characters. Helical sculpturing in vessels and vascular tracheid of the secondary xylem offers ancther distinctive feature. No prominent helical sculpturing occurs in spite eine The coarse band wall relief in Amphipappus is sufficiently distinct from the continuous groove pattern noted in Gymnosperma, Thuro- via, and certain species of Xanthocephalum. Somewhat high rays characterize all of the genera. Wide heterocellular rays are also characteristic. The fact that Thurovia lacks wood rays cer- tainly sets it apart from the other genera studied. This treatment of the genera is by no means conclusive. Additional bio- systematic studies of the taxa are needed to more accurately reflect their evolutionary history. However, the distinctiveness described above when coupled with the cytological, anatomical, and morphological evidence re- ported earlier (Ruffin, 1971) provides one more line of evidence supporting the distinctiveness of Gymnosperma, Amphiachyris, Amphipappus, and Thu- rovia and unifying reasons for merging Xanthocephalum, Gutierrezia, and Greenella. REFERENCES ANDERSON, L. C. 1963. Studies on ria (Compositae). Anatomy, cytology, taxon- omy. Trans, Kansas Acad. Sci. 66:632— BAILEY, I. W. 1953 ie anatomical a ek to the study of genera. Chron. Bot. M4s121-125 . 1957. The potentialities and limitations of wood anatomy in the study of phylogeny nd class is of an ae J. Arnold Arbor. 38:234—254. BARGHOORN,. E. S. 19 The oe development and phylogenetic specialization of ue in the xylem of Hioryledom Modific ation of the multiseriate and uniseriate rays J. Bot. 28:273- BE NTHAM, ‘ and J. D. ose 1873. Genera plantarum 2:553. London. CARLQUIS’ 1960. Wood anatomy of Astereae (Compositae). Trop. ee as 113:54-84, York. ee, oo ee plant anatomy. Holt, Rinehart, and ae on sO ———. 1962a. Wood anatomy of Susaaie es Sp ea Allis er : ee led . 1962b. A theory of 4 phosis in woods. Phytomorphology 12:30—45. ———. 1966. Wood anatomy of Compositae: A summary with comments on factors con- trolling wood evolu Aliso 6:25—44. CHALK, L. 1937. The Phylogenet value of certain anatomical features of dicotyledonous woods. Ann. Bot. 1:409—428, COMMITTEE ON NOME nc LATURE. 1957. sini nes iae Association of yee ei ists. International gloss ae of terms used in wood anatomy. Trop. Woods 107 CORRELL, D. S. and M. JOHNSTON. ae Manual of the vascular ce nae e. “Texas Texas feceare Ses Renner, Texas. GRAY, A. 1873. Notes on oe and characters of certain genera and species. Proc. Amer. Acad. Arts 8:632-63 NELSON, A. 1934. Rocky ae herbarium. studies, I]. Amer. J. Bot. 21:573—582. PORTER, C. L. 1943. The genus Aw phipappus Torr. and Gray. Amer. J. Bot. 30:481—483. ROSE, J. N. 1895. Thurovia, 1 genus of Compositae. Contr. U.S. Natl. Herb. 3:320—322. RUFFIN, J. 1971. serena and anato of genera Amphipappus RET Greenella, Gutierrezia, Gymnos perma, Se nrc and Thurovia (Compositae). The sis, Kansas State University, Moanhauere: Kansas. SASS, J. E. 1958. Botanical microtechnique. 3rd ed. Iowa State University Press, Ames. SHINNERS, L. 1950. Notes on Texas Compositae IV. Field & Lab. 18:25-32. SOLBRIG, O. ‘TV. 1960, The status of the genera Amphipappus, i toes byris, Greenella, Gutierresta, Gymnosperma and Xanthocephalum (Compositac). Rhodora 62:43—54. NOTES BAUHINIA ERYTHROCALYX (LEGUMINOSAE), A NEW SPECIES FROM MEXICO.