Taxonomy of the Genus Chloris (Gramineae)

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Brigham Young University Science Bulletin, Biological Series

Volume 19 Number 2 Article 1

3-1974

Taxonomy of the genus Chloris (Gramineae)

Dennis E. Anderson Department of Biology, Humboldt State College, Arcata, California 95521

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MAY271P74 HARVARD UNIVERSITAXONOMY OF THE GENUS CHLORIS (GRAMINEAE)

Dennis E. Anderson

BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2 MARCH 1974 /ISSN 0068-1024 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES

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TAXONOMY OF THE GENUS CHLORIS (GRAMINEAE)

Dennis E. Anderson

BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2 MARCH 1974 / ISSN 0068-1024

TABLE OF CONTENTS

ABSTRACT 1

ACKNOWLEDGMENTS 1

INTRODUCTION 1 NOMENCLATURAL HISTORY OF THE GENUS 2 TAXONOMIC RELATIONSHIPS OF CHLORIS 2 Cytology 3 Leaf Epidermis 5 Anatomy, Embryology, and Seedling Morphology 6 Relationships of Chloris and Other Genera 9 Relationships within Chloris 13

Chloris Swartz 15 Key to the species of Chloris 16

1. Chloris roxlmrghiana Schultes 22

2. Chloris puniculata Scribner in Robinson 24

3. Chloris herroi Arechavaleta 25

4. Chloris ciliata Swartz 27

5. Chloris lamproparia Stapf in Chevalier 30 6a. Chloris canterai Arechavaleta var. canterai 30

6b. Chloris canterai Arechavaleta var. grandiflora ( Rosengurtt & Izaguirre de Artucio) Anderson 32

7. Chloris dantUjiina C. D. Adams 34

8. Chloris cruciata (Linnaeus) Swartz 37

9. Chloris filiformis (Vahl) Poiret in Lamarck 39 10. Chloris suringari Hitchcock in Urban 39 11. Chloris scariosa F. von Mueller 40 12. Chloris prieurii Kunth 42

13. Chloris rohusta Stapf in Chevalier 43

14. Chloris mollis (Nees) Swallen 45 15. Chloris longiaristata Napper 46 16. Chloris quinque.sctica Bhide 48 17. Chloris wightiana Nees ex Steudel 48

18. Chloris montana Roxburgh 51 19. Chloris bournci Rangachariar & Tadulingam 53

20. Chloris inflata Link 53 21. Chloris formosana (Honda) Keng 57 22. Chloris pilosa Schumacher 58 23. Chloris virgata Swartz 60 24. Chloris gai/aiu/ Kiuith 65 25. Chloris castilloniatui Lillo & Parodi 69 26. Chloris orthonoton Doell in Martins 69 27. Chloris aristata (Cervantes) Swallen 70 28. Chloris hrandegei (Vasey) Swallen 72

29. Chloris chloridca ( Presl) Hitchcock 74 30. Chloris ventricosa R. Brown 76 31. Chloris truncate R. Brown 78 32. Chloris pumilio R. Brown 80 33. Chloris lobata Lazarides 82 34. Chloris divaricata R. Brown 83 35. Chloris pcctinata Bentham 86 36. Chloris crinita Lagasca 87 37. Chloris pluriflora (Foumier) Cla)'ton 89 38. Chloris cticullata Bischoff 91 39. Chloris verticillata Nuttall 93 40. Chloris andropogonoidcs Foumier 95 41. Chloris texensis Nash 103 42. Chloris sesqiiiflora Burkart 104 43. Chloris humhcrtiana A. Camus 104 44. Chloris amethi/stca Hoclistt-tter 106 45. Chloris subniutica Humboldt, Bonplantl. Kunth 106 46. Chloris somalewis Rendle 109 47. Chloris cleimntis Merrill 109 48. Chloris radiata (Linnaeus) Swartz 110 49. Chloris pycnotlirix Trinius 114 50. Chloris halophila Parodi 1 17

.51. Chloris ekmanii Hitchcock 119 52. Chloris arcmiria Hitchcock & Ekman in Hitchcock 119 53. Chloris sagnwuna Achille Richard in .Sagra 121 54. Chloris cubensis Hitchcock & Ekman in Hitchcock 123 55. Chloris mossambicensis Schumann 123 56. Chloris hurrnetuis Ander.son 125 UNIDENTIFIED, REJECTED, OR REASSIGNED NAMES 126 LITERATURE CITED 131 TAXONOMY OF THE GENUS CHLORIS (GRAMINEAE)

Dennis E. Anderson'

ABSTRACT

This study is a worldwide taxononiic treat- of unidentified or rejected names. Distributions ment utilizing endomorpliic characters such as and maps are given for each species, along with cytology, histology, and ciiihrNology, in addition lists of specimens examined dining studies at to the traditional cxoniorphic features. New various iu-rbaria. Reports of field studies in- chromosome numbers and histological informa- volving introgression among certain species are tion are reported. Relationships between Chloris also included. A new species, Chloris hurmen- and other genera in the Chlorideae are dis- sis, is described from Bumia, and a new com- cussed. A total of r6 species is recognized in the bination, Chloris canterai Arech. var. graiuliflora genus; each is described and illustrated. Com- ( Rosengurtt & Izaguirre de Artucio ) Anderson, plete synonymies are included as well as a list is also proposed.

ACKNOWLEDGMENTS

These studies were begun in the summers A special debt is owed to Dr. W. D. Clayton of 1963 and 1964 as part of a National Science of the Royal Botanic Gardens, Kew, England, Foundation program in l\esearch Participation who provided advice, cooperation and friend- for College Teachers sponsored b\' the Depart- ship, and thus made my stay at Kew most pro- ment of Botan\-, University of Texas. Additional ductive. financial support was provided b\' this program California State University, Humboldt, has through grants ( NSF GY-3002, NSF GE-69:8). generously provided released time and leaves in The assistance of Dr. Harold Bold, director of support of these studies, and the continued as- the summer program, is also gratefully acknowl- sistance of President Cornelius Siemens is ac- edged. Initialh', Dr. Walter V. Bro\\'n pointed knowledged with many thanks. out to me many of the biosystematie problems I would also like to thank Mr. Rupert C. of Chloris in Texas, and I am grateful to him Barneb\- for his aid in the preparation of the for his continued encouragement. Latin description of Chloris btirmsiisis. .additional direct assistance was received Michael Lazarides kindly let me examine his from the National Science Foundation in 1969 manuscript treatment of Chloris in Queensland, (NSF GB-7235), enabling me to visit a number prior to its publication. of major overseas herbaria and to do additional I am grateful to my wife, Nancy, for the end- field work. The curators were most helpful and less hours she contributed in filing and organiza- allowed me to borrow man\- specimens. Their tion, in establishing and verifying geographical cooperation is much appreciated. locations, and, finally, in cartography.

INTRODUCTION

With its delicate, digitate clusters of spikes, of Nebraska, from the Cape of Good Hope to Chloris must certainly be among the most strik- the sands of the Sahara, from Afghanistan to ing of grass genera. The distribution spans the island archipelagoes of the South Pacific, the warmer regions of most of the continents, and from the subcontinental islands of Ne.v from the pampas of Argentina to the sandhills Guinea and Borneo through the continent of

^Department of Biology, Humboldt State College, Areata, California 95521. Bricham Young University' Science Bulletin

Australia. Thus, it is likely that taxonomists in me to conclude that a classical studv was needed these areas have seen (me or more of the taxa and that experimental studies could then be in the genus. Though these attributes would made on a sounder foundation. seem to insiue popularity among agrostologists, Well over 15,000 herbarium specimens and Chloris nevertheless has not been subjected to many mass collections from hybridizing popula- any systematic review for nearly a centuiy. This tions were studied. The numbers of specimens has been an era characterized by extensive ex- available van' widely. Some taxa are repre- ploration and collection of hitherto unknown sented only by the holot\'pe. Others are known areas, as well as the development of revolu- from such large numbers of specimens that tionary systematic thought. population descriptions may be advanced with My interest in Chloris began with studies in confidence. Still others ha\'e been collected so hybridizing species in Texas; this interest spread thoroughly that the enormous complexit)' of the later to similar studies in the Southeast; and ulti- population has revealed the necessity for further mately it encompassed more classical investiga- intensive investigation. tions in manv of the major herbaria of the This study, then, attempts to bring the tax- United States, Great Britain, and Europe. Ini- onomv of the genus CliJoris up to date. It is tially, my interests centered in experimental largely morpho-geographical in nature, with oc- studies but the continuing frustration of not casional incursions into biosystematics as the being able to name species with confidence led nature and availability of materials allow.

NOMENCLATURAL HISTORY OF THE GENUS

Swartz first described Chloris in 1788, in- hitmhertiana (1950). Subsequently, Camus cluding two new species, C. ciliata and C. pc- (1957) elevated the latter subgenus to the ge- traea, and transferred into the genus three Lin- neric level. naean species: Agrostis criiciafa, Agrostis radiata, Generic conspeeti of Chloris have been writ- and Andropogon poh/dactylon. ten spor;idicallv, Iiut all are of the nature of With the description of the genus Eitstachijs catalogs rather than re\isions or monographs. by Desvaux (1810) and the transfer to it of Of these, the treatment by Steudel ( 1854) is the Chloris petraea Swartz, the basic nomenclatural most complete. In it he recognized 69 taxa and pattern at the generic level was established, included a s\noptic key as well as an enumera- though many authors now consider Eustachijs tion of species, each with a short description. as a subgenus of Chloris. Later listings by Regel ( 1862, 1863), based upon Other genera, based on various species of the specimens at Leningrad, included a total of Chloris, have been proposed: Rabdochloa Beau- only 21 taxa. vois (1812), including C. cruciata and several Regional revisionary studies restricted to species from other genera; Schtdtesia Sprengel Chloris are few, the most notable being those (1815) containing C. petraea only; and Phacel- of the North American species by Nash (1898), laria Willdenow ex Steudel ( 1840), including C. the treatment by Swallen (1939) in l^orth suhmutica. None of these genera have been ac- American Flora, and the publications bv Everist cepted by recent authors. ( 1935a,b, 19.37, 1938) on the Queensland species.

More recently, Camus proposed the subgen- SubsequentK , Lazarides (1972) treated Chloris era Monanthochloris, including within it C. per- in his stud\' of the tribe Chlorideae in Austra- rieri (1949), and Pterochloris, which contains C. lia.

TAXONOMIC RELATIONSHIPS OF CHLORIS

Most recent classifications of the Gramineae b\- the redefinition of the subfamily Festucoi- are based heavily upon cytological and ana- deae, which has had many of its genera distrib- tomical characteristics; in contrast, the older sys- uted to other subfamilies. tems were constructed primarily on the basis of All recent treatments have placed Chloris spikelet morphology. The higher categories rec- and most of the rest of the genera in the tribe ognized in the older classifications (especially Chlorideae in a separate subfamily, which is subfamilies) may be quite artificial. This is usualh' called the Eragrostoidcae or Chlori- reflected most strikingly in the modern systems doideae. Tlie number of tribes included in this ) ) ) )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chi.obis (GnAMiNEAE) subfamily has varied considerably; for example, Cytology

Gould 196Sa ) recognized seven. ( No single, extensive attempt to determine Throughout the development of the more chromosome numbers in Chloris has been made, modem classifications, the group of genera fonn- and the counts reported in Table 1 are largely ing the Chlorideae has remained relatively taken from incidental reports of chromosome stable, including such familiar genera as Bou- numbers of a variety of grasses. In some cases telotia. Biichlol'. Ctcnium. Cijnodon, Entcropo- I have been able to verify the identification of gon, Eustacliys, Gynmopogon, and Tetrapogon, the plant b\' examining the voucher specimen; among others. for these, the herbarium containing the voucher genera The taxonomic parameters of these is indicated in parentheses after the counter's have been widel)' accepted; yet intensive study name. Chromosome numbers followed by my of certain complexes immediately reveals the name are reported for the first time. Vouchers artificialit\' of man\' of the genera. Most inves- for these are deposited in the California State tigations of generic relationships have been con- University Herbarium at Humboldt. fined to the study of Narious species of selected From Table 1 it can be seen that chromo- genera; seldom have attempts been made to ex- some numbers have been listed for fewer than amine the majorit\' of species from a number half of the species in Chloris, and thus, only of viewpoints, especially for endomorphic char- limited conclusions can be drawn. Most spe- acters. cies have a base number of x=10 and onl\- in- A report of such an attempt for ChJoris fol- frequently have other numbers been noted. Rel- lows. The limited a\'ailabilit\' of material for atively few diploids have been identified; most manv taxa has frustrated certain phases of stud\', species are at various polyploid levels, with the most notabh' those requiring living materials. decaploid number of 2n = 100 being the highest Areas and species deserving further investiga- known in the genus. Several species have vary- tion will be immediateK' apparent. ing chromosome numbers. Cltloris gatjana is

The discussion is divided into three areas: noteworthy in this respect, with 2n = 20, .30, and cvtological, histological, and morphological evi- 40. dence. An attempt to s\nthesize these lines of There are few studies of reproductive mecha- evidence as a basis of classification of the spe- nisms. .Again, the best-known species is Chloris cies in the genus follows. gaijana. Moffett (1944) suggested that the spe-

Table 1. Cliromosome numbers in the genus Chloris.

Species Geographic origin Counted by

Chlom andropogonoides 40 Te.vas (without precise locality) Brown (1950) Texas: Nueces Co. (TAESI); San Patricio Co. (TAES!); Starr Co. (TAES! TEX! UC!) Gould (1960) Texas: Hays Co. Anderson 3,282 (HSC) C. aristata 100 Mexico: Oaxaca (USI) Tateoka (1962a) (as C. rufescens) Me.xico: Hidalgo (UC! US!) Gould (1965) (as C. rufescens)

Mexico: Queretaro Gould ( 1966) (as C. rtifescens) Mexico: Chiapas Gould & Soderstrom (1970) (as C. rufescens

C. Iicrroi 40 Texas : Brazos Co. ( as a weed in a grass nursery) Gould (1960)

C. bournei 40 India ( without precise locality Janaki-Ammal in Darlington & Janaki- Amma! (1945)

50 Without locahty Krishnaswamy ( 1940 C. canterai 72 Texas: Milam Co. (TAES! UC! US!) Gould (1958) (as C. pohjdactyla) C. chloridea 40 Mexico: Chiapas Gould & Soderstrom (1970) 80 Mexico: San Luis Potosi (US!) Gould (1965) C. ciliata 40 Texas: Wharton Co. (TEX!) Brown (1950) Texas: Walker Co. Gould (1958)

Botanical Garden, Lyon (cultivated) Singh & Godward ( 1960) Dominican Republic: Santiago Gould & Soderstrom (1967) Texas: Bee Co. (TAESI) Gould (1968b)

Texas: Cameron Co. Anderson 4,546 ( HSC C. crinita 40 Texas (without precise locality) Brown (1951)

Argentina : Cafayate Bowden & Senn (1962) Bricham Young University Science Bulletin

Table 1. (continued) C. cucullata 40

C. gaijana 20

30 40

C. halophila 80 C. injlata 20 40

ca 50

C. vilosa 20

C. plurifhra 80

C. pyciwthrix 30 36

C. radiata )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gr.\mineae)

Table 1. (continued)

C. tnincata 40 Australia ( without precise locality Avdulov (1928) Without locality Krishnaswamy (1940) Australia (seed from USPI, no. 212-389) Anderson (HSC)

C. verticillata n = )

Brigham Young Univehsity Science Bulletin

(Linnaeus) Swartz; (RR) C canterai Arechava- leta; (SS) C. amethystea Hoclistetter; (TT) C. lobata Lazarides; (UU) Eustachys tenera (Presl) A. Camus; (VV) Chloris pycnothrix Trinius; (WW) C. aristata (Cervantes) Swallen; (XX) C. virgata Oo o OoOopgoOO Swartz; (YY) C. pilosa Schumacher; (ZZ) C. gayana Kunth; (AAA) C. inflata Link; (BBB) C. or-

thonoton Doell in Martins. Arrangement is by overall similarity of microhairs. X 475.

raphy, from ab.sont to moderately abundant. 80888SRRSSOO Because of this random variabilit)', prickles were 0,0,0,8,^,^,0,0,0, 0,0.0, not emphasized as a taxonomic character. Of the otlier epidemial characters studied, 8§Mf 9 99" #tii Rogers concluded that the most stable were shape, size, and type of microhair and the shape ogggo 88g§ooq of the costal silica cells. Typical microhairs and costal silica cells are illustrated ( Fig. 1 ) for 40 species of Cliloris and for 14 other species 8 which were formerly placed in Chloris, but 8919998 @99 which are now placed in other genera. — Two types of microhairs, unicellular and bi- '^ ( C-J cellular, were taxa n n ^ —in d observed in the studied. Table 3 lists the species showing these two microhair t)'pes. One species complex (consist- ©eeoo V ing of C. aristata, C. gaijana, C. inflata, C. ortho- OQ R Qf^ noton, C. pilosa. C. jx/rnothrix, and C. virgata) On n 9 Q possesses vmieelliilar microhairs. The spikelets O 8 SyOo vv WW XX n^-^ZZ AAA^BI and inflorescences of these species are similar to one another, yet they are not unique in the Fig. 1. Microhairs (upper figure of each set) and genus, and several strongly resemble particular costal silica cells (lower figure) of Austrochlom, Chloric, Enleropogon, Eustachi/s, and Tetrapogon. species with bicellular liairs. The remaining (A) Austrochloris dichanthioides (Everist) Laza- species all have bicellular microhairs. While rides; (B) Chloris brandegei (Vasey) Swallen; several different shapes and sizes of hairs are Brown; submutica (C) C. divaricata R. (D) C. recognizable, these cannot be correlated with Humboldt-Bonpland-Kunth; ( E ) Eustachys petraea an\' macroscopic features. Swartz; (F) Chloris bournei Rangaehariar and Tadulingam; (G) Euatachijs distichnplujUa La- \'irtuall\' all of the taxa examined have

gasca; ( H ) Chloris scariosa F. von Mueller; ( I costal silica cells var\ing from cuboidal to Eustachys retusa Lagasca; Chloris halophila (J) saddle shaped (Fig. 1; Table 3). Only one spe- Parodi; (K) C. ventricosa R. Brown; (L) C. mos.^- cies differs from this pattern: C cruciata has flni/;iVt'ti.siv Schumann; (M) Tetrapogon macrnnlhns (Jaubert and Spach) Bentham; (N) Chloris pec- costal silica cells which tend to be more elon- tinaia Bentham; (O) C. robusta Stapf in Chevalier; gated and to ha\'e more deeply indented end (P) C paniculata Scribner in Robinson; (Q) C. walls than the remaining species. roxburghiana Schultes; (R) Tetrapogon spathaccus

( Hochstetter) Hackel ex Durand and Schinz; (S) Anatomy. Embryology, and Seedling Chlorii ciliata Swartz; (T) C. prieurii Kunth; (U) C sagracana A. Richard; (V) C. suriiigari Hitch- Morphology cock in Urban; (W) Tetrapogon tcncllus (Rox- Anatomical, embr\ological, and seedling burgh) Chiovenda; (X) Chloris chloridea (Presl) studies on a selected group of 28 taxa in Chloris Hitchcock; (Y) C. somalensis Rendle; (Z) Entero- and a]Ii(>d genera were completed by Jensen pogon dolichostachya ( Lagasca ) Lazarides; ( AA

Chloris dementis Merrill; (BB) C. radiata (Lin- ( 1969 ) . The species were selected to be repre- naeus) Swartz; (CC) C. texensii Nash; (DD) C. sentati\'e of as man\' different sections of the

mollis (Nees) Swallen; ( ) Tetrapogon villosus EE genus as possible in terms of spikelet morphol- Desfontaines; (FF) Eustachys paspaloidet (Vahl) og\'. Characteristics studied included leaf anat- Lanza and Mattei; (GG) E. glauca Chapman; (HH) Chloris ekmanii Hitchcock; (U) C. pumilio omy, embryo stnicture, root-hair position and R. Brown; (JJ) C cuhensii Hitchcock and Ekman; aspect, shape of the first seedling leaf, and t\'pe ( KK ) C. ilandi/ana C. D. Adams; ( LL ) Eiitero- of starch grains. Individual discussions of these pogon acicularls (Lindley) Lazarides; (MM) features follow. Eustachys neglecla Nash; ( NN ) Chloris truncata Jensen examined several anatomical features R. Brown; ( OO ) C andropogonoides Fournier; (FF) C. berroi Arechavaleta; (QQ) C. cruciate of leaf cross sections, including the shape of the Biological Series, Vol. 19, No. 2 Ta.xonomy of the Genus Chloris (Ghamineae) Brigham Young University Science Bulletin

keel, the disposition of vascular bundles, the tex- ture of the adaxial surface (ribbed or smooth), the extent of the sclerenchyma girder, the distribution of bullifomi cells, and the nature of the vascular bundle sheath. While individual species may show stable and recognizable patterns, few correlations between leaf anatomy and spikelet morphology could be made. Like- wise, there were no consistent correlations of anatomical features with either microhair or costal silica cell types. The classical study of Rceder (1957) in comparing anatomical features of embryos of

various genera of grasses is well known. Reeder separated the grass genera studied into essentially two groups, panicoid and festucoid, on the

basis of four embiTO characteristics: ( 1) whether the lower portion of the scutellum is separate from the rest of the embr)'0 (panicoid) or fused (festucoid); (2) the presence of an epiblast (festucoid) or its absence (panicoid): (3) whether both vascular traces diverge from about the same point (festucoid) or are separated by an elongated area of vascular tissue between points of divergence (panicoid); and (4) whether the margins of the embrvonic leaf overlap (panicoid) or just meet (festucoid). Jensen (1969) studied three of these characteristics for 25 selected species in C^'Inris and certain related segregate genera (Table 4). Typically, all species studied had the scutellum separate from the rest of embryo (panicoid), an epiblast present (festucoid), and vascular traces separated by an elongated area of vascular tissue (panicoid). Tlie fourth feature, whether or not the margins of the embr\'onie leaf overlap, was studied on 19 of Jensen's 2S taxa (Table 5). While both festucoid and panicoid types are represented among these species, consistent correlations with other anatomical features or with

Table 4. Ta.xa in CMoris and related segregate genera having panicoid scutellum type, an epiblast present (festucoid), and a panicoid arrangement of vascular tissue.

CMoris aristata C. herroi C. hournei Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae) of the limited availability of viable seeds, Jensen lated genera is presented in Fig. 2. The number was able to examine onh' ten species for this of taxa commonly accepted in each is indicated characteristic, all of which had the root hair pro- by the relative size of the circle. Also given jecting at an angle of about 60° near one end of arc those species in ClUoris that resemble the the cell. The species examined were: Chloris satellite genera, as well as those in the satellite canterai, C. gayana, C. inflata, C. pilosa, C. genera that have been included in Chloris. stil)mutica. Eiistaclnjs pijcnothrix. C. C. truncata, There is little infonnation, aside from spike- caribaca, E. distichopluiUa. E. petraca, and E. let moiphology, that is useful in generic delimi- rettisa. tation within this group. Chromosome counts The first seedling leaf in panicoid grasses is are mostly of the same basic number; leaf epi- relatively wide and curled; that of festucoids is dermal and anatomical characteristics do not long and narrow ( Stebbins, 1956 ) . The species appear uniciue to particular genera; genetic in- listed leaves, ac- above all had panicoid seedling fonnation is \irtuall\- nonexistent. Consequently, cording to Jensen. speculation on relationships, both intergeneric and interspecific, rests heavilv upon infomiation Relationships of Chloris and Other Genera gleaned primarily from the structure of the in- As with manv genera in the Gramineae, florescence and spikelet and, secondarily, upon generic delimitations in the Chlorideae may be data concerning vegetative features. However, difficult to assess; and they, of necessity, may the genera in this complex have been long be somewhat arbitrar\' and artificial. The com- recognized and have remained relatively un- plex that includes Chloris is no exception. While changed. The description of each of these, as most of the species have long been included commonly accepted toda}-, is given in Table 6, within the genus and most of the characteristics and their relationships to Chloris follow. recognized, some species have been placed in Eustachi/s. This genus was first described adjacent genera in the past, a few more recentK' in 1810 by Desvaux, but it has been treated so. A graphical conspectus of Cliloris and re- subsequently as a subgenus of Chloris bv the

UNISPICEUS A U S T R C H L R I S

E N T E R P G N

ACICULARIS ; EOSTAPFIELLA DOLICHOSTACHYA

D A K N P H L I S

NEGLECTA X'CYNOCHLOR I S PASPALOIDES

PETRAEA EU ST ACHYS RETUSA ) DISTICHOPHYLLA DELICATULA TENERA / FLORIDANA G YMNOPOGO ULIGINOSA y iSLAUCA

Fig. 2. Relationships of Chloris and allied genera. 10 Brigham Young University Science Bulletin

Table 6. Comparison of various characteristics of Chloris and related genera.

Genus Vegetative characteristics Inflorescence type Glumes Number

Chloris tufted, rhizomatous or sto- usually a single radiate se- both glumes simi- loniferous; sheaths usually ries of 4-10 spikes, occasion- lar, narrowly ovate overlapping at the base, ally 2-more approximate or to lanceolate; acute blades narrow to broad, not only slightly separated series to acuminate; awn- distichous of 10-20 spikes; rarely an indefinite number of terminal spikes or as few as 1-2 spikes

Cyrtodon rhizomatous to stoloniferous; a single radiate series of sheaths usually overlapping 4-9 spikes at the base, blades generally narrow, acute, distichous

Daknopholis prostrate, repent to stolonif- a single radiate series of erous; leaves basal, sheaths 2-4 spikes overlapping, blades small, subelliptic, obtuse

Enteropogon erect, tufted, sheaths usual- usually a single terminal ly not markedly overlapping spike, sometimes several to at the base, blades narrow, many in a radiate or sub- flat or convolute verticillate series Biological Series, 12 Bricham Young University Science Bulletin inforces the exomoiphic differences between the sheaths are not as strongly overlapping. The them. spikes are somewhat distant, but not as markedly

Tetrapogon. This Old World genus is best so as in Gymnopogon. Sterile florets in C. mollis separated from Chloris on the basis of having are well developed; in this respect they are unlike those in Gymnopogon. two ( occasionally three ) fertile florets per spikelet. Chloris regularly has a single fertile floret, Chloris dementis, C. ekmanii, C. pycno- though very occasionally individuals of certain thrix, and C. radiata all have mucli reduced species may produce a spikelet with two or more sterile florets; but in all other respects they fit fertile florets. In addition, both glumes in Tetra- Chloris well. pogon are thin, papery, or parchmentlike and Cynodon. Clifford and Everist (1964) de- (like the spikelets) are relatively large. The scribed a sterile intcrgeneric h\brid, which they glumes in species of Clihris are not thin and named Cynochloris macivorii, which was found papery, but are finner and relatively smaller. growing with Cynodon dactylon and Chloris Several species in Chloris strongly resemble divaricata on a lawn bowling rink at Ipswich, Tetrapogon. Chloris inossamhicetisis, an African Queensland. The identification of this plant species, is similar in many respects, but the bulk rests upon its intermediacv between the two of material which this author has examined has putati\e parents with respect to a number of spikelets with one fertile floret; only a few plants spikelet, habit, and behavioral characteristics, have spikelets with two or more florets. Chloris since experimental recreation of the hybrid has bournei, C. tcightiana, and, to a lesser extent, not been attempted. On the basis of the data,

C. quinquesetica—aW from India—are similar to there is little doubt as to the correctness of Clif- several Tetrapogon species, primarily in size of ford and Everist's inteipretation, however. spikelets. However, they have just a single fer- The existence of this bigeneric hybrid at- tile floret and are best retained in Chloris. tests to the genetic proximity of these two gen- The strikingly different Chloris scariosa from era; \et the two can ordinarily be separated Australia could easily be given generic status, readily by morphological features. Most species for its spikelets (with their several flabellate of Cynodon have obsolete or verv poorly de- sterile florets) arc unlike any other species in veloped sterile florets, while in Chloris they are Chloris. Chloris scariosa has relatively large, usually well developed. Likewise, in Chloris papery glumes, in this respect strongly resem- the callus is prominentK' short bearded, whereas bling Tetrapogon. I am maintaining this spe- in Cynodon it is less obviously pubescent. cies in Chloris largely because of its single fer- Daknopholis. Described in 1967 by Clayton, tile floret. If Chloris scariosa were to be given this genus was erected to contain several anom- generic rank, then a number of other somewhat alous species of Chloris: C. hoivinii, C. perrieri, unusual species are also deser\ing of such con- and C. ramosissima. Two features will separate sideration—a process that could hopelessly frag- the new genus from Chloris: ( 1 ) the lack of mentize the genera in this complex. a well-defined sterile floret and (2) the presence Gymnopogon. The most striking difference of a truncate and erose second glume. In over- between this genus and Cliloris is vegetative. all appearance Daknopholis is more suggestive

Qymnopogon is characterized by thick, sharp- of Cynodon than Chloris. The relationships pointed, distichous blades, the sheaths of which were discussed in some detail by Clayton are more or less cquitant and basal. Further- (1967). more, many species of Gymnopogon have the Entcropogon. Tliese Old World trooical pe- spikes well separated and spreading, making the rennials (as treated b\' Clayton [19 17], for ex- inflorescence quite open. Chloris generally has ample) are easily separated from Chloris in hav- broader, more lax blades and sheaths that only ing a single spike or, rarely, two; most Chloris partly overlap. While a few species have distant species have three or more .spikes. In addition, spikes, they are mostly borne in one or more the fertile lemmas of Entcropogon are strongly, radiating series. dorsally compressed and have a raised midnerve. Most species of Gymnopogon either lack I do not believe that this difference in num- sterile florets or they are very greatly reduced. ber of spikes accurately reflects the relationships.

In Chloris the sterile floret is generally well Two species with many spikes that are often developed. treated in Chloris— C. acicularis from Australia

Overall, Chloris mollis is probably most simi- and C. dolichostachya from southeastern Asia- lar to Gymnopogon, a fact reflected in its no- have strongly dorsally compressed spikelets with menclatural histoiy. While the leaf blades of a raised midnerve on the lemma. The similarity C. mollis are relatively short and sharp pointed. to Enteropogon is such that one would be hard )

Biological Series, Vol. 19, No. 2 T.\.\onomv of the Genus Chloris (Cramineae) 13 pressed to separate these two taxa of Chloris or with the single terminal sterile floret identical and various species of Enteropogon on the basis to, or strongly resembling, the basal fertile of spikelets alone. They could, of course, be floret. Additionally, the number of spikes (one separated readily on the number of spikes. But to three) is fewer than that generally found in the differences do not end here. Tateoka ( 1962b Chloris. Neostapfiella is probably more closely reported that starch grains were simple and an- related to Tetrapogon than to any species in gular in Enteropogon, while Chloris has only Chloris. compound grains. Both CJdoris acicularis and C. dolichostaduja have Enteropogon-ty^e starch Relationships within Chloris further reinforcing the removal of these grains, This synthesis is based primarily upon the species from Chloris. Accordingly, I have fol- gross morphologv of the inflorescence and spike- in removing these two lowed Lazarides (1972) let; but studies of the leaf epidermis, already En- species from Chloris and referring them to discussed, provided important additional infor- teropogon. mation. Chromosome number and structure have Trichloris. This genus has been character- contributed virtually nothing, for the genus is ized as having three prominent awns on the essentially monobasic and the chromosomes very fertile and sterile lemmas: one is an extension similar. other two are of lateral of the midnerve and the Some species complexes are well defined; i.e., relative length of these awns varies nerves. The there is a strong correlation of morphology, an- between species. atomv, and geographv. Other species may re- The prominently threc-awned lemmas of semble one another, but they are geographically virtual- Trichloris will distinguish it readily from isolated in such a wav as to preclude a direct Chloris in the Western Hemis- ly all species of cvolutionarv- connection. In still other complexes Chloris chloridca and C. halophila have phere. there may be little or no outward resemblance; these are somewhat bilobed lemma apices, but vet the formation of hybrid swarms recombines mucronate, at most. those highly divergent characters and indicates difference serves also for the This same awn strong cvolutionan' ties. Still other taxa are in Australia. There Eastern Hemisphere, except isolated entities, displaying no obvious relation- are several Australian species, too, that have ships with any other species or complexes. lateral nerves extended into awns (prominently Tlie formal recognition of a series of sub- so in C. lohata and C. punulio; less obvious in genera would be folly, not that some complexes C. ilivaricata and C. pectinata.) In all likelihood, are unworthy of it, but because the naming of of course, this resemblance is a result of parallel these as subgenera will necessarily create a resi- evolution and docs not reflect a close genetic due of species that cannot be grouped. The in- relationship. Nevertheless, it is ambiguous to dividual species in this remnant cannot be continue to recognize Trichloris in the New treated as a single subgenus, for such would World, while maintaining the three-awned spe- be blatantly artificial and misleading. The other cies from Australia in Chloris. Clayton (1967) extreme would be to treat each as the single recognized this ambiguity and treated the two member of its own subgenus, a practice leading species of Trichloris as members of Chloris. to rather hopeless fragmentation. I have tried More recently, Sanchez (1971) resurrected to compromise in the following discussion. Those Trichloris on the basis of anatomical studies of groups containing obviously closely related spe- leaves. According to Sanchez, the bulliform cies—whether the basis is morphological, geo- cells of most Argentine species of Chloris have graphical, genetic, or all three—are treated as achloroph\llous cells hing immediatelv below species complexes and discussed as such. The them in the mesophyll. Such achlorophvllous isolated species are discussed individually wher- cells are onlv rarely found in Trichloris. ever this author feels that they have importance. While there are recognizable tendencies in In addition, discussions of relationships, espe- the patterns of variation in these anatomical cially within the various complexes, follow the is spike- features, there no reinforcement from descriptions provided later. A graphical synop- let I following morphology. Consequently, am sis of relationships within the genus is provided the lead of Clayton ( 1967 ) in treating the two in Fig. 3. species of Trichloris as members of Chloris. Neostapfiella. This genus from Madagascar The Chloris inflate complex. is relatively poorly represented in most herbaria. Members of this complex are characterized The limited material available, however, shows by single-celled microhairs, by typical Chloris- the spikelets with either two basal fertile florets type silica cells, and, in most, by a prominent 14 Brigham Young Univebsity Science Bulletin

burmensts

scavvosa

robusta

anethystea

mossambicensis

somalensis

filiformis Qubmutica paniaulata

Fig. 3. Relationships of species and species complexes in the genus Chloris.

tuft of hairs on the upper margins of the fer- is well defined as a whole. All species (Chloris tile lemma. Included here arc five very widely berroi, C. canterai, C. ciliata, C. dandijana, and distributed species {Chloris gmjana, C. inflata, C lamproparia, are characterized bv having rel- C. pilosa, C. pi/cnothrix, and C. virgata), as well atively short-awned, elliptic fertile lemmas that as several more restricted species (C. aristata, are prominently spreading-ciliate along most of C. castilloniana, C. jormosana, and C. orthono- the margin. Epidermal patterns are typical of ton). Of these, Chloris jn/cnothrix is the most Chloris. Chloris sesquiflora is similar, but the

aberrant; if it were not for possession of unicel- marginal pubescence is strongly appressed.

lular microhairs, it would be placed confidently The Clitoris verticillata complex. in the following complex. Were it not for the extensive introgression Chloris radiata complex. The between Chloris andropogonoides, C. cucullata, Six similar species (C. arenaria, C. ctibensis, and C. verticillata, they would probably not be C. ekmanii, C. halophila, C. radiata, and C. sa- included in a single complex. Moi-phologically,

graeana) make up this complex. All have com- there is considerable divergence, especially in C. paratively narrow spikelets and reduced sterile cucullata, in which the broad fertile and sterile florets; relatively long awns are also a common lemmas are distinctive in the genus. While in- feature. The cpidennal pattern includes bicel- trogression involving the very rare C. texensis lular microhairs and cuboidal silica cells. While has not been demonstrated, in spikelet and in-

most of the species have inconspicuously ciliate florescence morphology it is obviousl)' similar to upper lemma margins, C cuhcnsis may approach C. andropogonoides. the inflata complex in having more prominent C In Chloris chloridea and C hrandegei there pubescence. Though Chloris cruciata is not in- is a strong tendency toward dorsal compression cluded here because of its somewhat unique of the spikelet. In other respects the plants are short cells, it is similar in overall spikelet struc- similar to C. halophila, though considerably ture. more robust. The development of underground The Chloris ciliata complex. spikelets in Chloris chloridea is unique in the While this group presents more interspecific genus.

taxonomic problems than any other group, it Four Australian species have bilobed fertile Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlouis (Ghamineae) 15 and sterile lemmas. In Chloris pumilio and C. size from only a few cm to over 2 m tall; epi- lohata, this lobing is very prominent; the other dermal cells of root tips slightly different in two, C. pectinata and C. cUvaricata, are less so. size, giving rise to root hairs which project The outward similaritv' to C. plurifhra and C. forward at about 60°; sheaths glabrous, scab- crinita, both formerly in Trichloris, an American rous, or villous; hgule often a ciliate crown, genus, is probabK- the result of parallel evolu- sometimes absent; blades narrow to wide, flat tion. or rolled, deeply keeled and V-shaped, or with Another pair of Australian species, Chloris rounded keels and with several furrows, scleren- triincata and C. ventricosa, is notable for having chyma girders complete or only on abaxial side, ver\' truncate or ventricose fertile lemmas. Chlo- bulliform cells generally large and penetraring ris triincata is strikingK similar to the North the mesophvll, bundle sheaths complete or in- American C. verticillata; again, the similarity complete; costal silica cells usually cuboidal- seems best explained by parallel evolution. saddle- or axehead-shaped, very rarely some- Chloris h>ngiaristata and C. prieurii, both what cruciate, stomatal apparatus medium- to primariK- African in distribution, have several triangular-dome shaped, bicellular microhairs sterile florets and relati\ely narrow, long-aw ned usually present, with relatively broad and short lemmas. Tlie glandular strip along the side of terminal cells, or, less commonly, single-celled; the lemma in C. prieurii is unusual in Chloris. spikes' usually 5 to 25 (occasionally as few as Clitoris hournei. C. (piincptcsetica, and C. 1 or 2, rareb' more than 25), frequentlv digi- wightiana all have relatively large spikelets with tateh arranged, occasionally in several verticils, obtuse lemmas, coarsely pubescent lemma mar- sometimes with a few isolated single spikes; gins, and several sterile florets. Thev bear a spikelets borne in two staggered rows on one strong resemblance to species in Tetrapogon side of the rachis, sessile to pedicellate, com- but ha\e just a single ferHlc floret, whereas monly rather densely imbricate and appressed, Tetrapogon has more than one. sometimes sparsely so; glumes usually unequal, The above discussion leaves unmentioned a lanceolate, often glabrous with scabrous nerves, rather large number of species which are quite mostly acute, rarely acuminate, shorter than florets; fertile isolated moiphologicalh' ( Fig. 3 ) . Unquestion- the lemmas one (very rarely ably, these are best retained as a part of Chloris, two), lanceolate, occasionally elliptic, three- for thev have all of the features characteristic of nerved, callus bearded, margins shortly to the genus; and yet, the elaboration of some in- prominently pubescent, intemerves usually di\idual structural feature causes them to stand glabrous, sometimes scabrous, rarely pilose, alone. None is trulv widelv distributed, and midnerv'cs scabrous or glabrous, rarely long- several are narrow endemics; this suggests adap- pubescent, midnerve usually extended as an tation to a particular climatic or edaphic regime. awn, either from the tip or from betv\'een two teeth, rarely only mucronate; palea shorter than Chloris Swartz the fertile lemma, with two scabrous nerves, otherwise glabrous; stamens three, anthers rela- Chloris Swartz, Prodr. Veg. Ind. Occ. 25. tively small; pistil one, with two stigmas; lodi- 1788. cules two; sterile floret usually one (rarely two Rabclochloa Beauvois, Ess. Agrost. 84, 158, or three, mostly barren, very rarely staminate 176. 1812. or perfect, varying widely in size and shape Phacellaria Willdenow ex Steudel, Nom. from highly reduced to well developed and Bot. ed. 2. 1:353. 1840. Herbarium name, given similar to the fertile floret, rudimentary to as a svnonym. cylindrical to obovoid, acute to truncate, awned Heterolepis Ehrenberg ex Boissier, Flora or unawned; caryopsis ovoid, elliptic to obo- Orientalis 5:554. 1884. Herbarium name, given void, round to trigonous in cross section, the as a synonym of Chloris. embryo relatively small, lower portions of Trichloris Fournier, Mex. PI. 2:142. 1886. scutellum separate from the rest of the embryo, Chloropsis Kuntze, Rev. Gen. PI. 2:771. 1891. epiblast present, vascular traces separated by Leptochloris Munro ex Kuntze, Rev. Gen. PI. an elongated area of vascular tissue, embryonic 2:771. 1891. leaf margins usually not overlapping, very Pterochloris Camus, Bull. Mus. Hist. Nat. rarely overlapping; starch grains of endosperm (Paris). Ser. 2. 29:349. 1957. compound; basic chromosome number x^9 Fibrous-rooted annuals or rhizomatous, (rarely), 10 (usually), the chromosomes rela- stoloniferous, or cespitose perennials, ranging in tively small.

^The inflorescence branches of the various species bear spikelets that range from virtually sessile to obviously pedicellate, and thus may be called fpikes or racemes, depending upon emphasis. In this study the main inflorescence branch is arbitrarily called a spike. 16 Brigham Young University Science Bulletin

Widely distributed in tropical, subtropical, spheres, both north and south of the equator or warm temperate regions of both hcmi- (Fig. 4).

Fig. 4. Worldwide distribution of the genus Chloris. { This map is a composite of the distribution maps of all of the species.)