—Two unusual collections of Bauhinia from the Yucatan Peninsula were examined during the course of a revision of the Mexican species which have proven to be of an undescribed species. BAUHINIA erythrocalyx Wunderlin, sp. nov. Bauhinia jenningsti P. Wilson affinis a differt foliis bilobatis, apice loborum obtusis, basi cordatis, 7-ner- vatis. Shrubs; branches strigose when young, soon glabrescent. Leaves ovate, bilobate for % or more their length, lobes divergent, 3.5-7.0 cm long, 4-6 cm wide, apex of lobes obtuse, base cordate, margins slightly crisped, char- taceous, glabrous above, strigose below, 7-nerved; petiole 1.5-3.0 cm long, slightly canaliculate, strigose: stipules lanceolate, ca. 1.5 mm long, cadu- cous; adpetiolar pair of intrastipular excrescences subulate, up to 1.5 mm long, others minute. Inflorescences racemose, terminal or subterminal and axillary, 10-to 20-flowered, rachis strigose, buds linear, 1.5-2.0 cm long, strigose, red, free tips minute; bract ovate-lanceolate, ca. 1 mm long; brac- teoles similar to bract although slightly smaller, subbasal; pedicels 1-2 long, strigose, reddish; hypanthium cyathiform, 2-3 mm long; calyx spa- thaceous at anthesis; petals 5, subequal, white or cream, ca. 1.5 cm long, blade linear-lanceolate to oblanceolate, ca. 1/3 the length of claw, 2-3 mm wide, sparsely pilose externally, claw glabrate; fertile stamen 1, slightly exceeding petals, filament slender, slightly arcuate, glabrous, connate with staminodes at base, anther oblong, apiculate at apex, emarginate, 3-4 mm long, red, glabrate; staminodes 9, alternate slightly shorter, 1.0-1.5 cm long, connate for 144 their length, pilose within at connate portion, abortive an- thers present or absent; gynoecium + equalling fertile stamen, slightly ar- cuate, ovary densely strigose, gynophore and style sparsely and loosely strigose, gynophore + equalling style, stigma terminal, slightly differenti- ated from style. Fruit an elastically dehiscent legume, linear, apiculate with persistent style, 7-10 cm long, ca. 1.5 cm wide, light brown, strigose to glabrate, gynophore ca. 2 cm long, glabrate; seeds: oblong, 5-8 mm long, 4-6 mm wide, dark brown, puncticulate, obscurely striate, funicular branch sears subequal, ca. 1.5 mm long. HOLOTYPE (F): MEXICO: Campeche: ‘‘Central Buenfil’ (? Puerto Buen fiel), 1 Jan. 1932, C. C. Lundell 1152. Isotypes at F, MIC ADDITIONAL ‘MATERIAL EXAMINED: MEXICO: aeons ‘“Colectado en el sur del estade en Noviembre de 1945,’ N. S. Novelo 404 (US). Mixed collection with Bauhinia ungulata L. Bauhinia erythrocalyx is a rare species endemic to the Yucatan Peninsula and known only from the above two collections. This species is most closely related to Bauhinia jenningsii from Cuba, the Yucatan Peninsula of Mexico, SIDA 5(5): 353. 1974. 304 Belize, and northern Guatemala, but differs from that species in having its leaves bilobate, lobe apices obtuse, base cordate, and seven nerves. Bauhinia erythrocalyx is so named because of its scarlet calyx.—Richard P. Wunderlin, Dept. of Biology, University of South Florida, Tampa, Florida 33620 HYDRILLA VERTICILLATA (HYDROCHARITACEAE): NEW TO LOU- ISIANA.—The submerged hydrophyte Hydrilla verticillata (L.f.) Caspary was first collected in Louisiana on 30 July 1973 by Robert Barry, Jr., Robert Barry III, and Hugh Hitt, a team surveying various bodies of water in the southern part of the state. Hydrilla is easily distinguishable from Egeria and Elodea by its sharply serrated (not serrulated) leaves and by the pres- ence of vegetative buds in the leaf axils and on the roots. Vouchers are deposited in the U.S.L, Ornamental Horticulture Herbarium and the U.S.L. Biology Herbarium (LAF). Duplicates have been sent to the Chief of Aquatic Plant Control, Washington, D.C. Collection data are: shallow water, Spanish Lake, Iberia Parish, 30 Jul 1973, Robert Barry, Jr. et al. S0730731.—Sigmond L. Solymosy, University of Southwestern Louisiana, sence 70501. TRANSCRIPTION ERROR—In Sida vol. 5 no. 4 page 205 and correspond- ingly on the upper map on page 204, Styrax americana a is attributed to “Pennsylvania, Lancaster Co.—Lancaster, Small, Jul. 1889 (F).’’ The spe- cies is however not included in the Flora of Lancaster County by Small and Carter, 1908, nor among the plants of that county in any other herbarium. Apparently the writer of the label on the F sheet copied the address of the collector, not the collection point—Edgar T. Wherry, University of Pennsyl- vania, Philadelphia, Pa. 19104. EULA WHITEHOUSE (1892-1974) Dr. Eula Whitehouse, botanist, artist, naturalist, and educator, died at Dallas on 6 September 1974 at the age of 82. Dr. Whitehouse was retired Curator of Cryptogams of the Southern Methcdist Herbarium. Prior to com- ing to SMU she had served on the faculties of Houston Municipal Hospital and the University of Texas College of Mines and was for several years curator of the Texas Memorial Museum in Austin. Dr. Whitehouse’s botanical contributions include several studies in Texas bryophytes and revisions in Salvia and Phlox. She also authored and illus- trated Texas Flowers in Natural Colors (1936), the first color-illustrated guide to Texas wildflowers.—Jerry M. Flook, SMU Herbarium. SIDA 5(5): 354. 1974. INDEX TO VOLUME 5 Names of contributing authors are in capital letters. New scientific names are in boldface. Page numbers in boldface indicate illustrations. Aconitum columbianum 292, 293 Adelobotrys macrophylla 38 Adnaria odorata 193, 203 Amarolea americana 281; megacarpa 281 Amauria rotundifolia 127 Ammannia coccinea 45 , 304, 306, 309, 314, 326, 330, 343, 345, 350, 351 Amphiachyris, Amphipappus, Green- ella, Gutierrezia, Gymnosperma, Thurovia, and Xanthocephalum (Compositae), A taxonomic re-eval- uation of. 301-333 Amphicarpaea africana 338, 339, 340; bracteata 338, 339, 340; comosa 338; edgeworthii 338, 339, 340 Amphicarpaea, Pollen morphology of. 338-340 Amphipappus fremontii 304, 306, 309, 314, 324, 330, 3438, 347, 350 Angiosperms, sexual terminology 59- D Anliot, Sture I*. 296 Apis mellifera 198 Aquilegia laramiensis 292, 293 Arachis hypogaea Artemisia absinthium 50, 51; ludo- viciana var. gnaphalodes 273, var. ludoviciana 273 Atriplex reptans 25 AVERETT, JOHN E. 3, 48 Ayuque, Balmea stormae, an endan- gered Mexican species. 