Key to the species of Chloris

1. Inflorescence of many (30 or more) short spikes (6 cm or less) arranged along a central axis which is at least 1 cm in length, the spikes not in verticils 2 Inflorescence of less than 30 spikes, the spikes arranged in one or more verticils 3

2. Sterile florets two or more; widely distributed in Africa, occasional in India C. rox])ur

3. Spikes two to four, forming a cylindrical, spikelike inflorescence; the spikes attached to one another at least at the base and often for the full length, separable only with difficulty; Argentina, Uruguay C. herroi, p. 25 Spikes neither attached to one another nor forming a cylindrical, spikelike inflorescence 4

4. Spikes one to three 5 Spikes four or more 14

5. Sterile florets two or three 6 Sterile floret one (See also Chloris suringari, which has a highly reduced upper sterile

.' floret.) - 10

6. Fertile lemmas 1.8 to 2.8 mm long 7 Fertile lemmas 2.7 to 4.2 mm long 8

7. Inflorescence a single spike; lemmas awned from about the middle of the keel; Cuba, Cura9ao) C. stirinp.ciri, p. 39 Inflorescence of more than one spike; lemmas awned subapically; southcentral United States, Caribbean Islands, Mexico, Central and South America C. ciliata, p. 27 Biological Series, Vol. 19, No. 2 T.jv.xonomv of the Genus Chloris (Gr.^mineae) 17

8. Combined length of fertile- and sterile -floret awns 6.5 to 9 mm, usually more than 7 mm; annuals; Africa C. lamproparia, p. 30 Combined length of fertile- and sterile-floret awns 4 to 8 mm, usually less than 7 mm; perennials; South America, Texas 9

9. Plants densely tufted, with basal, narrow, rolled, curved leaves; spikes 3 to 5 cm long C. canterai var. grandiflora, p. 32 Plants not densely tufted; leaves cauline, up to 5 mm wide, not curled or curved; spikes 4 to 14 cm long C. canterai var. canterai, p. 30

10. Spikelets diverging from the rachis at nearly right angles; spikelets obviously pedicellate; Culja, Haiti, Jamaica C. cruciata, p. 37 Spikelets appressed or cmly slightly divergent from the rachis; spikelets sessile or nearly so 11

11. Awn of fertile lemma less than 1.5 mm long; internerves appressed-pubescent; Mau- ritius C. filiformis, p. 39 Awn of fertile lemma more than 2 mm long; margins and keel spreading-pubescent; internerves glabrous or with inflated hairs 12

12. Awn of fertile lemma longer than 5 mm; Cuba, Jamaica, Hispaniola

_ C. ekmanii, p. 119

Awn of fertile lemma less than 4.5 mm long 13

13. Internc'i-ves glabrous; Cuba, Curasao C. surin^,ari, p. .39 Internerves densely covered with inflated hairs; Malagasy Republic C. humbertiana, p. 104

14. Sterile florets two or more, the uppermost floret sometimes represented only by the clavate rachilla joint 15 Sterile floret one 41

15. Sterile florets greatly modified, commonly o\er 3 mm wide, flabellate, margins scarious, more or less inrolled and enclosing one another; Australia C. scariosa, p. 40 Sterile florets neither greatly modified, flabellate, nor involute, seldom more than 1.5 mm wide 16

16. Fertile lemmas with a row of glands along the sides; Africa, India — C. prieurii. p. 42 Fertile lemmas without a row of glands, though sometimes appressed-pubescent with nonglandular hairs 17

17. Fertile lemmas with a short mucro (less than 1 mm long); Africa C. pilosa, p. 58

Fertile lemma with an awn more than 1 mm long .- 18

18. Sterile floret pubescent, either with a prominently appressed-pubescent callus, or densely pubescent on the midncrve, upper margins, or apex 19

Sterile floret glabrous or, at most, with a few ( 10 or so ) appressed hairs on the back 26

19. Callus of sterile floret bearded 20

Callus of sterile floret not bearded 21

20. Sterile floret unawned, acuminate; rachilla joint not obvious; culms stout, bamboolike; Africa C. rohusta, p. 43

Sterile floret obviously awned; rachilla joint prominent; culms not stout and bamboolike; New World C. mollis, p. 45

21. Leaf sheath, blade, and spike rachis densely pustulose-pilose; sides of fertile and sterile lemma prominently and densely pustulose; India C. wip.htiana, p. 48

Leaf sheath, blade, and spike rachis glabrous , scabrous, or sparsely pilose, but never pustulose; sides of lemma glabrous or occasionally appressed-pubescent, but not pustulose.. 22 )

18 Bricham Young University Science Bulletin

22. Culms densely appressed-pubescent for several mm just below the points of attachment of the spikes 23 Culms glabrous or scabrous below the points of attachment of the spikes 24

23. Upper margins of lowermost sterile floret ciliate; sterile florets usually four ( occasionally 3); uppermost sterile floret spherical and inflated; India ...C. quinqtiesetica, p. 4S Upper margins of lowermost sterile floret usually not ciliate, occasionally sparsely pubescent; sterile florets usually three (rarely two or four); upper sterile floret resembling the lower, neither spherical nor inflated; Ceylon, India, Thailand C. montana, p. 51

24. Awn of fertile lemma 2 mm long or less; Burma C. btinnensis, p. 125 Awn of fertile lemma more than 3.5 mm long 25

25. Midnerve of lowest sterile floret densely and prominently appressed-pubescent; fertile lemma longer than 3 mm; lowest sterile floret 1.5 mm or longer, its awn usually shorter than 4 mm; stoloniferous perennial; southern India C. hournei. p. 53 Midnerve of lowest sterile floret sparsely appressed-pubescent; fertile lemma less than 3 mm long; lowest sterile floret less than 1.5 mm long, its awn over 5 mm; annual or weak perennial, sometimes rooting at the lower nodes; pantropical C. infhita. p. 53

26. Awn of fertile lemma longer than 30 mm; Africa C. longiaristata, p. 46 Awn of fertile lemma less than 15 mm long 27

27. Sterile florets two or three, only the lowermost awned, the others awnless 28

Sterile florets two or three; all are awned 34

28. Fertile lemmas prominently ciliate along most of the margin, the cilia spreading at nearly right angles; lemmas broadly elliptic 29 Fertile lemma margins glabrous or with .strongly spreading hairs near the apex, but never along tlie lower margins 33

29. Fertile lemmas 1.5 to 2.8 mm long 30 Fertile lemmas 2.9 to 4.2 mm long 31

30. Spikes three to five (rarely six or seven), usually less than 8 cm long; Texas, Mexico, Central America, southern Brazil, Argentina C. ciliata, p. 27 Spikes 7 to 40 (usually 10 to 20), usually more than 8 cm long, occasionally as short as 5 cm; Florida, West Indies, South America C. dandijanci, p. .34

31. Spikes one to four (often two); combined length of awns of fertile and sterile floret 6.5 to 9 mm (usually about 7 mm); annuals; Afric;i C. himpwparia, p. .30 Spikes two to nine (usually three or more); combined length of awns of fertile and sterile floret 4 to 8 mm (usually less than 7 mm); perennials; South America, Texas .32

32. Plants densely tufted with basal, narrow, rolled, curved leaves; spikes 3 to 5 cm long C. canterai \ar. grandijUmi, p. 32 Plants not densely tufted; leaves cauline, up to 5 mm wide, not curled; spikes 4 to 14 cm long C. canterai var. canterai. p. .30

33. Annual plants arising from a fibrous root system, occasionally rooting at the lower nodes

if culms are decumbent; Africa C. pilosa, p. 58 Perennial plants, usually with prominent, aggressive stolons, occasionally tufted; widely distributed in the warmer parts of the world C. gaijana, p. 65

(NOTE: Occasional specimens of C/iZoris uenfn'ro.sr; having two sterile florets will - key to this point.

34. Awn of fertile lemma less than 2 mm long; Burma C. burmensis, p. 125 Awn of fertile lemma more than 3.5 mm long 35 )

Biological Series, Vol. 19, No. 2 T.^.xonomv or the Genus Ciii.oris (Cramineae) 19

35. Culms densely appressed-pubesceiit for several mm just below the points of attachment of the spikes; Ceylon, India, Thailand C. montana, p. 51 Culms scabrous or glabrous just below the points of attachment of the spikes 36

36. Upper sterile floret inflated, nearh spherical; widespread in wanri regions C. inflata, p. 53 Upper sterile floret cylindrical or very narrowly turbinate 37

37. Rachis pilose; eastern China, Hong Kong, North Vietnam, Taiwan ...C. formosana, p. 57 Rachis scabrous 38

38. Fertile lemma with three awns, the lateral about equal to the central, or shorter.... 39 Fertile lemma with only a single awn, lateral awns absent 40

39. Awns subequal, the lateral only slightly shorter than the central C. crinita, p. 87

Awns unequal, the lateral about one-tenth the length of the central ..C. pluriflora. p. 89

40. Fertile lemmas acute at the ape.x; margins not inrolled, densely ciliatc, especially on the upper margins; spikelets tawny: robust, stoloniferous perennials more than 50 cm tall; widespread in warm regions C. gmjana, p. 65

(NOTE: On rare occasions, specimens of C/i/o;/.s HKWsa7ni»!censis, from southern Africa, and C. sa'^raeana, from tlie Caribbean region, both rarely haying two sterile florets, will key to this point.

Fertile lemmas obtuse to rounded at the apex; margins inrolled and appressed- pubescent; spikelets dark brown to black at maturity; relatively short, tufted perennials; Australia C. truncata. p. 7S

41. Sterile lemma bilobed for upper one-third to nearly all of its length, the lateral lobes usually awned 42 Sterile lemma not prominently bilobed 47

42. Fertile lemma margins pubescent to ciliate on upper half, the hairs usually prominent and spreading, especially toward the apex 43 Fertile lemma margins glabrous, scabrous, or strongly appressed-pubescent, but upper margins without prominent spreading hairs 44

43. Fertile lemma margins pubescent mostly on upper half; sterile floret bilobed nearly to the base, the lobes usually widely divergent at maturity; sterile lemma lobes gradually tapering into an awn; fertile lemma awns subeciual; Australia C lohata, p. 82 Fertile lemma margins pubescent along most of length (the hairs spreading apically, appressed basally); sterile floret bilobed to about the middle, very occasionally to the base, the lobes not widely di\ergent at maturity; sterile lemma lobes usually truncate to somewhat rounded, the awn arising laterally; median awn of fertile lemma usually about twice the length of the lateral awn C. piimUio, p. 80

44. Lateral and central awns of fertile lem ma subequal to equal; southwestern United States, Mexico, and South America C. crinita, p. 87 Lateral awns of fertile lemma about one-tenth the length of the central awns 45

45. Spikelets densely pectinate, crowded, diverging at wide angles from the rachis; Aus- tralia C. pectinata, p. 86 Spikelets not densely pectinate, only slightly imbricate, appressed to the rachis 46

46. Spikes in two or more verticils; South America C. halophila, p. 117

Spikes in a single radiate series; Australia C. divaricata, p. 83

47. Fertile lemma awnless or with a short awn, generally less than 2 mm long 48 Fertile lemma prominently awned, the awn usually more than 3 mm long 52 20 Bricham Young University Science Bulletin

48. Sterile floret very narrowly cylindrical, usually about 0.2 mm wide, usually sparsely pilose near the apex; Baja California, Mexico C hrandegeu p. 72 Sterile floret broader, usually more than 0.3 mm wide; at most merely scabrous at the apex 49

49. Sterile floret inflated-spherical, about 1 mm wide, usually as wide as long; Texas, New Mexico, Oklahoma, northeast Mexico C. cticullata, p. 91 Sterile floret not inflated-spherical, less than 5 mm wide, usually at least twice as long as broad 50

50. Fertile lemma margin glabrous, scabrous, or appresscd-pilose for no more than one-half of its length; keel usually glabrous, occasionally scabrous near the apex, rarely sparsely appressed-pubescent; Mexico, Colombia C. sul>imitica, p. 106 Fertile lemma margin and keel appressed-pubescent for most of length 51

51. Sterile floret elliptical, flattened, about 1 mm long; fertile lemma prominently spreading-pilose on margins and keel; South America C. sesquiflora, p. 104 Sterile floret cylindrical to naiTowlv turbinate, inflated, ca 2 mm long; fertile lemma margins and keel appressed-pubescent; Africa C. ametht/stca, p. 106

52. Fertile lemma broadly elliptic, prominently long-ciliate along most of the margin, the cilia spreading at nearly right angles 53 Fertile lemma margins glabrous or with appressed to widely spreading hairs near the apex, but never along the lower margins 54

53. Fertile lemma more than 3.5 mm long, gibbous; awn definitely subapical; Ar- gentina C. castilloniana. p. 69 Fertile lemma less than 3 mm long, elliptic, not gibbous; arising only slightly subapi-

cally; Florida, Bahama Islands, Jamaica, South America . C. chmdyana, p. .34

54. Plants producing cleistogamous underground spikelets at the ends of thin rhizomes; southwestern United States, Mexico, Central America, and V'cne- zuela C. chloridea, p. 74 Plants not producing cleistogamous underground spikelets 55

55. Spikes naked 2 to 5 cm from the liase; endemic to eastern Texas; rare, possibly extinct C. texemis, p. 103 Spikes floriferous to near the base 56

56. Spikes borne in two or more verticils, these usually well separated. (See also C. virgata which may have two verticils, closely inserted.) 57 Spikes borne in a single terminal whorl 61

57. Sterile floret greatly reduced, usually less than 1 mm long; annuals; widely distributed in American tropics; Hawaii C. racliata, p. 110 Sterile floret longer than 1 mm, well developed; perennials, tufted or stoloniferous 58

58. Upper margins of fertile lemma with a prominent tuft of spreading white hairs, these usually longer than 1 mm; side of fertile lemma grooved, the groove

usually with many glandular hairs ( though these may be somewhat obscure or, rarely, absent); Brazil Ascension Island C. orthonoton, p. 69 Upper margin of fertile lemma scabrous or appressed short-pilose; side of fertile lemma without a groove or glandular hairs 59

59. Fertile lemma 2.0 to 3.5 mm long; widely distributed in central and southcentral United States C. verticiUuta, p. 93

(NOTE: Various introgressants of Chloris verticillata, C. cticuUata, and C. ondropof^o- noides may key to this point. See full discussion of this problem in the text.) Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 21

Fertile lemma longer than 3.8 mm 60

60. A\\'n of fertile lemma longer than 9 mm long; South America C. halophila, p. 117 Awn of fertile lemma shorter than 5 mm long; Baja California, Mex- ico C. hrandeiiei, p. 72

61. Fertile lemma with a dense tuft of spreading hairs on the upper margins, the longest usually more than 1.5 mm long (see also C. aristata and C. cubensis, which occasionally have rather long hairs near the fertile lemma apex) ^ 62 Fertile lemma without a dense tuft of long spreading hairs on the upper margins, though often with shorter (generall)' less than 1 mm), appressed hairs, occasionally scabrous or glabrous 64

62. Fertile lemma with a shallow lateral groove 63

Fertile lemma without a lateral groove; southern Africa C. mossamhicensis, p. 123

63. Annual plants; fertile lemma usually prominently carinate, gibbous; lateral groove of fertile lemma glabrous or appressed-pilose, not glandular; widely distributed C. virgata, p. 60 Perennial, stoloniferous plants; fertile lemm.x rounded on the back, neither carinate nor gibbous; lateral groove of fertile lemma usually with many glandular hairs (though these are sometimes obscure, or, rarely, absent); Brazil, Ascension Island C. orthonoton, p. 69

64. Spikelets bright green to olive green; Ethiopia, Somali Republic

-. C. soinalensis, p. 109 Spikelets pale brown, tawny to black, but not greenish, at least at maturity 65

65. Fertile lemma usually less than 2 mm long; Phillippini' Islands C. dementis, p. 109 Fertile lemma more than 2 mm long 66

66. Spikes naked 2 to 5 cm from the base; endemic to eastern Texas; rare, possibly extinct C. texensis, p. 103 Spikes floriferous to near the base 67

67. Sterile floret more than 0.6 mm wide, often nearly 1 mm wide 68 Sterile floret less than 0.6 mm wide 70

68. Upper fertile lemma margins densely ciliate or spreading-pilose, the margins not

inroUed; Central America . C. aristata, p. 70 Upper fertile lemma margins glabrous, scabrous, or appressed-pilose, the margins inrolled, especially below 69

69. Margins of fertile lemma glabrous or sparsely pilose; fertile lemma apex generally acute, occasionally somewhat rounded; fertile lemma tawny to purple tinged, seldom dark brown or black; Australia C. ventricosa. p. 76 Margins of fertile lemma appressed-pilose; fertile lemma apex broadly obtuse to rounded; fertile lemma dark bro\vn to black at maturity. Australia C. truncata, p. 78

70. Awn of fertile floret usually less than 5.5 mm long; southern Texas, northeastern Mexico C. andropogonoides, p. 95 Awn of fertile floret usually more than 6 mm long 71

71. Sterile floret less than 1 mm long 72

Sterile floret more than 1 mm long ; 74

72. Culms slender, wiry; blades filiform or narrow, less than 1 mm wide (frequently less than 0.5 mm wide); Cuba, Jamaica, Hispaniola C. ekmanii, p. 119 Culms stout, not wiry; blades (at least the wider) 5 to 10 mm wide 73 ) )

22 Bricham Young University Science Bulletin

73. Awn of fertile lemma 6 to 13 mm long; fertile lemma margins sparsely pilose; leaf blade apex acute to subacute; widely distributed in American tropics; Hawaii C. racliata, p. 110 Awn of fertile lemma 10 to 45 mm long; fertile lemma margin scabrous, rarely sparsely pilose; leaf blade apex obtuse, rarely acute; South America, Africa C. pipiothrix, p. 114

74. Culms slender, wiry; blades filiform or very narrow, usually less than 1 mm wide, frequently less than 0.5 mm wide; Cuba, Hispaniola, Jamaica C. ekmanii, p. 119

Culms stout, not wiry; blades, at least the larger, generally broader than 1.5 mm .-. 75

75. Blades usually densely pubescent on both upper and lower surfaces; awn of fertile lemma 20 to 24 mm long; sterile floret narrowly cylindrical, 1.6 to 1.9 mm long, 0.2 to 0.3 mm wide; Cuba C. arenaria, p. 119 Blades usually glabrous or scabrous, occasionally sparsely pilose near the base; awn of fertile lemma less than 13 mm long; sterile floret comparatively broader, 0.3 to 0.6 mm wide, 0.7 to 1.9 mm long 76

76. Spikes strongly divergent from the vertical axis, becoming horizontally spreading or even somewhat reflexed at maturity; keel of fertile lemma glabrous, occasionally very sparsely appressed-pubescent; longest callus hairs 0.3 to 0.8 mm long; marginal hairs of fertile lemma strongly appressed, shorter than 0.5 mm long; Caribbean Islands, Cuba, Haiti, Jamaica C. sagracaiui, p. 121 Spikes not divergent more than 45° from the vertical axis; keel of fertile lemma usually prominently appressed-pubescent, occasionally sparsely so, rarely glabrous; longest hairs of the callus I.O to 1.2 mm long; marginal hairs of the lemma somewhat spreading, especially toward the apex, the longest usually 1 mm or longer; Caribbean Islands, Cuba, Haiti, Jamaica .C. cuhemis, p. 123

I. CHLORIS ROXBURGHIANA Schultes, spikelets densely imbricate, ca 17 per cm of the Mantissa 2:3.39. 1824. (Based on Chloris rachis length; glumes narrowly lanceolate, gla- pohjstachija Roxburgh. See explanatory note brous except for the slightly scabrous midrib; in text.) Fig. 5, A-E. first glume 0.9 to 1.0 mm long, ca 0.1 mm wide;

_, -- second glume 1.9 to 2.2 long, ca 0.2 . mm , mm ^, , , , , Chlons pohjstochya Roxburgh Hortus ^^j^^. ^^^j,^. ,^^^,^^^^ ^^ ^^ 2.I mm long, ca 0.4 Bengalensis 82. 1814. Nomen nudum. Non ^^^ ^^.^^ ^^^^^^^ ^^ ^^^.^^.^ ^,^^„^^ ,,^,.^^^f^^, ^^,^^_ Lagasca, IHlb. ^^^^ glabrous to sparsely ciliate, especially Chloris pohjstacluja Roxburgh, Flora In- above; sides sparsely appressed-pilose to gla- dica 1:332. 1820. Non Lagasca, 1816. (See b^ous, apex acute, awn 10 to 15 mm long; sterile explanatory note in text. florets usually two (occasionally three) gla- Chloris myriostachija Hochstetter, Flora brous, shaped much like the fertile lemma, lower 38:204. 1855. sterile floret 0.7 to 1.0 mm long, 0.2 to 0.3 mm (HOLOTYPE: "Hb. abyss. Buch. nr. 1416," wide, upper floret(s) progressively smaller, awn not seen, but description is of this taxon. of lower sterile floret ca 10 mm long; caryopsis ca long, 0.4 wide, ellipsoidal, trigo- Chloris myriostachija viu. minor Chiovenda 1.1 mm mm nous; chromosome number 2n 20. in Pirotta, Annuario Reale 1st. Bot. Roma = Only C/i/on* paniculafa \ms as large a num- 8:54. 1903. (Description is of a small var- ^^^ °^ spikes as C. roxburghiana. These two are iant of the species ) most easily differentiated by the number of Perennial to 125 cm tall, arising from a sterile florets, C. roxburghiana always having at strongly stoloniferous, woody base; sheaths least two, while C. paniculata always has one. glabrous below, often pilose above; ligules Chh)ris roxburghiana is a stoloniferous perennial pilose; blades up to 30 cm long, 5 to 6 mm found in Africa and India. Chloris paniculata wide, glabrous or scabrous; spikes many (usu- has a unique growth habit, with well-developed, ally 50 or more) attached along a short axis at short, vertical rhizomes bearing the frayed rem- the culm apex and fonning a dense inflores- nants of previous years' growth as well as many eence up to 15 cm long and 4 to 6 cm wide; long, arching, basal leaves; it is endemic to Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 23

spikelet, partly Fig. 5. C/iforis roxhurghiana and C. paniculata. (A-E) C. roxburghiana. (A) habit, x 1/5; (B) dissected, x 15; (C,D) lower and middle sterile florets, respectively, x 10; (E) caryopsis, x 10. (F,G) C. paniculata. (F) habit, x 1/5; (G) spikelet, partly dissected, x 15. 24 Bricham Young University Science Bulletin

Cocos Island, which is off the western coast of Panama. Chloris mijriostachya has been used as a name for this species for many years, especially in African floras. Part of this erroneous usage can be traced to the difficulty of typifying the Roxburgh names. William Roxburgh first used C. pohjstachija in 1814 (Hortus Bengalensis

82.), publishing it without a description. Iden- tification of this name rests with its subsequent publication by Roxburgh (Flora Indica 1:332. 1820.), who included a fairly complete description. By that date, however, Lagasca y Segura (1816) had published the same name for a Mexican species (sec under C. suhimiticu). Shortly afterward, Schultes (Mantissa 2:.3.39. 1824.) published a new name, C. roxhurghkina. based upon C pohjstaclu/a Roxburgh. No plant specimen that mav be connected positively with Fig. 6. Distribution of Cliloris mxl>urnhiana. Inset Roxburgh's name can be located; however, im- A: India. portant clues may be obtained from the Rox- burgh drawings at Kew and from the descrip- Wei Wei 50 mi N of Kapenguria, Trelawnij AB tions given b>' Roxburgh, both those of the 4,325 (K,UC). MALAGASY REPUBLIC: Mom- published Flora Indica as well as the copy of bas, anon. (K). MALAWI: Lilongwe Dtr., Salu- manuscript of that work in the Li- the Kew beni 428 (K). MOZAMBIQUE: Louren^o brary. The drawings, labelled "C. pohjstachija," Marques Dtr., between Moambas and Sabie, show an inflorescence of sixteen spikes arranged Torre 2,237 (K). REPUBLIC OF THE along a short common axis, and a series of dis- CONGO: Plaine du Lac Edouard, Pare Nation- sected spikelets showing broadly ovate-elliptic ale Albert, Louis 4,787 (K, US). SOMALI RE- fertile lemmas and two similar sterile florets. PUBLIC: Boundaiy Pillar 93, 8°37'N, 45°9'E, While the inflorescence drawing does not Gillett 4.197 (K, US). SOUTHERN RHODE- portray accurately the more t\pieal specimens SIA: 30-60 mi S of Ft. Victoria, Rodin 4,251 (K, of this taxon from Africa ( though Indian col- UC, US); Sabi River Valley, Melsetter, Whellan lections bearing a strong resemblance are com- 1,021 (K). SUDAN: Equatoria Prov., Kidepo mon) the spikelet drawings leave no question resthouse, Mijers 11,236 (K). TANZANIA: as to the identity of Roxburgh's Chloris pohj- Tanga Dtr., Kange Estate, Faulkner 832 (K); stachija and thus, indirectly, C. roxhurghiana SW of Umba River, Kivingo, Greenuaij 1,996 Schultes. (BM, K); Lake Manyara National Park, Green- Chloris roxburghiana is common (sometimes wai/ and Kirrika 11,089 (K, US). UGANDA: dominant) in grasslands, brush, and disturbed K;inamugit, Eggeling E 2.946 ( K, US ) ; Turkana areas, often on sandy loams from low to me- Dtr., Oropoi Valley, Liebenberg 1,776 (K). dium elevations (Fig. 6). Associates include UNION OF SOUTH AFRICA: Kruger Natiimal Cenchrus sp., Pennisettim sp., Sporobohis sp.. Park, between Punda Maria and Pafuri (Mo- Acacia mellifera, Blepharis sp.. Commiphora sp., zamljique), Godfrey SH-1.729 ( K, US); be- and Euphorbia spinescetu. tween Beauty and Ellisras, Werdemann and Representative specimens examined: AN- Oberdicck 1,801 (K). GOLA: Mossamedes Dtr., Camucuio, Azancot de Menezes 363 (K). Maklautsi- BOTSWANA: 2. CHLORIS PANICULATA Scribncr in Rob- Rivers, s.n., Shashi Palapye, de Beer, 9 May inson, Proc. Amer. Acad. Arts Sci. 38:262. 1957 (K). ETHIOPIA: Harrar Prov., 40° 39' E, 1902. (HOLOTYPE: "Nuez Isl . . . an islet 10° 10' N, Burger 2,899 (K); Sidamo Prov., 20 near Cocos Isl.: Snodgrass and Heller, no. mi of Moyale (Kenya), Moonei/ 7,421 (K). N 968;" in GH. Not seen, but description is

INDIA: Nilgiri Dtr., ' Kitagin Ghat, Fischer clear.) Fig. 5, F and G. 2,086 (K). KENYA: Kibwezi Plains, Balhj 1,329, 8,090 (K); 25 mi SSW of Kitui, Bogdan AB Perennial to 70 cm tall, arising from an 5,126 (K,US); Lugard's Falls Road, Tsavo Na- upright, stout, underground stem bearing many tional Park, Greenwatj and Kanuri 12,636 (K); rootlets and shreds of leaf sheaths; sheaths Biological Seiiies, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae) 25 glabrous; ligule lacking; blades very long and and are the species as commonlv under- narrow, up to 50 cm long and 5 mm wide, arch- stood. The description and illustration ac- ing, glabrous except for the scabrous margins; companying the original description leave inflorescence paniclelike, made up of at least no (juestion as to the identity of the taxon,

50 spikes racemosely arranged on the upper 10 however. ) Fig. 7, A-D. to the culm; spikes about 5 cm long 12 cm of Chloris accumbens Hackel ex Arechavaleta, inflorescence, becoming pro- at lower part of Anales Mus. Nac. Montevideo 5:391. 1896. gressively shorter near the tip; glumes narrow Nomen nudum, pro sijn. C. berroi. to broadly lanceolate, glabrous except for the scabrous midnerve; first glume ca 1.2 mm long, Perennial from a fibrous root system, tufted, 0.2 mm wide; second glume ca .3 mm long, 0.4 15 to 80 cm tall; sheaths glabrous; ligule ciliate; mm wide; fertile lemma ca 2.6 mm long, 0.6 blades narrow, 3 to 15 cm long, 1.5 to 2.0 mm mm wide, broadly lanceolate, glabrous except wide, glabrous or sparsely pilose near the base; for the prominently bearded callus and scab- spikes two to four, 3 to 12 cm long, tightly rous keel, apex ± obtuse, awn 2.5 to 2.8 mm appressed, adherent, and forming a narrow, long; sterile floret one, ca 1.5 mm long, 0.3 mm cylindrical, spikelike inflorescence; spikelets wide, glabrous, awn 1.5 to 1.8 mm long. densely imbricate, 9 to 12 per cm of the scab- Only Chloris roxhurghiana has as many rous-hispid rachis; glumes glabrous, lanceolate; spikes as C. panicithta. Differences between first ghnne 1.5 to 1.6 mm long, ca 0.3 mm wide; these two species have been discussed under C second glume 2.1 to 2.6 mm long, 0.3 to 0.6 mm roxlmrghiana. wide; fertile lemma ovate, 2.7 to 3.5 mm long, to 1.2 wide, margins and keel ciliate Chloris paniculata is an endemic known 0.8 mm hairs to 2 long, awn 2.7 to 3.4 mm only from Cocos Island and its associated islets with up mm three, up to 1.9 —an archipelago lying about 300 miles south- long; sterile florets usually mm lowennost awned, upper unawned, gla- west of Costa Rica, to whom it belongs. Rob- long, long, 0.5 to 0.7 inson (1902) and Stewart (1912) both reported brous; caryopsis 1.2 to 1.8 mm wide, obovoid, trigonous; chromosome it as being abundant on rocky cliffs near the mm coast, both on the main island and on the islets. number 2n^40. Chloris berroi can be distinguished from Most of the vegetation of the island area is cylindrical made up of common Central American species all other species of the genus by its interweaving or pantropical weeds. Only eight endemics in inflorescence; this results from the adjacent spikes and, more a total of about a hundred species were re- of the spikelets on rachises. ported by Stewart. Fosberg and Klawe (1966) especially, from the hispid hairs of the past maturity, also report C. paniculata from Cocos Island. The spikes remain interconnected except for the very tips, and may be separated I cannot suggest a species in Chloris. nor from one another only by actually tearing them for that matter anv other genus in the Chlori- apart. deae, that might be said to be closely related Individual spikelets of Chloris berroi are to C paniculata. The previously discussed re- very similar to the other species in which the semblance to C. roxhurghiana is superficial, for lemnias have ciliate margins, especially those of there are manv spikelet details in which they C. ciliata and C. clanchjana. Other than the differ. Certain species of Leptochloa have a inflorescrnce differences previously mentioned, similar aspect because of the large number of there seem to be no other characteristics sepa- spikes. Leptochloa, however, has spikelets which rating them. unifonnh' have a larger number of fertile flor- Chloris berroi is restricted to the Rio de la ets, and the sterile floret is not as elaborated in of Argentina and Uruguay (Fig. structure and size as is generally the case in Plata region reported it to be abun- Chloris. 8), where Parodi ( 1919) Specimens examined: COCOS ISLAND: ex- dant in the campo. posed rocky cliffs near the ocean, Stewart 260 Specimens examined: ARGENTINA: Prov. (F,MO,NY,US). Buenos Aires; La Plata, Spegarrini 1,433 (NY); Pergamino, Niedfeld 38 (US). Prov. Entre Rios: Gualeguaychu, 3. CHLORIS BERROI Arechavaleta, Anales camino de Puerto Constanza a Mus. Nac. Montevideo 5:388. 1896. Burkart 10,529 (US); Dep. Federacion: Estan- (HOLOTYPE: "Berro, Uruguay, Estancia cia "Buena Esperanza," Pedersen 6,254 (US). Montevideo, de Soriano, Estancia de Vera." This speci- URUGUAY: Dep. Canelones: Mansa- men was not seen, though specimens at K! Sello s.n. (MO). Dep. Florida: Arroyo and US! are labeled essentiallv the same viUagra, Gallinal 5,790 (MO). Dep. Soriano, 26 Bricham Young UNivERSiTi' Science Bulletin

sterile Fig. 7, Chloris berroi and C. ciliata. (A-D) C. benoi. (A) h;ibit.. x 1/5; (B) spikelet, x 10; (C) lower floret, X 15; (D) upper sterile floret, x 15. (E-I) C. ciliata. (E) habit, x 1/4; (F) spikelet, x 10; (G) sterile florets, x 15; (H) upper sterile floret, x 15; (I) caryopsis, x 10. . ) )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gbamineae) 27

Chloris nashii Heller, Muhlenbergia 5:120. 1909. (Based on Chloris ciliata var. texana Vasey.

Perennial 25 to 60 cm tall, tufted, erect; sheaths glabrous, hgule absent or reduced to a short ciliate crown (Northern Hemisphere) or densely and conspicuously pilose (Southern Hemisphere); blades 10 to 20 cm long, ca 5 mm wide, long-acuminate, glabrous or scabrous; spikes three to five (rarely si.x or seven), 3.5 to 6.0 cm long (occasionally up to 8 cm) somewhat flexuous and spreading, glumes narrowly lanceolate, glabrous except for the scabrous midrib, becoming hyaline near the margins; first glume 1.3 to 1.7 mm long, 0.2 to 0.4 mm wide; second glume 2.0 to 2.5 mm long, 0.2 to 0.4 mm wide; fertile lemma 1.8 to 2.8 mm long, 0.8 to 1.1 mm wide, strongly flattened, elliptic, margins and keel strongly ciliate, the cilia 0.5 to 1.5 mm long, awn 0.9 to 2.7 mm long; sterile florets two, lowermost enclosing the upper; lower sterile floret 1.3 to 1.8 mm long, 0.8 to 1.8 mm wide, truncate, glabrous, awn 0.9 to 1.4 mm long, upper floret similar but smaller, 0.8 to 1.1 mm long, 0.9 to 1.2 mm wide, membranous, awnless; caryopsis ca 1.4 mm long, ca 0.7 mm

Fig. 8. DistribuKon of Chloric herroi (southern South wide, obovoid to ellipsoid; chromosome number America ) 2n=40. The species in this complex (Chloris herroi, Berw 6,385 (US). Dtp. Tacuarembo, Herter C. canterai, C. ciliata, C. dandijana, and C. 587a (MO). lamproparia) are characterized by elliptic to lanceolate lemmas, the margins of which are 4. CHLORIS CILIATA Swartz, Prodr. Veg. densely ciliate, usually along the entire length. Ind. Occ. 25. 17S8. (HOLOTYPE: Not Chloris herroi is easily distinguished, for it has seen. Swartz's original description in 1788 is spikes so interconnected by hispid hairs on the

clear; his amplification in 1797 leaves no rachises that it appears to have but a single doubt.) Fig. 7, E-I. spike. The remaining species, however, present a more complex pattern. While most collections Anchopoiion ptihescens Aiton, Hortns Ke- will fit a given species quite well, certain geo- wensis 3:423. 1789. (Based on Chloris cil- graphic areas (most often in South America) iata Swartz.) may have individuals which are somewhat inter- Ctinodon ciliatus (Swartz) Raspail, Ann. mediate with respect to the characteristics. Fig- Sci. Nat. Bot. (Paris) 5:303. 1825. (Based ure 9 illustrates selected features of a large on Chloris ciliata Swartz. number of specimens chosen at random from Chloris propinqua Steudel, Syn. PI. Glum. throughout the ranges of the species. 1:204. 1854. (HOLOTYPE: "Duchai.ssing While this diagram reveals no distinct sub- legit in Guadeloupe." A fragment in US!, populations, certain variation trends may be P, is badly from the Steudel specimen in discerned. One trend is characterized by indi- shattered but would appear to be Chloris viduals with relatively short lemmas, short Chloris ciliata. The description seems to fit marginal cilia on the lemmas, short awns, ciliata Swartz.) and' few spikes (six or less). These are the Chloris ciliata var. texana Vasev, U. S. Dept. plants commonly called C. ciliata and are repre- Agr. Div. Bot. Bull. 121. pi. 30. 1890. sented bv the open circles situated roughly in (HOLOTYPE: "Near Brownsville" US!) the lower left quadrant of the diagram. A sec- circles Chloris texana (Vasey) Nash, Bull. Torrev ond trend, represented mostly by open the upper Bot. Club 25:441. 1898. Based on Chloris with an upward line, is shown in had longer ciliata var. texana Vasey.) left quadrant. These collections 28 Bricham Young University Science Bulletin

3.5- t CO

46 A < 3.0- 9 f n: 6 4 44 4 4 < 44 44 4 4 o 4 o 4 2.5- 44 44444 po 44 oo^ i t 444444 A A LlI o4 444 44 ooo A A 4444 44 4 o o A 44444 44 2.0 4 4 4 44AA44o 4 4 4 4 44o44 44 44 44o4444 4 o o 4 44 4 o en 1.5 44 44oA4 4 o o o o 4 o < 4 o o o oo O NEW WORLD o oooooo o ooo • OLD WORLD o ooo oo o o Ll 6 4 SPIKES 7 OR MORE 1.0- o oooo ooo o oo o o O o ooo o oo O • SPIKES 6 OR LESS n o o ooo o oooo COMBINED LENGTHS OF AWNS OF I— STERILE AND FERTILE FLORETS: CD 0.5- O • MORE THAN 7.5 MM z O •LESS THAN 7.4 MM UJ

1.5 2.0 2.5 3.0 3.5 4.0 4.5

LEMMA LENGTH IN MM.

Fig. 9. Comparison of various morphological and geographical characteristics of Clitoris canterai var. canterai and var. grandiflora, C. ciliata, C. dandyana, and C. lamproparia.

lemmas and marginal hairs, a larger number of tics employed. However, it does not seem that spikes (often more than 20), but short awns; merging all of these into a single species is these are representative of plants commonly justified, for discernible variation trends are called C. damhjana. Extremes of these popula- evident. Thus four species are recognized, each tions are easily separated, though individual centering in one of the four trends discussed plants may diverge considerably from the above. "average" member of each trend. Certain other characteristics were of value The third and fourth trends overlap more in differentiating elements within these taxa, but with respect to the morphological characteris- did not clarify the relationships between them; tics analyzed and are most easily separated these are discussed individually under the ap- geographically. These two groups are inter- propriate species. All populations are deserving mingled in the upper right quadrant and differ of further study, especially in areas of sympatry. from both of the pre\ious trends in length of Mass collections as well as cytological exam- the lemma and of its marginal cilia. Within this ination and genetic analysis ^\ill be especially group, the New World collections are perennials valuable. with relatively short lemmas, marginal cilia, and Tabic 7 compares all of the species in this awns. These are the plants commonlv named complex witli respect to a variety of characters.

C. canterai. The remaining specimens are Old Figures 7, 11, 1.3, and 15 should also be used World annuals and have, generally, longer for comparison. lemmas, marginal cilia, and awns; these are There aie two recognizable geographic vari- commonly called C lamproparia. ants within Chloris ciliata: (1) a North Ameri- Well-defined discontinuities between the can population with nonciliate ligules and (2) various populations have not been demon- a South American group with prominently cil- strated, at least on the basis of the characteris- iate ligules. No other differences could be Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (CfRamineae) 29

Table 7. Comparison of Chloris berroi, C. canterai, C. ciliata, C. dandyana, and C. lamproparia. 30 Bricham Young University Science Bulletin

Leonard 12,934 (US). JAMAICA: Middlesex Annuals 10 to 60 cm tall, erect to decum- Co.: Clarendon Parish, Inverness, Harris 12,755 bent; stems usually branched at the base, these (F, MO, NY, US); Manchester Parish, Harris sometimes rooting at the lower nodes; sheaths 12,689 (F, MO, NY, US); Cornwall Co., St. glabrous; ligule a short ciliate crown; blades James Parish, Montego Bay, Hitchcock 9,671 5 to 20 cm long, 3 to 5 mm wide, sparsely pus- (US). LEEWARD ISLANDS: Antigua: Dead- tulose-pilose above, especially near the base, sands Bay, Box 31 (US); Guadeloupe Island, glabrous to scabrous below; spikes one to four Duss 31,587 (NY, US); St. Barthelemy, Questel (often two) ca 4 cm long, appressed or only 337 (US). MEXICO: Coahuila: Castaiios, slightly divergent; spikelets densely inserted oh Reeder, Reeder and Soderstrom 3,270 (US); the rachis, usually divergent, ca 10 per cm of Nuevo Leon: Monterey, Kenoijer 315 (MO); the long scabrous-hispidulous rachis; glumes Oaxaca: Toniellin, Hitchcock 6,220 (US); San lanceolate-ovate, thin, brown, glabrous (except Luis Potosi, 7.5 mi N of El Naranjo, McGregor for the scabrous midnerve); first glume 3.9 to et al. 813 (US); Tamaulipas: 22 km S of Vic- 4.4 mm long, 0.6 to 0.9 mm wide, apex acute; toria, Martinez and Lmjando F-2,283 (TEX, second glume 6.5 to 8.0 mm long, ca 1 mm wide, US); Yucatan: Chichen-Itza, Swallen 2,409 apex acute to acuminate; fertile lemma 3.2 to (US); Tizimin, Swallen 2,491 (US); Uxmal, 4.2 mm long, 0.8 to 1.7 mm wide, broadly ovate Swallen 2,612 (US); Peto, Swallen 2,672 (US); to elliptic, dark brown, keel appressed-pubes- Merida, Souza-Novelo 15 (US). PUERTO cent, margins with spreading ciliate hairs, 2 to RICO: Ponce, A. Chase 6,495 (NY, US). 3 mm long, internerves glabrous, apex acumin- SANTO DOMINGO: Prov. Santo Domingo: ate, awn 3.5 to 6.0 mm long; sterile florets two Llano Costero, Ekman 12,486 (US); Barahona, or three; lowest sterile floret 1.3 to 1.6 mm Fuertes 308 (US). UNITED STATES: New long, 1.0 to 1.5 mm wide, triangular, apex trun- York: Yonkers, mill yard, Nash s.n., 24 July 1898 cate, awn 2.3 to 4.7 mm long, upper florets (NY). Texas: Aransas Co., Rockport, Swallen similar in shape, but progressively smaller, un- 10,297 (US); Bee Co., Pettus, Tharp 43,071 awned; caryopsis 1.5 to 2.0 mm long, 0.8 to 1.0 (TEX); Bexar Co., 20 mi S of San Antonio, mm wide, ovoid to obovoid. Silveus 137 (TEX); Calhoun Co., 2 mi SW of Chloris lamproparia is the only species in Port Lavaca, Rowell and Webster 2,117 (TEX); the group with ciliate margins on the lemma Cameron Co., Laguna Atascosa Refuge, Fleet- that is distributed in the Old World, and it is wood 3,800 (TEX); De Witt Co., near Edgar, also the only annual. It is most similar to C. Owens 155 (US); jim Wells Co., 6 mi W of Alice, canterai, differing from it in having larger spike- Gotdd and Kapadia 8,854 (UC); Karnes Co., lets, though occasional specimens are inter- 5 mi S of Kennedy, Emery 773 (TEX); Kleberg mediate in spikelet size and can be assigned Co., King Ranch, M. C. Johnston 541,669 (TEX); most easily by their geographic origin and Matagordo Co., near Palacios, Rogers 6,607 habit. (TEX); San Patricio Co., 1 mi SE of Sinton, Chloris lamproparia is distributed across cen- Gould 9,903 (TEX); Starr Co., Fort Ringgold, tral Africa (Fig. 12), where it is reported to be E of Rio Grande City, Tharp and York 51-84 common in wet grasslands and savannas. (TEX); Travis Co., Zilker Park, W. V. Brown Representative specimens examined: CAM- s.n., 6 Oct 1957 (TEX); Wharton Co., 12 mi EROUN: Tchabaol, 45 km ENE of Maroua, N of El Campo, W. V. Brown 3,265 (TEX); Letouzeij 6,473 (K). MALI: Bore, Dcmange Willacy Co., Sauz Ranch, M. C. Johnston and 14/1.957 (K). NIGERIA: Domaturu, De Leeiiw Davis, 5,319.84 (TEX). URUGUAY: Dep. Mon- 1,097a (K); Zaria Dtr„ Tat/lor H. 2,624/32 (K). tevideo, Herter 78,127 (US). VIRGIN IS- SUDAN: Darfur Prov.: 3 km NE of Kes, Jack- LANDS: St. Croix: Bassin Yard, Ricksecker 42 son 4,040 (K); Jcbel Marra, Zalingei, Wickens (F, MO, NY, UC, US). WINDWARD IS- 2,304 (K); Equatoria Prov., 13 mi SW of Ka- LANDS: Barbados, Hitchcock 16,514 (US); poeta. Peers s.n., 29 Aug 1953 (K); Kordofan Grenada: Richmond Hill, Broadway 1.869 Prov.: El Muglad, Andretos 3,063 (K); near (NY); Martinique, Du.ss 1,273 (MO, NY. US). Hamadi, on El Obeid-Dilling road, Harrison 901 (K). TANZANIA: Tumba Rukwa, Michel- more Cl,557 (K). UGANDA: Moroto, Wilson 5. CHLORIS LAMPROPARIA Stapf in Che- 15 (K). VOLTA: Ouagadougou, Chev- valier, Bull. Bot. Soc. France Mem. IV. UPPER 24,674 8:220. 1912. (HOLOTYPE: Chad, "Bag- alier (K). uimii, emplacement de la ville de Massenia, 25-31 Aout 1903, no. 9633 bis." P! ISO- 6a. CHLORIS CANTERAI Arechavaleta var. TYPE: K!) Fig. 11. CANTERAI. Anales Mus. Nac. Montevideo )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gbamineae) 31

r~\

Fig. 11. Chloris lamproparia. (A) habit, x 1/4; (B) spilcelet, partly dissected; (C) lemma with one side removed, showing palea and sterile floret within; (D) sterile florets; (E-F) upper sterile florets; (G) caryopsis. (B-G) \ 10.

rolled 5:385. 1896. (HOLOTYPE: "Anderson . . . up to 7 mm long; blades narrow, often Montevideo." Not seen, but original descrip- and appearing filiform, 1 to 5 mm wide, up to tion clearly identifies the plant.) Fig. 13, 25 cm long, glabrous to slightly scabrous, occa- A, C-J. sionally sparsely long-pilose; spikes two to nine Chloris parodiana Rosengurtt and Izaguirre, (often three to six), 4 to 14 cm long, usually Bol. Soc. Argentina Bot. 12:128. 1968. more than 7 cm long, erect to curving; spikelets densely imbricate, appressed, light to me- (HOLOTYPE: "Paraguay, Paraguari . . . Rosengurtt B-5,979, MVFA." Not seen, but diimi brown, ca 11 per cm of the scabrous description and illustration are clear. rachis; first glume 1.6 to 2.4 mm long, 0.5 to 0.6 mm wide, narrowly lanceolate, markedly Perennial up to 1 m tall, tufted and erect; inequilateral; second glume 2.3 to 3.8 mm long, sheaths glabrous; ligule long-ciliate, with cilia 0.4 to 0.6 mm wide, narrowly lanceolate, equi- 32 Bbicham Younc University Science Bulletin

While one immediately suspects that an environ- mental factor may be influential, or that the cespitose individuals are merely more heavily

grazed, the morphological disccmtinuity is so well defined that it is felt that the population is worthy of being recognized taxonomically. Rosengurtt and Izaguirre de Artucio (1968) have given this population the name Chloris grandiflora. I feel that this overemphasizes the differences, however, and I have consequently

reduced it to a varietal rank. In addition, Rosengurtt segregated a second species from this variable population, naming it Chloris parodiana. It is based largely upon

Fig. 12. Distribution of Chloris lamproparia. the relative length of lemma cilia. I do not feel that the differences emphasized are stable, at lateral or inequilateral; fertile lemma narrowly least on the basis of the data summarized in ovate, 2.7 to 3.7 mm long, usually more than 3 Fig. 9; and I, consequently, have treated it as a mm, 0.6 to 1.1 mm wide, ovate, callus bearded, synonym under var. canterai. Further studies densely ciliate on keel and margins, marginal of a more experimental nature will be needed cilia 1.5 to 3.0 mm long, intcmerves glabrous, to detemiine the validity of all of these taxa. awn 2.4 to 5.5 mm long; sterile florets two While some specimens of Chloris canterai (sometimes three), similar, turbinate-flattened; and C. lamproparia may be difficult to separate, lower sterile floret 1.1 to 1.8 mm long, about there are ample differences. Chloris canterai is half as wide, glabrous; awn 1.5 to 3.5 mm long; a tufted. South American perennial (Fig. 13) upper sterile floret(s) similar to lower in shape, with relatively smaller and narrower spikelets but smaller, about 0.5 mm long, awnless; car- and more spikes per inflorescence than C. yopsis ovoid-ellipsoid, 1.3 to 2.0 mm long, 0.8 to lamproparia, which is an African annual (Fig. 0.9 mm wide; chromosome number 2n=36. 11).