268-270 Bacopa repens 45 Balmea stormae 269 Baptisia lecontei 30 Bauhinia erythrocalyx 353; jenningsii Betula alleghaniensis 277; lutea 277 BIERNER, MARK W. 45 T., JR. 271 Cacalia atriplicifolia 50, 51, floridana 30; plantaginea 286 Caesalpinia chicamana 43; drum- mondii 44; egena 44; glauca 43; peninsularis 43; pumilio 44; terna- ta 44: viscosa 44 Calyptocarpus vialis 52 Caperonia palustris 286 Carex bicknellii var. opaea 49; toniana 49; merritt-fernaldii 49 Caulinia guadalupensis 261 Centaurea americana 286 Chaenomeles lagenaria 271; speciosa brit- Chenopodium album var. stevensii 291; ambrosioides 290, 291; californicum 290, 291; carnosulum var. patagonicum 290, 291; gigan- teum 290, 291; gigantospermum 290, 291; glaucum var. glaucum 290, 291; hybridum 290, 291; macro- calycium 290, 291; polyspermum var. obtusifolium 290, 291; pumilio 290, 291; rubrum var. rubrum 290, 291 Chaenorrhinum minus 2 Chionanthus henryae 280, 281, 283; pygmaeus 280, 281, 283; virginicus 281, Chloris truncata 182 356 Chromosome numbers 3-7, 50-52, 290- 294 Clematis ligusticifolia 292 Clidemia ulei 38 Coldenia hispidissima 133 Cooke, John F., Jr. 297-300 Corallorhiza odontorhiza 130, 287 Coreopsis tinctoria 50, 51 CRAWFORD, DANIEL J. 292 Cuthbertia graminea 26, 28, f. ¢ minea 30, f. leucantha 28, 29, 30, 31; ornata 27, 30; rosea 26, 27 Cuthbertia, Field gages with description of new form. Cydonia japonica 271 Cyperus rotundatus 271 Danaus plexippus 198 Darlingtonia californica 193, 336; diviva 336, 337 Delphinium barbeyi 292, 293; geyeri 292, 293; nelsonii 292; occidentale 293; ramosum 293 Dirca palustris 132 Dopatrium junceum 45 DUNCAN, WILBUR H. 42 Duncan, Wilbur H. 190 UQUESNAY, DAMIAN 286 tenuiloba re- Dyssodia_ tenuiloba_ var. 130 Echinacea pallida 286 EIFERT, IMRE J. 43 Eleocharis atropurpurea 45; elongata 130; obtusa 45 Engelmannia pinnatifida 286 Eragrostis plana 182 Eriochloa contracta 48; gracilis 42 Eriogonum jamesii var. undulata 265-267; turneri 265-267; umbella- tum 2¢ Eriogonum from North Mexico, A new subfruticose. 265-267 Erodium cicutarium 272 Eryngium lemmoni 135 Euonymous fortunei 272; kiautschovi- ca 272 Eupatorium perfoliatum 50, 51 Euphorbia glomerifera 130; hyper- icifolia 130 Eurema nicippi 198 Eutetras palmeri 65 Ewan, Joseph 180-181 Fagus grandifolia 132 Fatoua villosa 286 FLOOK, JERRY M. 137, 296, 354 FLYR, DAVID 130 Flyr, Lowell David 54-58 inata 280; globularis 280; ligustrina 280; pinetorum 280; porulosa 280; pubescens 280; segre- gata var. pinetorum 280, 283, var. segregata 280 Formicarial Melastomataceae, Ob- servations on some upper Amazo- nian. 33-41 Forsythia suspensa 279; viridissima 279 FOSBERG, F. R. 268 Foveolaria ferruginea 192-193 Frankenia gypsophila 25; johnstonii 133; leverichii 132, 184; margaritae 266 Fraxinus americana 275-277, var. biltmoreana 276, var. microcarpa 276; biltmoreana 276; caroliniana 275, 276, var. cubensis 276, var. ob- lanceolata 276; catawbiensis 276; arlingtonii 277; michauxii 277; nigra 275, 276; pauciflora 276; pennsylvanica 275-277, var. subin- tegerrima 277; profunda 275-277; quadrangulata 275; smallii 277; to- mentosa 27 FRYXELL, PAUL A. 1 Galium ea 42 GARDNER, ROBERT C., 292 Geranium texanum 130 GONSOULIN, GE J 1 Gopherus polyphemus 284 Gossypium hirsutum 1; sandvicense 1, 2; taitense 1; tomentosum 1, 2 Gossypium, A reconsideration of the correct name for the Hawaiian. 