Chloris canterai var. canterai is restricted to northeastern Argentina, extreme southern Brazil, 6b. CHLORIS CANTERAI Arechavaleta var. GRANDIFLORA (Rosengurtt & Izaguirre Paraguay, and Uruguay ( Fig. 13, shaded circles), where it is reported as occurring on moist de Artucio) Anderson, comb. nov. Fig. 13, soils on the campo. Chloris canterai var. grandi- B-J. Chloris <:,ran(UfIora Rosengurtt and Iza- on is guirre de Artucio, Bol. Soc. Argentina Bot. flora, the other hand, found on dry sites, 12:127. 1968. (HOLOTYPE; "Uruguay, often rocky areas, most commonly in Argentina. A single specimen from BM! (collected by dcp. Florida . . . Gallinal et Rosengurtt, PE- Gardner in 1839) is labeled "Banks of the Rio 5,790 . . . MUFA." Not seen, but original description and illustration are clear.) Gurgueia, Piaui," an area of Brazil far to the north of any existing collections and probably Perennial, densely cespitose, 5 to 30 cm tall represents a labeling error. All North American (often about 20 cm) arising from a dense plants arc naturalized introductions. cluster of very narrow, often curved and rolled Representative specimens examined: CHLO- leaves; spikes three to five, ca 3 to 6 cm long; RIS CANTERAI var. CANTERAI. AR- other characteristics as in var. canterai, except GENTINA: Prov. Chaco: Tintina, Parodi 812 chromosome number unknown. (US); Prov. Corrientes: Mercedes, Parodi 6,242 The most marked difference within Chloris (US); Prov. Santa Fe: bajos del Rio Salado, canterai is a vegetative one. A group of speci- Burkart 9,077 (US); Prov. Formosa: Pilcomayo, mens, primarily from Argentina ( Fig. 14, open Morel, 6.569 (US). BRAZIL: Rio Grande do circles) is shorter than usual and has a well- Sul, Sao Gabriel, Rambo 25,567 (US); Uru- defined, cespitose growth habit. No spikelet guaiana, Poste Zootecnico, Stcallen 7,617 (US). distinctions could be found. A few of these PARAGUAY: Dep. Paraguari: Caapucu, Rosen- have been collected in the same precise loca- gurtt 5,405 (US); Dep. Presidentc Hayes: S of tion as specimens of var. canterai; but the Villa Hayes, Rosengurtt 5,631 (US). UNITED collection data indicate that they are found in STATES: Texas: Atascosa Co., 4.5 mi SE of dry, rocky areas as opposed to the lower, wetter Jourdanton, /. C. Johnson 2,044 (TEX); Jack- sites in which var. canterai is typically found. son Co., 4 mi E of Edna, Gould 6,781 (TEX, Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 33

Fig. 13. Chloris canterai var. cantcrai and var. grandiflora. (A) habit, C. canterai var. canterai, x 1/6; (B)

habit, C. cantcrai var. grandiflora, x 1/3; (C) spikelet, x 10; ( D.E ) sterile florets, partially dissected, x 15; showing variation, x 10. (F) uppermost sterile floret, x 15; (G) caryopsis, x 10; { H-J ) outlines of spikelets, ) ) ) ) )

34 Bricham Young University Science Bulletin

ed. 2. 2:1483. 1763. (See discussion of this name below.) Chloris pohjdactijla (Linnaeus) Swartz, Prodr. Veg. Occ. 26. 1788. (Based on An- dropogon polydactijlon Linnaeus. Chloris consanguinea Kunth, Rev. Gram. 1:89. 1829, pubhshed in key; Enum. Pi. 1:264. 1833, republished with full description. (HOLOTYPE: "Rio Janeiro," not seen,

but description is clear.

Chloris elata Desvaux, Opuscules 73. 1831. (HOLOTYPE: "Brasilio," not seen, but de-

scription is clear. Chloris arundinacea Nees ex Steudel, Syn. PI. Glum. 1:207. 1854. (HOLOTYPE:

• =C, CANTERAI VAR. CAMERA 1 "Sello. Brasil." Fragment of type in US! 0=C. CANTERAI VAR, GRANDIFLORA from B.) Chloris harhata (Linnaeus) Nash, Bull. Fig. 14. Distribution of Chloris canterai var. canterai Torrey Bot. Club 25:443. 1898. Non C. har- (dots) and var. grandiflora (open circles) in South America and Te.\as (Inset A). hata ( Linnaeus ) Swartz, 1797. ( Based on Andropogon harhatum Linnaeus, 1759.

UC); 14 mi NE of Port Lavaca, Gould and Chloris pohjdactijla f. stolonifera Parodi, Reeves 8,208 (TEX, UC); La Ward, Tharp et Rcvista Argentina Agron. 20:24. 1953. al. 51-1,603 (TEX); 4 mi SE of Morales, Turner (PARATYPE: US! A stoloniferous form.) 3,276 and Tharp 53-430 (TEX); Milam Co., Perennial to 135 cm tall, usually tufted, Buckholts, Gould 7,692a (TEX); Wharton Co., rarely stoloniferous, erect, arising from a thick 6 mi S of El Campo, Anderson 3,288 (HSC). fibrous root system; sheaths glabrous, becoming URUGUAY: Dep. Cancloncs: Pando, Montaro long-pilose at the ligule; blades up to 45 cm 709 (K); Dep. Cerro Largo: Rio Negro, Gal- long, 1.5 cm wide, usually glabrous with scab- linal et al. PE-5,531 (MO, UC, US); Dep. Flor- rous margins, occasionally scabrous on both ida: Arroyo Mansavillagra, Rosengurtt 5,852 surfaces; spikes four to twenty-eight (usually ( US ) ; Dep. Paysandu, without preci.sc locality, more than eight), flexuous, to 20 cm long, usu- Rosengurtt B-909 (US); Dep. Rocha: Camino ally more or less .spreading, sometimes even de las Indios, Rosengurtt B-6,938 (K); Dep. drooping; spikelets densely imbricate, averaging Salto: without precise location, Del Puerto s.n. 12 per cm of the scabrous to hispid-ciliate in LS74 (US). CHLORLS CANTERAI var. rachis; glumes pale, glabrous, with scabrous to GRANDIFLORA. ARGENTINA: Prov. Buenos short-hispid midnerves; first glume 1.0 to 2.5 Aires: Sierra de la Ventana, Bartlett 20,029 mm long, 0.2 to 0.4 mm wide, ovate-lanceolate; (US); San Nicolas, P«ror// 1,023 (US). Prov. second glume 1.9 to 3.5 mm long, 0.2 to 0.4 Corrientes: Dep. Mercedes, Estancia "Ita mm wide, narrowly lanceolate; fertile lemma caabo," Parocli 6,118 (US). BRAZIL: Rio 1.6 to 2 8 mm long, 0.5 to 0.9 mm wide, Grande do Sul, Uruguaiana, Poste Zootccnico, elliptic, prominently flattened, usually pale Swallen 7,656, 7,697 (US). URUGUAY: Dep. brown, margins prominently ciliate with hairs Colonia: San Juan, Herter 18,789 (US); Dep. at least 1 mm long and often up to 3 mm long, Florida: Arroyo Mansavillagra, Rosengurtt 5,790 keels densely appressed-pilose, awn 1.4 to 4.8 (US). mm long; sterile florets usually two, the upper hidden in the lower; lower sterile floret cvlin- CHLORIS DANDYANA CD. Adams, Phv- drical to narrowly turbinate, 1.0 to 1.6 mm long, tologia 21:408. 1971. (See discussion of this glabrous, awn 1.7 to 4.0 mm long; upper sterile

name below. ) Fig. 15. floret flattened and thin, 0.5 to 0.9 mm long, 0.3 to 0.7 mm wide, unawned; caryopsis ca 1 Andropogon harhatum Linnaeus, Svst. Nat. mm long, 0.5 mm wide, ellipsoidal. ed. 10. 2:1305. 1759. Non A. harhatum Lin- Because of a complicated nomenclatural naeus, 1771. (See discussion of this name problem involving Chloris dandijana and C below. inflata, it is necessary that a complete review Andropogon polydactijlon Linnaeus, Sp. PI. of the history of these taxa be made. BiOLOGiCAi, Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 35

M\miViM 4 ^!;:l^^»»»-s

Fig. 15. Chloris dandyana. (A) Inflorescence, x 1/3; (B) spikelet from specimen with long leimna margin hairs; (C) spikelet from specimen with short lemma margin hairs; (D) sterile florets; (E) caryopsis; (F-J)

outline of series of spikelets, showing variation. ( B-] ) x 10.

In 1759 Linnaeus described Andropogon given for the change. There is no question of barhatum (Syst. Nat. ed. 10. 2:1305), providing Linnaeus's intent, for he copied directly the a brief description and a reference to "Sloane. description accompanying the 1759 publication hist. t. 65. f. 2." The protologue is inconclusive. of Andropogon barbatum and referred to Brown The plate in Sloane is likewise not confirming, and Sloane, this time adding a direct reference but the description of a Jamaican plant accom- to "p. Ill" of the latter work. Reference was panying the plate (p. Ill) would seem to fit also made to the Amoenitates (1760). best the plant described above. Reference will In addition to the changes of names in the be made later in this discussion to a specimen written descriptions, corroborative evidence is labeled Aiidropogon barhatum in the Linnaean obtained from the Linnaean Herbarium, for

Herbarium. there (in the Andropogon folder) is a specimen Linnaeus again described Andropogon bar- of the plant described above with three epithets batum in 1760 (Amoen. Acad., p. 412), chang- written on it: "barbatum," "fasciculatum," and ing and amplifying the description and drop- "polydactylon." The first two names are both ping the reference to Sloane, citing instead lined through; the last is not. Nonetheless, An-

"Brown, jam. 364." The description again is not dropogon polydactylon is clearly a superfluous conclusive, but it would still appear to be the name and is illegitimate. above-described species. In 1771 Linnaeus again used the name An- In 1763 the above names, references, and dropogon barbatum (Mantissa, p. 302), this descriptions were tied together (Sp. PL, ed. 2. time providing a clear description of a plant

2 ; 1483 ) ; but the name of the plant was changed from the East Indies. There is no question to Andropogon polijdactijlon, no reason being about this description; it obviously refers to the .

36 Brigham Young Univebsity Science Bulletin

pantropical weed with inflated, truncate sterile • LEMMA MARGIN QUA MORE THAN florets, commonly called either Chloris barbata, 2.2 MM LONG •-LEMMA MARGIN CILIA LESS THAN C. paraguaijensis, or (as in this study) C. inflata. 2.1 MM LONG • LEMMA LENGTH MORE THAN 22 MM There are two specimens—one immature, the -• LEMMA LENGTH LESS THAN 21 MM other mature—of this ta.xon in the Linnaean • SPIKES EIGHT OR MORE Herbarium. 4 SPIKES SEVEN OR FEWER One is labeled "barbatum"; the • SPIKES MORE THAN 7.0 CM LONG f SPIKES LESS THAN 6.9 CM LONG name is not lined through. Again, there is no explanation for Linnaeus's second and different usage of Andropogon barbatum. The protologue is a description only; there are no references to previous publications. There is little question then about the species referred to; it is certainly not the same ta.xon which was given this name previously.

The transfer of Andropogon barbatum Lin- naeus (1771) by Swartz cannot stand, for the basionym is a later homonym. It did, however, occupy barbata in Chloris, so that Andropogon barbatum Linnaeus (1759)-the first name applied to the species with ciliate margins on the lemmas described above—cannot be used as was proposed by Nash ( 1898 ) Thus, all of the names used for this species are unacceptable and the new name Chloris dandtjana Adams is adopted.

Chloris dandijana is distinguished from other members of this group primarily in spike- Fig. 16. Geographic distribution and morphological characteristics of Chloris dandtjana. let size and lemma pubescence—characters in which it may approach C. ciliata, at least in South America. It is distinguished from C. canon this problem. Since there are no sharp dis- terai in usually having a relatively large num- continuities in the variation pattern and only ber of flexuous spikes. There are, however, slight correlations of the characteristics with some specimens that differ from this pattern— geographical distribution, I do not feel justified these plants usually having relatively short hairs in naming the variants. on the lemma margins and sometimes only Representative specimens examined: AR- seven to ten short spikes. In an attempt to GENTINA: Prov. Catamarca: Dep. Capital, learn more about this pattern of variation, many Choya, Castillon 42,882 (US); Prov. Chaco, specimens were scored for these and other without precise location. Asp 21 (US); Prov. characters and various correlations were at- Cordoba, Rio Tercero, Stuckert 13,689 (US); tempted. Fig. 16 shows the geographic origin Prov. Corrientes: Isla Corrientes en el Rio Uru- of some of these specimens and their character- guay, Cerca Monte Caseros, Rosengurtt B-3,679 istics. No sharply defined patterns emerged, (US); Prov. Formosa: Costa Alegre, N de la except that individuals with combinations of Primavera, Morel 7,039 (US); Prov. Jujuy: 15 short lemma hairs and short spikes occurred km SE of San Pedro, Eyerdam and Beetle randomly in South America but were neither 22,320 (UC, US); Prov. Misiones: San Javier, found in the West Indies nor in Florida. In Obcra, Schwindt 4.779 (US); Prov. Salta: Dep. South America, however, these plants have no Oran, Maldonado 790 (F, US); Dep. Cande- distinctive distribution pattern. If these variants laria, Ruiz de los Llanos, Montenegro 322. 325 were always found in areas of sympatry, a good (US); Rosario de la Frontera, Los Banos, Ven- case could be made for introgression with an- turi 7,974 (US); Prov. Santiago del Estero: be- other species; but the presence of aberrant tween Forres and Robles, Bartlett 17,944, 19,744 plants in Venezuela, for instance, where the (US); Beltran, Maldonado 510 (US); Prov. nearest sympatric populations are in southern Tucuman: Dep. Leales, Tres Pozos, Descole- Brazil, virtually rules out all but the most Schreiter s.n.. 5 Dec 19.38 (NY, US); Rio Chico, ancient or fast-moving gene flows. Further Escaba, Monetti 42,931 (US); Cruz Alta, studies, especially attempts at hybridizing var- Schreiter 3,987 ( US ) ; Burruvacu, Venturi 2,537, ious portions of the populations, may shed light 2,661 (US), 7,536 (F, US). BAHAMA IS- )

BiOLOciCAi. Series, Vol. 19, No. 2 Taxonomy of the Genu.s Chloris (Gramineae) 37

LANDS: Cat Island, Orange Creek, Britton and URUGUAY: Dep. Artigas: Santa Rosa del Milkpaugh 5J33 (F, NY); Eleuthera I.sland, Guareim, Herter 1,048 (US). UNITED Governor's Harbor, Hitchcock s.n., 14 Nov 1S90 STATES: Florida: Dade Co., Miami, Tracy (F, MO); Long Island, vicinity of Clarence 8,857 (F, MO, NY, TEX, US); Monroe Co.": Town, Britton and Millspaugh 6,245 (F, NY); Upper Matccumbe Key, A. Chase 3,914 (US); New Providence, Blue Hills, Wihon 8,247 (F, Long Key, Silveus 5,329 (TEX); Key Vaca, NY); Hog Island, near New Providence, Wilson Sicallen 5.189 (US); Plantation Key. Swollen 8,292 (F, MO, NY). BOLIVIA: Dep. Santa 5,210 (US); Big Pine Key, Swallen 14,461 (US); Cruz: among bushes in savanna, Cardenas 5,362 Ke\- Largo, Weatherwa.x 61,009 (UC). VENE- (US); Dep. La Paz: Coripata, Hitchcock 22,689 ZLTELA: Carabobo, Valencia, Carreno 8,236

(US); Dep. Cochabamba: without precise loca- ( US ) ; Distrito Federal, vicinity of Caracas, tion, Spiag,ai 26 (US). BRAZIL: Bahia: Cruz Bailey and Bailey 200, 554 (US); Merida, Tovar, das Almas, Pinto 746 (UC, US); Matto Grosso: Fendler 1.854.5 (US); Miranda, between Los Corumba, A. Chase 11,136 (US); Aquidauana, Tequcs and El Garrizal, Pittier 12,973 (NY, US); A. Chase 11,046 (US); between Campo Grande Tachira, La Mulera, Tamayo 2.296 (US); and Dourados, A. Chase 10,915 (F, NY, US); Trujillo, San Jacinto, Tamayo 1,698 (US); Zulia, Minas Geraes: Buritys, near Rio Sao Francisco, savannas of Lagunillas, Jahn 658 (US). A. Chase 10,460 (US); Priapora, Dorsett and Popenoe 3,436 (US); Pernambuco: vicinity of 8. CHLORIS CRUCIATA (Linnaeus) Swartz, Pernambuco, A. Chase 7,757 ( NY, US ; Rio de ) Prodr. Veg. Ind. Occ. 25. 1788. (Based Janeiro: San Pedro, NW of Gabo Frio, A. Chase on Agrostis cruciata Linnaeus. ) Fig. 17, 10,158 (US); vicinity of Sao Leopoldo, Leite A-E. 471 (NY); Geno Largo, near San Luiz, Ramho 53,116 (US); Santa Vitoria de Palmar, SwuIIen Agrostis cruciata Linnaeus, Svst. Nat., ed.

7,416 (US). BRITISH GUIANA: Essequibo Co., 10. 2:87.3. 1759. ("Sloan, jam. t. 69. f. I." Rupunini River, Loxton s.n., 26 Sep 1955 (K). cited by Linnaeus. The illustration cited is

COLOMBIA: Dep. Antioquia: 1 km E of Puente a good likeness of the species treated here. de Occidente, Pelaez and Barkleij 595 (US); A specimen, in LINN!, labeled Agrostis Dep. Bolivar: Tierrabomba Island, Cartagena cruciata is somewhat immature and does Bay, Killip and Smith 14,136 (NY, US): Dep. not ha\'e the spreading spikelets character- Torrecillo: near Turbaco, Killip and Smith istic of the species. In all other respects 14,639 (F, MO, NY, US); Div. Boyaca, Soata, the specimen fits the usual circumscrip-

Cuatrecasas 1,094 (F, US); Dep. Gundinimarca: tion. ) Route de Honda a Guaduas, Humbert 27,097 Rahdochloa cruciata (Linnaeus) Beauvois, (US); Girardot, Pennel 168 (NY); Dep. Huila: Ess. Agrost. 84, 158, 176. 1812. (Based on 3°23'N, 74°49'W, Fosherg 19,589 (US); Dep. Chloris cruciata [Linnaeus] Swartz.) Valle: del Rio Sanjunifiuin, Cnafreca.^as Hova Cynodon cruciafus (Linnaeus) Raspail, 15,366 (US); Dep. Magdalena: E of Los Ve- Ann. Sci. Nat. Bot. (Paris) 5:.30.3. 1825. nadas, 10°N, 73°43'W, ^ 5,829 Ditgand (US); (Based on Chloris cruciata [Linnaeus] Santander: Lati- Dep. 12 km from Bucaramanga, Swartz. genheim 3,198 (UC, US). ECUADOR: Prov. Chloris humholdtiana Steudel, Syn. PI. Imbabura: cntre Paramb;as y Pajon, Acosta- Glum. 1:205. 1854. ("C. cruciata H. B. [vix Solis 12,667 (F, US). GRAND CAYMAN: Swartz.] Ins. Cuba" cited. Description Hitchcock s.n., 19 Jan 1891 (MO). JAMAICA: clearly of C. cruciata as commonly under- Cornwall Co.: Santa Cruz Mts., Britton 1.172 stood. (NY); Middlesex Co.: Lower Clarendon Parish, ) Chloris hrevightma Wright, Anales Acad. Inverness, Harris 12.743 ( F, MO, NY, US ) ; Lee- Ciencias Habana 8:200." 1871. (HOLO- ward Islands: Antigua, road to Hermitage Bay, ." Box 26 (US). PARAGUAY: Dep. Boqueron: TYPE: "1848. Ganuela. . . is cited in pro- "1848" Puerto Casado, Hartley SH 141 (US); Puerto tologue. is evidently an error for "1548" for a photograph, in US!, of the Sastre, Ramirez 5, 19 (US); Mariscal Estigar- type in has the latter number. Isotypes, ribia, Rosengurtt B-5,813 (US); Dep. Goncep- GH also with the number "1548" are in BM! cion: Estancia "Fanciere," Ramirez 193 (US); and Gi.) Dep. Paraguari: Caapucu, Anderson 1155A

(US). PERU: Dep. Cajamarca: Jaen, Hutchin- Tufted perennial up to 35 cm tall; sheaths son, 1,422 (UC, US); Dep. San Martin: Puca- glabrous; ligule shortly pilose; blades very nar- yacu, 11 km S of Tarapoto, Ferretjra 7,725 (US). row, usually less than 0.5 mm wide, up to 6 38 Brighaai Young University Science Bulletin

$m^

Fig. 17. Chloris cniciata, C. filifoTmis, and C. suringari. (A-E) C. cruciata. (A) habit, x 1/6; (B) portion of single spike, x 2; (C) spikelet, partly dissected, x 10; (D) sterile floret, x 10; (E) caryopsis, x 10. (F-J) C. suringari. (F) habit, x 1/5; (G) portion of spike showing rachis and attached spikelets, x 10; (H) spikelet, partly di.ssected to show lowest sterile floret, x 10; (I) upper two sterile florets, x 10; (J) caryopsis, x 10. (K-L) C. filifonnis. (K) habit, x 1/3; (L) spikelet, partly dissected, x 10. cm long, involute; inflorescence of two or three caryopsis narrowly cylindrical, 1.4 to 1.5 mm radiate branches, spreading horizontally; spikes long, ca 2.0 mm wide. 1.0 to 2.5 cm long; spikelets somewhat ap- Chloris cniciata is the only species in the pressed when immature, eventually spreading genus with a tufted habit, very narrow leaves, at nearly right angles to the rachis ; spikelets two or three horizontally spreading spikes, and only slightly ( if at all ) imbricate, averaging spikelets that are divergent from the rachis at five or six per cm of the rachis length; first right angles (Fig. 17). Other species from the glume ovate-lanceolate, 0.7 to 1.3 mm long, Caribbean (especially C. sagraeana and C. ek- 0.1 to 0.3 mm wide, glabrous, midrib sometimes inanii) have similar, though appressed, spike- slightly scabrous; second glume narrowly lan- lets; they also lack the other features. ceolate, 1.6 to 2.2 mm long, 0.1 to 0.3 mm wide, Epidermal studies of this species ( Rogers, glabrous, midrib occasionally scabrous; fertile 1967, and those reported earlier in this paper) lemma pale, narrowly lanceolate, 2.2 to 2.8 mm revealed the presence of somewhat cruciate long, 0.3 to 0.5 mm wide, glabrous, occasionally short cells, a feature not found elsewhere in the inroUed hyaline margins with a few short Chloris. All inflorescence and spikelet character- hairs, callus bearded, awn 7.5 to 17.0 mm long; istics are typical of Chloris, however, and I do sterile floret one, 0.5 to 1.4 mm long, 0.2 to 0.3 not feel that ta.xonomic recognition of this spe- mm wide, narrowly truncate, borne on a fairh' cies as a distinct genus is warranted. long rachilla joint, awn 3.7 to 7.0 mm long; Chloris cruciata has been collected in stony ) ) . )

Biological Series, Vol. 19, No. Taxonomy of the Genus Chloris (Ghamineae) 39 soils (often of serpentine origin) from Cuba, Paris." Not seen, but description and ac- Haiti, and Jamaica (Fig. 18). companying plate are clear.

Representative specimens examined: CUBA: Stoloniferous or tufted perennial, 10 to 30 Prov. Habana: Guanabacoa, Lomas de Jatas, cm tall; sheaths glabrous to sparsely pilose, Casuora, Ekman 279 (US); San Miguel de compressed, keeled; ligule a very short ciliate Prov. Las Loma de la Vita, Leon 11,753 (US); crown, often with a few longer hairs near the 10 S of Santa Clara, Howard et al. Villas: km margins; blades narrow, 1 to 6 mm long, about 100 vicinity of Sancti Spiritus, Leon (UC); 1 mm wide, folded, often curved, sparsely pilose banks of 1,497 (US); Oriente Prov.: Bazate, near the base, becoming apically glabrous, HAITI: Rio Canto, Ekman 3,016 (K, NY, US). acute; spikes one or two, 1 to 3 cm long, erect; Massif du Nord, between Cerca-Carvijal and spikelets rather densely imbricate, ca 12 per cm (US). Bois-Charles, Ekman H607 JAMAICA: of rachis length, more or less appressed; glumes Troy, Harris 12,647 (F, K, MO, Cornwall Co., lanceolate, glabrous; first glume 1.7 to 2.0 mm Parish, between Ipswich NY, US); St. Elizabeth long, ca 0.4 mm wide; second glume 2.5 to 2.8 Black River, Hitchcock 9,597 (US); Middle- and mm long, ca 0.4 mm wide; fertile lemma 2.6 to vicinity of Mandeville, Britton 1,029 sex Co., 2.8 mm long, 0.5 to 0.6 mm wide, lanceolate- SurrcN' Co., St. Andrew Parish, Stonv (NY); elliptic, apex acute, callus bearded, internerves Clifton, Hill, Harris 'll,341 (F, K, NY, US); densely appressed-pubescent, awn 1.0 to 1.2 Harris 11,375 (F, NY, US); near Kingston, along mm long; sterile floret 1.3 to 2.0 mm long, 0.2 Stonv Hill Road, Hitchcock 9,276 (F, NY, UC, to 0.5 mm wide, narrowly lanceolate to linear, Yallahs Valley, Perkins US); Blue Mountains, acute, sparsely appressed-pubescent, awn 0.5 to 1,481 (US). 1.1 mm long; caryopsis not seen. Known only from Mauritius and nearb\- is- 9. CHLORIS FILIFORMIS (Vahl) Poiret in lets, Chloris filiformis is easily distinguished Lam. Encycl. Method. Bot. Suppl. 2:238. from the similar C. .suring,ari (from Cuba and 1811. (Perhaps based on Ctinosurus fili- Cura^'ao) by having considerably shorter awns formis Vahl.) Non Chloris filifortnis Poiret. and lemmas with appressed pubescence on the Fig. 17, K-L. internerves rather than spreading pubescence on the margins and keels (Fig. 17, K-L, F-J, Ci/nosurus filiformis Vahl, Symbolae Bot. respectively ) 2:20. 1790. (HOLOTYPE: Photograph, in US!, of type from Vahl Herbarium, C. Specimens examined: MAURITIUS: Mon- Chloris filiformis Poiret, in Lam. Encvcl. tague Longue, dry regions, Botiton s.n., without Meth. Bot. Suppl. 2:237. 1811. (TYPE date (K);"R()und' Island, rocky hillside, 520 ft., MATERIAL: P! There are several speci- //. //. Johnston s.n., 28 Nov 1889 (K); Round and mens at P, in the Herbarium Richard, Island, Wiene s.n., Nov 1937 (K). Hubbard labeled "Chloris filiformis Poiret. He. de Vaughn (1940) reported this species also from France." While Poiret did not designate a the islets of Gunners Quoin and Le Morne. ts'pe as such, they probably represent authentic material. Tlie specimens fit the de- 10. CHLORIS SURINGARI Hitchcock in Ur- scription closelv; and this is, unquestion- ban, Svmb. AntiU. 7:167. 1912. (HOLO-

ably, the same taxon first described by TYPE: "Curasao ad Brakkeput . . . W. F. R. \'ahl under Cynosurtis, though Vahl is not Suringar, Jan 14, 1885" Type fragment, in cited by Poiret in the protologue of this US!, from Krug and Urban Herbarium.) name. Fig. 17, F-J. Chloris distachya Kunth, Rev. Gram. 1:291. Densely tufted, very shortly stoloniferous, pi. 57. 1830. "(HOLOTYPE: "Chloris dis- diminutive perennial up to 15 cm tall; sheaths tachya Kunth, Insula franciae. Ex Museo very short, about 1 cm or less, striate, sparsely to densely pilose below, becoming glabrous above on some culms; ligule a short tuft of hairs; blades highly reduced, less than 0.5 mm wide and 2 cm long, usually somewhat curved; apex blunt to acute, glabrous to sparsely pilose; inflorescence a single spike 1.0 to 2.5 cm long; spikelets imbricate, about 15 per cm of the rachis; first glume 1.0 to 1.5 mm long, 0.4 to to the Fig. 18. Distribution of Chloris crucUita. 0.5 mm wide, lanceolate, appressed 40 Brigham Young University Science Bulletin rachis, one-nerved, glabrous, marginal areas thin Erect cespitose or stoloniferous perennial, and hyaline; second glume very narrowly lan- 25 to 50 cm tall; sheaths glabrous to very ceolate, diverging from the rachis at about 70°, sparsely spreading pubescent; ligule a short 2.3 to 3.0 mm long, ca 0.3 to 0.4 mm wide, one- ciliate crown; blades up to 20 cm long, 5 to 8 nerved, glabrous, margins hyaline; fertile lemma mm wide, glabrous to scabrous or very sparsely ovate-lanceolate, 1.7 to 2.5 mm long, 0.4 to 0.6 pilose; spikes four to six, 3 to 5 cm long, virgate mm wide, densely pubescent on the callus, to somewhat spreading; spikelets imbricate, ca margins and keel with hairs about 0.7 mm long, 7 per cm of the scabrous rachis; glumes un- avvned from about the middle of the keel, the equal, purplish, membranous, glabrous except awn 2 to 3 mm long, straight or slightly genicu- for the scabrous midrib; first glume elliptic to late; sterile florets three, the lowest about 1 mm linear, 4.0 to 4.7 mm long, ca 0.5 mm wide, long, 0.3 to 0.4 mm wide, avvned from about rounded at the apex; second glume 5 to 7 mm the middle, the awn 1.5 to 2.0 mm long; upper long, ca 1 mm wide, narrowly oblanceolate to sterile florets much reduced, awned; caryopsis obovate or elliptical, apex bifid; fertile lemma ca 1.2 mm long, 0.5 mm wide, ovoid, flattened highly modified and differing from any other to slightly trigonous. species in Chloris, callus narrow and elongated This unique species, endemic to Cuba and at the base of the lemma, 1.0 to 2.5 mm long, Cura9ao, may be immediately separated from densely bearded, sharp-pointed, body of lemma 2.5 long, 0.5 to the similar species, C. filiformis, in having rela- ovate to ellipsoid, 1.0 to mm 1.2 wide, margins membranous, prom- tively long awns and a single spike ( Fig. 17, mm inently winged, glabrous below, becoming F-J). Additional notes on the relationships of these species have been presented under C. densely spreading ciHate above, keel sparsely below, 5.8 to filiformis. to densely pilose, especially awn 8.5 long; sterile florets three to five, broad, Collection labels indicate that Chloris sur- mm membranous margined, with prominent veins; ingari grows on limestone benches and rocks lowermost floret broadly cordate or even am- not far from the seashore. Only four specimens plexicaulous at the base and partly surrounding were seen in all herbaria consulted. the corky rachilla, tapering to a narrow tip Specimens e.xamined: CUBA: Oriente Prov., above, ca 4.5 mm long, ca 5 mm wide, awn 5 El Guanal, Leon 11,564, Jan 1940 (US). CURA- mm long, upper sterile florets progressively re- SAO: without precise location, Bohlinp^h 9.281. duced; caryopsis 1.3 to 1.5 mm long, 0.8 to 1.0 1909-1910 (US); limestone bench near Willem- mm wide, broadly obovoid-ellipsoid, trigonous; Britton and Shafcr 3,058, 20-27 Mar 1913 stad, chromosome number 2n=^40. (US); Kleine Berg, Mordant 251, 23 Nov 1966 No other species in the genus has such (P)- highly modified spikelet parts as Chloris scariosa. The broadly winged fertile lemmas and 11. CHLORIS SCARIOSA F. von Mueller, sterile florets at once set this species apart from Fragm. Phyt. Australiae. 6:85. 1867. all others. This unique character was recognized MEL, not seen. There are (HOLOTYPE: by Thellung (1919), \\ho erected the section two specimens at Kew, collected by von Hackelochloris to include it. Mueller in 1879, both of this species as Furthei-more, there is a strong overall re- commonly understood. One is labeled semblance between C. scariosa and various spe- "Chloris scariosa F. Muell. Gracemere. cies of Tetrapogon. Certainly, the two genera Bowman. Herb. F. Mueller. 1879." The approach closely at this point and must be other is labeled "Chloris lativalvis Muell. rather arbitrarily separated on the basis of fer- Sturt's Creek. M. Gregory." Lazarides tile floret number. Both epidermal and anatomi- [1972] says that the latter specimen is an cal features have been studied; no significant isotype of the holotype at MEL. The pro- differences between the two genera can be tologue. which is very short, refers to demonstrated. On these bases it does not seem "glumis latis scariosis insignis." Since von desirable to erect a new genus for this single Mueller, as evidenced by other descriptions species. on the same page, was familiar with the Chloris scariosa has been collected from a species of Chloris in Australia, there is no variety of soils, most commonly in arid regions doubt as to the taxon to which he referred of Queensland and Northern Territory, Aus- and Lazarides 's designation of the Gregory tralia (Fig. 20). specimens as the type is reasonable. Chloris lativalvis was apparently never published.) Representative specimens examined: AUS- mi S of Yam- Fig. 19, A-D. TRALIA: Northern Territory: 5 BiOLOCicAi, Series, Vol. 19, No. 2 Taxonomy of the Genus Cm,onis (Gramineae) 41

Fig. 19. Chloris scariosa and C. prieurii. (A-D) C. scariosa. (A) habit, x 1/4; (B) spikelet, partly dissected, X 5; (C) sterile florets, partly dissected, x 5; (D) caryopsis, x 10. (E-H) C. prieurii. (E) habit, X 1/4; (F) spikelet, partly dissected, x 10; (G) sterile florets, partly dissected, x 15; (H) caryopsis, x 10. 42 Bricham Young Universit-i- Science Bulletin

Dr. W. D. Clayton, in a personal commu- nication, says that the type cannot be found

at CN. Under these circumstances, it must be presumed lost. The description, while not detailed enough for positive identification, suggests strongly that the plant re-

ferred to is C. prieurii. ) Chloris multiradiata Hochstetter, Flora 38:204. 1855. (The original description seems to refer to C. prieurii. There are possible isotypes in K! and P! The K specimen is C. virgata; the P specimen, C. prieurii. )

Stoloniferous perennial up to 80 cm tall; Fig. 20. Distribution of Chloris scariosa. sheaths glabrous, occasionally becoming pilose near the ligule; ligule variously short-ciliate to bah Station, Perry 3,368 (K); near coast, 60 mi long-pilose; blades narrow, 10 to 14 cm long, up N of Wollogorong Station, Pern/ 1,218 (K, US); t(j 5 mm wide; lower surface glabrous, upper 8 mi N of Tennant Creek, Perry 551 (K); Burt- surface scabrous to pilose; spikes appressed to well, Cleland 349 (K). Queensland: Burke Dtr., slightly di\'erging, three to seven, 6 to 11 cm Cloncurry, Allan 20 (K); Forest Home Station, long, spikelets imbricate, 8 to 11 per cm of Brass 1,790 (K); near Normantovvn Township, the scabrous-hispidulous rachis; glumes thin flood flats of Norman River, Lazarides 4,248 and delicate, narrowly lanceolate, glabrous with (K, US); Charters Towers, Michael 1,299 (K); slightly scabrous midnei-ves; first glume 2.1 to Antil Plains, near Townsville, White 8,909 (K, 2.2 mm long, 0.2 to 0.3 mm wide; second glume NY, US); Warrego Dtr., claypans near Cun- 3.7 to 4.0 mm long, 0.2 to 0.3 mm wide; fertile namulla. White 11,566 (K, US). South Austra- lemma nairowly elliptic, 3.3 to 4.7 mm long, lia: Cordillo (Downs), Clehnul s.n., 5-24 (Ma\-, 0.4 to 0.7 mm wide, with a row of clavate 1924?). Western Australia: between Gasco\ne glandular hairs along the middle of the lemma and Fortescue River, King s.n. 1885 (K). side, awn 10 to 18 mm long; sterile florets SWITZERLAND: Derendingen, Kammgarn four or five, the lower successively enclosing Fabrik, Prob.st s.n., 25 Oct 1919, adventive (K). the upper and becoming progressively more reduced; lowemiost sterile floret cylindrical, 1.5 to 2.5 long, 0.2 to 0.4 wide, awn to 12. CHLORIS PRIEURH Kunth, Rev. Cram. mm mm 8 17 mm long, uppermost sterile floret more or 1:89. 1829. (HOLOTYPE: "... He St. less flabellate, ca 0.3 long, 0.1 Louis [Senegal] Roger Dedit, Major 1S25." about mm mm wide, awn 1.5 to 2.0 long, intermediate P! ISOTYPE: K! Fig. 19, E-H.) mm sterile florets gradualh' changing from cylin- Chloris drical to flabellate; caryopsis 2.0 to 2.5 cryptostachya Steudel ex J. A. mm Schmidt, Beitr. Flora Cap Verdischen In- long, 0.5 mm wide, ellipsoid, strongly flattened, " seln 148. 1852, (HOLOTYPE: . . . C. light brown.

punctulata Hochstetter hrbr. u. it. iiubic. nr. (July two species of Chloris have a row of 23 Cordofan." P! ISOTYPE: K!) glandular hairs on the side of the lemma—C. Chloris punctulata Hochstetter ex Steudel, prieurii and C orthonoton. The two are easily Syn. PI. Glum. 1:205. 1854. (HOLO- separated. C. prieurii is limited to the Old

TYPE: "Hrbr. un. it. nubic. nr. 2.3." P! \\'orId ( Fig. 21 ) and has four or five sterile ISOTYPE: K!) florets (Fig. 19, E-H); C. orthonoton is, on has Chloris subtriflora Steudel, Syn. PI. Glum. the other hand, from South America and 1:208. 1854. (HOLOTYPE: "LaPrieur in a single sterile floret (Figs. .33, A-D; 39). Senegambia." No collection that can be Representative specimens examined: BOTS- identified positively as the holotype can be WANA: between Kapupahedi and Tamso, Oka- found, though there are specimens in the vango National Territory, anon. (K). CHAD: Herbarium Richard in P! that were col- Bilthie, Ro.ssetti 39 (BM). CAPE VERDE IS- lected by Le Prieur in Senegal in 1829. LANDS: Boa Vista, Chevalier 44,423 (K). These are not labeled as Chloris triflora, ETHIOPIA: Nario Dikeno, Schimper 1,607 and there are no other data on the labels. (MO). GHANA: Accra, Irvine 3,023 (MO). Biological Series, Vol. 19, No. 2 Taxonomy ok the Genus Chlohis (Gramineae) 43

to twenty-eight, 6 to 10 cm long, erect, somewhat flexous; spikelets imbricate, ca nine per cm of rachis length; first glume ovate-lanceolate, glabrous except for the slightly scabrous midnerve, 1.6 to 2.5 mm long, ca 0.2 to 0.3 mm wide; second glume linear-lanceolate, arcuate, glabrous except for the scabrous midnerve, 4.0 to 5.5 mm long, 0.4 to 0.5 mm wide; fertile lemma 3 to 4 mm long, 0.4 to 0.5 mm wide, narrowly elliptic, apex long-acuminate, tapering into the awn, upper % of the margin with long hairs, these up to 2.5 mm long; callus bearded, awn ca 0.5 to 1.0 mm long; sterile florets two; lower sterile floret similar to the fertile floret in shape, staminate, often with a rudimentary pistil, 2.5 to 3.5 mm long, ca 0.3 mm wide, margins with cilia up to 2.5 mm long, callus bearded, upper lemma sides sparsely pubescent, palea present, about as long as the sterile lemma; upper sterile floret clavate, less than 1 mm long, pilose. Caryopsis not seen.

Chloris rohusta is unique in many respects, Fig. 21. Distribution of Chloris prieurii. Inset A: including the bamboolike growth habit, pref- India. erence for aquatic to subaquatic habitats, and spikelets with sterile florets that are regularly INDIA; Rajasthan Prov., Jaipur, Raizada 2/53 staminate. These features, together with the (K). MALI: San.sancling, Chevalier 24,958 (P); overall morphological aspect of the spikelets, Nara, Chudeau sm., 26 Aug 1918 (P); Tim- suggest that this species should be separated buktu, Hagerup 239 (US). NIGERIA: Bomu from the rest of Chloris. Epidennal and ana- Prov., 48 mi N of Damaturu, De Leeuw 1,150 tomical characters, however, are those typically (K). SENEGAL: Kaolack, Berhaut 456 (P); found in other species of Chloris; and, there- Mbambev, Chevalier 33,795 (P); Pointe Louis, fore, there seems to be little justification for Roberfi/' 149.16.2 (K). SUDAN: Blue Nile segregating it. Prov., Bagein, 50 mi S of Khartoum, Jackson collected throughout 2,336 (K); Darfur Prov., ]chv\ Mana, Blair 140 This species has been Its tall, bamboo- (K); Khartoum Prov., Khartoum, Schweinftirth equatorial Africa (Fig. 23). colonies along 813 (US); Kordofan Prov.: El Muglad, An- like, elastic culms form large drews 3,062 (K); El Obeid, Harrison 76 (K). and in rivers and streams. Associated grasses UNITED STATES (adventive): Alabama: Mo- include Phragmites communis and species of bile, Mobile and Ohio wharf, Mohr 44 (US); Andropogon. North Carolina; Wilmington, McCarthi/ 90 Representative specimens examined; CAM- (US). UPPER VOLTA; Gourma, Rossetti 110 EROUN; riverain forest of Sanaga River, near (K). Goyoum, 20 km W of Deng Deng, Breteler 980 (K). DAHOMEY: entre de Lac Ozri et Zagnando, Chevalier 23.054 (K). GHANA; 3 13. CHLORIS ROBUSTA Stapf in Chevalier, mi from Yendi, Kufapindi River, Adams and Bull. Soc. Bot. France Mem. IV 8:221. Akpahla 4,101 (K); 30 mi from Navrango, 1912. (DUPLICATES OF SYNTYPES; Road, Adams and Akpabla 4,361 (K); "Haut-Chari, Ndelle, 20-2.5 Dec 1902, Chcx- Tumu Sakogu-Shishe Road, old Morago River bridge, alicr, 6.991; Nicreria du Nord, Borgu. Bar- Ankrah 20.507 (K); Daka River near Yendi, ter, S7S." K! ) Fig. 22. A-D. Brand 444 (US); Weija, near Accra, Hall 2,552 Mor- Robu.st perennial to .3 m tall, aquatic to (K); \\'enchi, between Banda and Menji, subaquatic with very strong, heavy culm bases, ton GC 25,256 (K); near Daboya, Thorold 288 and often dense rooting at the lower nodes, (K). I\'ORY COAST: Marabadiassa, along stoloniferous with stolons up to 7 m long; Bandama Blanc River, As.si 7.178 (K); entre sheaths glabrous; ligule a minute naked crown; Sanlo and Kalepui, Assi 9,291 (K); bords du blades up to 40 cm long and 20 mm wide, Bandama River, Dume, Roherty s.n. (MO glabrous and glaucous or scabrous; spikes seven 1640174). NIGERIA: between Oyo and Iseyin, 44 Brigham Young University Science Bulletin

Fig. 22. Chloris robusta and C. Mollis. (A-D) C. rohmta. (A) lower and middle portions of stem, \ 1/3; (B) upper stem and inflorescence, x 1/4; (C) spikelet, partly dissected, x 10; (D) sterile florets, x 15. (E-J) C. mollis. (E) habit, x 1/4; (F) spikelet from type of C. mollis, partly dissected, x 10; (G) floret from type of C. anlsopoda, x 10; (H) sterile florets from spikelet with two sterile florets, x 20; (I) sterile floret from spikelet with single sterile floret, x 15; (J) caryopsis, x 5. )

Biological Series, Vol. 19, No. 2 T.^.xonomv of the Genus Chlobis ( Gr.a.mineae) 45

TYPE: "Serra do Borborema, State of Para- hvba do Norte, Brazil, April, Lutzelburg, No. 12451" US!) Chhris anisopoda Scribner in Robinson, Proc. Amer. Acad. Arts Sci. 38:118. 1902. ISOSYNTYPES: [Galapagos Islands] "Charles Isl., Baur, 333; Snodgrass and Heller, 459" US! SYNTYPES; GH, not seen. Chloris angustiflora Arcsehoug, Kongl. Sven- ska Fregatten Eugenics Bot. III. 118. 1910. (TYPE: not seen, but description of specimen, collected on the island of Puna, by

Andersson is conclusive.) Fig. 23. Distribution of Chloris robusta (equatorial Chloris leptantha Hitchcock e,\ Urban, Africa). Symb. Antill. 7:166. 1912. (HOLOTYPE: "Hab. in insulis orae \'enezuelensi adjacen- Brenan and Keaij, 8,959 (K); Yola, Dalziel 269 tibus Bonaire, 19 Feb 1885. W. F. R. Sur- (K); Lagos, Foster 2 (K); Jebba, on Niger ingar." US!) River, Uagerup 692 (K, US); Onitsha Prov., near Nzam Anambra Forest Reser\'e, Oramili Chloris rupestris (Ridlev) Hitchcock, U. S. 243:126. 1936. Creek, Jones 6,888 (K); Oyo Prov., Dtr. Ibadan, Dopt. Agric. Misc. Publ. rupestre Ridley.) Awba Hills Forest Reserve, Onochie and Jones (Based on Gijmnopogon 14,666 (K); bv Chinchaga River, near Muina, Annuiil 40 to 60 cm tall from a fibrous, Meikle 736 (K); Kaiama, Ward 35 (K, US). rather poorlv developed root system; sheaths REPUBLIC OF THE CONGO; Prov. Equa- usually denseh- appressed- to spreading-pilose, teur, Bangui, Ubangi River, MeruiUan 209 (K). especialK near the apex, occasionally i-nclosing SIERRA LEONE: Moa River, Mafindo Falls, cleistogamous spikelets; ligule long-pilose; near Kailahun, Deifihton 4,004 (K, US). SU- blades variable in size, ranging from short and DAN; Equatoria Prov., 40 mi NW of Yei, Myers verv narrow up to -30 cm long and 1 cm wide, 7,865, 8,496 (K); Bahr Chazal Prov., Mvolo. appressed-pilose on both upper and lower sur- Schwcinfurfh 2,843 (US). UGANDA: Kitgum, faces; inflorescence of three to seven spikes, 6 Cluia. Thomas 4,346 (K). to 9 cm long, mostly radiate at the apex of the culm, sometimes with one or two single spikes 14. CHLORIS MOLLIS (Nees) Swallen, North inserted separately just below; spikelets imbri- AiiKT. Flora 17:596. 19.39. (Based on Ct/w- cate, seven to ten per cm of rachis length; nopo^on mollis Nees.) Fig. 22, E-J; 24. glumes thin, transparent, narrowly lanceolate, glabrous except for the scabrous midnerve; first Gi/mnopogon mollis Nees, Agrost. Bras. glume 3 to 4 mm long, 0.2 to 0.3 mm wide, 427. IS.3,3. (FRAGMENT OF TYPE?: US! Labeled "G\ninop(>gon mollis Nees. Type. Campi Mimoso dicit. Prov. Piauhy." The name of the herbarium from which the fragment was obtained is illegible. While the infomiation on the label is sketchy, it does match exactly the collection location noted by Nees. This, together with the very complete original description, leave no question as to the typification of this name.) Gijmnopogon rupestre Ridley, Jour. Linn. Soc. Bot. (London) 27:73. 1891. (HOLO- TYPE: BM! ISOTYPE: US! "Fernando do Noronha. Cliffs on Portuguese Bay. Ridley, Ramage, Sept. 14,1887.") Fig. 24. Distribution of Chloris mollis. Insets: (A) Chloris luetzeUmrgii Hitchcock, Proc. Biol. northern South America. (B) eastern Cuba, Ja- Soc. Washington 36:197. 192.3. (HOLO- maica, Hispaniola. (C) Galapagos Islands. 46 Brigham Young University Science Bulletin

glabrous; second glume 3.1 to 4.6 mm long, ca ZIL: Bahia: Juazeiro, Dorsett and Popenoe 0.3 mm wide, glabrous; fertile lemma very 411h (US); Ceara: Crateiis, Swollen 4,481 narrow, 2.5 to 6.8 mm long, ca 0.3 mm wide, (US); Iguatu, Swollen 4,399 (K. US); Fernando glabrous e.xcept for the prominently bearded de Noronha: Morro Francez, Simmons 9,263 callus and the short-ciliate upper margins, awn (US); Maranhao: Municipio de Loreta, be- 4.0 to 7.5 mm long; sterile floret usually one tween Balsas and Paniaiba Rivers, Eiten and (occasionally two), and then the lowermost Eiten 4,503 (K, US); Paraiba: Pombal, Picket bearing rudimentary flower parts; lowermost 3,804 (US); Soledade, Pickel 3,851 (US); Per- sterile floret cylindrical, 1.2 to 1.6 mm long, nambuco: Serra do Genipapo, A. Chase 7,686 0.1 to 0.2 mm wide, awn 3.7 to 5.8 mm long, (US); Pcsqueira, Pickel 1,662, 3,675 (NY, US); callus prominently pubescent with spreading Piaui: Fazenda Nacional, Sivallen 4,179 (NY, hairs, margin ciliate above; upper sterile floret, US), 4,296 (K, US); Picos. Sivallen 4,228 (US); when present, ca 1 mm long, 0.1 mm wide, Rio Grande do Norte: Santa Cruz, Swollen callus bearded, awn ca 2.5 mm long; caryopsis 4,558 (US); Angicos, Swollen 4,714 (K, US). ca 3.3 mm long, ca 0.5 mm wide, narrowly COLOMBIA: Bolivar Div.: Cartagena, Hitch- cylindrical. cock 9,906 (US); Magdalena Div.: Santa Marta,

ChloTts mollis is unique in the genus in hav- H. II. Smith 2,752 (F, K, MO, NY, US). COS- ing sterile florets with a prominently pubescent TA RICA: Boruca, Pittier 3,675 (BR). CUBA: callus and long awns. Oriente Prov.: Caimenera Naval Station, Hiorom and Ramsden s.n., 6 1919 (US). As may be seen from the synonymy, this Feb CU- near Willemstad, Britton species has been placed in both Gymnopogon RASAO: and Shafer 2,906 (US). DOMINICAN REPUBLIC: Prov. and in Chloris. In some respects it does have Barahona: Salinas, 1,921 characters of both genera. Many species of Las Fuertes (US). Prov. Guayas: Chanduy, in littore Gymnopogon are characterized b\' having short, ECUADOR: Maris Pacific!, Spruce 6,436, Apr 1862 (K, NY, stiff, sharp-pointed leaf blades, which appear Salinas, Hitch- crowded at the base of the culm because of US); between Guayaciuil and cock 20.056 A.^plund their short, overlapping sheaths. Chloris mollis (US);' Chongon, 7,671 has these characters, except that the leaves are (US). GALAPAGOS ISLANDS: Charles Is- land: Black Beach, .9,.3.55 not crowded at the base. Gt/mnopogon tends near Howell (NY, 184 Post Office Bay, to have inflorescence branches somewhat stag- US), Svenwn (US); Za- gered at the culm apex rather than in the ver- Howell 8,801 (US). GUATEMALA: Dep. capa: I of Tcculutan, Stei/ermark 29,192 ticillate arrangement so common to Chloris. In mi W (US). HAITI: Anses-a-Pitre, Ekman 6,999 (G, this respect, Cliloris mollis is intemiediate between the two genera, with the upper spikes K, US); jean Rabel, Leonard and Leonard 12,581 Harris more or less radiate but occasionally with a few (US). JAMAICA: Gordontown, 11,515 (F, US); near Kingston, Harris remote lower spikes. In Gi/mnopogon the sterile K, MO, 12,212 (K, US). LESSER ANTILLES: Bon- floret is usually much reduced, often being rep-

aire Island, Boldingh 7,208B ( ; Aruba, Sur- resented only by a swollen area at the junction US ) ingar 12 Prov. Lambayeque: en- of rachilla and awn. One species, G. foliosus, (US). PERU: however, does have a reasonably well-developed tre Motupe y Olmos, Lopez 1,238, 2,463 (US); Prov. Gato, sterile floret and bears a remarkable resem- Tumbes: entre Pocitos v Una de Fcrreyra al. 10,6.34 blance to Chloris mollis. Essentially then, we et (US).' VENEZUELA: are faced with the problem of which characters Dep. Federal, Gran Roque, Gines 4,615 (US); Anzoatequi, 14 km S of San Mateo, Garroni 53 to accept. I believe that by emphasizing primarily the rather unusual vegetative character- (US). istics and inflorescence of such species of Gt/m- nopogon as G. floridamts, G. cliapmanniamis. 15. CHLORIS LONGIARISTATA Napper, Kir etc., that the highest degree of homogeneit\' kia 3:117. 1963. (ISOTYPE: "Tangain ika, may be established in the resulting group. In Ngudu. R. R. Staples. 382. 1/4/1935."' K!) such a system, Chloris mollis is still essentially Fig. 25, E-H.

unclaimed, for its outward vegetative appear- Stoloniferous perennial 30 to 70 cm tall; ance and inflorescence type do not fit well with siieaths somewhat compressed, glabrous; ligule Gymnopogon. However, the well-developed a short ciliate cro\vn; blades up to 15 cm long, sterile florets of C mollis are certainly more 3 to 6 mm wide, acute, glabrous below, sparsel)' like the majority of Chloris species rather than long-pilose above and on the margins, especially most of the Gymnopogon taxa. near the base; spikes three to five. 3.5 to 12.0 Representative specimens examined: BRA- cm long, appressed to slightly divergent; spike- Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Ciiloris (Gramineae) 47

habit, x (B) spikelet, Fig. 25. ChloTis quinquesetica and C. longiaristata. ( A-D ) C. quinquesetica. (A) 1/4; longiaristata (E) habit, x partly dissected, x 10; (C) sterile florets, x 10; (D) caryopsis, x 10. (E-H) C. . 1/4; (F) spikelet, partly dissected, x 5; (G) sterile florets, x 5; ( H) caryopsis, x 5. ):

48 Bricham Young UNrv'EBsrrv Science Bulletin lets slightly divergent, ca four per cm of the cent, sometimes glabrous, margins densely scabrous rachis; glumes very narrowlv lance- spreading-pubescent with hairs about 1 mm olate, pale, membranous, tapering to a delicate long, callus densely bearded, awn 4 to 8 mm point or a short awn; first glume 3.7 to 4.9 long; sterile florets three or four; lowest sterile mm long, 0.2 to 0.4 mm wide; second glume floret tiuncate, 1.3 to 1.6 mm long, 0.5 to 0.6 5.5 to 6.7 mm long, 0.3 to 0.5 mm wide; fertile mm wide, glabrous below but with a prominent lemma dorsally compressed, 7.5 to 8.1 mm long. fringe of hairs along the upper margin, awn 0.8 to 0.9 mm thick, 1.0 to 1.5 mm wide, nar- 3.5 to 6.0 mm long; upper florets enclosed by rowly lanceolate to elliptic, callus well devel- the lower and progressively reduced, becoming oped, tapering, shaip pointed, densely bearded, more inflated; carvopsis 1.4 to 1.6 mm long, margins, sides, and keel glabrous except coarse- 0.5 to 0.7 mm wide, ellipsoid. ly scabrous near the apex, awn .30 to .35 mm Onlv two species of Chloris—C. (fuitvjuese- long, subtended by two short teeth; sterile tica and C. montana—\\Ave a dense fcltlike pu- florets two or three, lowermost sterile floret bescence just below the points of attachment narrowly cylindrical, 2.0 to 2.7 mm long, 0.3 of the spikes. Chloris quinquesetica usuallv has to 0.4 mm wide, glabrous below, sparsely four sterile florets (occasionally three); the scabrous above, gradually narrowing into the lowermost sterile floret is ciliate along the awn, awn 20 to 35 mm long, subtended by a upper margin, and the upper sterile floret is pair of deltoid teeth, rachilla joint well de- spherical and inflated (Fig. 25, A-D). On the veloped; upper sterile florets progressively re- other hand, Chloris inontana usually has three duced; car\()psis ca 3 mm long, ca 0.7 mm sterile florets. These are conunonK' all glabrous, wide, dorsal!)- flattened, shallowly trigonous. though occasionally sparsely pilose; and the

Chloris longiaristata is most closely related upper sterile floret resembles the lowest and to C. prieiirii, from which it mav be separated is not spherical and inflated (Fig. 27, F-I). A by its ven,- long awns, longer than those on comparison of C. qiiinqiiesetica and similar any other species of Chloris, and lack of ciliate species is shown in Table 8. hairs on the upper lemma margins. Specimens examined: INDIA: Kistna Dtr. Specimens examined: TANZANIA: Mwanza, Vijayarion, Barber 8,243 (K); Kannefalli, Lloyd 4 (K). Bourne s.n., without date, (K); Juhi, near Cawn- porc, Diithie 7,7.59 (K); Cawnpore. Makldom- pur, Mohbs s.n., Aug 19.33 (K); Sindi, Hooper 16. CHLORIS QUINQUESETICA Bhide, Jour. s.n., without date (K); Madh Island, Santapau Proc. Asiatic Soc. Bengal n.s. 8:311. pi. 24. 21,284 (K); Andra Pradesh, Secunderabad, 1912. (HOLOTYPE: "Papadi Bassein, without collector or date (K). growing on the bounds of rice fields in semisalt lands. R. W. Bhide. .30-8-191 1." 17. CHLORIS WIGHTIANA Nees ex Steudel, K! ) Fig. 25, A-D; ,30. Syn. PI. Glum. 1:206. 1854. (ISOTYPES:

Perennial, culms eri'ct to strongh stolonif- There are three sheets with apparentK' four different collections at erous and rooting at the lower nodes, to .50 mounted upon them K!, cm tall; sheaths glabrous; ligule a tuft of hairs; among which are undoubtedly duplicates of the synt\pes cited Nees and, blades to 15 cm long, 2 to 4 mm wide; upper bv surface glabrous to scabrous, lower surface subsequently, Steudel. Among these is a glabrous to loosely pilose; spikes three to eight specimen labeled "Herb. Wight. 1766. (rarely two, occasionally to eighteen, as up Chloris Nxightiana NE. Wall s.n. .3809." The indicated in original description), only slightK' label corresponds in many respects to the divergent, 1.5 to 5.0 long, cm decunent at the specimen cited in the original description. base united into a axis and common bearing Fig. 26. a few spikelets; culms villous for 5 to 6 mm Chloris incompleta Wight ex Steudel, Svn. below the spikes; spikelets densely imbricate, PI. Glum. 1:206. 1854. Pro sijn. Non'C. averaging ca 20 per cm on the prominenth incompleta RotJi, 1821. hirsute rachis; glumes linear-lanceolate to ovate, membranous, glabrous except for the scabrous Tufted to somewhat stoloniferous perennial, midnei-ve; first glume 1.5 to 1.8 mm long, 0.2 10 to 30 cm tall; sheaths densely pustulose- to 0.3 mm wide; second glume 2.2 to 2. .5 mm pilose; ligule a densely ciliate crown; blades up long, 0.5 to 0.6 mm wide; fertile lemma 2.0 to 4 cm long, ca 4 to 6 mm wide, densely pustu- to 3.1 mm long, 0.8 to 0.9 mm wide, ovate, lose-pilose on both surfaces; spikes three or four, keel and lateral nerves usually appressed-pubes- 1.0 to 2.5 em long, appressed or onlv slightly Chloris Cr.^mine.\e) 49 Biological Series, Vol. 19, No. 2 Taxonomy of the Genus (

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Fig. 26. Chlom wightianii. (A) habit, \ 1/4; (B) leaf (left) and enlargement of blade (right) showing dorsal pustulose-ba.sed hairs; (C) spilcelet, partly dissected, x 10; (D) sterile florets, x 15; (E) caryopsis, view (left) and lateral view (right), x 10. 51 Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Ciiloris (Gramineae)

perennial divergent; spikeli't.s clen.sel\- imbricate, .spread- Stoloniferous, occasionally tufted, ing widely from the pii.stulose-pilose rachis, ca from 20-50 cm tall; sheaths glabrous; ligule a 12 per cm of rachis length; glumes broadly short ciliate crown; blades up to 12 cm long, lanceolate, thin, membranous, glabrous except 2-4 mm wide, glabrous below, glabrous to scab- for the midnei-ve, which is sometimes scabrous; rous above; spikes two to five, 1.5 to 6.5 cm first glume 2.0 to 2.5 mm long, 0.2 to 0.3 mm long, slightly divergent; upper culms puberu- wide; second glume 2 to 3 mm long, 0.3 to 0.4 lent; spikelets densely inserted, appressed, av- mm wide; fertile lemma 2.7 to 3.1 mm long, eraging ca 15 per cm of the scabrous to sparsely 0.9 to 1.0 mm wide, elliptic, callus, margins, pilose rachis; glumes narrowly lanceolate, pale, and keel denseh' and prominently spreading- translucent, glabrous except for the scabrous pilose, lemma sides pustulose, awn 7.5 to 10.0 midnerve; first glume 1.5 to 1.7 mm long, ca mm long; sterile florets three or four, lowest 0.2 mm wide; second glume 2.2 to 2.7 mm long, sterile floret broadly triangular, 1.4 to 1.5 mm 0.2 to 0.4 mm wide; fertile lemma 2.1 to 3.5 long, 1.9 to 2.0 mm wide, pustulose, midrib mm long, 0.5 to 0.7 mm wide, elliptic, upper densely pilose, awn 6 to 7 mm long, upper margins ciliate with hairs up to 1 mm long, sterile florets progressively reduced; caryopsis keel glabrous, sides glabrous to sparsely ap- ca 1.3 mm long, ca 0.9 mm wide, ca 0.5 mm pressed-pilose, callus bearded, awn 4.2 to 9.5 mm flattened. thick, ovoid-elliptic, strongly dorsally long; sterile florets usually three (rarely two or Ciiloris wigJitiami may be easily separated four); lowest sterile floret 0.9 to 1.7 mm long, from all other species in the genus by its pustu- 0.4 to 0.6 mm wide, truncate, usually glabrous, lose pubescence; this is especially noticeable on rarely with a few appressed hairs in the mid- the sheaths and blades. Only Ciiloris arcnaria nerve region, awn 3.9 to 9.5 mm long; upper is as pubescent, but it is not pustulose. A com- sterile florets progressively reduced, temiinal parison with other similar species is presented sterile floret elongated to subspherical, usually in Table S. only slightly inflated; caryopsis 1.7 to 2.0 mm Only two collections in addition to the iso- long, ca 0.3 to 0.4 mm wide, ellipsoid, trigonous. types mentioned above were studied in the Some specimens of Chloris montana have the herbaria. One label indicated the plant was sterile lemma apex quite ciliate, though not as collected on alkaline soil. densely as C. quinqttesetica, which they may Specimens examined: INDIA; Peninsula In- resemble in other respects (Figs. 27, F-I; 25, diae Orientalis, Wig/if without precise localit)', AD). date, or collection number (K); Tinnevelly Dtr., Occasional specimens may approach C. Tamparaparani Ri\er delta, Lumlararaj v.n., 28 inflata. particularly in the number of sterile flo- Dec 1943 (K). rets. Chloris montana usually has three sterile florets, though sometimes only two. Likewise,

IS. CHLORIS MONTANA Roxburgh, Flora (.'. inflata usualh has only two sterile florets, Indica ed. Care>- 1:331. 1820. (T\pe not though some plants from the South Pacific found. There is, however, a colored plate island archipelagoes regularly have three. How-

at K!, number 882 in the Roxburgh Flora ever, C. montana is a stoloniferous perennial with Indica drawings labeled "Chloris montana puberulous upper culms and without inflated Roxb." This plate clearly shows a villous upper sterile florets (Fig. 27, F-I). Chloris in-

upper culm and a good spikelet dissection, flata is usualh' a tufted annual though oceasion- leaving no question about the plant to which alh stoloniferous, and has glabrous upper

Roxburgh was referring.) Fig. 27, F-I, 28. euims and inflated upper sterile florets ( Fig. .30, Andropo'^on tefru-aristiitiis Roxburgh, ex A-M). Additional differences between these Hooker, ' Flora British India 7:292.' 1896. and other similar species are presented in Pro si/n. Table 8. Representative specimens examined: INDIA: Chloris montana Roxburgh var. glatica Nellore Dtr., Tada, Bourne 2,471 (K); Kistna Hooker f. in Trimen, Flora Ceylon 5:276 Gamble 17,396 (K); 1900. (ISOTYPE?: "from the herbarium. Dtr., Manilipatain, Madras, Dtr.: Dugarazpatam, Gamble 20,389 Ceylon. Jaffna, abundant. H. Trimen 22. 2 Madras (K); Adzar, Gamble 20,784 (K); Chingleput ii 90." US! The collection data correspond Chitoor with those given in the original description, Dtr., Aradi, anon. IS Sept 1917 (K); or date but no collection number or date was gi\en. Dtr., Panappakkam, without collector s.n., (K). The plant is small, possibly only a seasonal (K); Pondicherry, Perrottet 1839 dwarf; no spikelet differences were noted.) MAURITIUS: Round Island, Barkley?, without 52 BniGHAM Young University Science Bulletin

partly dissected, 17. Chloris bournei and C. montana. ( A-E ) C. bouniei. (A) habit, x 1/4; (B) spikelet, habit, 10; (C,D) sterile florets, showing variation, x 10; (E) caryopsis, x 10. ( F-I ) C. montana. (¥) 1/4; (G) spikelet, partly dissected, x 10; (H) sterile florets', dissected, x 10; (1) caryopsis, x 15. I

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae) 53

(Table 8; Fig. 27, A-E). Also, Chloris bournei

is a stoloniferous perennial, while C. inflata is a tufted or stoloniferous annual. Representative specimens examined: INDIA: Coimbatore Dtr.: Talamalai Forest Reserve, Jacob 100 (K); Coimbatore Botanical Garden, Janaki-Ammal 1,172 (K); Bombay, Hubb, Koelz 19,901 (US); Arcot Dtr.: Ammapettai, banks of Uppanera, Narayanaswami 4,171 (K); Tanjore Dtr.: Ponnakani Meda, Narayanaswami 5,141 (K); Bellary Dtr.: Tornagallu, Rangachari s.n., II Aug 1901 (K); Madras Presidency: near Ah- mednagar, Shattuck s.n., (K, US 879426); prope Palamaltah, Wight 3,316 (US); Godaveri Dtr.: Samalkot, anon., 19 Jan 1918 (K).

20. CHLORIS INFL.\TA Link, Enum. PI. 1:105. 1821. (FRAGMENT OF TYPE?: "Chloris

Fig. 28. Distribution of Chloris montana. Inset A: inflata Link. Hb. Link. Sem. ex California." Mauritius, Reunion, and Round Island. US! It is uncertain whether this is a fragment of the type, but the original descrip- collection number or date (K). THAILAND: tion does refer to seed having been sent by HuaHin, Kerr 13,477 (K). Dr. Chamisso and that the origin was Cah- fomia. The locality must be an error. For- 19. CHLORIS BOURNEI Raiigachariar and tunately, the original description clearly ap- Tadulingam, Jour. Indian Hot. Soc. 2:189. plies to the species described below.) Fig. 1921. (TYPE: MH, not seen. Original 30, A-M. description and accompanying illustration Andropogon barJiatum Linnaeus, Mantissa arc clear. Fig. 27, A-E. 302. 1771. Noil A. barbatum Linnaetis Syst. Nat. ed. 10. 2:1305. 1759. (For a full ex- Stoloniferous perennial up to 1 m tall, with planation, see below and also under Chloris geniculate culms ascending from a tufted l)ase; (landi/ana.) sheaths glabrous, ciliate near the membranous ligule; blades up to 10 cm long, 5 mm wide, glabrous, scabrous, or sparsely pilose above, pilose on lower surface, especially at base; spikes three to nine, 6 to 9 cm long; spikelets imbricate, ca six per cm of the scabrous rachis; glumes broadly lanceolate to ovate, glabrous and membranous except for the scabrous to hispid midnerve; first glume 1.8 to 2.5 mm long, 0.3 to 0.4 mm wide; second glume 3.2 to 3.7 mm long, 0.4 to 0.5 mm wide; fertile lemma elliptic to obovate, 3.0 to 3.7 mm long, 0.8 to 1.3 mm wide, densely appressed-pubescent along callus, margins, and keel, awn 3.5 to 7.0 mm long; sterile florets two or three; lowest sterile floret obovate, 1.5 to 2.8 mm long, ca 1 mm wide, glabrous to sparsely pubescent on the midnerve, • = BOURNEI somewhat inflated, often containing rudiments A=QUINQUESETICA of the palea, stamens, and pistil, occasionally apparently fertile, somewhat inflated, awn 3 to 4 mm long; caryopsis ca 1.7 mm long, 0.7 mm wide, ellipsoid-ovoid; chromosome number 2n =40. ChloTis bournei and C. inflata are similar, Fig. 29. Distribution of Chloris bournei (circles and though C. bournei has much larger spikelets C. quinquesetica (triangles). 54 Bricham Young University Science Bulletin

Fig. 30. Chloris inflata and C. formosana. (A-M) C. inflata. (A) hubit, x 1/3; (B) spikelet, partly dissected, X 10; (C-G) fertile florets, showing variation, x 10; (H-L) sterile florets, showing variation, x 10; (M) caryopsis, x 10. (N-Y) C. formosana. (N) spikelet. partly dissected, x 10; (O-S) fertile florets, showing variation, X 10; (T-X) sterile florets, showing variation, x 10; (Y) caryopsis, x 15. )

Biological Series, Vol. 19, No. 2 Ta.xonomv of the Genus Chloris (Gramineae) 55

Chloris harbata (Linnaeus) Swartz, Flora tin

Annual 15 to 95 cm tall, erect to decumbent comparison of this and other related species. and rooting at the lower nodes; sheatlis gla- Considerable confusion has been evident as brous; ligule short, pilose, occasionally mcreh- to whether Chloris harbata (Linnaeus) Swartz, shortly cihate; blades flat, up to 15 cm long, C. inflata Link, or C. paraguatjensis Steudel 0.3 to 0.6 mm wide, glabrous, occasionally pi- should be the accepted binomial. Certainly, the lose at the base; spikes seven to fifteen, 3 to 8 last name is predated by the other two and must cm long, digitate, ± erect, often somewhat be rejected. A full discussion of the relationship fk'xuous and purplish; spikelets densely imbri- of Andropogon harhatum Linnaeus (both 1759 cate, about 14 per cm of the scabrous rachis; and 1771), Chloris polydactyla (Linnaeus) glumes lanceolate to narrowly ovate, hyaline, Swartz, and Chloris harbata (Linnaeus) Swartz, glabrous except for the scabrous midnerve; has been presented in the treatment of C first glume 1.2 to 2.1 mm long, 0.1 to 0.3 mm dandt/ana and should be consulted (p. 34). wide; second glume 2.3 to 2.7 mm long, 0.2 to Essentially, I am rejecting C. harbata (Lin- 0.3 mm wide; fertile lemma 2.0 to 2.7 mm long, naeus) Swartz, for the basionym (Andropogon 0.4 to 0.7 mm wide, oxate to elliptic, callus and barbatum Linnaeus, 1771) is a later homonym. upper margins spreading to appressed-pilose, Swartz clearly referred to Andropogon barbatum with hairs up to 1 mm long, keel glabrous to Linnaeus, 1771, when making the transfer; and pilose, sides of lemma occasionallv sparsel\ ap- thus, C. harbata cannot be accepted as a new pressed-pilose, awn 4.0 to 7.7 mm long; sterile name as proposed by previous authors. The florets usuall)' two (rarel\- three), lowest sterile next available name is Chloris inflata. floret 0.9 to 1.3 mm long, 0.4 to 0.9 mm wide, Chloris inflata has been collected in a wide narrowly turbinate, tnmcate, inflated, usually variety of habitats throughout the warm temper- glabrous, occasionalh' sparsely appressed-pilose ate, subtropical, and tropical regions of the on back, awn 5 to 7 mm long; upper sterile floret world (Fig. 31). Chloris inflata is often a obovoid to subspherical, markedly inflated, \veed in waste areas, but it may be found also about % the lengtli and width of the lower; in cultivated fields, along beaches, etc. It ap- caiyopsis 1.1 to 1.4 mm long, 0.4 to 0.5 mm pears to be very salt-tolerant: one gathering wide, ellipsoid to obovoid; chromosome num- from La Salinas, Dominican Republic {Howard ber 2n = 20, 40, ca 50. and Howard 8,411, NY! US!), was growing on Chloris inflata varies greatly in several char- pure salt rock. Still other collections are from acters, including size, growth habit, spikelet sand dunes, loams, or limestone derivatives. pubescence, and number of sterile florets. Ni'i- Representative specimens examined: (Note:

ther discernible geographic patterns nor con- Chloris inflata is very well represented in major sistent correlation of characters can be dis- herbaria and over 700 individual specimens 56 Brigham Young Universiti- Science Bulletin

Fig. 31. Distribution of Chloris inflata. Inset A; Hawaiian Islands were examined. The following list represents namo Bay, Britlon 2.0.32 (NY, US); Isla only a small fraction of these collections. A com- de Pinos, Curtiss s.n., Apr. 1904 (NY). plete list is available from the author in mimeo- CURASAO: near Willemstad, Curran and Ila- graphed form.) ADMIRALTY LSLANDS: Bis- man 63 (NY, US). DOMINICA ISLAND: near marck Arch., Goodenough Island, Milne Bay Morne Daniel, Hodge 3.893 (NY). DOMINI- Dtr., Brass 24,419 (K).' ANTIGUA: near St. CAN REPUBLIC: Prov. Barahona: Neiba Val- John, Rose et al. .3,377 (NY, US). ARGEN- ley, Howard and Howard 8,360 (NY, US); Prov. TINA: Buenos Aires, Isla Manel, Parodi 740 Monte Cristo: Guayubin, Abbott 876, 878 (US). AUSTRALIA: Northern Territory: near (US); Prov. Santo Domingo: vicinity of Ciudad Darwin, Black 1:270.200 (K); Queensland: Trujillo, A//«rf/ 14,626 (NY). FERNANDO DE South Kennedy Dtr.: 1.3 mi SE of Eaglefield NORONHA: Quixaba, Cutjler 9,395 (US). FIJI Station, Adams L079 (K). BAHAMA IS- ISLANDS: Viti Levu Island, Greenwood 1,213 LANDS: Giaiul Turk Island, Millsixiii^h and 1.308 (US). FRENCH GUIANA: Cayenne, Millspaugh 9,002 (F, NY); Inagua, Hitchcock lloock s.n., 2 Jun 1962 (NY). GHANA': near s.n., 3 Dec 1890 (F, MO). BARBADOS: Bridge- Labadi, Adams 3,835 (K). GRAND CAYMAN: ton, F. W. Jolnison 1,194 (NY). BORNEO: without precise locatitm, Moggs 61 (K). GREN- Balikpapan, Frost i (K). BRAZIL: Bahia: 46 ADINES: Petit Martinicjue, Howard 10.917 km VVSW of Joazeiro, A. Chase 7.939 ( F, US); (NY). GUADELOUPE: Basse Terre, Dms Pernambuco: Bello Jardim, Serra do Genipapo, 3,158 (NY, US). HAITI: Gonave Island, vicinity A. Chase 7.698 (F, MO, US). BRITISH of Etroite, Leonard 3,339a {¥, NY, US); Tortue GUIANA: Georgetown, Graham 345 (K). BUR- Island, vicinity of LaVallec, Leonard and MA: Mandalav, White 132-237 (US); Sagaing, Leonard 11.262 (UC). HONDURAS: Dep.

Huk s.n., 5 July 1890 (K). CANTON ISLAND: Atlantida, Standleij 56,587 ( F, US). INDIA: without precise location, Degener and Hathaway Madura Dtr.: Pulnev Hills, Dindigal. Anglade 21,251 (MO, NY). CELEBES ISLAND: Mana- 1.109 (K); Chingleput Dtr.: Madnis, Gamble s.n., 18.36-.38 do, Egvjan 1.732 ( K, US). CEYLON; Trincoma- 16,415 (K); Calcutta, Heifer (BR, lee coast, Kingdon-Ward 23,064 (K). COCOS NY); Coimbatore Dtr.: Anai Katti, Rapi and KEELING ISLAND: St. John 26,465A (K). Naganathan 4,802 (K). IVORY COAST: COLOMBIA: Dep. Atlantico: entre Palmar de Abidjan Plateau, Adjanohoun 428A (K). Varelay Pondera, Dttgand 4.42S (US). CUBA: JAMAICA: between Portland Point and Havana Prov.: without precise location. Baker Rocky Point, limestone plain, Britton 1.903 1,441 (F, US); Oriente Prov.: Guanta- (NY); Kingston, Harris 9,049 (F, K, NY, US). BiOLOcicAi. Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 57

JAVA: Ambarawa, Brooks s.n., Mar 1909 (BM). TRINIDAD: Woodbrook, Broadwai/ 7.177 ( F, KENYA: Voi Dtr., Tsavo National Park, E of MO). UNITED STATES: Hawaii: Oahu, N of Lugard Fall.s, Grceiucai/ ami Kamiri 12.S74 Koko Head, Degener 10,835 (NY); Texas: (K); Kwale, SW of Mombasa, Bogdan AB Hidalgo Co., McAllen, roadside, Anderson 3,194 (K). LINE ISLANDS: Christmas Island, 3,875 \USC). VENEZUELA: Lara, Palmasola, Pearson P22 (K). LIBERIA: Cape Palmas, Pittier 6,385 (US). VIRGIN ISLANDS: St. Hale 157 (US). MACAU: vacant lot, Hitchcock Croix, Thompson, 5 Oct 1921 (NY, US); St. 19,153 (US). MALAGASY REPUBLIC: No,ssi Thomas, Hitchcock 16.320 (US); Tortola, Velez Be, Hildchramit 2.939 (US), 2,932 (K). MA- 3.123 (K). WAKE ISLAND: Peale Islet, Fos- LAYSIA: Singapore, Bor S61 (K); Selonger berg 34.928 (US). WEST IRIAN (IRIAN State, Port Swettenham, BuckiU 12,839 (K). BARAT): Merauke River, Kelapi Lima, McKee MARSHALL ISLANDS: Guam, Fosberg 25,334, 1.705 (K). ZANZIBAR: Cooke s.n.. 1861 (MO). 31,223 (US); Pagan Island, Bonham 33 (US); Saipan Island, Fosberg 31,287 (US); Tinian Is- 21. CHLORIS FORMOSANA (Honda) Keng, land, Fosberg 24,798 ( US ) ; Bikini Atoll, Taiilor Claves Gen. Sp. Gramineamm Primarum 1,500 (UC); Eniwctok Atoll, Fosberg 24.407 Sinicanun 197. 1957. (Based on Chloris (US); Kwajalein Atoll, Fos]>erg 31,202 (US); barbata Swartz var. jormosana Honda.) Fig. Majura Atoll, Fosl>rrg 31.193 (US). MAR- 30, N-Y. TINIQUE: St. Anne, S of Dunkertiue- Swartz \ar. formosana Catherini' mangrove, Ei^lcr 39—73 (NY). Chloris harhata Bot. 40:4.37. 1926. MAURITIUS: Battc-rie Dauphine, Saner 2.701 Honda, Mag. Toho 'Takao, G. Naka- (F). ME.\IC:0: Tamaulipas: N of Ciudad (SYNTYPES: "Taiwan: 1905." Mante, Beetle M-521 (UC); Yucatan: Me- hara, no. 6.35, anno "Baehu, B. Ha- anno 1919." Not seen. Description re- rida, Swallen s.n.. Aug 1932 (US). MOLLUCA yata ISLANDS (HALMAHERA): Morotai, Main fers to species as commonl)' understood.) Aden 1.477 (US). and NEW CALEDONIA: Perennial often stoloniferous, 30-75 cm tall, Anse Vata, 4.063 McKee (K). NEW PROVI- erect to decumbent; sheaths glabrous, rarely Nassau, 61 DENCE: Matthews (K). NIGERIA: sparsely pilose; ligule a very short ciliate fringe; Lagos, Ward 9 (K). Ara- PALAU ISLAND: i)lades up to 20 em long, usuaii\' folded, gla- kabesan Island, Ilosaka Peleliu Is- 3.354 (US); brous, acute; spikes five to nine, 4 to 8 cm long, land, Fosberg 31,952 (US). PANAMA: near erect to somewhat lax; spikelets densely imbri- Miraflores, Locks, Allen 1,752 (F, K, MO, NY). cate, ca fourteen per cm of the pilose rachis; PAPUA: Daru Island, Brass 6.403 ( BM, US); glumes lanceolate, glabrous, except for the Fairfax, 9°55'S, 147°20'E, Gillison 22,211 NGF scabrous midnerve; first glume 1.0 to 1.5 mm (K); base of Mt. Lawes, Eucalyptus savan- SW long, ca 0.2 mm wide, acute; second glume nah, Sclwdde 2.644 ( US). IS- K, PEMBA 2.2 to 2.5 mm long, 0.2 to 0.3 mm wide, apex LAND: Vaughn 252 IS- (BM). PHILIPPINE obtuse, mucronate; fertile lemma 2.3 to 3.0 mm LANDS: Luzon: Manila, Clemens s.n., 1924 Sep long, 0.5 to 0.7 mm wide, with a lateral groove, (UC, US); Corregidor, Cunung 696 (K, MO). which is occasionally appressed-pilose, upper RICO: Island! Sardinera, PUERTO Mona Brit- margins denseh' pilose with hairs up to ca 1 ton et al. 1.831 (NY, Isla US); de Culebra, mm long, keel glabrous, slightly gibbous, awn Brown and Wheeler 141 (NY, US); Guanica 2.0 to 5.7 mm long; sterile florets two; lower Bav, A. Chase 6,529 (US). RYUKYU ISLANDS: sterile floret oblanceolate to truncate, 1.6 to Okinawa, Amano 6,800 ST. (US). KITTS: 1.9 mm long, 0.6 to 0.7 mm wide, flattened or Hitchcock 16,369 (US). ST. LUCIA: Brooks 20 only slightl\- inflated, awn 2.3 to 5.0 mm long; (US). SAMOA EAST: Tutuila Island, Collarnio upper sterile floret of same shape as lower, but 508 (US). SERAM (CERAM) ISLAND: Am- smaller, 0.5 to 0.7 mm long, 0.4 to 0.5 mm wide, boina, Robinson 1,647 (BM). SEYCHELLES glabrous, awn 1.5 to 2.5 mm long; car\'opsis ISLANDS: Mahe Island, Boivin s.n., 17 Nov 1.2 to 1.6 mm long, 0.3 to 0.4 mm wide, ellip- 1855 (K). TAIWAN: Chiayi, Devol 7,095 (UC). soid. TANZANIA: Dar Es Salaam, Hitchcock 24.424 Chloris formosana is most easily confused (US); Tanga Pro\-.: Sakura Sec: Panxani with C. inflata. The following key will separate Dtr., Tanner 2,027 (K, UC). THAILAND: the two: Bangkok, Kerr 3,761, 7.853 (BM); Wangka, Kwae Noi River basin, Wichian 330 (K)."tI- 1. Lower sterile floret 1.6 to 1.9 mm long, MOR: Kupang, R. Brown s.n.. Apr 1S0.3 (BM). flattened or only slightb' inflated; upper TOBAGO: Spexside, Purseglove P. 6,358 (K). sterile floret about one-third the size of )

58 Brigham Young University Science Bulletin

the lower, scarcely inflated; upper Chloris breviseta Bentham in Hooker f., sheaths, ligules, and Iowit portions of Niger Flora 566. 1S49. (SYNTYPES: "Cape leaves glabrous, rarely sparsely pilose; Coast, Don" BM! "Cape Coast, Vogel" K!) perennials. Chloris nigra Hackel. Bol. Soc. Broteriana C. fonnosana (Fig. 30, N-Y) 21:179. 1908. (TYPE: "Insula S. Jacobi Prom Viridis, pr. Trinidade, leg. A. Bar- 1. Lower sterile floret 0.9 to 1.3 mm long, jona" not seen, but description is clear.) usually inflated; upper sterile floret two- var. breviseta (Ben- thirds to aljout the same size as the Chloris virgata Swartz lower, usually inflated and spherical; up- tham) Pilg(>r e.x Peter, Beih. Repert. Sp. Cliloris per sheaths, ligule, and lower portions of Nov. 40:262. 1931. (Based on the leaves usuallv loosely pilose, though breviseta Bentham. sometimes sparsely so; annuals, occasion- Annual often weakly rooted, sometimes ally rooting at the lower nodes. shortly stoloniferous, erect or in stoloniferous C. inflata (Fig. 30, A-M) specimens somewhat decumbent, usually 30 to 70 cm tall, occasionalK' to 2 in; sheaths gla- Other species that might be confused with brous to denscK' pilose; ligule densely and prom- C. formosana are compared in Table 8. inently pilose; blades 2 to 10 mm wide, up to Collected from sandy or gravelly soils, often .30 cm long, scabrous; spikes five to nine, 3 near the ocean in China, Hong Kong, Taiwan, to "^j cm long; spikelets loosely imbricate, ca five and Vietnam (Fig. 32). to seven per cm of the scabrous rachis; glumes Representative specimens examined: CHINA: membranous, une(jual; first glume 1.1 to 1.6 Fukien Prov.: Amoy, Chung 6,055 (US), Price mm long, 0.2 to 0.3 mm wide, narrowly lanceo- 1,363 (K); Kwangtung Prov.: Taio, Chun 3,- late, glabrous except for the scabrous mid- 114B, 3,123 (NY); Pakhoi, Hitchcock 19,255 nerve; second glume 1.9 to 2.3 mm long, 0.3 to (US); Hainan Island: Hoihow, Hitchcock 0.4 mm wide, glabrous with scabrous midnerve, 19,547 (US), McChire 10,451 (BM, K), Tsan^i rounded at the apex and bearing an awn ca 0.3 986 (NY). HONG KONG: Bor H.K. 76 (K), mm long; fi'rtile lemma 2.3 to 3.5 mm long, Ford s.n., 25 Aug 1893 (K, NY), Hance 1.943 0.7 to 1.5 mm wide, broadly ovate or elliptic (BM); Quarry "Bay, Sampson 851 (BM). in overall outline, prominently gibbous on the NORTH VIETNAM: Tonkin, route de Haiphong keel; var\ing from pale to dark gra\ or black, a Doson, bords des chemins, Petelot 425 (NY, often mottled when mature, callus bearded, US). TAIWAN: Takao, Henri/ s.n., 1S93-4 sides with a prominent glabrous or appressed- (BM), Henry 1,023 (K), Plaijfair' 53 (K); with- pilose furrow, apex acute to blunt, mucronate, out precise location, Tonaka 10.352 (BM NY or with an awn up to 6 mm long; steiile florets UC). two (very rarely one) glabrous, becoming scabrous at the apex, lower sterile floret 1.5 to 22. CHLORIS PILOSA Schumacher, Beskr. 2.2 mm long, 0.7 to 1.1 mm wide, narrowed at Guinciske Planter 75. 1827. (PHOTO- the base, becoming inflated at the truncate GRAPHS OF HOLOTYPE: K! US! Type apex, glabrous below, apex scabrous, mucronate, specimen, in Isert and Thonning Herbarium, or \\ ith an awn up to 3.0 mm long, upper sterile C, has mucronate lemmas.) Fig. .33, E-K. floret a triangular inflated rudiment less than 1 mm long, ca 1 mm wide, usually hidden in the lower sterile floret, awnless; anthers ca 0.4 to 0.5 mm long; caryopsis 1.3 to 1.5 mm long, ca 0.5 to 0.6 mm wide, obovoid, trigonous; chromosome number 2n = 20, 30. Awn length in this species varies widely. Most specimens fall in one of two groups, those with awns less than 1 nun long, generally only mucronate (represented on the map in Fig. 34 by unshaded circles) and those with longer awns, usually from 2 to 6 mm (indicated on the map by shaded circles). No strong correlations of awn length can be made with geo-

Fig. 32. Distribution of Chlaris formosana (southeast graphic origin, nor with other morphological China, Hainan, and Taiwan). characters. While the variation in awn length BiOLOCiCAi. Series. Vol. 19. No. Taxonomy ci- the Genus Ciilohis (Gramineae) 59

Fig. 33. Chloris cnstilloniuud and C. pilosa. (A-D) C. castilloniami. (A) habit, x 1/5; (B) spikelet, partly dis-

sected, .\ 10; (C) sterile floret, \ 15; (D) caryopsis, \ 10. (E-K) C. pilosa. (E) habit, .x 1/3; (F) spikelet of awned variant, partly dissected, x 10; (G) spikelet of unawned variant, x 10; (H) sterile florets of a\vned variant, x 15; (I) sterile florets of unawned variant, x 15; (J) upper sterile floret, x 15; (K) caryopsis, x 10. ) )

60 Bbicham Vounc University Science Bulletin

produces plants with somewhat diverse appear- Prov. Oyo; Ibadan, Brenan 9,601 (K, US); ances, tlieir nomenclatural recognition is not Prov. Zaria: Samaru, Freeman S. 116 (K); Jebba,

warranted. llagerup 739 ( Plateau, along De- K, US ) ; Jos Chloris pilosa most closely resembles C. vir- limi River, Naraguta, Laivlor and Hall FHl

gata, from which it may be separated in having 46,540 (K); Toro, 20 mi S of Jos. Semple 176 prominently pilose ligules and broader lemmas (US). REPUBLIC OF THE CONGO: Musa, which have shorter marginal cilia. De Giorgi 1,299 (K); Eala, Germain 1,695 (K); Kasenye, Johnston 1,042A (K); Bukama, Widely distributed in equatorial Africa, Shantz .565 '(K). SUDAN: Cordofan Prov.: El Chloris pilosa has been collected in weedy areas Obeid, Harrison 75 (K); Darfur Prov.: Jebel and as a native in savannahs. It is occasionalh' Marra, Wickens 2,279 (K); Equatoria Prov.: cultivated as a forage grass. The natural distri- Lado Dtr., Cartwright 27 (K); Upper Nile bution is shown in Fig. 34. Prov.: Zerat Island, Harrison 1,005 (K). SIER- Uepresentative specimens examined: BEL- RA LEONE: Mano, Deighton 674 (K); Musaia, GIUM (adventive): Dunquenjue, anon. (US). Deighton 5,386 (K); Freetown, Deighton 2,029 CAMER(HTN: Boro, 35 km ENE of Maroua, (K). SENEGAL: Dakar, Baldwin 5,714 (K). Bounougou 13 (K); Darga, 30 km ESE of Ma- TANZANIA: Tanga Dtr.: Kange Estate, Faulk- roua, Koechlin s.n., May 1965 (K); Victoria Dtr.; ner 808 (K); Rufiji Dtr.: Utete boma, Milne- roadsides, weedy, Maitland 9 (K). CAPE Redhead and Taylor 7,537 (K); Morogoro Dtr.: VERDE ISLANDS: San Nicalao, Loive s.n., 22 Duturi, Nicholson 29 (K). TOGO: Palime, Feb. 1864 (BM); Sao Tiago, Grandvaux-Bar- Stage 73 (K). UGANDA: Buganda Dtr., hosa, 5,755 (K). CONGO: LeopoldviHc, Fas- Hitchcock 24,934 (K); Lango Dtr.: Amugo, seaux 1,019 (K). DAHOMEY: Parakou, Ris- ]ohn.ston 1,037 (K); Ankole Dtr.: Kiruhara, opoulos 1,197 (K). ETHIOPIA: Serilia, Ghattas. Thornton 39 (K); near Mt. Elgon, Snowden ^chucinfurth 2,283 ( K, US). FERNANDO PO; 1,207 (K). Mann 118 (K). GAMBIA: Sankuli-Kunda, Pirie s.n.. Sept 1933 (K). GHANA: Nungua. Accra Plains, 5°40'N, 0°06'W Ankrah 20,150 23. CHLORIS MRGATA Swartz, Flora Indiae

(K); 2.5 mi SW of Kpetchu Ferr\ . 7°55'N. Oce. 1:203. 1797. ( Holotype not .seen, but 0°22'E, Ankrah 20,387 (K); Baikpa, 6°51'N, description clearly refers to the species as 0°26'E, Rose-Innes 31,189 (K); Axim, Vigne 395 commonh' understood.) Fig. 35. (NY, US). GUINEA: Nzerekore, Bald- win 9,722 (K, US); Kouroussa, Pobequin 546 ? Chloris pubescens Lagasca, Var. Cienc. Lit. Artes (Madrid) 4:143. 1805. Descrip- (K). IVORY COAST: Kouiblv, Adjanolwun tion recopicd in Lagasca, Gen. Sp. Pi. 5 413A (K). LIBERIA: Grand Ba.ssa Co.: Timbo, Nov. 1816. (A specimen, at G!, labeled Baldivin 11,223 (K); 10 mi inland from River "Chloris pubescens Lagasca, ex Peruvia, La- Cess, Baldwin 11.228 (K); Sinoe Co.: Sasstown, gasca 1807" is Chloris virgata. The original Baldwin 11,603 (K). MAURITANIA: Dahr, description is not conclusive, and this name 16°30'N, 7°W, Rossetti 61/172 (K). NIGERIA: is included here provisionall)'. Rahdochloa virgata (Swartz) Beauvois, Ess. Agrost, 84, 158, 176. 1812. (Based on chloris virgata Swartz, but Rahdochloa not x'ahdly published.) Chloris elegans Humboldt, Bonpland,

Kunth, Nov. Gen. .Sp. PI. 1:166. /)/. 49. 1816. (HOLOTYPE: "Mexico, Queretaro. " • SPIKELETS AWNFD f:; • -C v.^ '"Af M..\. Bonpland n. 4194." P! Fragment in O SPIKELETS AWNLESS/ A- '\) °»T^ ORMUCRONATE \ ii<.„_ US! C/(/,)r;.s nlba Prc-sl, Rel. Ilaenke 1:289. 1830.