1-2 Greenella arizonica 304, 306, 308, 309, 314, 319, 330, 331; ramulosa 304, 313, 332 Gutierrezia californica 304, 306, 307, 314; mandonit ssp. mandonii 4, 314; ocephala 302, 305, 307-310, 312, ae 323, 347, 349, 351; sarothrae 3 ; 314, Ce opens. nee 305, 307- 310, 314, 325, 330, 333, 343, 345, 350 Halesia parviflora 203 Haplopappus johnstonii 133 HARDIN, JAMES W. 274 Hedeoma bicolor 16; glabrescens 18; greggii 16; incanum 8, 20; longi- florum 16; molle 10, 21; montanum ix 272 Heleniastrum amphibolum 47; oocli- niu m 47 Helenium amphibolum 47; apterum 7; chihuahuensis 45, 46; elegans var. eauibeiia 47; microcepha- lum var. ooclinium 47; ooclinium 47; scorzoneraefolium 47; stenop- terum 47 Helianthus petiolaris 186 HERMANN DERICK J. 49 Hesperozygis marifolia 9, 21 Heteranthera limosa 45 Heuchera sanguinea 135 Hoffmanseggia falcaria 43; glauca 4 Hydrilla verticillata 354 Hypochoeris brasiliensis 287, 289; mi- crocephala var. albiflora 287, 288, 289; radicata 287, 289 307 Ilex, International Checklist of Cul- tivated 136 International Botanical Congress XII IRVING, ROBERT S. 8 Isocoma gypsophila 24, 25, 266; halo- phytica 23; pluriflora 23; veneta 24, 25 Isocoma, Two new species from North Central Mexico. 23-25 Isoetes engelmannii var. caroliniana 129: leuisianensis 129; melanopoda Jasminum amplexicaule 279; azori- 278, 279; undulatum JONES, S. B., Judd, John Bagless 295 KEENER, CARL S. 290 Kentucky plants, Herbarium and field studies of. 271- LAKELA, O L megane GARRIE 128 lata 15: ari- congesta 100; 103: fastigiata 95; gilensis 103; gra- cilis 95; grandifolia 124; halimifo- lia 115; intricata 94; inyoensis 96; laciniata 116; lemmonii 106, var. lindheimeri 113; me- . intricata 94; palmeri 99, 100, peninsularis 127; rotund saxicola 102; scopulorum 127; stansburii 85; staurophylla 117; tenella 99; toumeyi 100; villosa 98 Lapsana communis 271, 273 Larrea glauca 43; peninsularis 43 LAWSON, CHERYL A 308 Leptochloa uninervia 42 Leptopharynx dissecta 110; gilensis 103; grandifolia 124; lemmonii 107 Leucophyllum flyrii 56, 57 Liatris squarrosa 186 Ligustrum amurense 282; japonicum 282; obtusifolium 282: ovalifolium 282; quihoui 282; sinense 282; vul- gare 282 Liriodendron tulipifera 132 Ludwigia decurrens 45 Lygodesmia juncea var. rostrata 183, 186 MAHLER, WILLIAM F. 180 Mahler, William I*. 53, 190 Maieta guianensis var. guianensis 33, 35, 38-40, var. leticiana 33, 34, 35, ; neblinensis 38; poeppigii 33, 35, 38-40 MECOM, JOHN O. 295 MEESON, BLANCHE W. 338 Melastomataceae, Observations on some upper Amazonian formicari- al. 33-41 Monarda fistulosa 286 Mcnothrix congesta 100: 95; intricata 94; rostrata 186; fastigiata megacephala 89: palmeri 99; stansburii 85; stans- buryana 85; toumeyi 100 Morisonia imrayi 335 MORTON, GARY H. 50, 131 Najas ancistrocarpa 259, 260; con- ferta 259; guadalupensis 259, 261, var. floridana 262, 263, 264; 261; wrightiana 259-261, 262 Najas, Notes on. 259-264 Oleaceae of the So: aaa United States flora. 274-2 Orontium pace lates a Osmanthus americanus var. canus 281, var. 283; floridana 281 Ossaea bullifera 33, 35, Ottelia alismoides 260 Pericome caudata 65, 75 minor ameri- megacarpus 281, 38, 39, 40 Perityle aglossa 122; angustifolia 64, 66, 68, 72-75, 80, 108, 115, 117, 119, 120, 122; bisetosa 64-68, 72-75, 77, 79, 84, 111, var. appressa 82-85, var. bisetosa 82, 83, var. sealaris 82-84, var. spathulata 82, 83, 85, 86; castillonii 64, 67, 73-75, 77, 80, 100, 108, 109, 111-113, 122; cernua 65; ciliata 70; coahuilensis 62, 63, 65-67, 73-75, 81, 108, 119, 120, 121, 12 ; ¢ i 23 congesta 62, 64, 73, 74, 76, 81, 88, 93, 96, 97, 100-102: cordifolia 69, 77, 125; coronopifolia , 124, 127; disseeta 64, 68; 126; gilensis 66, 67, 69, 103, 119, var. gilensis 82, var. Salensis 62, 82, 103, 106; gracilis 62, 65, 73, 76, 81, 88, 95, 96, 102, 119; grandifolia ee 63, 66, 67, 73, 74, 79, 82, 124, 125 mifolia 115; inyoensis 61, 62, 64, 66, 73, 74, 76, 80, 88, 90, 91, 93, j jaliscana 65; laciniata 117; lemmonii 64, 65, 67, 73-75, 80, 106, 108-111, 118, 123-125, 135; lep- toglossa 69, 125; lindheimeri 64, 66-68, 72-75, 79, 113, 115, 117, 119, var. halimifolia 108, 113, 115, var. lindheimeri 108, 113, 115; megalo- cephala 61, 66, 68, 69, 73, 74, 89, 97, 125, var. intricata 62, 66, 76, 81, 88, 89, 93-95, var. megaloce- phala 81, 88-91, 93, 94, var. oligo- phylla 62, 66, 76, 81, 88, 89, 91, 92, 93, 102; palmeri 69, 97, ‘ 125; cara 64, 65, 69, 112; queflora 64, 119; 80, 82, 102, 103, 106, 119; stansburii 61, 62, 67, 68, 73-76, 80, 84, 88-90, 95, 96, 125; staurophylla 61, 65-67, ; tenella 125; vitreomontana 73, 77, 84, 111, 118; warnockii 64, 67, 68, 72-75, 77, Perityle Section Laphamia, Taxon- omy of. 61-128 Phoebanthus grandiflora 30 Physalis crassifolia var. crassifolia 3, var. infundibularis 3; glabra 3; glutinosa var. glutinosa 3; hederae- folia var. cordifolia 3, var. hede- raefolia 3; lobata 4, 6, 7; sordida 4; vestita 5; viscosa var. cine- rascens 5; wrightii 5 Physalis and Solanum, Chromosome numbers in. 3-7 Pinus palustris 30 Poliomintha bicolor 8, 16; conjunc- rix 8, 13-16, 19; glabrescens 8, 12, 13, 16, 18; greggii 16; incana 8, 13-16, 19-21; longiflora 8, 9, 11, 13, 17, var. congesta 12, 16, 18, var. longiflora 12, 16; marifolia 8, 21; mollis 21 Poliomintha, A revision of the genus. 8-2 Pollen morphology of Amphicarpaea 338-340 Polygala rimulicola 135 Polytaenia nuttallii 286 Potamogeton pusillus 129 POWELL, A. MICHAEL 3, 61 Quercus laevis 30 Quincula lobata 6 Ranunculus acriformis 293; alismae- folius var. montanus Reeves, Robe REVEAL, aad L. 265 Rotala indica 45; ramosior 45 ROWELL, CHESTER M. 295 RUFFIN, JOHN 301, 341 309 Russell, Norman H. 190 Sartwellia mexicana 25 Senecio salignus 135 Setaria faberii 42 sexual terminology sperms 59-60 SHI ene. LLOYD H. 182 Shinners, Lloyd H. 53, 137-181, 190 Shinnersoseris rostrata 183, 184, 185, 186, 187, Sida lindheimeri 182 Silene plankii 135 Silphium laciniatum 286 Solanum, Chromosome numbers in Physalis and. 3-7 Solanum amazonicum 5; in the angio- elaeagni- folium 5; hindsianum 5; madrense Do; nigrum 5; rostratum 5; triflo- rum 6 Solidago bicolor 5, 51; brachyphylla 50, 51; gattingeri 50, 51; Southeastern United flora Studies o V. Oleaceae. 