( Description clearly refers to Chloris virgata. Scribner, 1899, examined and described the presumed type in the Bemhardi Herbarium, saving it was the same as C Fig. 34. Distribution of Chloris pilosa. Dots represent elegans. ) specimens with awned lemmas; open circles represent specimens with unawned or merely mucronate Chloris caudata Trinius ex Bunge, Mem. lemmas. Savans Etr. Acad. Petersburg 2:144. 1835. BiOLOCiCAi. Sehies, Vol. 19, No. 2 T.•\xONo^I^ (U- the Genus Chlouis (Gramineae) 61

Fig. 35. Chloris virgata. (A) habit, .x 1/4; (B) spikelet. partly dissected; (C-F) fertile lemmas, showing lower sterile floret (left) variations; (G-1) lower sterile florets, showing variation; (J) rudimentary palea of and attached upper sterile floret (right); (K) caryopsis, lateral view; (L-N) caryopses, dorsal views, showing variations. (B-N) x 10. ) ) ) ) ) )

62 Brigham Young Universiti' Science Bulletin

(Description, combined with geographical ata (Hochstetter) Chiovenda in Pirotta, citation, "Hab. propre Pekinum," seems Annuario Rc-ale 1st. Bot. Roma 8:54. 1903. to refer to C. virgata. A specimen in the (Based on C. multiradiata Hochstetter). Cosson Hb., at P!, collected by Bunge in Chloris gahriclae Domin, Biblioth. Bot. 85: 1835, and labeled "Chloris caudata," per- .368, Fig. 83. 1915. (HOLOTYPE: "Queens- haps in Bunge script, is C. virgata.) land; am Flinders River bei Hughenden,

haufig, Domin, II. 1910." K!, on extended Chloris penicillata Willdenow ex Steudel, loan from PR.) Nom. Bot. ed. 2. 1:353. 1840. Pro si/n. Chloris rogeoni Chevalier, Rev. Bot. Appl. Chloris decora Nees ex Steudel, Syn. PI. Agric. Trop. 14:127. 19,34. (SYNTYPES: Glum. 1:205. 1854. First published as "Dire. 43,871, El Oualadjii 4,3,937 et 42,- novien nudum by Nees in Royle, Illust. ,357." All except the last at P! Fragments Bot. Himalayan Mts. and Flora Cashmere of last two at K! 1:416. 1840.' (HOLOTYPE: "Rovle Hrbr. Chloris tibestica Quezel, Bull. Soc. Hist. nr. 87" K! Afrique Nord 48:84. 19,57. (TYPE: "Emi Chloris meccana Hochstetter ex Steudel, Koussi," not seen. Description and subse- Syn. PI. Glum. 1:205. 1854. (ISOTYPES: quent illustration in Quezel, 1958, leave no "802. Chloris meccana Hochst. ex Steudel doubt as to identity.) ... d. 11 Febr. 36, leg. W. Schimper" G! US!) Annual, extremely variable in size, ranging from a few centimeters to over a meter; culms Chloris iiwdagascariensis Steudel, Syn. PI.

". usually tufted, several arising from one root Glum. 1:203. 1854. (HOLOTYPE: . . Le ." system, occasionalU' stoloniferous; sheaths usu- Jolis legit 1849 . . P! ally glabrous, occasionalK' rather densely pilose Chloris notocoma Hochstetter, Flora 38:204. toward the apex; ligule glabrous, or with a cili- 1855. (ISOTYPES: "Hb. abyss. Hohenacker ate fringe of hairs up to 4 mm long; blades up nr. 2125" G! P! to 30 cm long and 1.5 cm wide, usually gla- Chloris nmltiradiata Hochstetter, Flora 38: brous witli scabrous margins, occasionally pilose; 204. 1855. (Description is of C. virgata. spikes four to twenty, 5 to 10 cm long, usu- Type, "Hb. abyss. Buch. nr. 486" in P! is ally more or less erect; spikelets densely imbri- C. prietirii, a duplicate at K! is C. virgata. cate, averaging 10 per cm of the scabrous or Chloris alba Presl var. aristulata Torrey, hispid-ciliatc rachis; glumes pale brown, lanceo- Rept. Expl. Survey Mississippi River Pacific A late, glabrous with scabrous midnerves; first

4:155. 1857. ( ' short-awned variant.) glume 1.5 to 2.5 mm long, 0.2 to 0.4 mm wide; Chloris hrachijstacltiis Andersson in PetiTs, second glume 2.5 to 4.3 mm long, 0.3 to 0.5 Naturwiss. Reise Mossambique 556. 1864. mm wide; fertile lemma 2.5 to 4.2 mm long, 0.7 (Based on Chloris alba Presl.) to 1.3 mm wide, pale brown, occasionally Agrostomia barl)ata Cervantes, Naturaleza puiplish or blotched with dark brown spots, (Mexico City) 1:.346. 1870. (TYPE: Cuer- keel usually prominently gibbous, keel, mid- na\aca, Mexico, not seen, but description ner\'es, and lower margins glabrous to promi- fits pilose, long-ciliate, C. virgata. ) nently margins with spread- Chloris barbata var. decora (Nees) Ben- ing hairs near the apex, awn 2.5 to 15.0 mm tham. Flora Australiensis 7:613. 1878. long, usualh' more than 5 mm long; sterile flor- (Based on C. decora Nees.) et one, on robust specimens, occasionally two; lower sterile floret 1.4 to 2.9 mm long, 0.4 to Chloris alberti Regel, Acta Horti Petropoli- 0.8 mm wide, awn ,3.0 to 9.5 mm long; upper tani 7:6,50. 1881. (TYPE: "Mongolia occi- sterile floret, when present, usually borne on a dentali prope Takiansi, A. Regel," not seen. short rachilla and greatU' reduced, sometimes Geographic origin and description seem to absent, leaving only the naked rachilla; caryop- suggest C. virgata.) sis 1.5 to 2.0 mm long, ca 0.5 mm wide, ellip- Chloris tetrastachtjs Hackel ex Hooker f., soid-trigonous; chromosome number 2n=20, Flora British India 7:291. 1896. Pro syn 26. .30, 40. Chloris virgata var. elegans (Humboldt Without doubt, Chloris virgata is the most Bonpland Kunth) Stapf in Thistleton-Dver, variable of all of the annual species in the Flora Capensis 7:642. 1900. (Based on C. genus. Great variation is shown in many traits, elegans. including such vegetative features as height, leaf Chloris pohjdactijla Swartz ssp. nmltiradi- size, tomentum, and habit, as well as spikelet Biological Series, Vol. 19, No. 2 T.axonomy of the Genus Chloris (Gramineae) 63 characters (especially various pubescence on to typical specimens of C. virgata and is consid- the lemma, such as presence or absence of hairs ered here as merely an aberrant individual. on the keels, midnerves, and margins). In addi- Chloris virgata closely resembles C. gaijana tion, the lemmas themselves ma\ be variously and C. pilosa in spikelet characters. It may be gibbous, keeled, or merely folded at the mid- easil)- separated from the former in being an nerve. Only two attributes seem constantly cor- annual and from the latter in having relatively related: (1) the conspicuous tuft ot spreading long awns. hairs on the upper lemma margins and (2) the CIdorif; virgata lias Ijcen widel\- collected appressed to erect or only slightly divergent from many habitats, primarily in warm temper- spikes. ate to tropical areas, but extending well into No well-marked patterns of variation can be temperate areas where hot summers are com- identified, though several somewhat cr)'ptic mon (Fig. 36). Over 1,300 different specimens populations are noticeable. Most specimens were examined in this study. The following list from Australia and some from Asia and Africa represents only a small fraction of the total. have relativel)' narrow lemmas that are not con- Representative specimens examined: AF- spicuously gibbous, though other collections GHANISTAN: Helmand River Valley, irrigated from the same general area may show the field. Long 326 (US). ANGOLA: Kaoko veld, typical prominently gibbous keel. Some of the N of Ohopoho, ca 1S°E x 14°S, de Winter and populations, especially those from Tibet and Leistncr 5,177 (K). ARGENTINA: Prov. Nepal, ma\' eventually prove to be distinct; but Buenos Aires: Pergamino, Parodi 834 (US); presently, so few specimens are available for Piov. Gatamarca: 6 mi SW of Andalgala, Bartlett

study that it seems advisable to treat the entire 20,226 (US); Prov. Cordoba: Dep. dc Santa variable population as a single specie's. By con- Maria, Ihmziker 1.294 (MO); Prov. La Rioja:

trast, western hemisphere material seems to be Patguia, Laliitte ij Castro 2 (US); Prov. Men- quite uniformly gibbous, usualh' conspicuously doza; Santa Rosa, Jen^en-Haartip s.n., 1904-05 so. (US); Prov. Salta: Dep. Rosario de Lerma, Las Occasionalh' novelties, such as a specimen Tres Acquias, Montenegro 455 (US); Prov. Tu- collected by Dr. T. L. Bancroft at Falm Island, cuman: Dep. Trancas, Rio Tipainago, Venturi Queensland (K!), may be very different. This 4,354 (NY, US). AUSTRALIA: Queensland: particular specimen was unique in having dense- Darling Downs, Ashcroft s.n., May 1916 (K);

ly pilose glumes. In other respects it is similar Petrie, Blake 170 (K); Western Australia: Wan-

Fig. 36. Distribution of Chloris virgata. Inset A: Hawaiian Islands 64 Brigham Younc University Science Bulletin

dagee, Minilya River, Gardner 3,234 (K). BO- Lvr\be, Dieterlen 6,326 (K). MEXICO: Aguas- LIVIA: Cocliabamba Dep.: without precise lo- calientes: 2 mi S of Aguascalientes-Zacatecas cation, Btichtien 2,512 (US); Dep. Tarija, near line, Emery 182 (TEX); Baja Cahfornia: Magda- Entre Rios, West S,250 (MO, US). i«)TSWA- lena Island, Brandegee s.n., 21 Jan 1889 (UC); NA: Kgalagedi, without precise location, Leist- 57 km NW of San Ignacio, Carter, Alexander, ner 3,103 (K); Ngwato, Bokalaka area, Francis- and Kellogg 1,950 (K, UC, US); Chihuahua: 2 town, McClintock s.n., 18 Dec 1966 (K). BRA- mi N of Lucero, Weber and Charette 11,604 ZIL: Bahia: near Rio Sao Francisco, Joazeiro, (UC); Coahuila: W of Puerto de las Monjas, A. Chase 7,946 (F, US); Ceara: Campo Salles, I.M. Johnston 8,640 (MO, US); Colima: Alzada, Swallen 4,2HH (US, Piaui: between Floriano and Hitchcock 7.054 (US); Distrito Federal: Ixtapa- Oeiras, Swallen 4,171 (US); Rio Grande do lapan, Matuda 25,685 (MO); Durango: 3 mi N Norte: Santa Cruz, SuaUcn 4,849 (US). BUR- of Donato Guerra, Emery 337 (TEX); Jahsco: MA: Mingaladon, Pokhant 1,375 (K). CHINA: Guadalajara, La Barranca, M.E. Jones s.n., 19 Honan Prov.: Cheng-Chou, Chiao 18,500 (F); Nov 1930 (MO, UC); Guanajuato: Acambaro, Hopeh Prov.: near Peking, Chcn^ 2.038 (US); Hitchcock 6.932 (US); Guerrero: near Iguala, Inner Mongolia: Ordos, Tapiichan, Hsiu II 3,799 Canon de la Mano Negra, Rose 9,385 (NY, US); (US); Kansu Prov.: Yao Kai, near Lichen, Ching Michoacan: 26 mi SE of Sahuayo, Pratt 699 258 (US); Kiangsu Prov.: Nanking, Chiao (TEX); Morelos: Cuernavaca, Hitchcock 6,835 12,916 (K, NY, UC); Manchuria: Darien, /»//«;!« (US); Navarit: Tepic, Acaponeta, Rose 14,248 59a (NY); Harbin, Dorsett and Dorsett s.n.. (NY, US)'; Oaxaca: I.xtepec, Fisher 35,311 (F, (US); Shungari River, Komarov 167 (BM, K); MO, NY); Puebla: Tehuacan, Hitchcock 6,073 Shansi Prov.: Sui Guan Dtr., I Ian Ngan 32 (US); Queretaro: 5 mi from Queretaro, Pratt (UC); Shantung Prov.: Cliingtao, Li Chuan, 774 (TEX); San Luis Potosi: 6 mi S of Mate- Chiao 2,864 (F, K, NY, UC, US); Shensi Prov.: huala, McGregor et al. 516 (US); Sinaloa: San Chou-chih-hsien, Knng K3,734 (US); Yunnan Ignacio, Ortega 4.468 (US); Sonora: 25 mi W Prov. La-Kov, Mairc Apr 1914 (BM, F). CU- of Angostura, Santos 1,831 (F, K); Tamaulipas: RAgAO: Sovonet, Suringar 9 (US). ECUA- near Aldoma, Martinez and Luyando F-2,177 DOR: Prov. Guavas: W of Cluavacjuil, .\sphin'J (TEX); Veracruz: 6 to 7 km N of Tierra Blanca,

' 15,369 (K); Puntilla, Salinas, 2 IS'S, 81 W, Santos 3,313 (NY); Yucatan: Izamal, Gaumer Svenson s.n.. 1 Mar 1941 ( F, MO); Prov. Loja: 1,085 (F, MO, NY, US); Zacatecas: 30 mi N of between La Tonia and Loja, Hitchcock 21,403 Zacatecas, Emery 307 (TEX). MADEIRA IS- (US). EL SALVADOR: vicinity of San Salva- LANDS: near Funchal, Sledge s.n., 14 Apr dor, StancUcij 19.234 (NY). GALAPAGOS IS- 1949 (BM). MALAWI: Zomba, Cormack 176

LANDS: Charles Island, Post Office Ba\ , lloucll (K); Domasi, between Zomba and Ncheu, 8,838 (US); Chatham Island: Wreck Bav, Ilou- Jackson 132 (K). MARIANAS ISLANDS: Sai- eU 8..'i95 (US), Stewart 1,254 (K, MO); Indefati- pan, Kagman Peninsula, Fosberg 31.288 (NY,

gable Island, Howell ; 9,917 ( US ) North Sc'smour US). MAURITANIA: Atar Parielle, Adam

Island, Howell 9.967 ( ; US ) South Se\'mour Is- 21.799-1 (K). MOgAMBIQUE: Prov. Mo9am- land, Howell 9,924 (US). GUATEMALA: near bique: Mocuba, Namozoa, Faulkner 29 (K); Jalapa, Kellerman 7,975 (F, NY); vicinitv of Lourenyo Marques Dtr.: near Costa do Sul, Zacapa, Standlei/ 74.606 (F). HONDURAS: 5 Gomes e Sowm 3. 438 (K). MONGOLIA: Sha- km N of Talaiiga, Johanncssen 892 (UC); Dep. harakh Usu, Chancy 407, 537 (NY, UC, US); Morazan: vicinitv of El Zainorano, Standlei/ s.n.. llOmiNWofHaHin, EnTcsson 478 (US). NE- 27 June 1949 (F). INDIA: Allahabad, Dudgeon PAL: Rohagan, Suli Gad, Polunin. Si/kcs and Wil- s.n., 1 Nov 1919 (MO); Bombay, Gamble 21,- liams 3,37-3. 3.376 (BM). NICARAGUA: road to 028 (K); Central Pro%'.: Nagpur, Haines 3.609 Granada, Hitchcock 8,706 (US). NIGERIA: (K); Assam, 21 27° mi marker on Ledo road, Bornu, Gudumbali, Rains s.n., Oct 1961 (K). 20'12"N, 96°2'55"E, Juan L8C93 (US); Bihar PAPUA: 9 mi NW of Ore Bay, Reeder 831 Prov.: Ranka Koelz 18,971 (K, US); Shi- (US). PARAGUAY: chaco boreal, Ro/os 7,724 moga, Mysore, Meehold 10.495 (K); Gwalior (US). PERU: Prov. Apurima: 20 km N of St.: Wisner 35 (US). KENYA: Esageri Abancay, Stork et al. 10,539 (UC, US); Prov. Station, Nakuru to Eldama Ravine Road, Lima: Huarochiri, Weberbauer 5,287 (F); Prov. Bogdan AB2,014 (K); 12 mi N of Magadi, Avacucho: entire Pucjuio v Nazca, Fcrreyra Clayton DC56 (K); 20 mi E of Isiolo, Stewart .5,52.3 (K, US); Prov. Piura:'l km W of Talara, 474 (K); Nairobi, Webster K33 (K). KOREA: Beetle 26,197 (F, K, UC, US); Prov. Tumbes: in campis Chinampo, Faurie 1,244 (BM); Pyeng- Contramirante Villar, Ferreyra 12,214 (US). yang. Smith s.n., 5 Sept 1935 (US). LESOTHO: REPUBLIC OF THE CONGO: Kasenye, )

Biological Series, Vol. 19, No. 2 Taxonomy ov the Genus Chloris (Gramineae) 65

Johnston 1.042, 1,084a (K); Katanga, Tumbwe, 12,677 (US); Kansas: Stevens Co., E of Hu.go- Stjmoens 8,382 (K). SAUDI ARABIA: SO to ton, McGregor 17,016 (KANU); Louisiana: 90 km N of Aden, Wahah 1,904 (US). SENE- beach. Lake Ponchartrain, C.A. Brown 2,387 GAL: without precise location, Adanson 74 (US); Maine: North Berwick, wool waste, Par- (BM). SEYCHELLES ISLANDS: Long I.sland, lin 1.516 (US); Maryland: Canton, chrome ore Sqxiihbs Gil (K). SOCOTRA ISLAND: Bal- piles. Reed 32,714 (US); Massachusetts: Mill- four 341 (K). SOMALI REPUBLIC: Ca'ad burv, wool waste. Gates 31,785 (US); Missouri: Forest Reserve, 7 mi W of El Afwein, Boaler Courtney, Bush 9,733 (MO); Nevada: Lincoln B 130 (K); Erigavo Dtr.: Hubcra, McKinnon Co., between Crystal Springs and Ash Springs,

S249 (K, US). SOUTHERN RHODESIA: Shan- Train 2,415 ( NY)'; New Mexico : Grant Co., Man- gani Dtr.: Gwampa Forest Reserve, Goldsmith gas Canyon, 16 mi WNW of Silver City, Barkletj 48,828 (K); Lower Sabi Dtr.: Rattraij 1,230 14,732 (NY); Sierra Co., Animas Creek, Met- (K); Miami Dtr.: termite mound, K.34 Experi- calfe, s.n. (MO, NY); New York: Yonkers mental Farm, Wild 1,773 (K). SOUTHWEST wool mill, Bickncll. .v.n., 4 Sept 1898 (NY); AFRICA: Avis Dam, Windhoek, Liehenberg North Carolina: Guilford Co., W of Greens- 4,446 (UC, US); Grootfontein, Schoenfelder boro, Blomquist 1,962 (F); North Dakota: Nor- 88 S494 (K). SUDAN: Darfur Prov.: Nyorlete, ton Co., Mandan, Stevens 2,649 (UC, US); 12°5S'N, 24°4'E, Blair 11 (K); Wadi Toro, Oklahoma: Payne Co., Stillwater, Henson 343 13°6'N, 23°56'E, Blair 144 (K); Jebel Mara, (UC); South Carolina: Myrtle Beach, Blom- Zalingei, Wickens 1,786 (K); Kassala Prov.: be- quist s.n., 22 Aug 1930 (US); Texas: Brewster tween Berber and Suakin, Schiieinfurth 399, Co., 9 mi S of Marathon, Ferris and Duncan 572 (US). TANZANIA: Tumba, Bullock 2,415 2,841 (MO); Hemphill Co., 5 mi E of Canadian,

(US); Mwanza Township, Cannidiael 1,160 Rouell, Jr. 4,266 (TEX); Jeff Davis Co., 8 mi (K); Mbulu Dtr.: near Mdala River, Lake Man- S of Ft. Davis, W.V. Brown 3,269 (TEX); Utah: yara National Park, Greenuaij and Kanuri 11,- 0.5 mi N of south entrance, Zion National Park, 239 (K); Dar-es-Salaam, Marshall 50A (K); Harrison 11,092 (UC); Whipple Exped., Camp Monik Plateau, above Lake Natron, Newbould 60, Ft. Smith to Rio Grande, Bigelow s.n., 6,206 (K): 7 mi NE of Old Shimanga, Welch 1853-4 (NY); Wright Exped., western Texas 62 (K). TIBET: Lhasa, Richardson s.n., autumn to El Pa.so, Wright 762 (MO, NY, US). VENE- 1946 (K). UGANDA: Jinja, Hitchcock 24,954 ZUELA: vicinity of Caracas, Bailey and Bailey (K, US); Karamoja, 16 mi W of Moroto, Lang- 201 (NY, US).' ZAMBIA: Zambra, Namuala- dale-Brown 1,586 (K). UNION OF SOUTH Pemb road, Astle 2,107 (K); Abercorn, Siame AFRICA: Sunday River, N of Monke\' Ford, 608 (K), Trapnell 1,735 (K); Monze, Lochinvar Burchell 2,862 (K); Zoutpansberg, Kruger Na- Ranch, van Rensburg 1,238 (K); 10 mi NE of tional Park, Codd 5,407 (K); 4 mi N of Kroon- Serenje, Vasey-Fitzgerald 2,956 (MO). ZANZI- stad, 27°3.5'S, 27°5'E, Schecpers 1.266 (K); IS BAR: Hitchcock 24,460 (US). mi E of Pietersburg, van Viewen 1,612 (K). UNITED ARAB REPUBLIC: Jebel Elba, Wadi 24. CHLORIS GAYANA Kunth, Rev. Gram. Kasisrob, Shabetai 6 (K). UNITED STATES: 1:89. 1829, in key; redescribed 2:293, /;/.

Alabama: Cliambers Co., Langsdale, Banks .58. 1830. ( Holotypc not seen, but Kunth's 306 (US); Arizona: Chiricahua Mts., Paradise, complete description and plate are of the

Blumer 1,715 (K, MO, NY); Sulphur Spring Val- species as commonly understood.) Fig. .37, ley, Forbes 1,634, 1,643 (NY, US); Toro'" Can- 38. yon, Baboquivari Mts., Gilman 9 (NY); Pima C'hloris ahi/ssinica Hochstetter ex Achille Co., Wilmont Road, Tucson, Ginter s.n., 22 Ricliard, Tentamen Florae Abyssinicae Sept 1943 (UC); Santa Cruz Co., Patagonia, 2:406. 1850. (DUPLICATE OF SYN- Hitchcock 3,645, 3,664 Ajo, (US); M.E. Jones TYPE; "Schimperi iter Abyssinicum, sectio

24,795 (MO, NY); California: Fresno Co., along • tertia, ISOO . . . =nr. 79. Sect. I, prope Hwy 180, 5 mi of W Sanger, Bacigahipi 2,487 Djeladcranne in vallibus. V.i. 1844." BM! (UC); Merced Co., S of Newman, Mason and K!) Smith ( 8,223 UC ) ; Riverside Co., Ford Dry Lake, Chloris glahrata Andersson in Peters, Nat- 24 mi W of Blythe, Grinncll 1.081a Yolo (UC); urwiss. Reise Mossambiquc Bot. 2:557. Co., 3 mi N of Davis, along HWY 99 W, Crainp- 1864. ( Holotype not seen, but description ton 3,140 (UC). Colorado: Powers Co., road- clear. side near Holly, Harrington 740 (UC, US); Chloris gayana ssp. Chi. [sic] oligostachys Hawaii: Oahu, Honolulu, Kaimuki, A. Chase Barratte and Murbeck in Murbeck, Acta ) )

66 BiucnAM Young Univehsity Science Bulletin

Fig. 37. Chloris gatjana (A) habit, x 1/6; (B) spikelet, partly dissected; (C-F) florets, showing variation in size and shape of fertile lemma and size, shape, number, and sexual condition of sterile florets; (G) lowermost sterile floret; (H) middle sterile floret; (I) uppermost sterile floret; (J) caryopsis. (B-J) x 10.

Univ. Lund II. Sect. 2. 36:8. pL 13. 1900. Chloris gaijami Kunth, but basionym not (The authors clearly refer to this as a new cited. subspecies, yet they interjected abbre- an Chloris gayana f. oligostachys (Murbeck) viation of the generic name, thereby chang- Maire and Weiller, Flore Afrique Nord ing the relative order of taxa. The descrip- 2:204. 1953. (Based on Chloris gayana tion and plate are of a variant witii small ssp. oligostachys Murbeck.) spikclets.) Clitoris gayana f. genuina Maire and Weil- Eustacliys gayana Mundy, Rhodesia Agric. ler, Flore Afrique Nord 2:204. 1953.

Jour. 14:142. 1922. (Presumably based on ( Based on Chloris gayana Kunth. Biological Series, Vol. 19, No. 2 Ta.xonomy of the Genus Chloris (Gramineae) 67

Perennial up to 3 m t;xll, usually stolon- ous and nonstoloniferous plants are found, with iferous, sometimes tufted, erect; sheaths gla- the former much more common. Individuals brous to scabrous, often ciliate apicallv; ligule vary greatly in size, though plants less than ciliate; blades up to .30 cm long and 1.5 cm 0.5 m are uncommon. Spikelet moipholog)' is broad, scabrous; spikes nine to thirty, 8 to 15 inconstant. Lemma pubescence varies from cm long, usualh' somewhat divaricate; spikelets glabrous to densely pilose. The number of ster- pale, rather densely imbricate, ca 10 per cm ile florets ranges from two (veiy rarely one) of the scabrous rachis; glumes lanceolate to to four, with the lowest "sterile" floret usually narrowly ovate, scabrous, especially on the staminatc, occasionally sterile. The second low- nerves; first glume 1.4 to 2.8 mm long, 0.3 to est floret is usually sterile; upper florets are 0.4 mm wide, acute to awn-tipped; second always so. Twenty-one spikelets randomly glume 2.2 to 3.5 mm long, 0.3 to 0.6 mm wide, chosen from one collection (Wickens 2,715) acute to awn-tipped; fertile lemma 2.5 to 4.2 showed five different combinations of floret mm long, 0.7 to 1.0 mm wide, ovate to obovate number (two or three) and sexual condition to elliptic, pubescence variable, margins usually ( male or barren). with a prominent tuft of hairs near the apex, A number of variants have been named. Of occasionally appressed-pilose for full length, in- these Chloris ahijssinica Hochstetter ex Achille terner\es usually glabrous, sometimes scabrous Richard has been mostly widely used, though or appressed-pilose, lateral nerves usually gla- geneialK not recenth'. Duplicates of a syntype brous, occasionally appressed-pilose, somewhat were examined (BNI, MO, K); these did not gibbous, callus bearded, awn 1.5 to 6.5 mm differ markedly from the variation spectrum long; sterile florets two to four (rarely one). exhibited by the remaining specimens that were the lowest 2.2 to 3.2 mm long, 0.5 to 0.8 mm examined. Stapf (19.31) provided a short key wide, similar to the lemma, but more cylindri- to Chloris aht/ssinica and C. gaijona, separating cal, staminatc or barren, awn 0.8 to 3.2 mm them on the presence or absence of stolons and long; upper florits awnless or awn-tippcxl, the length, sexuality, and number of sterile progressi\'elv smaller \\ith the terminal reduced florets. These characters were studied on a rep- to a turbinate cup; earvopsis 1.0 to 1.5 mm resentatixe number of specimens collected long, 0.5 mm \\'ide, ellipsoidal; chromosome throughout Africa, but neither discontinuities number 2n = 20, 30, 40. in measurements nor major patterns could be This species is the most variable of the distinguished. perennial species in the genus. Both stolonifer- There is a possibilit\ of a very casual cor-

Fig. 38. Distribution of Chloris gaijana. Inset A; Hawaiian Islands. ;

68 Bricham Young University Science Bulletin relation with ecological conditions, for tnfted Chikwawa Dtr.: Bra.^s 17,996 (MO, NY, US); plants were frecjuently, but not alwa\s, col- Kasita River near Ekwendeni, ]ack.son 366 (K); lected troni salt pans. Port Herald Dtr.: Phipps 2,606 (MO). MAU- Representative specimens examined: (A RITIUS: Curepipe, Vaughan 1,844 (K). MEX- total of over 800 different specimens of this ICO: Morelos: Cuenievaca, Lijonnet 2,807 species were studied. The following list repre- (US). MOgAMBIQUE: Nya.ssa Dtr.: Ama- sents only a small portion of this group.) AN- ramba, 22 km de Nova Freixo, Torre and Paiva

GOLA: Huila Dtr. : Namaculungo, Teixera 10,624 (K). NEW ZEALAND: near Tauranga, 1,407 (BM). ARGENTINA: Prov. Cordoba: Hodgkins 70,536 (US). NIGERIA: Prov. Cruz del Eje, ViUafarie 2,413 (US); Prov. Cor- Bornu: Lake Chad, Daggash 24,970 (K); Ngala rientes: Dep. Lavalle, Estancia "La Pastrol" Dtr.: Wulgo, Daveij 2 (K); Rann Dtr., Daveij Pedersen 3,864 (NY, UC, US); Prov. Salta: s.n., 17 Apr 1947 (K); River Kalkala, SW corner Metan, Cabrera 3,089 (NY); Prov. Tucuman: of Lake Chad, Golding 76 (K). REPUBLIC Trancas, Venturi 6,480 (US). AUSTRALIA: OF THE CONGO: Kasenve, Johmton 1,057 New South Wales: Kenebri, 11 mi N of Bara- (US); Albertville, Under s.n., 23 Jan 1927 (K). dine, Constable 15,978 (UC); Northern Terri- RWANDA: Pare National Kagera, Lcbnm 9.482 tory: near Darwin, Allen 67 (K); Queensland: (K); Biumba, Mutara, colline Nyaka Kanga, Conjuboy, Blake ami Lazarides s.n., 26 Feb Troupin 8,813 (K). SIKKIM: Gangtok, Dr. 1954 (US); Kenmore, Clemens s.n., 18 Ma\ Bors collector s.n.. 15 Jul 1945 (K)."sOMALI 1943 (US); Western Australia: Latham, Moore REPUBLIC: Farquahorson s.n.. Sept 1931 (K). K197 (K). BOTSWANA: Koudoum Valley, 30 SOUTHERN RHODESIA; Katasi Reserve, km E of Southwest Africa border. Wild and Bikita, Cleghorn 177 (BM). SOUTHWEST Drummond 7,056 (BM). BRAZIL; Minas AFRICA: Omuramba Khaudum, ca 18M°S, Geraes: Lavras, A. Chase 8,827 (F, VIO, NY, 20'i°E. de Winter and Marais 4.667 (K); Oka- US); Rio Grande do Sul: Mono Santa Ana, vango, at Bagani pontoon, Kaprivi, de Winter Rambo 41,982 (US). COLOMBIA: Pasto, and Wiss 4345 (K). SUDAN: Zalingei, Robert-

Medcllin, Rivera 131 ( K ) ; vicinity of Medellin, son 10 (BM); Dafur Prov.: Nyertiti, Robert- Toro 299 (NY, US). COSTA RICA: Turrialba, son 22 (BM); Jebel Marra, Dariba Crater, Leon 286 (MO). CUBA: Prov. Ilerradura: Robertson 140 (BM, K); Equatoria Prov.: Nim- Pinar del Rio, Ekwan 14,108 (NY). EASTER ule, Shantz 907 (US). SYRIA: Ali al Gharbi, ISLAND: Mataveri, waste land, Chaplin 1,041 Graham 360 (BM). TANZANIA: Mbeye Dtr.: (NY). ECUADOR: Prov. Guayas: Milagro, Rujewa, Anderson 1,117 (K); Musoma, Em.

Acosta-Solis 70,693 (US); Prov. Pichincha: 232 ( US ) ; Ngari Roiigi plain, western slopes of Valle del Guaillabamba River, Acosta-Solis Mt. Kilimanjaro, Grecnwatj 6,741 (K); Ai-usha 21,259 (US), A.'iplund 7.036 (K). ETHIOPIA: Region, Ol Doingo Sambu, Greenwaij 4,413 Ocule Casai, Pappi 2,081 (MO, NY, UC); 10 (K); Moshi Dtr.: Sanya River, Haarer 1,097 mi N of Assale, Se/nple 2 ( US ) ; Metahara. (K); Rukwa, Lea LR8 (K); Banagi, Serengeti Semple 10 (US). GUATEMALA: near Chimal- National Park, Lcippert S603 (K);' 45-65 mi S tenango, Standlei/ 80,483 (F, K). HAITI: plaine of Dodoma, Lijnes 76a (US); Selous Game Re- du Nord, Ekman H. 9,890 (K, US). HONDU- serve, Nicholson 48 (K); Ngorongoro Crater, RAS: Dep. Morazan: Zamorano, Rodri'^nez 103 Paulo 317 (K), Piemiesel and Kepliart 506 (F), 3,277 (F, MO, US); Standlei/ 1,767a, (US); Kilwa, Rifiji, Schlieben 2.541 (K); Lindi 13,035, 21,041 (F), 18,979 (F, NY, US); Wil- Dtr.: Lutamba, Schlieben 5,880 (K); Monduli, liams and Molina 12,216 (F). INDIA: Mysore, near Arusha, Semple s.n., Mav 1945 (US); Bangalore, Koelz 19,898 (UC); Salem Dtr.; Olduvai, Williams 681 (K). THAILAND: Hua Hosen Cattle Station, Naratjanaswami 3.028 Hin, lones s.n., June 1931 (BM). TONGA- (K). JAMAICA: St. Andrew Parish, Richmond TAPU ISLAND GROUP: 'Eua, Houma, Set-

Park, Harris 12,707 (F, K, MO, NY). KENYA: chell and Parks 15,431 ( UC, US). UNITED Klboko, Bogdan AB 2,239 (UC); Longonot, STATES: Arizona: Yuma Valley, Fochtman 70 Hitchcock 24,828 (K, US); Nakuru, Hitchcock (NY); California: Merced Co., N of Los Banos, 25,075, 25,082 (K, US); between Oljoro-o-Nyon Mason and Smith 8,226 (UC); Florida: Her-

River and Lake Naivasha, Mearns 636 ( US ) nando Co., Brooksville, McFarlin 6,072 (US); between Narok River and Lake Naivasha, Hawaii- Kiholo, A. Chase 12.688 (US); Mis- Mearns 628 (NY, US). LORD HOWE IS- sissippi: Lumberton, Real s.n., 10 Jun 1920 LAND: Binoleij 25 (K, UC). MALI: El Oual- (US); Texas: Port Lavaca, Allen 17 (US); Kle- adji. Chevalier 43,927 (K); Tombouctou, Olaf- berg Co., .5.5 mi S of Kingsville, Corij 51.303 sen 205 (US). MALAWI: Southern Nvasa Dtr.: (NY, UC); Hidalgo Co., 0..5 mi N of Edinburg, Dambo, Namwera Road, Ballard C.h.7 (K); Ferris and Duncan 3.064 (MO, NY, US); Rock- )

Biological Series, Vol. 19, No. 2 Ta.xonomy of tiie Genus Chloris (Gramineae) 69 port, Fisher 41.145 (US); Kleberg Co., King 33, A-D; 35). Chloris virgata is an annual with Ranch, Lundell 14.877 (US); Cameron Co., Port the cilia restricted to the upper portion of the Isabel Road, Rimi/on 4,138 (MO, NY). lemma margin. UGANDA: Old Entebbe, Darker 257 (UC); The few specimens that have been collected Luzinga, Ditmmer 4,351 (US); Mbulamuti, are from sandy savannas in Argentina (Fig. Hitchcock 24,975 (US); Tororo, Swamp, Hitch- 39, triangles). cock 24,980 (US); vicinity of Kabula Miiliro, Specimens examined: ARGENTINA: For- Mearns 2,532 (US); Mbalc, near Mt. Elgon, mosa Territory: orillos del arroyo Ferreyra, Smusden 1,098 (US); S of Namasagali to Ka- Asp 20 (US); Las Lomita, Parodi 8,455 (NY, muli Road, Bugabula City, Wood 948 (MO). US); Prov. Catamarca: Mollecito, 20 km S of UNION OF SOUTH AFRICA: between Sellen- Andalgala, Bartlett 20,239 (US); Dep. Belen, dam and Riversdale, Godfreti SI I 1,312 (VC. Las Faldas, Lillo 45,625 (NY, US); Prov. La US); near Pietermaritzburg, thornveld, Godfreij Rioja: Patquia, Lahitte y Castro 3 (US). SH 1,912 (US); Mposa, Godfreti and Baijer SH 1,463 (US). URUGUAY: Dep. Montevideo: 26. CHLORIS ORTHONOTON Doell in Mar- Osten 19.923 (US). VENEZUELA: Merida, La tins, Flora Brasiliensis 23:64. 1878. (ISO- Mesa, Uanhurti-Tracij 16 (K). ZAMBIA: TYPES: "Glaziou n. 3610" BR! K! US!) Mweru Wantipa swamp, 8°50'S x 29°40'E, Fig. 39; 40, A-E. Tijrer 206 (BM). Chloris guaranitica Parodi, Revista Argen- tina Agron. 28:106. Fig. 2. 1961. (ISO- 25. CHLORIS CASTILLONIANA Lillo and TYPE: Argentina: Corrientes, Dep. Mbu- Parodi e.\ Parodi, Pli\sis 4:176. 191S.

rucuya, Estancia Santa Maria . . . Pederson (TYPE: not seen. OriginalK collected by no. 3742." K! Castillon in the Calchaqui Valley, Argen- aeeom- tina. The original description and Stoloniferous, erect perennial to 70 cm tall; the panving illustration positi\'el\ id(ntif\' sheaths glabrous, occasionally pilose near the plant.) Fig. 33, A-D. ligule; ligule prominently pilose; blades 0.5 to

Caespitose perennial up to 1 m tall; she iths glabrous; ligule short-ciliate; blades 10-20 cm long, 4 to 5 mm wide, glabrous; spikes eight to fifteen, 9 to 11 cm long, apprcssed to strongly ascending; spikelets densely imbricate, appressed, about eight per cm of the scabrous rachis; glumes lanceolate, thin, membranous, glabrous except for the scabrous midnerve; first glume 1.6 to 2.0 mm long, 0.3 to 0.4 mm wide; second glume 2.9 to 3.9 mm long, 0.4 to 0.6 mm wide, sometimes deHcately mucronate; fertile lemma 3.5 to 4.7 mm long, 1.0 to 1.3 mm wide, lanceolate-elliptic, rather prominently gibbous, callus flattened and elongate, densely bearded, lemma margins densely ciliate with hairs 1 to 2 mm long, keel glabrous, occasionally groove present along the A

Chloris castilloniana is most closely related to C virgata, from which it may be separated its habit well in by perennial as as having promi- Fig. 39. Distribution of Chloris orthonoton (dots) and nent cilia along most of the lemma margin ( Fig. C. castilloniana (triangles). 70 Brigham Young University Science Bulletin

0.7 mm wide, up to 25 cm long, pilose near the Swallen 4,641 (US); Minas Geraes: Lavras,

( base, otherwise glabrous to scabrous; spikes Black 887B US ) ; Juiz de Fora, A. Chase 8,625 four to eleven, 4 to 10 cm long, in several verti- (US); Sao Miguel, NW of Fomiiga, A. Chase cils near the culm apex; spikelets eight to ten 10,544 (US); between Sucupira and Omega, A. per cm of the scabrous rachis, densely imbricate, Chase 11,162 (US); Paraiba: Soledade, Cesan appressed; glumes lanceolate, acuminate, gla- 692 (US), Pickel 3,827 (US); Pernambuco: Bel- brous except for the scabrous midnerve; first lo Jardim, A. Chase 7,700 (F, US); vicinity of glume 1.6 to 2.4 mm long, 0.2 to 0.3 mm wide; Recife, A. Chase 7,720 (F, NY, US); Tapera, second glume 2.3 to 3.6 mm long, 0.3 to 0.5 mm Pickel 81 (F); Rio de Janeiro, NW of Cabo wide; fertile lemma 3.0 to 4.5 mm long, 0.7 to 1.6 Frio, A. Chase 10,122 (US); Niteroi, A. Chase mm wide, narrowly lanceolate to elliptic, mar- 10,037 (US); Rio Grande do Norte: Estremoz gins long-ciliate, especially near the apex, keel to Natal, Swallen 4,773 (US); Nova Cruz to glabrous to appressed-pubescent, internerves Montanhas, Stcallen 4,822 (US); Santa Cruz, with a longitudinal groove bearing one or more Sirallen 4,861 (US); Rio de Janeiro, C.laziou rows of dark glandular hairs, callus bearded, 17,363, 1888 (NY, US). awn 7.5 to 20.0 cm long; sterile floret one, 1.5 to 2.2 mm long, 0.3 to 0.5 mm wide, cylindric- 27. CHLORIS ARISTATA (Cervantes) Swallen, turbinate, glabrous, awn 3.5 to 9.0 mm long; North Aim-r. Flora 17:,596. 19.39. (Based anthers ca 0.4 mm long; caryopsis ca 1.7 mm on Agrostomia aristata Cervantes. ) Fig. 40, long, 0.4 mm wide, narrowly ellipsoid, trigonous. F-J. Chloris orthonoton is closely related to C. aristata Cervantes, Naturaleza aristata. Table 9 and Fig. 40 show the main Agrostomia "San differences between the two. (Mexico City) 1:345. 1870. (TYPE: Angel. San Agustin de las Cuevas." Not Occasional specimens of Chloris orthonoton seen, but description is clear.) may resemble C. vir^ata. The latter, however,

is a fibrous-rooted annual, lacks glandular hairs Stoloniferous perennial with erect culms to in the lateral groove of the lemma, and has a 80 cm tall; sheaths glabrous; ligule a short- prominent tuft of cilia only along the upper por- ciliate crown; blades up to 15 cm long, 5 mm tion of the lemma margin. wide, usualK glabrous or scabrous, occasionallx' A variant of this species, lacking pubescence the upper surface pilose; spikes four to eight, on the midnerves of the lemmas and having 4.0 to 8.5 cm long, erect, somewhat appressed, somewhat coarser spikes, has bei'ii named radiate; spikelets appressed, densely imbricate, Chloris isiiaranitica Parodi (1961). In all other ca five to eleven per cm of scabrous rachis; respects the plant is identical with typical C. glumes narrowly lanceolate, glabrous t-xcept for orthonoton, and I do not feel it wortlu of the scabrous midnerve; first glume 1.8 to 2.3 mm nomenclatiual recognition. long, 0.2 to 0.4 mm wide; second glume 3.0 to Representative specimens examined: AS- 4.1 mm long, 0.2 to 0.5 mm wide; fertile lemma CENSION ISLAND: roadside near airfield, 3.4 to 4.5 mm long, 0.8 to 1.0 mm wide, lanceo- Duffey 1/83 (US). ARGENTINA: Prov. Cor- late to elliptic, apex acute, margins appressed- rientes: Dep. Mburucuya, Parodi 3,742 (BR). pilose below, the hairs slightly spreading above, BRAZIL: Prov. Bahia: Baliia, A. Chase 7.871. keel usualK' glabrous, occasionally with a few 8,033, 8,051 (US); Feira de Santana, A. Chase appressed hairs, midnerves glabrous, callus 8,060 (US); Prov. Ceara: Acude, Municipio de bearded, awn 6.0 to 12.0 mm long; sterile floret Maranguape, Drouet 2,180 (F, US); Camocim one, 1.6 to 24 mm long, 0.6 to 0.7 mm wide, to Granja, Swallen 4,634 (US); Martinopolis, awn 5 to 7 mm long; caryopsis 1.5 to 1.9 mm

Table 9. Comparison of characteristics of Chloris orthonoton and C. aristata.

Species Ligule Lemma margin pubescence Lemma internerves

C. orthonoton long-ciliate ( ca 2 cilia more than 1.5 mm long; ap- lateral groove glandular-pubes- mm long) pressed basally, becoming spreading cent; otherwise glabrous apically

C. aristata short-ciliate ( less cilia less than 1 mm long; appressed lateral groove and internerves than 1 mm long) most of length or only slightly spread- glabrous, occasii)iuillv ap- ing apically pressed-pubescent, but not glandular )

Biological Seiuks, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gbamineae) 71

Fig. 40. Chloria orthonoton and C. aristiita. (A-E) C. orthonoton. (A) habit, .x 1/4; (B) spikelet, x 5; (C) florets, ,x 5; (D) sterile floret, x 5; (E) caryopsis, x 7.5. ( F-J ) C. arhtata. (F) habit, x 1/4; (G) spikelet, X 5; (H) florets, x 5; (I) sterile floret, x 5; (J) caryopsis, x 7.5.

long, ca 0.5 nini wide, ellipsoid, trigonou.s; tification. Accordingly, this name is rejected

chromosome number 2n = 100. ( see under excluded names. Differences between Chloris aristata and C. Chloris aristata has been widely collected orthonoton have been discussed previously. from prairies and grasslands throughout much This species has occasionally been called of Central America (Fig. 41). Chloris nifescens Lagasca, a name which can- Representative specimens examined: COSTA not be typified and the original description of RICA: ccrca de San Ramon, Brenes s.n., 3 which is not complete enough for positive iden- Jan 1935 (F); San Jose, Cooper 5,997, Hitch- ) )

72 Bbicham Younc University Science Bulletin

Diplachne hrandegei Vasey ex Brandegee. Fig. 42. Diplachne hrandegei Vasey ex Brandegee, Proc. California Acad. Sci. H. 2:213. 1889. (HOLOTYPE: "Magdalena Island. T. S. Brandegee 18 January 1889." US! Leptochloa hrandegei (Vasey ex Brande- gee) Hitchcock, U.S. Dept. Agric. Bur.

Plant IndustiN Bull. 33:21. 1903. ( Ba.sed on Diplachne hrandegei Vasey ex Brande- gee.)

Fig. 41. Distribution of Chloris aristata. Gouinia hrandegei (Vasey ex Brandegee) Hitchcock, U.S. Dept. Agric. Bur. Plant In- cock 8,469 (US), Valerio i04 (F, K); San Rafael dustiy Bull. 33:21. 1903. (Based on de Cartago, Pittier 7,109 (US); between San Diplachne hrandegei Vase>- ex Brandegee.) Pedro de Montes de Oca and Curridibat, Perennial to 1 m tall, densely tufted from a Standley 32,817 (US). GUATEMALA: moun- coarse, fibrous root s\stem; lower sheaths tloc- tains E of Quiche, Grout 640 (US); Guatemala cose, becoming glabrous above; ligules pilose, City, Hitchcock 9,039 (US); La Aurora, Ritana the lower more so than the upper; blades scab- 699 (US); Dep. Alta Verapaz, near San Juan rous; inflorescence with up to 20 spikes ar- Chamelco, StamUcy 92,435 (US); 3 km S of ranged in a series of somewhat distant sub- Huehuetenango, Williams et al. 22,109 (F, US). verticils, occasionally the spikes single at a HAITL W of Palmiste, Leonard and Leonard node; spikes 5 to 10 cm long; spikelets distant, 12,538 (NY). HONDURAS: Dep. Choluteca: ca three or four per cm of the spike length, vicinity of San Marcos de Colon, Standley strongly appressed; glimies markedly unequal, 15,765 (F); Dep. Morazan: vicinit\- of El Zanio- narrowly lanceolate, thin, hyaline, glabrous ex- rano, Standlcii 1501. 26,327 (F); Sicallen cept for the slightly scabrous midnerve; first 10,832 (US). MEXICO: Aguascalientes, Hitch- glume 1 5 to 1.8 long, ca 0.3 wide; cock 7,454 (US); Chiapas: Monserrate, Pur- mm mm second glume 3.3 to 4.0 mm long, ca 0.5 mm pus 388 (F); Coahuila: Parras de la Fuentes, wide; fertile lemma 4.7 to 5.5 mm long, 0.5 to Palmer 450 (F, MO, NY); Saltillo, Hitchcock 0.8 widi', strongly dorsally compressed, 5,620 (US); Durango: Duiango, Hitchcock 7,657 mm iiarrowK' elliptic to linear in side view, more (US); Guanajuato: 2.5 km N of San Fran- broadh' elliptic in lace view, apex acute to cisco del Rincon, Sohns 355 (US); Irapuato, shortly acuminate, margins, keels, and inter- Hitchcock 7,412 ( US ) ; Acamboro, Hitchcock nerves scabrous to hirsute, callus bearded, awn 6,927 (US); Hidalgo: Jaeala, V. //. Chase 7,098 1.0 to 4.5 mm long; sterile floret one, ca 1.5 mm (US); La Placita, 10 mi S of Jaeala, Gould long. 0.3 mm wide, ver\- narrowly c\'1indrieal to 10,389 (US); Jalisco: 15.5 mi E of Guadalajara, ellipsoid, scabrous to sparsely appressed-pilose, Leavenworth 1,852 (US); San Nicolas, Hitch- awn 0.7 tn 1.9 mm long; earyopsis 2.5 to 3.0 mm cock 7.193 (US); Michoacan: vicinity of Mo- long, ca 0.7 mm wide, ca 0..3 mm thick, strongly relia, Arsenc 3,134 (MO, NY, US); 'Uiuapan, dorsally compressed, ellipsoid, slightly trigonous. Hitchcock 6,965 ( US ) ; Nuevo Leon, Monterrey, Chloris l)randegei is similar to and probably Hitchcock 1,263 (MO, NY); Galeana, V. H. closely related to C. chloridea, though distin- Chase 7,738 (US); Oaxaca: Cuicatlan, San Juan guished from it by veiy short awns on the lem- Bautista, Conzatti 4,024.5 (US); Cd. Oaxaca, mas and a lack of underground spikelets. Hitchcock 6,110, 6,129, 6,170 (US); Puebla: Chloris hrandegei is endemic to Baja Cali- Tehuacan, Hitchcock 6.033 (US); Queretaro: fornia Sur, Mexico (Fig. 43, triangles), where Queretaro, Hitchcock 5,804 (US); San Luis it grows on rocky slopes and in arroyos. Potosi: Cardenas, Hitchcock 5,749 (US); Ve- Specimens examined: MEXICO: Baja Cali- nado, Whiting 624. 711 (US); Veracruz: Jalapa, fornia Sur: La Purisima, Brandegee L91 (US); Hitchcock 6,692 (US); Orizaba, Hitchcock San Jose del Cabo, Brandegee s.n., 6 Oct 1890 6,336 (US). (NY); Sierra de la Giganta, 25°58'N, 111°.32/'2'W, on road from Loreto to San Javier, 28. CHLORIS BRANDEGEI (Vasev) Swallen, Carter 5.014 (UC); Cerro Teombo, 25"51'N, Amer. Jour. Rot. 22:41. 19.35. (Based on 111°25'W, N of Fortezuelo de Gabilan, 1100 m. BiOLOGiCAi, Series, Vol. 19, No. 2 T.\xonomv of the Genus Chloris (Gramineae) 73

Fig. 42. Chloris hmndegei (A) habit, x 1/4; ( B,C,E ) .spikelet, partly dissected; ( D,F,G ) florets, showing variation; (H) caryopsis. (B-H) x 5. ) ) )

74 Brigham Young University Science Bulletin

Chloris clandestina Scribner and Merrill, U.S. Dept. Agric. Div. Agrost. Bull. 24:25.

1901. ( Based on Gymnopogon longifolius Fournier.

Perennial to 1 m tall, tufted, erect, bearing cleistogamous spikelets at the tips of long, scaly, branched rhizomes; each cleistogamous spikelet consists of a series of three to five overlapping bracts enclosing a saclike structure, which in turn encloses a pistil and three stamens; mature cleistogamous fruit broadly ovoid-ellipsoid, up to 4 mm long and 2.5 mm wide, brown; sheaths mostly glabrous, occasionally be-

Fig. 43. Distribution of Chloris brandegei (triangles) coming sparsely long-pilose near the ligule; low- and Chloris suhmutica (dots). er ligules with a prominent tuft of hairs, upper ligules often reduced to a low crown; blades Carter 5,064 (UC); 37 km SE of San Ignacio, up to 30 cm long and 1 cm wide, scabrous, oc- among lava rocks in giavcllv aiTO\o, Carter et al. casionally pilose, innovations often sparsely pi-

1,977\\JC, US); Siena de la Giganta, Arroyo lose; spikes three to fifteen ( usually less than del Cajon de Tecomaja, SW of Puerto Escon- 10), usually 6 to 10 cm long, in several well- dido, 25°43'N, 111°20'W, Carter and Kellogg separated verticils, occasionally borne singly at 2,897 (UC, US); El Triunfo, A/. E. Jones 27,658 a node; spikelets rather distant, ca four per cm (UC); Carmen Island, Palmer 8,621, 1 to 7 Nov of the scabrous rachis, appressed except for the 1920 (US); Punta San Lorenzo, ca 25 mi E of divergent awns; glumes niarkedh' unequal, gla- La Paz, Wiggins 15,638 (UC). brous except for the scabrous inidnerve; first glume 1 to 2 mm long, ca 0.4 nun wide, lanceolate; second glume 2.0 to 3.5 mm long, ca 0.3 29. CHLORIS CHLORIDEA (Presl) Hitchcock, mm wide, narrowly lanceolate; fertile lemma 4.5

Proc. Biol. Soc. Washington 41:162. 1928. to 7.5 mm long, ca 1 mm wide, eu 0.5 mm (Based on Dincha chloridea Presl.) Fig. 44. thick, dorsally compressed, linear to narrowly lanceolate in side view, lanceolate in top view, Dineha chloridea Presl, Hel. Haenke 1:291. apex acuminate, callus densely bearded, mar- 1830. seen, there is a frag- (TYPE: Not but gin ciliate, lemma otherwise glabrous, awn 6.5 ment in US!, from Haenke Herbarium, PR, to 15.0 mm long, often arising from between two

labeled: "Mexico . . . Presl." This the and setae; sterile floret one, 1.4 to 3.0 mm long, less description clearly identify the name.) than 0.3 mm uide, often somewhat flattened, Etitriana chloridea (Presl) Kunth, Rev. scabrous to short-pilose, awn 2 to 8 mm long; Gram. Supplenda, p. 23. 18.34. (Based on car\opsis ca 4.5 mm long, 0.8 mm wide, some- Dineha cidoridea Presl. what (lattened, long-acuminate; chromosome number 2n 40, SO. Gymnopogon virletii Fournier, Mex. PI. The imderground cleistogamous spikelets of 2:144. 1886. First published as nomen Chloris chloridea make it one of the most un- nudem in Hemslev, Biol. Centr. Amer. Bot. usual grassi's. These spikelets are borne on a 3:560. 1855. (TYPE: "San Luis de Potosi. rather intricately branched rhizome system ( Fig. Virl. n. 1441." There is a fragment in US!, 44, A), the branches of which are brittle and from P, labeled "Mexi(jue. Prov. de San often are not collected. The cleistogamous spike- Luis . . . \'irlet d'Aoust 1851," which is lets on the specimens studied set a high pro- C. cidoridea. ) portion of seed, while most of the above ground Gymnojwgon longifolius Fournier, Mex. PI. spikelets were sterile. Whether this is due to 2:144. 1886. (FRAGMENT OF TYPE: the immaturity of the specimens examined, en- ". US!, originally from P, labeled . . Mex- vironmental factors, or is an indication of a more prov. \'era 52.") ique Cruz. Gouin. 1867. complicated reproductive pattern, must await Chloris longijolia (Fournier) Vasey, Contr. further field studies.

U.S. Nat. Herb. 1:284. /;/. 7.9. 1893. Non Chloris chloridea is quite variable in spike- Steudel, 1854. (Based on Gymnopogon let size, and some plants may approach C. bran- longijolia Fournier. degei in lemma length, but C chloridea will al- BiOLOciCAi. Series, Vol. 19, No. 2 Taxonomy ok the Genus Chloris (Gramineae) 75

Fig. 44. Chloris cMoridea. (A) habit, x 1/3; (B) spikfli t. partly dissected. .\ 7. .5; (C,D) florets, showing

variation, .\ 7. .5; (E) carvopsis from aboveground spikelet, x .5; (F) underground spikelet, x 5; (G) flower of underground spikelet, x 15; (H) caryopsis from underground spikelet, x 5. )

76 Bricham Younc University Science Bulletin ways have relatively long awns and, of course, 30. CHLORIS VENTRICOSA Robert Brown, the underground spikelets. Prodr. Flora Nov. Holl. 186. ISIO. (HOLO- "1. This species is widely distributed in the TYPE: Chloris ventricosa Richmond" southwestern United States, Mexico, Honduras, BM!) Fig. 46. and Venezuela and has been collected in grass- Chloris sclerantha Lindley in Mitchell, Jour. lands, brushy areas, and old fields (Fig. 45). E.xped. Interior Australia '31. 1848. (TYPE: "New South Wales; near Muddal banks of Bogan River, Mitchell" CGE. Not seen. Fragment of type at K! Chloris ventricosa var. tenuis Bentham, Flora Australiensis 7:613. 1878. (SYNTYPE: "Rockhampton, O'Shanesy. Herb. F. Muel- ler 1879 [?]" K!)

Perennial to 1 m tall, stoloniferous, erect; sheaths glabrous, scabrous or sparsely pilose; ligule area scabrous to prominently pilose; blades glabrous, scabrous, or sparsely pilose; spikes two to nine, 5 to 11 cm long; spikelets Fig. 45. Distribution of Chloris chloridea. densely imbricate, ;i\x'raging 10 per cm of the scabrous racliis; glumes scarious, narrowK lan- Representative specimens examined: HON- ceolate to ovate, acute to acuminate, glaI)rous ex- DURAS: Dep. Morazan; El Zamorano, Rod- cept for the scabrous midnerves; first glume 1.2 riguez 3,516 (F, US), StaiuUeij 13,076 (F, to 2.3 mm long, ca 0.5 mm wide; second glume NY, US), 14,454 (F), 14,H96 ("f, UC, US), 2.5 to 4.1 mm long, ca 0.8 mm wide; fertile 16,073 (F), 3,9.36 (US), Suallen 11,156 (US). lemma 2.0 to 5.4 mm long, 0.6 to 1.2 mm MEXICO: Baja California Sur: El Triunfo, M. E. wide, elliptic to obovate, sometimes ventricose, Jones s.n., 6 Oct 1930 (US); Guanajuato: 6 km S acute to obtuse, usuallv glabrous except for the of Ltoii, Soltm 384 (MO, US); Jalisco: Tu,\pan, scabrous midncrve, occasionally scabrous on the A/

Nue\() Leon: Colonia del \'alle, Monterrex , F. F. to 11 mm long; sterile floret one (very rarely Smitli M267 (TEX); Oa.xaca: Tomellin,' //(7r/(- two), 1.0 to 2.6 mm long, 0.3 to 1.0 mm wide, cock 6,226 (US); San Luis Potosi: Las Canoas, cylindric to narrowly turbinate, apex obtuse to Hitchcock 5,766 (US); Sinaloa: Topolobampo, truncate, awn 0.5 to 7.5 mm long; anthers 0.7 Palmer 2.38, Sept 1897 (K, US); Sonora: Hermo- to 1.4 mm long; caryopsis 1.5 to 2.1 mm long, sillo, Hitchcock 3,572 (US); Taniaulipas: 3 mi N ca 0.4 mm wide, narrowly obovoid, ± trigonous. of Santo la Marina on road to Jimenez, Crtitch- This species is quite variable, mostly with fiehl and Joltn.ston 4,972 (TEX); Veracruz: 5 mi respect to the height and apparent vigor of the E of San Luis Potosi border, M. C. Johnston plant, the size of spikelet parts, and the shape 6,106 (TEX, UC, US). UNITED STATES: Ari- of the sterile florets. Attempts at correlating zona: Pima Co., W of Topawa, Coodding 194-45 various features of about 75 specimens were (NY, US); Texas: Cameron Co., Laguna Atas- made, but no conspicuous patterns could be de- cosa Refuge, Fleetwood 3,857 through 3,859, tected. This corroborates an earlier study by 3,861 (TEX); Brownsville Municipal Airport, Everist (193.5b), in which he found many in- Rum/on 2,928 (US); Brownsville, Sdvcus 379 ( K, termediate plants between various extremes. US, TEX); 1 mi N of Combes, Silveus 1.583. Certain of these have received names. Chloris 1,586 (MO); 0.5 mi of N Conches, Silveus 6.781 ventricosa \ar. tenuis Bentham is based on a (TEX); 5 mi S of San Benito, Silveus 7,432 small plant with relatively small spikelets, while (TEX); Kleberg Co., Kingsville, Diaz 1,951 a jjomewhat larger variant was called C. scle- (US); 4 mi S of Kingsville, Meyer s.n., 24 Oct rantha Lindley. 1958 (US); Nueces Co., ca 5 mi S of Banquete, Chloris ventricosa and C. truncata are closely M. C. Johnston 542,273 (TEX); San Patricio allied. Most specimens can be differentiated by Co., Odem, Silveus 2,408 (TEX). VENEZUE- the length of the spikes, but some specimens of LA: Estado Lara, La Ruczga, cerea dc Bar- C. truncata have abnormally short spikes. The quisimeto, Tanuiijo 4,230 (US). following key will separate most specimens: Biological Series, Vol. 19, No. 2 T.\.\onomv of the Genus Chloris (Gramineae) 77

1. Lemma margin.s apprcssod-pilosc, rarely than 8 cm long; lemmas 0.6 mm wide or

sparsely so; awn of sterile floret ( the lower, more, usually about 0.8 to 1.0 mm; spike- if two are present) 3.1 to 16.0 mm long, lets tawny, rarely dark brown. usuallv 4 mm or more; spikes 5 to 23 cm, C. ventricosa, Fig. 46. usually 10 cm or more in length; lemmas Chloris ventricosa is widespread in eastern 0.7 mm wide or less, usually ca 0.5 mm Queensland and Ni'w South Wales ( Fig. 47, wide; spikelets usually ver\' dark, black or B ) , where Everist ( 1935b ) reported that it ap- nearly so. C. tnmcata. Fig. 4S. pears to prefer heavier and richer soils. 1. Lemma margins usually glabrous or Representative specimens examined: AUS- scabrous, not appressed-pilose, rarely veiT TRALIA: New South Wales: Gravesend, Break- sparsely pilose; awn of sterile floret (the well s.n., May, 1913 (F); Warialda, Hadleij lower, if two are present) 0.5 to 7.5 mm s.n., Apr 1908 (F); Hunter Valley, 9 mi NW of long, usuallv about 2 mm long; spikes gen- Scone, Story 6,932 (K); beween Windsor and erally short, 5 to 11 cm long, usually less Kingswood, Vickenj and Constable 16,121 (K,

Fig. 46. Chloris tculricosa. (A) hahit, x 1/6; (B) spikelet, partly dissected, x 15; (C-F) florets, showing

variation, .\ 15; (G) sterile floret, x 15; (H) caryopsis, .\ 10. A ) )

78 Bricham Young University Science Bulletin

Fig. 47. Distribution of Chlork truncata (.M and C. ventricosa (B).

MO, US); Svdnev, Wilkes Exped s.n., 1832-42 31. CHLORIS TRUNCATA Robert Brown, (US); Queensland: Chinchilla, Beasleij 128 Prodr. Flora Nov. Roll. 186. 1810. (HOLO-

(K); near Wandoan, Behon s.n., 1930 (K); TYPE: "Port Jackson . . . R. Brown" BM! Miles, Belson s.n., 1930 (K); Wallumbilla, Bel- ISOTYPE: K!) Fig. 48. son s.n., 1930 (K); Downfall Creek, near Gilgul- Chloris elon^ata Poiret in Lamarck, Encycl. gil, Behon s.n., 1930; near Petrie, Blake 24 (K); Suppl. Meth. Bot. 2:236. 1811. (HOLO- Allora, Darling Downs, Blake 294 (K); Dray- TYPE: Photograph and fragment, from ton, Blake 5,177 (K); Mundubbera, Blonsome Desfontaines Herbarium at FI, in US! 19, 54 (K); Laidley, Clemens s.n., 11 May 1943 Chloris metcast acluja Sehrad. ex Schultes, (US); Kenmore, Clemens s.n., 18 May 1943 Mantissa 2:.340. 1824. (TYPE: "In Nova (US); Crow's Nest, Clemens s.n., 2A Feb 1944 Hollandia." Not seen, but description is (US); Yanaman Forest Reserve, Clemens s.n.. clear. 2 Aug 1944 (NY, UC, US); Moreton Dtr., Mt. Chloris truncata R. Br. f. ahhreviata Thel- Coolon, Clemens s.n.. Mar 1945 (K); Jericho lung, Viertelijahr. Naturforsch. Gesell. vicinity, Clemens s.n.. Mar-Apr 1946 (US); Cal- Zurich. 64:706. 1919. lide Cotton Research Station, Biloela, Coudrtj (ISOTYPE: Switzer- land: Derendingen, Probst 27 Sept 1917. 55 (K); Maranoa Dtr., Cumming 20 (NY); K!) Rockhampton, Davies s.n., 23 Apr 1932 (K); Gatton, Davies s.n.. 1932 (K); May Maranoa Perennial, 30 to 50 cm tall, stoloniferous; Dtr., Boatman Station, Everist 2,779 (K, US); sheaths glabrous, iigules short-ciliate; blades 9 Darling Downs Dtr., Benandri, 35 mi SE of to 17 cm long, 0.2 to 0.3 mm wide, glabrous to Texas, Everist 7,137 (US); Mt. Gravatt, Jack- scabrous; spikes five to thirteen, 5 to 23 cm son s.n.. Mar 1931 (K); near Mt. Sturgeon Sta- long, radiate; spikelets appressed, rather distant, tion, Lazarides 3,644 (K, US); Leichhardt Dtr., ca six per cm of the scabrous rachis; glumes 2 mi NE of Moura Township, Lazarides 6,910 narrowlv lanceolate, thin, hsaline, glabrous ex- (K); Moonie River, Roe 822 (US); Burnett, cept for the scabrous midrib; first glume 1.4 to Kingaroy, Smith 3,042 (K); Toowoomba, White 2..3 mm long, 0.2 to 0..3 mm wide; second glume 6,642; Wondai, White 7,214 (K); Dalbv, White 2.8 to 4.2 mm long, 0.2 to 0.4 mm wide; fertile s.n., Apr 1916 (K). lemma l.S to 4.5 mm long, 0.2 to 0.7 mm wide. Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 79

Fig. 48. Chloris tnincata. (A) habit, x 1/3; (B) spikelet, partly dissected; (C-E) florets, showing variation;

(F) sterile floret; (G) caryopsis. ( B-G ) \ 10. ) ) ) )

80 BniGHAM YouNO Univkhsity Science Bulletin narrowly elliptic, glabrous except for tlie ap- bet 42 (K). UNITED STATES: Hawaii, Ft. pressed-pilose margins, often becoming black at Shafter, Hitchcock 13,849 (F). maturity, apex truncate, callus bearded, awn 3.1 to 16.0 mm long; sterile floret one, rarely two, 32. CHLORIS PUMILIO Robert Brown. Prodr. 1.3 to 3.5 mm long, 0.5 to 0.9 wide, apex mm Flora Nov. Holl. IS6. 1810. (LECTO- truncate, awn 3.1 to 12.5 long; anthers ca mm TYPE: BM! Additional specimens K! BM! 0.6 mm long; cai-yopsis 1.7 to 2.2 mm long. See discussion of nomenclature below. ) Fig. 0.3 to 0.5 mm wide, ellipsoid to narrowh' obo- 49, J-P; 50, B. vate, trigonous; chromosome number 2n=40.

Chloris truncata is most closely related to Chloris pallida Hackel, Bot. jahrb. 6:244. C. ventricosa. Differences between the two have 1885. (HOLOTYPE: "In Australia boreali- been discussed under C. ventricosa. occidentali Ig. Pemberton-Walcott." W, at K! Everist (1935b) reported that this species Fragment seems to favor "black soil open downs, edge of Chloris ptimilio (Robert Brown) Kuntze, red soil country, and the edges of brigalow and Rev. Gen. PI. 2:771. 1891. (Based on C. balah country." Fig. 47—A illustrates the dis- pumilio Robert BrowiL tribution in Australia. Chloris ruderalis Domin, Biblioth. Bot. Representative specimens examined: AUS- 85:365. 1915. (SYNTYPES: "Queensland: 1910." ". TRALIA: Capital Terr.: Canberra, McKee 7,712 Prope opp. Atherton legi II. . . in (US), 8,931 (K), Evans 2,608 (K); New South xerodrymio prope opp. Mareeba legi II. ". 1910." . Wales: Nyngan, Baker s.n.. Dec. 1899 (US); . prope opp. Charters Towers Hastings River, Berkler s.n. (K); Tocumwal, legi II. 1910." All at K!, on extended loan Black m (US); Ballina, Black s.n.. Feb 1S94 from PR.)

(US); Singleton, Boorman s.n.. Nov 1914 (K); Chloris ruderalis f. biaristulata Domin, Menindee Dtr., Quondang Tank, Constable Biblioth. Bot. 85:365. 1915. (HOLO- 4,716 (K, US); Warialda, Hadlet/ s.n., Apr 1908 TYPE: A specimen at K!, on extended loan (F); Narromine, Helms 31,321 ". (K, NY); Guym, from PR, lalieled: . . Queensland apud McKee 646 (K); Stephen's Creek, Morris 489 Mungana prope opp Chillagoe legi II. 1910" (K); near Sydney, Vickenj 88 (K); Warren to consists of se\eral plants, one of which Collie Road, Vickenj K26 (K); Coonamble, fits the description in having two ver\' short Vickenj K27 (K); Windsor to Kingswood, lateral teeth at the base of the sterile floret

Vickertj and Constable 16,08.5 (K); Belltrees awn and is possiblx the plant upon which via Scone, White 8 (US); Queensland: Brisbane this fonn is based. River, Bick and White s.n.. May 1916 (K); Chloris ruderalis f. rolmsta Domin, Bib- Petrie, near Young's Crossing, Blake 48 (K); lioth. Bot. 85:.365. 1915. (TYPE: Ap- Wallangarra, Blake 4.422 (K); Cambooya, paientlv one of the specimens on the sheet Cashmore 808 (US); Dalveen, Clark s.n.. Mar discussed above, but no particular plant 1916 (K); Darling Downs Dtr., Ballandean, can be assigned positively to this name. Clemens 44,480 (K); Brisbane, Turbot Street pavement, Clemens s.n., 18 Mar 1942 (US); Annual 60 to 90 cm tall, tufted, or rooting Stanthorpe, Clemens s.n., 13 Nov 1943 (US); at the lower nodes, from a fibrous root svstem; Bald Mt., Clemens .s.n, 9 Nov 1944 (US); 35 sheaths glabrous except pilose near the apex; mi SE of Texas, Everi.st 7,140 (K, US); Cun- ligule a short-ciliate crown with a few long namulla, Gilnith Plains, Hartlei/ A18.5 (US); hairs; blades 7 to 12 cm long, ca 3 mm wide, Roma, White 9,573 (K); Darling Downs, be- scabrous abo\e with a few long hairs near the tween Inglewood and Milmenan, White 9,778 base, glabrous below except for the scabrous (K); Dalby, White s.n., Apr 1916 (K); Noondee midrib; spikes fom' to twelve, 4 to 8 cm long. Station near Warwick, Young 12 (K); South erect to slightlv divergent; spikelets densely Australia: Koonamore Vegetation Reser\-e, Pal- inserted. a%eraging 9 to 16 per cm of the scab- tridge 22 (K); Victoria: 2 mi S of Bacchus rous rachis; glumes narrowK lanceolate, gla- Marsh, Aston 282 (US); Rochester, Davies s.n., brous except for the scabrous midnerve, mem- 28 Jan 1932 (K); Melbourne University ground. branous; first glume 1.5 to 2.5 mm long, ca Sonenberg s.n., Jan 19.35 (K); Western Aus- 0.3 to 0.4 mm wide; second glume 2.5 to 5.0 tralia: Merredin," Allbani 11.48 (US): Carnar- mm long, with a short hairlike awn; fertile von, red muddy soil, Gardner G 3,003 (K); lemma .'3.7 to lO.O mm long ( including the Yundamindera Run, 120 mi NW of Kalgoorlie, lateral awns). 0.4 to 0.8 mm wide, narrowly Pearse 1,931 (K). FIJI ISLANDS: Suva, Ti/r- lanceolate-elliptic, callus bearded, margins in- BioLOGiCAi, Series, Vol. 19, No. 2 Ta.xonomv of the Genus Chloris (Gr.^mineae) 81

Fig. 49. Chloris lohata imd C. pumilio. (A-I) C. lohata. (A) habit, x 1/4; (B) spikelet, partly dissected, x seniilateral 5; (C-E) florets, showing variation, X 5; (F) sterile floret, ventral view, x 5; (G) sterile floret, pumilio. view, X 5; (H) group of sterile florets, showing variation, x 5; (I) caryopsis, x 10. (J-P) Chloris showing variation, x 5; (L) lower sterile (J) spikelet, partlv dissected, x 5; (K) group of fertile florets, x floret, lateral view', x 5; (M) upper sterile floret (not always present), x 5; (N) sterile floret, ventral view,

( of sterile florets, showing variation, x 5; ( ) caryopsis, x 5. 5; O ) group P 82 Brigham Young University Science Bulletin rolled, appressed short-pilose below, becoming Brown." The latter sheet is taken here to be spreading long-pilose above, otherwise miniitelv the lectotype, for it is the particular sheet se- scabrous, apex bifid, the lobes gradually taper- lected and annotated by Brown. All of the ing into awns up to 7 mm long, midnerve ex- specimens of the Bennett distribution at the tending as an awn 7 to 16 mm long, almost British Museum and Kew match the lectotype alwa)'s at least twice the length of the lateral as well. awns; sterile floret one (rarely two), 1.3 to 4.0 The totalit) of this evidence leads to only mm long (including the lateral awns), 0.3 to one conclusion: the plant (commonly called 0.7 mm wide, glabrous to slightly scabrous, apex Chloris rudcralis for many years) must now be minutely ciliate, divided to as much as halfway called C. pumilio, leaving the plant commonly into two lobes, the lobes asymmetrical and with called C. pumilio without a name. This was lateral awns 1.0 to 3.5 mm long, central awn correctl)' recognized by Lazarides ( 1972 ) , who 3 to 15 long, apex minutely ciliate; ear\ opsis mm provided the new name C. lohata for it. 1.2 to 3.3 mm long, ca 0.5 wide, ellipsoid, mm Specimens examined: AUSTRALIA: North- trigonous. ern Territory: Katherine River levee, 14°20-2rS, Chloris piiinilio and C. hihata are very simi- E of 132"E,' Black 17,414 (K, US); Queensland: lar. The best characters to use to distinguish Stannary Hills, Bancroft 1,90S: Forest Home C. pumilio are: lemma margins pilose for full Station,' Br«.9.v 1,821 (K); Gilbert River, Cum- length, sterile floret bilobed onl\ halfwa\ to berland, Brass 8,857 (K); Gilbert River, with- the base, and the asymmetrically awned lobes out precise location. White 1,489 (K); Stanley, of the sterile floret. near Townsville, Chisholm-Ellis 16 (K); Towns- Robert Brown (ISIO) described Chloris ville. White s.n., 11 Feb. 1918 (K); Thursday pumilio on the b;isis of specimens collected on Island, Ilockings s.n., Apr 1931 (K); 2 mi S islands off the northern coast of Australia. The of Nonnanton Township, Lazarides 4,272 (K); description is as follows: Hayman Island, White 10.222 (K); 'Western Australia: De Grey River, Mulgee Station, An- "4. C. pumilio, spicis 2-4 erectiusculis, derson 1,898 (K); North Kiniberle\- Dtr., May glumis bifloris: valvuhs acuminato-aristatis, River, Ba.sedow 75 (K); Broome, ' Bf/^es 7, .902 perianthiis eiliatis lanceolatis, triaristatus. (K); Holmes 1270.210 (US); Wandagee, Mouri- aristis lateralibus \alvula brevioribus: inter- v." bandy, Gooch s.n., 16 Jul 1937 (K); Kimmaya, media elongata. ( T. ) v. Holmes s.n., Apr 1942 (US); Whim Creek, This description was compared with the original Mitchell, without precise location or date (K); Brown manuscript notes at the librar\ of the Muhie, Roebounie, Morrison s.n.. Mar 1899 British Museum (Natural History). While many (K)'. of the words in this manuscript description are illegibly written, it seems to matcli Brown's 33. CHLORIS LOBATA Lazarides, Australian published description in the important points. Jour. Bot. Supplement 5. 20. 1972. ( HOLO- ". The phrase . . arista lateralibus valvula TYPE: "Queensland: Burke District; 4 ." brevioribus . . suggested something (juite dif- miles east of Nonnanton Township. 6. iii. ferent from the taxon that has been called C. 1954. Lazarides 4.289." CANB. ISOTYPES: pumilio for many years—the plant treated under K! US! ) Fig. 49, A-I. C. lohata in this paper. The latter species has Chloris pumilio auct. non Robert Brown. lateral awns that are clearly equal or nearly equal to the central awn and much longer than Annual 15 to 45 em tall, ert-et or geniculate, the lemmas. As with many of Brown's speci- often branching from thi' lower nodes; sheaths mens, typification is not straightforward, since glabrous, often sparseK' pilose toward the sum- many were distributed and confused after his mit; ligule a densely ciliate crown with a few death. In the case of this plant, however, there long hairs intenningled ni'ar the edges; blades can be little doubt; for in the British Museum up to 15 cm long, 0.2 to 0.6 mm wide, glabrous (and also at Kew), there are several sheets of to scabrous above ;md below; spikes three to Brown's specimens containing many plants— all nine, erect to somewhat divergent. 2.5 to 6.0 fitting the above description. Most of these have cm long; spikelets appressed and imbricate, ca the typical blue label of the Bennett distribution 8 to 15 per cm of the scabrous raehis; glumes of Brown's specimens and, as such, are not narrowly lanceolate, acuminate, membranous, necessarily strictly authentic. One sheet, how- glabrous except for the scabrous midnen-e; first ever, has on the back the handwritten nota- glume 1.6 to 2.5 mm long, 0.2 to 0.4 mm wide; tion "Nova Hollandia Ora Scptentrionalis. R. second glume 2.6 to 3.9 mm long, 0.2 to 0.3 mm )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 83 wide; fertile lemma narrowly elliptic, 5.5 to 11.0 River, Else\- Station, E of Mataranka, Blake mm long (including the lateral but not the 17,517 (K, US): Kathenne, Tijack-Bake, May middle awn), 0.3 to 0.6 mm wide, callus white 1943 (UC). Queensland: Forest Home Sta- bearded, margins glabrous to scabrous below, tion, Gilbert River, Brass 1.704, 1,787 (K); Gil- rather prominently long-ciliate near tin- apex, bert River, without precise location. White 1.492 remainder of lemma prominently scabrous antl (K); Nonnan Ri\'er, Gulliver, without precise punctate, lateral nerves of lemma prolonged collection number and date ( K, US); near Nor- into two awns 2 to 6 mm long, midnerve pro- manton, Lazarides 4,243 (K, US); Mt Molloy, longed into an awn 3.0 to 10.5 mm long; sterile McKee 9.121 (K); 39 mi E of Lawn Hill Sta- floret one, 3.3 to 7.5 mm long (including tlu' tion, Pernj 1.115 (K, US); between Townsville lateral awns), 0.4 to 0.5 mm wide, divided and Rollingstone, White 8,895 ( K, US). West- nearly to the base into two flattened lanceolate em Australia: 10 mi SE of Mt. House Station, lobes, each with an awn ca 2 to 4 mm long, Lazarides 5,157 (K, US). middle awn 3 to 8 mm long; car)'()psis 1.7 to 2.4 mm long, ca 0.4 mm wide, ellipsoid, 34. CHLORIS DIVARICATA Robert Rrown, trigonous. Prodr. Flora Nov. Holl. L186. 1810. As has been discussed under the previous (LECTOTYPE: 3. Chloris divaricata, Port species, this taxon has for many years been ." I. Keppel Ra\. Whorlwater Ray . . BMl mistakenly called Chloris pumilio. See additional nomenclature note below.) Collected on heavy soils on banks, water- Fig. 51, K-R. holes, and alluvial flats, mostly in Queensland, Chloris ct/nodontoidcs Balansa, Bull. Soc. Australia, but also occasionally found in North- Rot. France 19:318. 1872. (HOLOTYPE: ern Territorx and Western Australia. ( Fig. "Bords des Chemins, Nouvelle Cale- .50, C). 18-70" donie . . . Paneher, P! Representative specimens examined; .A,US- TRALIA: Northern Territory, on levee of Roper Perennial 20 to 50 em tali, tufted to shortly stoloniferous, sometimes apparently niat-fonning; sheaths glabrous; ligule a short-ciliate crown; blades up to 15 cm long, generally 5 to 9 em, 1.0 to 1.5 mm wide, glabrous to scabrous; spikes three to nine, 4 to 17 cm long, divaricate, becoming horizontal; spikelets sparse-

1\- inserted, about three to seven per cm of

tlie scabrous rachis, appressed, only the awns divergent; glumes very narrowly lanceolate, thin and membranous, glabrous except for the scabrous midnei-ve; first glume 0.9 to 1.8 mm long, ca 0.3 mm wide; second glume 2.0 to 2.9 mm long, ca 0.3 mm wide; fertile lemma linear to narrowly lanceolate, 2.9 to 4.0 mm long, in- eluding the teeth at the apex, callus bearded, margins glabrous, scabrous or with a few appressed siiort hairs near the apex, keel scabrous, internei-ves scabrous, awn 7.5 to 17.0 mm long, sterile floret one, 1.2 to 1.9 mm long, ca 0.3 mm wide, narrowly elliptic, glabrous, apex

acute, bilobed K to 'A of its length; awn 4.5 to 9.5 mm long; car\'opsis ca 2.2 mm long, 0.4 mm wide, narrowl)' ellipsoid. Brown's original description listed the sym- bol 'T.v.\-." as the place of collection. Accord- ing to studies of original expedition charts by Shield Burbidge ( 1956), this is taken to be Cape near Groot Eylandt in the Gulf of Caqjcntaria.