274-285 Sparganium americanu 2 Sphenoclea — 45 Stone, Witmer 295 Strigilia Rr 223; racemosa 192, 193 STROTHER, JOHN L. 52 Strotheria gypsophila 266 STUCKEY, RONALD L. 297 STUESSY, TOD F. 52 Styrax agrestis 193; americana 193, 198, 203, 210, 354, r. americana 192, 196, 197, 199, bane 203, 205, 206, ; , 237, , var. pulverulenta 192, 195, 196, 199, 200, 203, 205, 206, 208, 236, 238, 254; americanum 334, f. pulverulentum 206; americanus f. genuinus 203; argenteus 192, 194, 196, 197, 201, 223, 225, 230, 233, var. argenteus 196, 223, 225, 229, ar. hintonii 195, 196, californicum cavalerici 208; contermi- 229, 230, 248, 256; 227; foveolaria 195, 197, 202, 230, 233, *. micranthus 233; oo pecen: 199, 202, 220, 222, var. glabrescens 220, 222, 244, 255, 220, 222, 244, 256; 336, 337; nus 195, 197, 202, cyathocalyx 225, 192, 193; ever var. pilosus 196, glabrum 203, 334-336; grandifolia 192, 194-196, 198-200, 202, 208, 210, 222, 236, 239, 254; guatemalensis 222: guianensis 234; hintonii saeGene 195-197, 202, 219, macranthus 217; magnus 191, 226; micranthus 227; myristicifolius 223; obassia 199, 200, 217; obtusifolia 195, 197, 201, 230-232, 249, 258: oc- cidentalis 233, 335; ochraceus 195, 197, 201, 230-232, 250, 258; drum 203; odoratissimus 217; cinale var. rediviva 337; officinalis 192, 201, 202, 208, 210, 215, 217, 220, var. californica 197, 199, 200, 216- 219, 242, 255, 337, var. fulvescens 195, 199, 200, 216, 218-220, 242, 255, jaliscana 219; orizabensis 227; pilosus 222; platanifolia 196, 202, ar. platanifolia 196, 210-213, stellata 195, 196, 210, var, 195, 197, 198, 201, 230, 232, 251, 258, 335; pulverulentum 206; punctatum 223; racemosa 192, 193; ramirezii 225, 227; rediviva 336, 337; texana 195, 202, 212-215, 241, 255; toucha- nensis 208; vulcanicola 191, 226; warscewiczii 227; youngae 195, Styrax, Further comments on. 334-337 Styrax in North America, Central America, and the Caribbean, A re- vision of. 191-258 Synedrella nodiflora 52 Syringa vulgaris 279 TAKHTAJAN, A. 136 Taub, Stephen 295 TERRELL, EDWARD E. 48 Thalictrum sparsiflorum 293, 294 THIERET, JOHN W. 45, 59, 129, THOMAS, R. DALE 132 Thrips tritiea 198 Pnurovis triflora a0 307, 308, 310, 314, 329, 330, 286 Gc 40; , 40; juruensis 33, nee 40; occidentalis 40; ee eens 33, 35, 36, 40; ulei 33, 35, 36, 38-40 TODSEN, THOMAS K. 135 TOMB, A. SPENCER 183, Tradescantia rosea 26, minea 26, 28, var. r, rosea 26 Tremanthes ferrugineum 193 Trifohum vesiculosum 286 Tripogandra rosea a0; 27 287 27, var. era- ornata 26, 27, ~) 3. cn 132 Vernonia er 44; WENTZ, W. ALAN 259 WHERRY, EDGAR T. 354 WHIFFIN, TREVOR 33 Whitehouse, a 3 olffia papulifera 271 Wood anatomy of Amphiachyris, Am- phipappus, Thurovia, Gymnosper- ma, and the Xanthocephalum com- plex 341-352 WUNDERLIN, RICHARD P. 353 Xanthocephalum centauroides crinita 44 305, 307, 308, 310-312, 330, 342, 344, 349, 349: rendpnii 344; microcephalum 342, 344, 246, 361 351; ramulosum a 342, 351; saro- 2, 344, 348, 350: wrightii 301, 303, 306, 307, 310, 311, oa 320, 330, 331, 332, 344, 347, 348, 2 sh pe eee li beelt—C tC (atst””:C he lO tC