I could not find specimens labeled as such at the British Museum, but there were five speci- Fig. 50. Distribution of Chloris pectinata (A) East Coast," pumilio (B), and C. lobata (C). mens labeled "Chloris jmrpuram. Young University Science Bulletin 84 Bnir.HAM

G *^H "I y\

dis- pcclinala. (A) habit, x 1/4. (B) spikelet, partially Fie. 51. Chloris pectimta and C. divaricata. (A-J) C. ^ dorsal view, partia ly flattened, ;

The label of the last specimen is handwritten plants with relatively large numbers of long and has two slits cut in it—a feature thought spikes are intermingled with tufted plants with to indicate that this is the specimen selected fewer, shorter spikes. Plants showing different by Brown from the bulk of the collections and combinations of these features or showing inter- studied b\' him on his journey home from Aus- mediate values may also be found. Until addi- tralia. Fortunately, all specimens are of the tional field studies are made, I feel that recog- same taxon. nition of distinct varieties is not warranted. The two collection sites mentioned, the one CIdoris divaricata most closely resembles C. in the description and the second on the vari- pcctinata, though it differs in having sterile ous Brown specimens at the British Museum, florets 1.2 to 1.9 mm long (as opposed to 1.7 are widely separated—Cape Shield being on the to 2.9) and only slightly imbricate spikelets. west shore of the Gulf of Carpentaria and "Port The spikelets of C. pectinata are strongly ." I. Keppel Bay . . on the east coast of Queens- imbricate.

land. The latter specimen was obviousl) seen Everist ( 1935a ) reported that this species and studied by Brown; and even though its col- occurs in "practicalK' all types of soil, from lection site is at variance with that cited in the heavy black soils to light sandy loams." It has description, it seems best to designate it as the been collected in eastern Queensland and New leetotype of the species. South Wales, Australia; Fiji Islands; and Tonga Cliloris divaricata from Australia and C. Islands (Fig. 52). Specimens from Hawaii, adventives, as are prob- cijnodontoidcs from Fiji, Tonga, and New Cale- Texas, and France are donia have commonly been treated as distinct ably those from New Caledonia. species, though, more recently, Lazarides Representative specimens examined. AUS-

( 1972 ) treated them as varieties of C. divari- TRALIA: New South Wales: 6 mi NW of Mus- cata. The priman' differences emphasized are wellbrook, Storij 7.079 (K); Warialda, Vickenj that C. divaricata is not stoloniferous and has 18.028 (K). Queensland: Brisbane River, on ielati\'el\- long spikes, while C. cijnodontoidcs lidges, Badey without collection number or is stoloniferous and has short spikes. Table 10 date (K); Brisbane, Bailey without collection presents the results of measurements of 56 dif- number or date (US 879,412); Dalby, Beiers ferent specimens from all of the geographic 47 (K); near Wallumbilla, Belson s.n., 1930 ariMS represented. (K); Petrie, near Young's Crossing, Blake 49 As can be seen, onlv minor statistical dif- (K); Dravton, Blake 5,176 (K); Mundubbera, ferences exist between the two populations. Blonwme'41 (K); Mt. Molloy, Brass 2,405 (K); While average spike number and length mav South Brisbane Park, Cdemens s.n., 20 Feb 1943 differ, the ranges overlap greatly. Likewise, the (US); near Mt. Gravatt, Clemens s.n., 11 Mar

presence or absence of stolons is not sufficienth 1943 (US); Kenmore, Clemens s.n., 18 May exclusive to be used in separating the two 1943 (US); Rosewood, Clemens s.n., 18 Jan groups. 1944 (US); Bunya Mts., Clemens s.n.. Mar 1944 Lazarides (1972) has mapped the two vari- (US); Bauple, Wide Bay Dtr., Clemens s.n.,

10-20 ( ; Charleville, Dtr., eties, showing var. ci/nodontoides in inland Jun 1945 K ) Warrego areas of southern Queensland and coastal onl\' Clemens s.n., 9 Nov. 1945 (K); Mt. Fox, Clemens in the north. Collections of var. divaricata may s.n., Sept-Dec 1949 (K); Callide Cotton Re- also be noted from these regions but are much search Station, Biloela, Cowdrij 1,930 (K);

Table 10. Comparison of spike length, spike number, and growth habit of plants called C. divaricata and C. cijnodontoidcs.

Stolons Geographic Number of Number of origin of Number of spikes Length of spikes plants having plants lacking plant Range Average Range Average stolons stolons

Australia 2^9 5.7 4.5-16 11.1 16 27

Fiji, Tonga, and New CiJedonia 3-9 5.2 4-8 6.3 11 86 Bricham Young University Science Bulletin

near Nasinu, Greenwood 226D ( US ) ; Viti Leon, near Casey, Suva, TothiU 43 (K); Totoya Is- ^D land, Tothill 201 (K); Matuka Island, Tothill 291 (K); Kondavu Island, Turbet 8 (K). FRANCE: Dunquerque, port area, anon. (K). NEW CALEDONIA: Noumea, Bernier 812 (K); foot of Ouen Toro, McKee 11,499 (K. US). TONGA ISLANDS: Tongatapu, Kolomotuia, .-f*" Sokai 843 (K). UNITED STATES: Hawaii: Honolulu, near Bishop Museum, Neal s.n., 7 Sept 1934 (K); Niihau, St. John 23,623 (K); University of Hawaii campus, St. John 24,832 (K); WjWer .s.n., 19 Jan 1931 (K). Texas: Bexar Co., U.S. Experiment Station, San Antonio, Sil- veus 2,564 (TAES); Nueces Co., near Tuloso- Midway High School. Hwv. 9, between Corpus ,o Christi and Calallen, W. V.' Brown 755 (TAES); , ^S:> Refugio Co., 2 mi SE of Woodsboro, along D Road 1360, Waller et al. 2,118 (TAES). 35. CHLORIS PECTINATA Bentham. Flora Australiensis 7:612. 1878. (SYNTYPE: ^ "Charlotte Waters. Giles" K!) Fig. 51, A-J.

E Chloris divaricata var. muelleri Domin, Re- pert. Sp. Nov 9:553. 1911. (ISOTYPE: Fig. 52. Distribution of Chloris clivaricata. (A) east-

"Sturts Creek. Feb 56 . . . Mueller." K!) em Australia. ( B ) Fiji Islands. ( C ) New Caledonia. (D) Hawaiian Islands. (E) Tonga Islands. Chloris j)ectin(ita var. tt/pica Domin, Bib- lioth. Bot. 85:366. 1015." (Based on C. pec- Rosedale, Dovey 439 (K); Boatman Station, tinata Bentham.) Everist 2,778 2,931 (K), (K, US); Benandri, 35 Chloris pectinata var. fallax Domin, Bib- mi SE of Texas, Everist 7,139 (K, US); Bishop lioth. Bot. 85:366. 1915. (HOLOTYPE: Island, mouth of Brisbane River, Everist and "Queensland . . . Rolling Downs . . . Domin Blake 367 (K); Macalister, Harris 5.505 (K); II. 1910." K!, on extended loan from PR.) 10 mi S of Miles, Johnson 719 (K); Rcedsdale, Cldoris divaricata var. minor Black, Flora 5 mi NW of B\ mount, Johnson 2,251 (K); South Australia Pt. 1. 83. pi. 4. 1922. Gympie, Kenny without collection number or ." (ISOTYPE: "Oodnatta . . . Miss Staer . . date (US 1,386,466); Conjubov Station, Laza- K!) rides 4,137 (K); 1 mi NE of Niall Station, Lazarides 4,630 (K, US); Moreton Dtr., Water- Annual 20 to 75 cm tall, erect, often branched ford, Mead 7 (K); Jandowae, NE of Dalby, above, especially on taller specimens; sheaths Moore 36 (K); King's Creek, E of Bowman, glabrous; ligule a dense short-ciliate crown, oc- Mueller s.n. 1870 (K); Mitchell Dtr., Brixton, casionally with a few longer hairs intermixed; Ratcson s.n., 13 Nov 1930 (K); Holcombe, via blades up to 15 cm long, 2 to 5 mm wide, gla- Tara, Salter s.n., 21 Jun 1930 (K); near Lake brous to scabrous; spikes four to thirteen, 5 to Elphinstone. 28 mi VVNW of Nebe Township, 11 cm long, erect at first, later becoming widely

Story and Yapp 85 ( K ) ; St. George. White 7S9 divaricate to horizontally spreading at maturity; (US); Rockhampton, White 3,375 (K, US); spikelets densely inserted on the rachis, spread- Clermont, White 3,416 (K, US) Murgon, ing, ca 10 to 14 per cm of the scabrous rachis; White 7,230 (K); Rolleston, White 8,830 (K,, glumes narrowlv lanceolate, membranous and Darling Downs, between Inglewood and Mil- glabrous except for the scabrous midner\('; first merran. White 9,790 (K); Wellington, White glume 1.4 to 2.5 mm long, 0.1 to 0.3 mm wide; s.n., 17 Mar 1916 (K); Mareeba, White s.n., second glume 2.9 to 4.3 mm long, 0.2 to 0.3" mm Jan 1918 (K); Laidley, White s.n., 17 Mar 1921 wide; fertile lemma 3.0 to 6.2 mm long (includ- (K). FIJI ISLAND: Suva, Greenwood 226A ing lateral teeth), 0.4 to 0.6 mm wide, linear to (K); Viti Levu, Nandronga, Singatoka, Green- narrowly lanceolate, becoming dark gray to wood 226C (K, NY, US); ViH Levu, Nafasiri, black at maturity, callus bearded, margins ) )

Biological Series, Vol. 19, No. 2 T.v.xonomy of the Genus Chloris (Gramineae) 87 glabrous, scabrous, or appressed short-pilose nudem; Nov. Gen. et Sp. 5. 1816. The (the hairs less than 0.2 mm long), sides of first cited publication has not been located lemmas sparsely to densely glandular and scab- in any United States library, but Mrs. Ag- rous, occasionally glabrous, lemma teeth 0.5 to nes Chase found the series in the British 10.0 mm long, awn 6 to 37 mm long; sterile Museum and placed copies of Lagasca's floret one, 1.7 to 2.9 mm long, 0.2 to 0.3 mm protologues on file at US! The name was wide, flattened, bilobed no more than halfwav apparently first used without a description; to the base, glabrous, awn 4 to 10 mm long; later Lagasca provided a description in the anthers ca 0.6 mm long; caryopsis ca 2.3 mm second work mentioned above. Parodi long, 0.3 mm wide, narrowly ellipsoid, trigonous. ( 1947 ) states that no specimen with the

Chloris pectiimta is most closely related to appropriate number could be found among C. divaricata. Differences have been discussed the types at Madrid, but that a figure bear- under the latter taxon. ing Lagasca's handwritten annotation is of

This species is widely distributed in central the species as commonly understood. Fig. and western Queensland and also in Northern 53, D-F. Territory and South Australia (Fig. 50, A). Chloris mendociim Philippi, Anales Univ. Everist (193S) reported that it occurs in a Chile 36:208. 1870. The original descrip- variety of habitats but develops best in low- tion cites a collection by Mendoza. A speci- lying clay soils. men in P! labeled "Phillipe pi. Representative specimens examined: AUS- chilense Mendocina. Chloris mendocina Phil. Ipse TRALIA: Northern Territoiy: 26 mi NNW of Mendoza" may be the type. It and the Brunette Downs, Blakc 17,819 (K, US); Mac- description are Chloris crinita. Donald Downs, Chalmers 32 (K); 22 mi NE of Argadargada Station, Lazarides 5,258 (K, Trichloris fasciculata Fournier v\ Bentham, US); Barkl)' Highway, James River crossing, Jour. Linn. Soc. Bot. (London) 19:103. Pernj 710 (K); Queensland: Eulo, near Cunna- 1881. Nomen nudem; Fournier, Mex. PI. mulla, Allan 10.948 (US); Forest Home Sta- 2:142. 1886. (HOLOTYPE: "San Luis de Hon, Gilbert River, Brass 1,822 (K); Charleville Potosi. Virl. n. 1440" P! Dtr., Warrego, Clemens s.n., 7 Nov 1945 (K); Trichloris hlanchardiana Fournier ex Scrib- Jericho, Clemens s.n., 6 Mar 1946 (US); ncr, Bull. Torrev Bot. Club 9:146. 1882. Mitchell Dtr., Springvale, Clemens s.n., 2 Apr (HOLOTYPE: "Santa Cruz Valley near 19" 1946 (US); Maranoa Dtr., Boatman Station, Tucson . . . Pringle, May US!) Everist 2,778 (US); Cook Dtr., road from Taba- Chloropsis hlanchardiana (Fournier) cum to Springmount, Goodall s.n., 2 Apr 1961 Kuntze. Rev. Gen. PI. 2:771. 1891. (Based (US); Hughenden, Hawthorn s.n., Jun 1919 on Trichloris hlanchardiana Fournier.) (K); 27 mi of Cloncuru', Lazarides 4.308 NW Chloropsis crinita (Lagasca) Kuntze, Rev. (K, US); 3 mi SW of Malbon Township. Gen. PI. 2:771. 1891. (Based on Chloris Lazarides 4,401 (K, US); Toorak Experiment crinita Lagasca.) Station, Lazarides 4,446 (K, US); Adovale, CIdoropsis fascictdata (Fournier ex Ben- MacGiUivraij 1,029 (K); Cunnamulla, McKee tham) Kuntze, Rev. Gen. PI. 2:771. 1891. s.n., 11 Apr 1963 (K); 25 mi NNE of Cam- ( Based on Trichloris fasciculata Fournier ooweal. Perry 955 (K, US); Blackall, Ranking, ex Bentham. 13 (K); Longreach Dtr., Thomas s.n.. Mar 1927 Leptochloris (K); Clermont, White 3,422 (K); Antil Plains crinita (Lagasca) Munro ex Kuntze, Rev. Gen. PI. 2:771. 1891. (Based near Townsville, White 8,893 ( K, US); War- on Chloris crinita rego Dtr., Carbeen, near Cunnamulla, White Lagasca.) 11,556 (K); Mitchell Dtr., Tower Hill, White Trichloris verticillata Fournier ex Vasey, s.n., Apr 1919 (K); Quilpie, White s.n.. 20 Mav U.S. Dept. Agric. Div. Bot. Bull. 12. Pt. 1928 (K); South Australia: Cordillo Downs, II: pi. 25. 1890. "Chloropsis hlanchardiana Cleland s.n.. May 1924 (K); Oodnadatta, Staer Gav in Herb." is cited; description and illus- s.n., Jan 1913 (K); Western Australia: Wan- tration are of C. crinita. dagee, Minilya River, Gardner 3,236 (K); Wan- Chloropsis inendocina (Phihppi) Kuntze, dagee Station, near Caniar\'on, Meadh/ A/77 Rev. Gen. PI. 3:.348. 1898. (Based on (K); Warrambic, Roebourne, Meares 345 (K). Chloris mendocina Philippi.)

Trichloris mendocina f. hlanchardiana 36. CHLORIS CRINITA Lagasca, Var. Cienc. (Fournier ex Scribner) Kurtz, Bol. Acad. Lit. Arts (Madrid) 4; 143. 1805. NoHien Nac. Ciencias Cordoba (Rep. Argentina) 88 Bricham Young University Science Bulletin

Fig. 53. Chloris phiriflora and C. crinita. (A-C) C. jih.trijlora. (A) habit, x l/."); (B) .spikelet, partly di.s- sected, x 10; (C) caryopsis, x 10. (D-F) C. crinita. (D) habit, x 1/5; (E) spikelet, partly dissected, x 10; (F) caryopsis, x 10. )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 89

16:270. 1900. (Based on Trichloris blanch- tes: Dep. Mburucuya, Estancia Santa Teresa, arkina Fournicr c.x Scribner. Pedersen 63 (US); Prov. Fonnosa: Guayculea, Chloris trichodcs Lagasca ex Parodi, Revis- Jorgensen 3,330 (US); Prov. Jujuy: Laguna de Fries Prov. Mendoza: Vallee ta Argentina Agron." 14:62. 1947. Citation la Brea, 115 (US); Wilczek 526 (US); of an herbarium name. de Rio Atuel, St. Raphael, Prov. Salta: Coronel Moldes, Htinziker 1,110 Tricltloris crinita (Lagasca) Parodi, Re- (US); Prov. San Juan: Colonia Fiscal, Sar- vista Argentina Agron. 6.3. 1947. (Based iniento, Bartlett 20,522 (US); Prov. Santiago del on Chloris crinita Lagasca.) Estero: Rio Huayco Hondo, Bartlett 19,754 Stoloniferous or tufted perennial up to 1 (US); Prov. Tucuman: Trancas, Tapia, Venturi m tall; sheaths glabrous to sparsely hirsute; 2,345 (US). BOLIVIA: Mataral, anon., 1804 ligule ciliate with cilia up to 3 mm long; blades (US). MEXICO; Chihuahua: 100 km W of up to 20 cm long, 5 to 10 mm wide, scabrous Cuarto Cienegas, Harvey 1,255 (US); Durango: above and below; spikes six to twenty, up to Torreon, Hitchcock 7.724 (US); San Luis Potosi: 15 cm long, erect, arising in several close ver- 20 km NE of Rio Verde, Rzedowski 5,157 (US). ticils, spikelets densely arranged on the rachis, PARAGUAY: Dep. Chaco: Loma Para, Rojas averaging ca seven to nine per cm of the scab- 2,751 (US). UNITED STATES: Arizona: Ben- ovate, acumi- rous rachis; glumes lanceolate to son, Griffiths 1,969 (US); New Mexico: Organ for the scabrous nate to awned, glabrous except Mts., W of Dona Ana, Wooton and Standletj s.n.. midrib; first glume 0.8 to 1.1 mm long, ca 0.3 Sept 1907 (US); Texas: vicinity of El Paso, mm wide, linear to narrowly lanceolate; second Chase 5,899 (US); Fort Worth, Tracy 8,208 glume 2.0 to 2.5 mm long, ovate, with an awn (US); Pecos, Silveus309 (US). up to 2 mm long; fertile lemma 2.4 to 3.S mm long, ca 0.5 wide, dorsally flattened, nar- mm 37. CHLORIS PLURIFLORA (Fournier) Clay- rowly lanceolate to elliptic, scabrous, especialh ton, Kew Bull. 21:102. 1967. (Based on three-awncd, central above the middle, apex Trichloris pluriflora.) Fig. 53, A-C. awn S to 12 mm long, the lateral usually somewhat shorter; sterile floret usually one (occa- Trichloris pluriflora Fournier in Hemsley, sionally two), greatly reduced, cylindrical, the Biol. Centr. Amer. Bot. 3:560. 1885. floret proper ca 1.0 to 1.5 mm long, gradualh' narrowing at the apex into three subequal awns, the awns ca 5 to 7 mm long; second sterile floret (when present) similar to the first but smaller; caryopsis 1.7 to 2.3 mm long, ca 0.6 mm wide, ca 0.3 mm thick, strongly dorsally flattened; chromosome number 2n^40. Onh three species in Chloris have three long awns on the feitile and sterile florets— C. crinita, C. lobata. and C. pumilio. Chloris crinita is a New World tufted perennial with the lemma apex tapering abraptly to the lateral awns. Both Chloris lobata and C. pumilio are Aus- tralian annuals in which the upper lemma is split, the lobes gradually narrowing into the awns. There are manv additional, subtle, quali-

tative differences; and I believe the resemblance

is largelv superficial.

Chloris crinita is widelv distributed in the south central to southwestern United States, northern Mexico, and, rarely, has been collected in South America (Fig. 54). Representative specimens examined: AR- GENTINA: Prov. Buenos Aires: Dep. Villarino. 60 km E of Rio Colorado, Eijerdam et al. 2.3,526 (US); Prov. Catamarca: Santa Maria, Castillon 3,290 (US); Prov. Chaco: Siena Blanca, Jor-

^ensen 2,878 (US); Prov. Cordoba: Reinedias, Fig. 54. Distribution of Chloris crinita. Inset A: South- Niedfeld s.n., 17 Apr 1920 (US); Prov. Corrien- ern United States and Mexico. 90 Bricham Young University Science Bulletin

Nomen nudem; Fournier Mcx. Pi. 2:142. Austi-alian species has the sterile floret with a 1886, with description. (HOLOTYPE: bearded callus. "Karw. in Herb. Petrop." Fra^nent of tvpe The presence of the lateral awns in Chloris in US!) pluriflora and in some Australian species would Chloropsis pluriflora (Fournier) Kuntze, suggest a close relationship, but there are many Rev. Gen. PL 2:771. 1891. (Based on features that are not shared; and I believe the Trichloris pluriflora Fournier.) similarits' is superficial. Chloris mollis is very similar phenetically, Trichloris pluriflora f. macra Hackcl in lacking only the well-developed lateral awns; Stuck., Anales Mus. Nac. Hist. Nat. Buenos I believe that it and C. pluriflora closely Aires 14:116. 1911. Notnen nudem. are related. Though this name is without a description, For many years Chloris pluriflora has been a dupHcate of a specimen in US! from the placed in Trichloris, along with C. crinita. Ar- Hackel Herbarium is t)'pical Chloris pluri- guments for the merger of the two genera were flora. presented by Clayton (1967). Earlier in the Trichlori.

Chloris pluriflora is most similar to C. utollis (a Central and South American species) and to C. pumilio and C. lobata (both from Aus- traha). It may be separated from all three in having usually two or more sterile florets. In Fig. 55. Distribution ot Chloris pluriflora. Inset A: addition, the presence of the lateral awns will Southern United States, Mexico, Central America, differentiate it from C. mollis. Neither of the and Caribbean Islands. );

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae) 91

niontes, anon., K. Pflanz no. 2,097 (US). in pure form, it differs markedly from these two CUBA: Prov. Habana: Casa Blanta, Ekman species (see Table 11 and Fig. 56, A-K, L-U; 16,879 (US). ECUADOR: Prov. Loja: benvccn 58, D-K). A complete discussion of the hybrid- La Toma and Loja, Hitchcock 21,400 (US). ization and introgression between these species GUATEMALA: Peten, Tikal National Park, follows the description and discussion of C. Lundell 16,066 (US). MEXICO: Coahuila: andropogonoides (p. 95).

Sabina.s, Nelson 6,827 (US); Neuvo Leon: 12 Chloris cucullata is widely distributed

mi S of Montcrrev, Mueller 375 (US); Oaxaca: throughout much of Texas and is also found Canon df Santa Catarina, Conzatti 3,985 ( US ) in adjacent areas of New Mexico and Mexico Tamaulipas: Podilla, Swallen 1,734 (US). ( Fig. 57, shaded circles ) . It is a common road- PARAGUAY: Asuncion, Puerto Casado, Rajas side or waste area weed in many areas. Cuzco, Gunther 66.5 2,307 (US). PERU: Representative specimens examined: NOTE: (US). UNITED STATES: Texas: Cameron Additional citations may be found in the Co., Resaca del Rancho Viejo, 6 mi NE of list of hybrid populations studied following Brownsville, Cory 51,411 (US); Duval Co., 2 mi the discussion of introgression and hybridization of Benavides, Harvey 895 (US); Kleberg SW of C. cucullata, C. verticillata, and C. andro- of Kingsville, Sicallen 10.258 (US); Co., W pogonoides.) MEXICO: Nuevo Lchju: below Webb Co., Laredo, Griffiths 6,542 (US); Wil- La Boca Canyon, R. F. Smith M596 (TEX). lacy Co., E of Ravmondville, Swallcn 10,100 Tamaulipas: .34 mi S of Matamoras, Crutch- (US). field and Johnston 5,475 (TEX); 5 km S of Nuevo Laredo, Dominguez 53 (TEX); 24 km 38. CHLORIS Bischoff, Ann. CUCULLATA S of Nuevo Laredo, IDominguez and McCart 1853. Sci. Nat. Bot. (Paris) III. I9:.3.57. 8,208 (TEX). UNITED STATES: New Mex- (TYPE: "Provincia Tamaulipas prope Mat- ico: Chaves Co., 21 m E of Roswell along Hwy. 1849". amoros . . . Engelmann, Not seen, .380, Anderson 4,740 (HSC); Dona Ana Co., Las but description elcarl\- refers to this taxon. Cruces, Plank sji., July 1894 (NY); Eddy Co., Fig. 56, A-K. 21 mi NW of Carlsbad, Gould and Kapadia Co., mi Perennial, tufted, erect. 15 to 60 em tall; 9,543 (TEX). Texas: Andrews 8 NW Armstrong sheaths glabrous; ligule a crown of very short of Andrews, Rowell 5,879 (TEX); 5,Z7 scabrous midnei-ve, membranous; first mi E of Buchanan Dam, mi E of glume 0.5 to 0.7 mm long, 0.4 to 0.5 mm wide, Celarier 5,476 (UC); Caldwell Co., 8 ovate to broadly lanceolate; second glume 1.0 Luling, M. C. Johnston 6,215 (TEX); Cam- Wildlife Refuge, to 1.5 mm long, 0.4 to 0.6 mm wide, obovate; eron Co., Laguna Atascosa

3,472 ( ; Colorado Co., Colum- fertile lemma 1.5 to 2.0 mm long, 0.7 to 1.0 Fleetwood TEX ) Co., E of mm wide, broadly elliptic, apex obtuse, callus bus, Anderson 3,282a (HSC); Comal An- short-bearded, keel and marginal nerves ap- New Braunfels, near Mission Valley Mills, pressed-pilose, otherwise glabrous, awn 0.3 to derson 3,888 (HSC); Duval Co., E of Crestonia, Co., 18 mi L5 mm long; sterile floret one, LO to 1.5 mm Barkley 13,003 (TEX); Ector W Frio Co., 4 long, 1.0 to 1.5 mm wide, markedly inflated of Odessa, Warnock 7,902 (TEX); C. with the upper margins inrolled, unawned or mi E of Zavala Co. line on Hwy. 76, M. of Post, with a short awn up to 1.5 mm long; canopsis Joluuton 6,192a (TEX); Garza Co., E 51-1,562 (TEX); Gon- 0.9 to 1.2 mm long, ca 0.5 mm wide, obovoid, Tharp and Gindnede s.n., 23 rounded to trigonous; chromosome number zales Co., Athens, Tharp and Higdon 2n=:40. Apr 1938 (TEX); Hall Co., near Estelline, Co.: McAllen, Chloris cucullata is quite distinct from most Gould 7,721 (UC); Hidalgo Santa Ana other species in the genus. It hybridizes readily vacant lot, Anderson 3,876 (HSC); with C. andropogonoides and C. verticillata, but. National Wildlife Refuge, M. C. Johnston Bbicham Young University Science Bulletin 92

A) h.hit x 1 /4 .erticiUan.. (A-K) C. c^:M.a. , Fig. 56. Ckloris cucutlata and C. :^^'fZ chssecte^^^^^ ^^^^^l^^^ ^ (right) x 4; (C) spikelet, partly (left) and lower side verticillata. (L) habit, showing variation, x 10; (K) ^^^^P'"'.!/,lW-V)C. above, X 10; (F-J) florets, '^ ^m) partly dissected, x 4. ( I\ .spiKeier,spikelet pa lower side (>'ght , ) y 1/4- portion of spike, upper side left and X (M) caryopsis. x 10. florets, showing variation, x 10; (U) X 10 (O) sterile floret, x l6; (P-T) Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohls (Ghamineae) 93

Canyon, Reed -3,984 (TEX); San Patricio Co., Welder Wildlife Refuge, near Sinton, Gould and Hijcka 8,020 (UC); Starr Co., 6 mi NW of La Gloria, Sanchez et al. 8,3-33 (TEX); Stonewall Co., Double Mt., Tharp and Gim- brede 51-1,551 (KANU, TEX); Tarrant Co., Ft. Worth, McMullen s.n., Aug-Oct 1927 (TEX); Taylor Co., Abilene State Park, Hen- derson 6.3-899 (TEX); Terrell Co., 2.5 mi S of Sheffield, Gould 9,699 (TEX, UC); Travis Co.:

Camp Mabry, Austin, Tharp s./i., 21 Apr 1938 (TEX); near Bee Caves, jet. of Hwys. 620 and 71, Anderson 3,273 (HSC); east end of Mans- field Dam, Anderson 3,274 (HSC); Val Verde Co.: 9 mi S of Del Rio, along Hvvy. 277, An- derson 4,466 (HSC); 35 mi N of Del Rio, near Loma Alta, Warnock and McBryde 15,154 (TEX); Ward Co., 1 mi NE of Sand Hills, Rose-Innes and Moon 183 (TEX); Webb Co.: Laredo, Martinez and Trevino 3 (TEX); 40 mi NW of Laredo, Emery 668 (TEX); Wichita Co.: Wichita Falls, McKce s.n.. Sept 1927 (TEX); Red River, above Burkburnett, Tharp o=C VERTICILLATA •=C.CUCULLATA s.n., 20 Julv 1921 (TEX); Willacv Co.: 2.5 mi o =C. ANDROPOGONOIDES S of Ytturia, Anderson 3,311 (HSC); Redfish Bay, Tharp s.n., 5 Mar 19.34 (TEX); 7 mi N of Ravmondville, Tharp and Brown 3,2-32

(TEX)'; Wilson Co., Parks -s.n., 25 Dec 1946 Fig. 57. Distribution of Chloris cucultata (shaded (TEX); Zapata Co., 19 mi S of Laredo, Mar- circles), C. verticillata (dotted circles), and C. tinez and Trevino 37 (TEX). andropogonoides (open circles).

39. CHLORIS VERTICILLATA Nuttall, Trans. .5.3-674 (TEX); Jim Hogg Co., 30 mi S of Heb- Amer. Philos. Soc, n.s. 5:150. LS37. bronville, Tharp 5,912 (TEX); Wells Co., Jim ". (HOLOTYPE; . . Arkansas. Wm. Nuttall" Romarsid Ranch, Freeborn ami Freeborn 524 Photograph of tvpe, from PH, in US! ISO- (TEX); Karnes Co., Karnes Citv, ] C. Johnson TYPE: K! ) Fig. 56, L-U. 813 (TEX); Kenedy Co., 2 mi 'S of Sarita, An- derson 3,305 (HSC); King Ranch, Norias Divi- Perennial 14 to 40 cm tall, tufted, erect or sion, M. C. Johnston 5.319.77 (TEX); Kleberg decumbent, sometimes rooting at the lower Co., King Ranch, Santa Gertrudis Division, nodes; sheaths glabrous; ligule a ciliate crown M. C. Johnston 5.319.76 (TEX); King Ranch, with some hairs up to 2 mm long; blades to Laureles Division, M. C. Johnston 5.319.81 15 cm long, 2 to 3 mm wide, glabrous to scab- (TEX); 2 mi N of Kingsville, Swallen 10,085 rous; spikes ten to sixteen, 5 to 15 cm long,

(US); La Salle Co., ,3 mi S of Dille\-, Painter usuallv in several verticils, these separated by and Barkley 14,310 (TEX); Live Oak Co., 5 mm or more, inflorescence terminating in a Tharp and Gimbrede 52-5.52 (TEX); Llano vertical spike; spikelets sparsely inserted, aver- Co., Enchanted Rock, Web.ster 47,261 (TEX); aging ca four to seven per cm of the scabrous McMullen Co., 10 mi S of Tilden, Alvarez et al. rachis, appressed; glumes lanceolate, membra- 7,714 (TEX); Mason Co., 18 mi N of Fredricks- nous, glabrous except for the scabrous midnei-ve; bm-g, Emert/ 814 (TEX); Midland Co., Mid- first glume 2 to 3 mm long, ca 0.3 mm wide; land, Tracy 7.962 (TEX); Nolan Co., 10 mi second glume 2.8 to 3.5 mm long, 0.3 to 0.4 S of jet. of Hms. 126 and 53, Anderson 4,651 mm wide; fertile lemma 2.0 to 3.5 mm long, (HSC); Nueces Co., ca 1 mi N of Kingsville, 1.5 to 1.9 mm wide, elliptic to lanceolate, callus

Anderson -3.302 ( HSC ) ; Corpus Christi. ^Heller bearded, keel glabrous to appressed-pilose, s.n., 14-21 Mar 1894 (UC, US); Mustang Is- margins appressed-pubescent, apex acute to land, /4 mi S of Port Aransas, Anderson 3,293 somewhat obtuse, awn 4.8 to 9.0 mm long; (HSC); Pecos Co., 6 mi N of Ft. Stockton, sterile floret one, 1.1 to 2.3 mm long, 0.5 to Warnock T464 (TEX); Randall Co., Palo Duro 1.0 mm wide, oblong, somewhat inflated, trun- :

94 Brigham Young University Science Bulletin cate, glabrous, avvn 3.2 to 7.0 mm long; caiyop- and McGregor 3,785 (KANU, UC); Meade Co., sis 1.3 to 1.5 mm long, ca 0.5 mm wide, ellip- County Park, Horr 3,272 (KANU); Montgom- soid, rounded to trigonous; chromosome num- ery Co., 1 mi N of Independence, along Hwy. ber 2n = ca 28, 40, 63. 7.5, Anderson 2,761 (HSC); Morris Co., 4 mi Chloris verticillata resembles the Austrahan SE of Council Grove, Richards 1,788 (KANU); C. trtincata; it differs in having a single sterile Morton Co., 7.3 mi N of Elkhart, Richards floret (rarely two). Chloris verticillata also has 2,492 (KANU); Neosho Co., 5 mi E of Erie, several somewhat remote verticils of spikes Holland 488 (KANU); Osage Co., Lyndon, terminated by a single erect spike, while C. Anderson 2,766, 2,767 (HSC); Pottawatomie truncata has a single verticil and lacks a ter- Co., without precise locality, Imler s.n., 13 Jun minal spike. 1929 (KANU); Republic Co., T4S, R2W, sec. Morley Chloris verticillata, C. ondropogonoides, and 22, 1,227 (KANU); Riley Co., without precise location, Gates C. cucullata introgress in Texas. A discussion of 12,801 (UC); Shawnee this follows the treatment of C. ondropogo- Co., near Wakarusa, Anderson 2,768, 2,769 noides. (HSC); Stafford Co., 8 mi NE of Hudson, Mc- Gregor 10.515 (KANU); Wichita Co., grass- Chloris verticillata (commonly called "wind- land, Agrelius .'i.n., 12 Aug 1912 (KANU); mill grass") is widely distributed in the central Wilson Co., 8 mi N of Neodesha, along Hwy. United States, where it is commonly found as 75, Anderson 2,762, 2,763 (HSC); Woodson a weed of roadside, lawn, or waste areas ( Fig. Co., T24S, R14E, sec 21, Lathrop 471 (KANU). 57, dotted circles). Disturbance, cultivation, and Missouri: Atherton, Bush 7,352B (TEX). overgrazing have largely eliminated much of Nebraska: Cherry Co., Valentine, Tolstead the native vegetation in this area, but earl\ 4-463 (NEB); Dundy Co., 3 mi S of Benkel- records (Barkley, 1965) indicated that windmill man, Andcr.wn 2,774, 2,776 (HSC); Hitchcock grass was found in generally low areas of the Co., Swanson Reservoir, Anderson 2,772 (HSC); native prairie. Nemaha Co., Auburn, Andenson 2,770 (HSC); Representative specimens examined: UNIT- Nickolls Co.. Superior, Bates 6,635 (NEB); ED STATES: Arizona: Gila Co., 7 mi W Redwillow Co., Indianola, Anderson 2,771 of Young, anon. (UC). Colorado: Baca Co.. (HSC). New Mexico: Cuny Co., 14 mi W of Sand Canvon, 25 mi S of Pritchett, C. L. Por- Melrose, along Hwy. 60, Anderson 4.663 (HSC); ter 4,272 (TEX); Che\enne Co., near Cheyenne San Miguel Co., Las Vegas, Whitehouse s.n., Wells, Harrington and Smith 384 (UC); Crow- 3 Sept 1929 (UC). Oklahoma: Cimarron Co., ley Co., Rocky Ford, Harrington 685 (UC); 6 mi E of Kenton, Rogers 5,933 (TEX); Cotton Powers Co., 10 mi W of Lamar, Harrington 724 Co., 5 mi N of Burkbumett, Rose-Innes and (UC). Iowa: Story Co., Ames, /. P. Smith 2,597 Moon 973 (TEX); Muskogee Co., rockv soil, (HSC). Kansas: Anderson Co., 3 mi S of Gar- Behh s.n.. H Sept 1940 (TEX); Payne Co., W nett, Horr and McGregor 3,591 (KANU, TE.X); of Stillwater, E.stes 57 (TEX). Texas: Ander- Barber Co., 1 mi W of Medicine Lodge, Mc- son Co., Engeling Wildlife Management area, Gregor 14,029 (KANU); Bourbon Co.,1 mi S Calley and Mar.^i 13 (TEX); Archer Co., 7 of Uniontown, Ungar 111 (KANU); Chase Co., mi S of Windthorst, Anderson 3.921 (HSC); 4 mi E of Marion, Horr 3.497 ( KANU, TEX ) Bell Co., old Tennessee Valley community site, Chautauqua Co., roadside grassland, Horr 1,540 York and York 54,447 (TEX); Brewster Co., (KANU); Clav Co., waste ground, Imler s.n., 24 mi E of Alpine, along Hwy. 90, Ander.wn 14 Jun 1929 (KANU); Coffee Co., Burlington, 4.447 (I ISC); Altuda Pa.ss, Glass Mts., Warnock Ander.wn 2,765 (HSC); Dickinson Co., without W362 (TEX); Burnet Co., Longhoni Caverns precise location, Imlcr s.n., 15 Jun 1929 State Park, near Marble Falls, Silvetis 7.642, (KANU); Ellis Co., 6 to 7 mi SVV of Hays, 7,643 (TEX); Clay Co., Prospect Hill, Silveus Rijdberg and Imler 1,248 (KANU); Ellsworth 117 (TEX); Comanche Co., DeLeon, Ander.wn Co., 1 mi E of Terra Cotta, McGregor 9,212 3,919 (HSC); Culberson Co., McKittrick Can- (KANU); Finney Co., Garden City, Kellerman von, Guadalupe Mts., Fischer s.n., 20 Jul 1950 s.n., 20 Aug 1882 (KANU); Ford Co., 1 mi (KANU); Dallam Co., 21 mi NW of Dalhart, NW of Ford, Hulhert 3.207 (KANU); Gear)- Rowell 5,388a (TEX); Dallas Co., Dallas, com- Co., waste land, Imler s.n., 15 Jun 1929 mon on prairie, Bii.^li 1.164 (US); Garza Co., (KANU); Grant Co., Thompson 55 (UC); 3 mi E of Post, Tharp and Gimhrede 51-1,558 Greenwood Co., 3.5 mi W or Woodson Co. line on Hwy. 54, Lathrop 3.186 (KANU); Harvev (TEX); Gray Co., 5 mi W of Pampa, Brown Co., 8.7 mi E of Newton, Harms 2,192 (KANU, 3,382 and Tharp GR 239 (TEX); Hays Co., W UC); Kiowa Co., 7 mi SE of Haviland, Horr of Wimberly, /. C. Johnson 258 (TEX); Hemp- )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 95 hill Co., 5 mi E of Canadian, Rowell 4.078, lemma 1.9 to 2.7 mm long, 0.5 to 0.6 mm wide, 4,174 (TEX); Howard Co., Big Spring, Hitch- narrowly lanceolate to elliptic, apex acute, callus cock 13,366 (US); Hudspeth Co., 4 mi NW of bearded, margins and keel appressed-pilose, Sierra Blanca, Anderson 4,420 (HSC); Jones otherwise glabrous, awn 1.9 to 5.2 mm long; Co., eastern part of county, Tharp and Ginihrede sterile floret one, 0.9 to 1.7 mm long, 0.25 to 51-1,556 (TEX); Lampasas Co., N of Lampasas, 0.5 mm wide, narrowly cylindrical, apex obtuse, Silveus 7,540 (TEX); Menard Co., Ft. Mc- glabrous, awn 2.5 to 3.5 mm long; caryopsis Cavett, Tharp 5,224 (TEX); Mills Co., Priddy, 1.3 to 1.4 mm long, ca 0.4 mm wide, ellipsoid, Anderson 3,918 (HSC); Nolan Co., 10 mi S of rounded to trigonous; chromosome number jet. Hvv)'s. 126 and 53, along .53, Anderson 2n=40.

4,650 (HSC); Palo Pinto Co., 0.5 mi N of Chloris andropogonoides is similar to C. vct- Stravvn, Anderson 3,920 (HSC); Parker Co.. ticillata and C. texensis. It may be separated SW of Weatherford, Silveus 7,543 (TEX); from the former by its single verticil of spikes, Pecos Co., 5 mi E of Brewster-Pecos Co. line, generally shorter awns, and narrower sterile along Hwy. 90, Anderson 4,460 (HSC); Potter florets (Figs. 56, L-U; 58, A-C, D-K). Addi- Co., N of Amarillo, Brenckle 48,302 (US); tional differences are given in Table 11. Randall Co., near Palo Duro Canvon, young Chhris andropogonoides is generally smaller Runnells Co., 0.5 mi s.n.. Sept 1917 (TEX); in spikelet dimensions than C. texensis but dif- of Ballinger, 4,641 (HSC); San N Anderson fers more obviously in having spikes that are 3,333 Saba Co., Richland Springs, Anderson floriferous to near the base, while the spikes 58,249 (HSC); 2.1 mi SE of San Saba, Cory of C. texensis are naked for several cm above (UC); Sherman Co., 18 mi S of Stratford, the base. without number or Weaver and McLaughlin, As discussed in the following section, Chloris park, date (TEX); Sterling Co., Sterling Citv, andropogonoides, C. cucullata, and C. verticil- Co., near Ft. Anderson 3,002 (HSC); Tarrant lata are involved in extensive h\bridization and Worth, Ruth 302 (NY, UC); Taylor Co., Camp introgiession in Texas. Berkeley, Tolstead 7,445 (UC); Travis Co., Bee Chloris andropogonoides is restricted primar- Caves, Anderson 3,272 (HSC); Wichita Co., ilv to grassy roadsides and prairie relicts of the Burkburnett, Anderson 3.923 (HSC); Wichita coastal plain of southern Texas and northeastern Falls, McKee s.n., June 1927 (TEX); William- Mexico (Fig. 57, open circles). Occasional very son Co., N of Leander, Johnston and McCart wideh' separated populations may be found far- 5,279 (TEX). ther inland, as at Rock Springs, Texas (Fig. 64). Representative specimens examined: MEX- 40. CHLORIS ANDROPOGONOIDES Four- ICO: Coahuila, Musquiz Palm Canyon, Marsh 1 Ejido nier, Mex. PI. 2:143. 1886. (HOLOTYPE: 984 (TEX); Tamaulipas: mi E of de "Mexique: Prov. de San Luis. Virlet d'Aoust. San Lazaro, 24°.35'N, 99°13'W, Graham and Ranch, 24 mi 1851. 1462." P! The specimen is depau- Johmton 4,291 (TEX); El Canelo perate, probably from overgrazing, but the N of San Fernando, M. C. Johnston 4,880 spikelets are those of the species as com- (TEX). UNITED STATES: Texas: Aransas monly understood.) Fig. 58, D-K. Co., Rockport, A. Chase 6,067 (US); Atascosa Co., 3 mi S of Hindcs, M. C. Johnston 6,197 Chloris tcmiispica Nash, Bull. Torrev Bot. (TEX); Bee Co., 2.6 mi N of Skidmore, An- Club 25:436. 1898. (HOLOTYPE: derson 2,699 (HSC); Bexar Co., 7 mi N of San ". . . Nealley . . . 1889." US! Antonio, M. C. Johnston 2,409 (TEX); Cald- Perennial 10 to 40 em tall, tufted to shortb well Co., near Luling, Silveus 4,170-A (TEX); stolon iferous; sheaths glabrous; ligule a short, Cameron Co.: 10 mi E of Brownsville on Hwy. naked crown; blades ca 1 mm wide, to 1.5 cm 4, Anderson 3,318 (HSC); Laguna Atascosa Na- long, glabrous to scabrous except basally sparse- tional Wildlife Refuge, Fleetwood 3.374 (TEX); ly pilose; spikes six to thirteen, 4 to 15 cm long, levee of Resaca de la Cringa, M. C. Johnston usually radiate in a single series, occasionally 542.186 (TEX); Comal Co., E of New Braun- a second poorh' de\eIoped verticil produced fels, near Mission Valley Mills, Anderson 3,889 above; spikelets distant, ca four to seven per (HSC); Edwards Co., substation no. 14, Cory cm of the scabrous rachis, appressed; glumes 52.467 (US); Hays Co., 2 mi W of San Marcos, narrowly lanceolate, acute, thin, glabrous ex- Emery 835 (TEX); Hidalgo Co., Santa Ana cept for the scabrous midnei-ve; first glume 2.0 National Wildlife Refuge, Fleetwood 3,161, to 2.3 mm long, ca 0.2 mm wide; second glume 3,373 (TEX); Jim Wells Co., King Ranch, Santa 3.0 to 3.3 mm long, ca 0.3 mm wide; fertile Gertrudis Division, M. C. Johnston 542,104 96 Bbicham Young University Science Bulletin

Fig. 58. Chloris texemis and C. andropogonoides. (A-C) C. 'texcnsis. (A) habit, x 1/4; (B) spikelet, partly dissected; (C) sterile floret. (D-K) C. andropogonoides. (D) habit, k 1/3; (E) spikelet, partly dissected; florets, showing variation; (F-H) fertile lemmas, showing variation; (I) sterile floret; (J) group of sterile

(K) caryopsis. (B, C, E-K) .x 10. BiOLOCiCAi. Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 97

Table 11. Characteristics differentiating Chloris cucullata, C. verticillata, and C. andropogonoides.

Characteristic )

Brigham Young University Science Bulletin from populations containing only one of the three species. In the case of C. verticillata, this presented httle problem as the natural range of this species extends far to the north of the ranges of the other two (Fig. 57). Tlius, the samples of C. verticillata used in Fig. 59 are representative of populations from Oklahoma, Kansas, Nebraska, and Colorado. The examples of C. cucuUata and C. ondropogonoides chosen for Fig. 59 were selected from those popula- • KASON CO tions in Texas that had plants of only one ZiLKER PWK^PATTEBSm PARK # EDWARDS CO •* species present and in which the other species •EASTROP ST PARK • SAN P*RCOS were not present nearby.

From Fig. 59 it may be seen that the three species are well defined. Chloris cucullata • SKtCrDRE stands well apart from the other two taxa on the basis of all six characters used. Chloris verticillata and C. ondropogonoides are somewhat RIO GRANDE C|T> closer but are easily separated by the simultane- ous use of sterile floret width and lemma Fig. 60. Location of collecting areas sampled for in- awTi length (see Fig. 56, L-U, 58, D-K). In trogression studies. addition, C. andropogoiwides usually has longer and narrower lemmas than C. verticillata. The those plants in which virtually all of the pollen dashed lines drawn around each of the three mother cells examined showed normal meiotic species groupings in Fig. 59 represent the figures. FulK shaded s([uares () depict those approximate range of variation of lemma ami plants in which nearly all of the pollen mother length and sterile floret width on the specimens cells or their products showed meiotic irregu- studied; these will be reproduced as frames of larity, including lagging chromosomes or reference in the scatter diagrams to follow. chromosome fragments, micronuclei, multiva- Chromosome numbers of 2n = 40 have been lents, or excluded chromosomes. Half-shaded rejTorted several times for all three taxa. A scjuares ([J) exemplif}- those plants in which chromosome number of 2n^ca 2S has been less than 20 percent of the PMC's or their reported once by Gould ( 1960). products showed meiotic aberrations. At least The collection sites sampled for cytok)gicaI 25 PMC's were examined in detemiining the and mass collections of h\'brid populations are relative amount of meiotic regularity or irregu- shown on the map in Fig. 60 and listed in Table larity of each plant plotted in the diagrams. 12. For convenience of discussion, these areas The following discussion of populations rep- are given the names of nearby landmarks or resents the different combinations of the three population centers. species involved: the three possible pairs of

In the illustrations of individual populations species and the combination involving aJll three following, unshaded squares ( [U ) represent species.

Table 12. Populations utilized in introgression studies {all from Texas)

Name of population Species involved Fig.

Bastrop State Park anomalous 66 Edwards Co. ( Rock Springs C. ondropogonoides x C. verticillata 64 Kinney Co. anomalous 68 Mason Co. C. cucuUata x C. verticillata Nolan Co. C. cucullata x C. verticillata Patterson Park C. cucullata x C. verticillata 61 Pawelekxille C. cucullata x C. andropogonoides Pettus C. cucullata x C. andropogonoides Raymondville C. cucullata x C. andropogonoides 63 Richland Springs C. cucullata x C. verticillata Rio Grande City C. cucullata x C. andropogonoides San Marcos anomalous Skidmore anomalous 67 Zilker Park C. cucullata x C. andropogonoides x C. vcrticiUata 65 Biological Series, Vol. 19, No. Taxonomy of the Cenu.s Chloris (Gramineae) 99

Chloris cuctiUata x C. verticiUata This Inbiid t\pe is well represented in Pat- M4^^:^4^ terson Park, a city park in the northeast section of Austin, Texas (Fig. 60). In this area Chloris cucuUata and C. verticiUata and their introgressants fonn extensive colonies in distinbed areas, along roadwa)'s, paths, and in neglected sec-

I ' 1 ) t tions of the park. This population is illustrated i 1 f diagrammatically in the scatter diagram shown in Fig. 61 and in the photograph in Fig. 62. The scatter diagram of the Patterson Park population shows a number of plants corre- sponding in all major characters with the "pure" Fig. 62. Silhouette photograph of representative in- populations of C. cucuUata and C. verticiUata. florescences of Chloris populations from Patterson In addition, there are man\- intennediate forms Park, Travis Co., Te.xas. that variously recombine the characters of these two species. In those specimens studied cyto- mon in this area but are not involved in the logicallv, the plants lying within the range of parentage of any of the hybrids. expected variation of the pure species have nor- Again, the intermediate plants show a high mal meiotic behavior, while those studied from frequency of meiotic irregularity. Unfortunate- the intermediate group show, for the most part, ly, preserved anthers of the parental species high percentages of irregular meiotic figures. were too young for meiotic study. Several of

The Patterson Park population is just one of the intermediate plants studied, however, several showing this pattern of introgression. showed higher chromosome numbiTs than the Others with similar characteristics, when repre- 2n— 40 reported for the parental species. Dip- sented in a graph, include Mason Co., Nolan loid counts of 2n = ca 60 were most frequent, the high degree of irregularity made Co., and Richland Springs ( Fig. 60). though exact counts impossible. Plants with 2n = ca 60 Chloris cucuUata x C. andropogonoides could ha\e resulted from the combination of This population, represented graphical K in reduced and unreduced gametes.

Fig. 63, is found in a relatively stable grassland In this population the intennediates are community along a railroad right-of-wa\- near well represented in the lower left quadrant Raymond\ille, in southern Texas (Fig. 60). of the graph. By comparison, the Patterson

Many individuals of Chloris andropogonoides Park diagram ( Fig. 61 ) shows a poor represen- and C. cucuUata, as well as intermediate plants, tation of intennediates in this section. This shift

are found in this area. Two other species of is largely influenced by variation in awn length Chloris, C. gatjana and C ciliata, are also com- and is to be expected in view of the differences

IRREGULARITY MECTC iRREGULAR.T> ALMOST ALLCELLS EXAMINED I ALMOST ALL CELLS E (AMINEO HmEOSiS NORMAL BmEOTIC B OPLANTS NOT STUOED criOLOGICALLT OF CELLS EXAMItCO Q PLl NTS NOT STUDIED CYTOLOGfCAL LV SPWELETS PER CM OF SPIKE Q SPLKELETS PER CM OF SPIKE Q Q 16 9-13

LEN3TM OF SPIKES t i LENGTH OF SPIKES (I CMJ o o 6 DISTANCE BETWEEN LiPPERmOST O Q- .O— DISTANCE BETWEEN UPPeRMOST O VERTICIL O-lO U-2t 22-32 AND LOWERMOST VERTICIL O 144 AND LOWERMOST o : CM.) (IN CM.) (IN LEMMA length/width LEfvWA LENGTm/wiDTM <> ,8'^2 13^.7 l.8"^2 : coo! -i

5 1,00 glOO ---'o^

-f^^f^^+ ^O.^-^ T- 'A

20 30 40 50 60 20 30 40 50 60 AWN LENGTH (IN MM) LEMMA AWN LENGTH (IN MM) LEMMA

Fig. 61. Scatter diagram of population of Chloris citnil- Fig. 63. Scatter diagram of populations of Chloris lata X C. verticiUata from Patterson Park, Austin, cucuUata x C. andropogonoides from near Ray- Te.xas. mondville, Texas. 100 Bricham Young University Science Bulletin in awn length of the putative parents (see SP«ELf TS PEn CM OF SPIKE O O O M-16 9 '3 4-8 Table 12). 175 LENGTH OF SPKESdNCM) O 6 O Several additional collection sites had popu- 3-^ 6.8 9-11 DISTANCE BETWEEN UPPERMOST O O O- ANO LOWERMOSTVERTlCiL 040 11-21 ??02 lations with intiogression patterns similar to 2,50- (IN CM) LEMMA LENGTh/wCTH O "O —O ' 8-12 tJ-17 18-22 that of Raymondville, including Pavvelekville, 2 5 1254 Pettus, and Rio Grande City (Fig. 60).

glOO- Chloris andropogonoides x C. verticillata ul- The collection site for this population is in O 75 -I Edwards County, Texas, on the grounds of the Li. Texas A and M Experimental Station about 25 ^ 50H miles northwest of Rock Springs (Fig. 60). While "good" Cliloris andropogonoides and C. verticillata arc present in abundance, C. ciicul- lata was not observed in the area. The mass 2 30 40 50 60 LEMMA AWN LENGTH (IN MM) collection of plants made in this area is represented in Fig. 64. Fig. 64. Scatter diagram of population of Chloris Unlike previous scatter diagrams, the inter- andropogonoides x C. verticillata from the Texas mediate populations shown in Fig. 64 are not A and M Experiment Station near Rock Springs, of obvious parentage. On the basis of the Texas. species present, it would be expected that the In Fig. recombinants would fill the areas between C. comparing 65 with the diagrams of Patterson Park RaMiiondville Fig. verticillata and C. andropogonoides in the and ( 61 and graph, though certainly more long-awned inter- 63), one can plainly see a number of mutual mediates similar to C. andropogonoides are features. The Zilker Park population shows present than in either of the previous two popu- many plants in the lower left and left center lations discussed. The unexpected position of portions of the diagram, in this respect similar the remaining intennediate plants might be the to the Raymondville population, and indicates result of the presence at some time in the past a strong influence of C. andropogonoides. It is also strongly expressed in right- of C. cucuUata in the area, still showing its in- the upper fluence on the remaining population. A more hand portion of the recombinant cluster, a fea- tiu-e it shares with the Patterson Park popula- likely possibilit\', however, is that the intermediates merely represent recombinants of tion. By contrast, the Raymondville population is not well expressed in some of the more extreme morphological \'ar- the upper right-hand iants of C. andropogonoides and C. verticillata. portion of the diagram, nor is the Patterson This population was collected when the Park population well represented in the lower plants were in seed and cytological studies were left. Thus, the Zilker Park plants seem to repre- not possible. sent various recombinations of all three species. I have been unable to find additional populations that I feel are representative of this Dmeosis normal HMEiOTIC iRREGULARITT MEIOTIC IRREGULARITY IN ALMOST ALL CELLS EXAMINED in all cells IN ^cr^ or less „ examined OF CELLS EXAMINED PLANTS NOT STUDIED CYIQLOGKTAllt species combination. Chloris cucullata is nearly SPKELETS PER CM OF SPIKE O Q O ubiquitous in areas in which C. andropogonoides 9 13 o LENGTH OF SPKESdNCMl O 6 6 and C. verticillata arc sympatric and usually 3-5 6-8 9 11 DISTANCE BETWEEN UPPERMOST O O O- enters into the 2 -0 O ' ANO LOWERMOST VERTICIL O-IO 11-21 22-32 introgression pattern when 2 "r-,0; LEMMA length/width O O —O present. 2 125 e-12 13-17 18-22

Chloris cucidlata x C. andropogonoides x - -ft-a C. verticillata a

The population represented in Fig. 65 is „f^^ from a grassy field utilized intensively as a recreation area in Zilker Park, a citv park in the southwest part of Au.stin, Texas. Plants that can be referred without hesitation to C. cucullata are in area. common the Others are VO 2.0 30 40 50 60 clearly close to C. verticillata and C. andro- LEMMA AWN LENGTH (IN MM) pogonoides but are somewhat taller than ex- Fig. 65. Scatter diagram of population of Chloris pected. An abundant population of inter- cucuUata x C. andropogonoides x C. verticillata mediates is also present. from Zilker Park, Austin, Texas. Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 101

As with the otlier species combinations ex- SPiKELETS PER CIV OF SPIKE o Q O amined cvtologically, the intemiediate popula- 14-18 9-13 4-6 tions in Zilker Park show a high degree of LENGTH OF SPIKES (IN CM) O O O 5 1.50 3-5 6-a 9-11 meiotic irregularity. Only those plants assigned 2 DISTANCE BETWEEN UPPERMDST O O O- ANO LCWERMOST VERTICl z o-lO 1 »-2I 23-32 I op to C. cucullata show little or no meiotic aber- - 125 I ' O rancy. o <

The Skidmore population ( Fig. 67 ) was River bridge northeast of San Marcos, Texas, collected in a low roadside ditch in Bee Co., and about two miles east of the eastern edge Texas (Fig. 60). It is similar to the one at of the Edwards Plateau. This population shows well-defined Chloris cucullata and C. andro-

QMEOSrs E>AMl^ 102 Brigham Young University Science Bulletin

DmEOSIS normal flMEfOTC IRREGU.AR1TV mcOTIC iRREBULARITT IN ALMOST ALL CELLS EXAMINED meiosis, producing few stainable pollen grains. 111 Ai I .-CI I c lu on-L no I F^^ PLANTS NOT STUDIED CYTOLOGICAlly O It would be expected on this evidence that the Sf^KELETS PER CM OF SPIKE 9 9 rate of seedset would be relatively low. This, 9-13 -1-8 however, is not always the case. Plants from {IN 2 150 ,0 LENGTH OF SPIKFi CM 1 O 6 i ' 1 2 3-5 6 - B 9 io aSTANCe BETWEEN UPPERMOST O o o- the intermediate populations may actually have AND LOWEFJMOST VERTICIL 0-TO 1.1-21 2 2-3.2 'O (IN CM ) good seedset—in some cases, a high percentage. LEMMA length/width O Such plants are frequently associated with stands that are quite uniform morphologically. The combination of highly aberrant meiotic be- :t havior, good seedset, and extreme uniformity ^?m^ suggests that an apomictic mechanism may be in operation.

Nomenclature The wide array of variable intermediate 20 30 40 50 60 LEMMA AWN LENGTH (IN MM) plants in this complex would thus seem to be the product of extensive hybridization and in- Fig. 69. Scatter diagnim of anomalous Chloric population from near San Marcos, Texas. trogression, possibly still further complicated by apomixis. The intermediate plants will key pogoiwides, as well as many heterogeneous, to Chloris suhdolichostachtja and C. latisqua- cytologically aberrant intermediates. The posi- mea in many standard grass floras but frequently will not fit the descriptions of these taxa tion of the intermediate plants is similar to the precisely. It seems likelv that these names, in Zilker Park population (Fig. 65), suggesting reality, have applied to that all three putative parents were present at been segments of the much larger hybrid populations and do not one time. I searched the area many times over a period of several seasons and could not find represent biological species in the usual sense. well-defined C. verticillata, although plants The extreme variabilit\' of the intennediate similar to that species were found about a mile populations presents a difficult noinenclatural problem. The wide variety of morphological north of the site. Since this collection area is close to the limestone-rich Edwards Plateau, types represented certainly cannot be individu- this saturate litera- the seemingly preferred habitat of C. verticil- ally named, for would the biological lata, one would expect the latter species nearby. ture with names that carry little meaning and might apply to plants of low Discussion. genetic stability. The problems become addi-

Ecologically, populations with large num- tionallv compounded in that there is no way bers of intennediate plants are most frequently of telling parts of the hybrid population of C. associated with highly disturbed areas. The cucuUata x C. andropogonoides from parts of best collecting sites, therefore, are roadside the C. cucuUata x C. verticillata populations ditches, waste places, and city parks. None (compare Fig. 61 and 63) and from the popu- of the species involved will persist in cultivated lations involving all three species (Fig. 65). or grazed areas, nor will they compete well To use the existing names C. latisquamea and with an established native flora or such ag- C. suhdolichostachtja for the intermediates is

gressive weeds as Johnson grass. Before the unsatisfactory, for there is no way of deter-

excessive disturbance caused by man, it is likely mining which, or how many, of the three species that the three species involved were only oc- are involved in the ancestry of their types. In casionally found, and then only in such areas short, these names cannot be precisely defined. as blowouts, slipouts, and banks and alluvial Likewise, the usage of a single name for a com-

areas along creeks and streams. There is little plex of morphological forms obscures their bio- information about the ecology of these species logical nature. before the rampant habitat destruction by man; I feel that the only solution is to designate however, a report of a grass-collecting trip the intermediate plants by a formula based upon through the Kansas prairies B. parentage. Thus, a population by J. Norton their probable in 1898 (Barkley, 1965) indicates that Chloris suspected to be the result of C. cucuUata intro- verticillata grew along the sides of sloughs and gressing with C. andropogonoides should have

on low ground but did not compete well with its intermediate plants designated by the for- upland grasses. mula name C. cucuUata x C. andropogonoides. Cytological examination of the intennediate While this will not satisfy those who want a plants shows them to be highly aberrant in particular name for a particular plant, it does . . .

Biological Sehies, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 103

satisfy what is, to me, a more important aspect C. cucullata x C. andropogonoides x C. ver-

of taxonomy that is seldom reflected in existing ticillata. TRAVIS CO.: Austin, Zilker Park

nomenclature: it helps to define the reasons for playing field, Anderson 3,011-3.090, 6 June 1964 the existence of a taxonomic problem. (mass collection): 2,603-2,628, 28 June 1963; The original descriptions of the following 2,650-2,664, 3 July 1963; .3,262-3,268, 25 June names seem to fit the characteristics of hybrid 1964 (all cytovouchers). plants. Anomalous populations. BASTROP CO.: Chloris vei-ticiUata var. aristidata Torrey and just outside W entrance of Bastrop State Park, Gray in Whipple, U.S. Kept. Expl. Pacific Anderson 3.128-3.138, 10 June 1964 (mass col- 2:176. 1857. lections), 3,116-3,127, 10 June 1964 (cytovouch- Chloris siihdolichostachtja C. Mueller, Bot. ers). KINNEY CO.: along Hwy. 693, 1-3 mi Zeitung (Berlin) 19:341. 1S61. E of its junction with Hwy. 277, Anderson Chloris verticillata var. intermedia Vascy in 4,479, 26 June 1968 (mass collection), 4,470- Coulter, Contr. U.S. Nat. Herb. 2:528. 1894. 4,478, 26 June 1968 (cytovouchers). HAYS Chloris hrevispica Nash, Bull. Torrev Bot. CO.: near bridge over Blanco River, 1 mi NE Club 25:4.38. 1898. of San Marcos on Hwy. 21-80, Anderson 3,276, Chloris latisquaniea Nash, Bull. Torrev Bot. 28 June 1964 (mass collection), 3,278-3.282. 28 Club 25:439. 1898. June 1964, 4,617-4,619, 30 June 1968 (all cytovouchers). CO.: 2.6 mi N of Skidmore, Specimens and populations examined (all at BEE Anderson 2.698-2.699. 13 July 1963 (mass col- HSC): C. cucullata x C. verticillata. TRAVIS 2.691-2.697, 13 July 1963 (cyto- CO.: Austin, Patterson Park, along Aiiport lections), x'ouchers ) Blvd., 0.5 mi SE of its junction with Hwy. 35, Anderson 3,336, 12 July 1964 (mass collection), 4L CHLORIS TEXENSIS Nash, Bull. Torrey 3,338-3.361, 12 Julv 1964 ( cytovouchers ) 23:151. 1896. (HOLOTYPE: MASON CO.: along Hw)'. 29, 2.5 mi E of the Bot. Club Nealley." US! ) Fig. 58, A-C. junction of Hwys. 29 and 377, 3 mi E of Grit, "Texas. G. C. Bull. Torrey Bot. Anderson 4,633, 2 Julv 1968 (mass collections). Chloris nealleyi Nash, 1898. (Based on Chloris NOLAN CO.: along' Hwy. 53, 9 mi S of the Club. 25:435. junction of Hwys. 126 and 53, Anderson 4,653. texensis Nash.) 2 July 1968 (mass collecHons). SAN SABA Perennial 30 to 45 cm tall, cespitose; sheaths CO.: Richland Springs, along Hw>'. 190, road- glabrous to sparsely pilose; ligule a low mem- side ditch near cit)' water tower, Anderson branous crown; blades narrow, ca 4 mm wide, 3,333, 9 Jul\- 1964 (mass collections). up to 15 cm long, scabrous; spikes eight to ten, C. cucullata x C. andropogonoides. WIL- up to 20 cm long, radiate, occasionally in two LACY CO.: grassy roadside, 1 mi N of Ray- verticils, bearing loosely imbricate spikelets only mondville, along Hwy. 77, Anderson 3,308. on distal half, naked at base; spikelets ca three 3,316, 1 July 1964 (mass collections), 3,.m9- or four per cm of the scabrous rachis; glumes 3,315, 1 July 1964, 4,564-4,572, 28 June 1968 lanceolate, glabrous except for the scabrous mid- (cytovouchers). KARNES CO.: 6 mi S of the nei-ve; first glume 2.7 to 3.0 mm long, 0.4 junction of Hwys. 123 and 889 and the village mm wide; second glume 3.5 to 3.8 mm long, of Pawelekville, Anderson 4,611, 30 June 1968 0.5 mm wide; fertile floret 3.7 to 4.3 mm long, (mass collection), 4,595-4,610, 30 June 1968 0.7 to 0.8 mm wide, lanceolate to narrowly (cytovouchers). BEE CO.: 3.3 mi N of Pettus, ovate, callus bearded, upper margins sparsely 1.6 mi N of the junction of Hwys. 181 and 798, appressed-ciliate, otherwise glabrous, apex Anderson 2,781-2,789, 13 Julv 1964 (mass col- acute, awn 7 to 11 mm long; sterile floret lections). STARR CO.: 8 mi W of Rio Grande 2.0 to 2.5 mm long, 0.5 mm wide, narrowly City, along Hwy. 83, Anderson 3,330. 2 Julv elliptic, apex acute, glabrous, awn 4.5 to 6.5 1964 (mass collection), 3,323-3,329 (cytovouch- mm long; caryopsis ca 2.3 mm long, 0.5 mm ers); 1 mi S of Rio Grande City, Anderson 4.524. wide, ellipsoid, trigonous. 27 June 1968 (mass collection); ca 1 mi SE of Arrovo Los Olomos bridge, ca 3 mi SE of Chloris texensis is most closely allied to C. Rio Grande City, Anderson 4,542, 27 June 1968 verticillata and C. andropogonoides (Fig 56, (mass collection), 4,525-4,541 (cytovouchers). L-U; 58, D-K). It differs from C. verticillata in C. andropogonoides x C. verticillata. ED- usually having a single verticil of spikes and WARDS CO.: Texas A and M Field Station. relatively narrow sterile florets. It may be sepa- 25 mi NW of Rock Springs, Anderson 3,903, 12 rated from C. andropogonoides by the rela- longer spikes. Chloris July 1965 ( mass collection ) tively larger spikelets and 104 Bricham Voung University Science Bulletin

texensis lacks spikelets on the lower portion of mm long, 0.3 to 0.5 mm wide; second glume the spikes, whereas both C. verticillata and C. 2.7 to 2.9 mm long, 0.4 to 0.5 mm wide; fertile andropogonoides are floriferous to the base. lemma 2.1 to 2.9 mm long, 0.9 to 1.1 mm Chloris divaricata, a similar Austrahan spe- wide, ovate to elliptic, apex acute, callus short- cies, is separable in having deeply lobed sterile bearded, keel and margins densely appressed- florets, scabrous fertile lemmas, and spikes that pilose, awn 0.7 to 1.0 mm long; sterile floret are floriferous to the base (Fig. 51, K-R). one, ca 1 mm long, 0.3 mm wide, strongly Chloris texensis has sterile florets that are mere- flattened, membranous, elliptic, awn 0.3 to 0.5 ly acute, not deeply lobed, glabrous fertile mm long; anthers ca 0.7 mm long. lemmas, and spikes that are naked at the base Only two specimens of Chloris sesquiflora, (Fig. divaricata is 58, A-C). Chloris apparently including the type, were seen. While it re- an escape in Texas and has been confused with sembles outwardly some of the species with C. texensis. ciliate lemmas (especially C. ciliata), the flat- Only six specimens of the species, including tened, poorly developed single sterile floret and the type, have been found in herbaria con- the appressed pubescence of lemma margins sulted. All are Brazoria, from Brazos, or Harris and keels separate it easily. It is probably not County, Texas, except the type, for which a closely related to C. ciliata. precise collection locality was not provided. Specimens examined: PARACUAY: Plaine On two separate occasions I searched the de Paraguari, plaques desseckles, dans les ar- area of two collections, Silveus 423 and 427, giles impermeables, Balansa 264 (BR). from the A and M Field Station near Angleton.

Most of the land in the area is either culti- 43. CHLORIS HUMBERTIANA A. Camus, vated, heavily grazed, or in stages of old-field Bull. Soc. Bot. France 91:63. 1944. succession; and I could not locate either C.

(HOLOTYPE: "Madagascar . . . Mananara texensis or any native vegetation that might

. . . pentes orientales du Vohipaly . . . Hum- support it. Because neither the plant nor native bert, no. 141.50." P! ISOTYPE: K!) Fig. vegetation could be found, and because few 70, D-F. specimens have been collected since 1931, I

feel that it is quite likely that the species is Pterochloris humhertiana (A. Camus) A. in danger of becoming extinct. Camus, Bull. Mus. Hist. Nat. (Paris) Ser. Specimens examined: UNITED STATES: 2. 29:.349. (Based on Chloris humhertiana Texas: Brazoria Co., Columbia, Bush 1,361 (MO, A. Camus.) NY); experimental farm near Angleton, Sil- veus 423 (TEX); open sandy woods and Perennial 20 to 30 cm tall, stoloniferous, with swamps, Angleton, Silveus 427 (TEX); Brazos erect culms arising at intei"vals from strong stolons; leaves basal; sheaths flattened, Co., without precise location, Malone s.n., Oc- equitant, glabrous; tober 1938 (TAES); HaiTis Co., exposed clay ligule a very short-ciliate crown; lowland, 3 mi S of Alvin, Parks 117 (TAES). blades 1.5 to 2.0 em long, 3 to 5 mm wide, glabrous, folded, slightly arcuate, apex obtuse; spikes two or three, 3 to 4 cm long, partially 42. CHLORIS SESQUIFLORA Burkart, Bol. enclosed in upper sheaths Soc. Argentina Bot. 12:287. 1968. (TYPE: when young, becoming exserted at maturity; spikelets "Dep. Concep. del Unig., bajos cerea del imbricate, ca five to seven per rio Cualeguaychu, vecindades del Parque cm of the glabrous laehis; Unzue, A. .' glumes laneeolate-ovate, thin and mem- Burkart. 25,676 . . . SI, not branous, glabrous except for the scabrous mid- seen. ISOTYPE: HSC!) Fig. 70, AC; 71, triangles. nei-ve; first glume 1.0 to 1.5 mm long, ca 0.3 mm wide; second glume ca 1.5 mm long, Stoloniferous perennial up to 45 em tall; 0.2 to 0.3 mm wide; fertile lemma 2.3 to 2.5 sheaths glabrous; ligule a short-eiliate crown; mm long, 0.7 to 1.2 mm wide, body of lemma blades to up 20 cm long, 4 to 5 mm wide, elliptic, with a promini'iit wing at the apex, glabrous to scabrous; spikes four or five, 6.0 making the lemma appear narrowly tnnicate, to 6.5 long, em ascending, somewhat arcuate; callus bearded, margins scabious below, gla- spikelets rather densely inserted, appressed, brous above, internerves densely pubescent vvith imbricate, averaging about 11 per cm of the prominent, inflated hairs, awn 2.4 to 3.5 mm scabrous racliis; glumes lanceolate to narrowly long; sterile floret ca 1 mm long, ca 0.5 mm ovate, acute to acuminate, glabrous except for wide, elliptic, flattened, apex acute, awn 3.0 the scabrous midnerve; first glume 2.1 to 2.2 to 3.8 mm long, arising below the apex. Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Cm.onis (Gramineae) 105

Fig. 70. Chloric se.squiflora and C. humbertiana. (A-C) C. sesquiflora. (A) habit, .x 1/3; (B) spikelet, partly dissected, x 10; (C) sterile floret, x 10. (D-F) C. humbertiana. (D) habit, x 1/3; (E) spikelet, partly dis-

sected, X 10; ( F ) sterile floret, x 10. )

106 Bhigham Young University Science Bulletin

Chloris humbertiana is a very distinct spe- Stoloniferous perennial up to 80 cm tall; cies; its strongly winged lemmas, with inflated sheaths glabrous below, becoming loosely pilose hairs on the intemerves, are unique in the toward the ligule, especially on the margins; genus. ligulc a short-ciliate crown, often with longer

Emphasizing these characters, Camus ( 1950 hairs near the edges; blades up to 10 cm long, proposed the subgenus Pterochloris to include ca 5 to 6 mm wide, glabrous with scabrous mar- Chloris humhertitina. Later (1957), Camus ele- gins and midnerves; spikes four to six, 5 to 9 cm vated Pterochloris to generic rank, retaining C. long, ascending when young, becoming more humbertiana as its single species. loosely spreading at maturity, usually radiate,

While Chloris humbertiana is unusual in sometimes in two verticils; spikelets appressed, Chloris, it is no more so than a number of other ca 7 per cm of the scabrous rachis; glumes un- species and I am maintaining it in Chloris. equal, glabrous except for the scabrous mid- 1.5 to 2.5 long, 0.2 to 0.5 Only four specimens of this species were nerve; first glume mm wide, lanceolate, apex acute; second glume seen in the herbaria consulted and all are from mm xerophytic bushland in the Malagasy Republic 2.5 to 3.8 mm long, ca 0.4 mm wide, linear, (Madagascar). apex acute to mucronate, sometimes somewhat lacerate, glabrous except for the scabious mid- Specimens examined: MALAGASY REPUB- nerve; fertile lemma 3.5 to 4.5 mm long, 0.7 LIC: Pateau Mahafetz, Bosser 15,698 (P); Cap to 1.2 mm wide, elliptic to obovate, often pur- Sainte-Marie, Humbert and Capuron 29,264 plish tinted, callus densely bearded, margins (P); d'Amparihy a rembouchuie de la Mcna- densely appressed-pilose, keel sparsely to dense- randra, Humbert and Capuron 26,347 (P); ly appressed-pilose, apex acute to obtuse, usu- environs d'Ampandrandava, Setjrig 809 (P). ally awnless, occasionally with a short awn less than 1.7 mm long; sterile floret one, 1.4 to 2.5 44. CHLORIS AMETHYSTEA Hochstetter, mm long, 0.6 to 1.2 mm wide, narrowly turbi- Flora 38:205. 1855. (ISOTYPE: "... Abys- nate, apex truncate, somewhat inflated, awn- ." sinia leg. Schiinper recu 1855. 1267 . . less; caryopsis ca 2.2 mm long, 0.6 mm wide, G!) Fig. 72, A-K. trigonous, ellipsoidal. Chloris amethijstea seems most closely related to C. i^aijana and C. mossambiccnsis, differing from both in having short awns. While relatively few specimens are available, C. amethijstea appears to be restricted to Ethiopia and Kenya (Fig. 73, triangles), occurring in montane scrub and grasslands at the forest edge on white or yellowish sandy loams and at elevations of 3,000-6,000 feet. Associated plants include Ficus, Acacia, Comhretum, and Com- miphora. Specimens examined: ETHIOPIA: Territory Agow, Dschascha, Hohenacker 2,247 (K, US); • = HALOPHILA A = SESQUIFLORA between Assala and Awash Rivers, Semple 19 (US). KENYA: Mt. Marsabit, Adam.wn E. A. H. 11,666 (K, US), Bo<^dan A. B. 4,774 (K); Stony Athi River, 20 mi SE of Nairobi, Bogclan AB .3,1.52 (K); Moyale, 3°32'N, .39°03'E, Gillett 12,852,14,065 (K).

45. CHLORIS SUBMUTICA Humboldt, Bon- pland, Kunth, Nov. Gen. Sp. 1:167. pi. 50.

1816. (There is a fragment, in US!, from

P, that may be from the tv'pe. It is labeled "Chloris submutica HBK mihi n. "4182. Mexique." The original description gives "Tezcucensi" as the collection site, but no

Fig. 71. Distribution of Chloric halophila (dots) and other p:.'rtinent information is given. The C. sesquiflora (triangles). detailed original description and accom- Biological Series, Vol. 19, No. 2 Taxonomy „i. thf. Genus Chloius (Chamineae)

^' '"'^ "^^ (A-K) C. arncthystea. (A) habit, .x (B) dl^sJ'd" 75 IPvT, .'"'""f^"- 1/6; spikelet, partly i" ^P'^'''}/h?r"S ^^,"«""" ^'^vn length, .x 10; (F-J) sterile florets, showing S^n V 7.5;7'f' Kf^caryopsis, ™°"' -^ .x 5. (L-V) W C. sub,„uUca. (L) habit, ( x 1/4; ) spikelet, partly X M f l- dissectecf /..b, (N-1) sterile florets, showing > / variation, x 7.5; (U) caryopsis, x 5. ) ) )

108 Brigham Young University Science Bulletin

glabrous with scabrous midnerves; first glume 1.5 to 3.2 mm long, 0.2 to 0.9 mm wide; second glume 2.5 to 3.4 mm long, 0.3 to 0.7 mm wide; fertile lemma 2.8 to 3.7 mm long, 0.6 to 1.1 mm wide, pale, broadly linear to elliptic, callus bearded, rest of lemma glabrous except for the appressed-pilosc margin, apex obtuse, unawned or with a very short mucro; sterile floret one, 1.4 to 2.2 mm long, 0.3 to 0.9 mm wide, truncate, awnlcss or occasionally with a very short mucro; anthers ca 0.5 to 0.9 mm long; caryopsis 1.7 to 2.3 mm long, 0.5 to 0.6 mm wide and thick, ellipsoid, trigonous; chromosome number 2n=ca 65, 80. Fig. 73. Distribution of Chloris amethystea (triangles) Chloris suhmutica outwardly resembles and C. somalensis (circles). species in Eustachys, primarily because of the appressed, awnless spikelets. Tlie second glume, panying illustration leave no doubt as to however, lacks the rounded apical lobes sub- the taxon described.) Fig. 72, L-V. tending the short awn, a characteristic of Eu- Chloris pohjstacluja Lagasca, Elenchus stachys. Also, C. suhmutica lacks the conspicuous Plantarum 59. 1816. (A fragment of a equitant leaf sheaths of Eustachi/s. specimen, in US! from "Hb. Madrid" Chloris suhmutica Humboldt, Bonpland, labeled "Chloris polystachya Lag. Nee' iter Kunth and C. polystachi/a Lagasca were both Hb Madrid" may be from the type. The published in 1S16. According to Stafleu (1967), original description is inconclusive. the particular section of the Nova Genera et Eustachijs suhmutica ( Humboldt, Bonpland, Species Plantarum, containing the original de- Kunth) Roemer and Schultes, S)St. 2:614. scription of C. suhmutica, was published in 1817. (Based on Chloris suhmutica Hum- Mav 1816; while C. poh/staclu/a Lagasca, pub- boldt, Bonpland, Kunth.) lished in Elenchus Plantarum, was probably Chloris inermis Trinius, Gram. Uniflorus published in June or Jul\' 1816. In addition to 232. 1824. (Based on Chloris pohjstachija the slightly earlier date of publication, C. .suh-

Lagasca. mutica is clearly identifiable and has been Eustachijs polystachya (Lagasca) Kunth, widely accepted, far more so than C. poly- Rev. Gram. 1:88. 1829. (Ba'scd on C/i/orf.? stachya. For these reasons C. suhmutica is polystachya Lagasca. maintained here. Phacellaria panicea VVilldenow ex Steudel, Cliloris suhmutica has been extensively col- Noin. Bot. ed. 2. 1:353. 1840, pro syn. lected in Mexico, occasionally in Columbia, Guatemala, New Mexico, and Venezuela ( Fig. Chloris suhhijlora Steudel, S\n. PI. Glum.

43, circles ) . Most specimens are from eleva- 1:206. 1854. (HOLOTYPE: P! A fragment tions of 3,000 to 6,000 feet and on a variety of an additional Steudel specimen, labeled of soil types. by him as C. suhhijlora, is in US!, originally from CN.) Representative specimens examined: CO- LOMBIA: Dep. Cundinamarca: Sabana de Chloris pratensis Lesible ex Steudel, Syn. Bogota, cerca de Madrid, Tarragon et al. IS Cu PI. Glum. 207. 1854. Pro syn., C. suhhi- 104 (US). GUATEMALA: Quezeltenango, jlora. Chiquilaga, de Koninck 11 (US). MEXICO:

Perennial 30 to 75 cm tall, erect, usually Aguascalientes : Aguascalientes, Hitchcock 7,453 tufted, occasionally short-stoloniferous; sheaths (US); Chiapas: N of San Cristobal de Las glabrous below, becoming ciliate at the ligule Casas, Breedlove 6,759 (F); Chihuahua: Ma- with hairs up to 2 mm long; blades up to 20 jalca, Pilares, 2,080 m, Harvey 1,483 (US); San cm long, ca 5 mm wide, the surfaces and mar- Juanito, Knohloch 5,425 (F); Coahuila: Saltillo, gins scabrous, occasionally long-pilose at base 5,300 ft, Fisher 57 (US), Hitchcock 1,261 (NY, of blade; spikes 5 to 17, up to 7 cm long, usu- UC) 5,58.5 (US); Durango: Tepehuanes,. F(.s/ier ally erect when young, becoming somewhat re- 44,221 (MO), Palmer ^262, 4-25 June 1906 flexed at maturity; spikelets rather densely im- (MO, NY, UC, US); Otinapa, Pahner 341, 25 bricate, averaging about twelve per cm of the June-5 Aug 1906 (MO, NY, UC, US); Dls- scabrous rachis; glumes lanceolate, acute, pale, trito Federal: Lomas de Mixcoac, Lyonnet 2,701 Biological Sehies, Vol. 19, No. Taxonomy of the Genus Chloris (Gramineae) 109

( US ) ; pyramid of Cuicuiico Tlalpan, McDcin- Included here would be Glover and Gilliland ieh 47 (F); Guanajuato: 6 km S of San Frlipe, 755 (BM!).

Sohns 405 (US). Hidalgo: Pachua, 8,000 ft, Onl\- six specimens were available in all Hitchcock 6,748 (US). Jali.sco: Zapotlan, herbaria consulted. Tlicse were collected in

Hitchcock 7,129 (US); Michoacan: 5 mi N of Ethiopia and the Somali Republic ( Fig. 73,

Patzcuaro, Webster 2,700 (US); Navarit: Tepic, circles ) in open, grassy savannahs with Jiini- Santa Teresa, Rose 2,143 (US); Nuevo Leon: periis and Dodonaea. Soils were partly lava with Galeana, Hacienda Pablillo, Tmjhr 127 (F, black clay and partly sandstone with red, sandy MO); Puebla: vicinit\' of Puebla, Arscne 330 soil in areas with moisture derived largely from (MO, NY, US); Queretaro: Cd. Queretaro, low clouds. Hitchcock 5,818 (US); Sonora: Rio de Bavispe, Specimens examined: ETHIOPIA: Hamasen, El Picaco de Pilar, Santos 2,168 (US); San Luis Nefarit, Baldrati s.n., 7 Mar 1916 (K); Sidamo, Potosi: Cd. San Luis Potosi, Schaffner 130 (NY, Mega, 4°5'N, 38°20'E, ca 2,200 m, GiUett 14,296 US); Tamaulipas: 6 km W of Miquihuana. 2 940 (K, US), Moonetj 5,565 (K). SOMALI RE- m, Stanford et al. 737 (MO, US); Tlaxcala: be- PUBLIC: Golis Range, Baldostie Codie, Drake- tween San Cristobal and Calpulalpan, Sohns 580 Brockman 480 (K); Iva Makarai, 9°56'N, 45°E, (US); Zaeatecas: 15 km W of Concepcion del Drake-Brockman 481 (K). Oro, near Zacatecas-Coahuila border, Stanford et al. .5.58 (MO, NY, UC, US). UNITED STATES: 47. CHLORIS CLEMENTIS Merrill, Philippine New Mexico: Dona Ana Co., State College, Jour. Sci. 40:181. 1929. (HOLOTYPE:

Norris 1,026 (US), Parker 459 (US). VENE- "Luzon: Pangasinan Province, Calasiao . . . ZUELA: Estado de Merida: between Mucaruba Clemens 17,267." UC! ISOTYPES: BR! F! and Mueliachies, Rudd 541 (US). NY! US!) Fig. 74, A-E.

Perennial 20 to .30 cm tall, with delicate, 46. CHLORIS SOMALENSIS Rendle, Jour. erect culms arising from the prominent stolons; Bot. (London) 37:66. 1899. (IIOLO- sheaths very short, glabrous below, becoming TYPE: "Wagga Mountain, Somaliland. somewhat ciliate along upper margins; ligule a Mrs. E. Lort' Phillips. 1897." BM!) Fig. short membranous ring; blades 1.0 to 2.5 cm 74, F-L. long, 2 to 4 mm wide, obtuse, slightly apiculate, Perennial to 70 cm tall, cespitose, erect; glabrous; spikes two to five, 2 to 3 cm long, sheaths glabrous, overlapping; ligule a short- radiate; spikelets appressed, imbricate, ca five pilose crown; blades 15 to 30 cm long, to 7 per cm of the scabrous rachis; glumes lanceolate mm wide, glabrous; spikes three or four, ca to ovate, membranous, glabrous except for the 8.5 cm long, erect when young, later spreading; scabrous midnerve; first glume 0.5 to 0.6 mm spikelets appressed, dense, ca ten per cm of long, ca 0.2 mm wide; second glume 0.7 to 0.8 the scabrous rachis; glumes unequal, linear- mm long, ca 0.2 mm wide; fertile lemma 1.5 to lanceolate, thin and glabrous to slightly scab- 1.8 mm long. 0.4 to 0..5 mm wide, lanceolate rous, especially on the midnerse; first glume to elliptic, callus bearded, margins sparsely 1.4 to 3.0 mm long, 0.2 to 0.4 mm wide; second ciliate apically, scabrous below, keel sparsely glume 3.2 to 5.0 mm long, ca 1.0 mm wide; scabrous, otherwise glabrous, awn 5.5 to 6.5 mm iFertile lemmas 3.5 to 5.2 mm long, 0.8 to 1.0 long; sterile floret one, ca 0.3 mm long, 0.1 mm wide, lanceolate, olive-green, callus beard- mm wide, much reduced, flattened, often con- ed, margins glabrous below, becoming spar- cealed within the lemma, awn ca 1 mm long; ingly appressed-pilose near the apex, otherwise car\opsis ca 1.2 mm long, 0.3 mm wide, nar- glabrous, awn 7 to 12 mm long; sterile floret rowly ellipsoid. one, 0.5 to 2.0 mm long, 0.1 to 0.5 mm wide, Chloris dementis resembles several other usually greatl\- flattened and reduced, some- species in gross appearance, but differs mark- times slighth' inflated, scabrous, awn 1.5 to 3.5 edly in detail. Its strongly stoloniferous habit mm long: anthers 1.0 to 1.5 mm long; car\'opsis is similar to that of C. divaricata as well as 2.0 to 2.2 mm long, 0.5 to 0.6 mm wide.' ellip- to EustachijS tenera and to various species of soid, trigonous. Cijnodon. It differs from Chloris divaricata in

In general the olive-colored spikelets and having much smaller sterile florets ( Figs. 73, greatly reduced sterile florets readily separate A-E; 51, K-R). Sterile florets in Cynodon are Chloris somalensis from related species. Occa- either greatly reduced or absent. Etisfachijs is sional specimens have exceptionally large rudi- characterized b\' having bilobed second glumes. ments and mav eventually prove distinct as Other species (such as Chloris ekmanii, C. better series of specimens become available. pijcnothrix, C. radiata, and C. sagraeana) may )

no Bbigham Young University Science Bulletin

Fig. 74. Chlorii dementis and C. somalensis. (A-E) C. dementis. (A) habit, x 1/3; (B) spilcelet, partly dissected, X 10; (CD) sterile florets, showing variation, x 30; (E) caryopsis, x 10. (F-L) C. somalensis. (F) habit, X 1/3; (G) spikelet, partly dissected, x 10; (H-K) sterile florets, showing variation, x 10; (L) caryopsis, x 7.5. have similar spikelets but are tufted rather than Agrostis radiata Linnaeus, Syst. Nat. ed. 10. stoloniferous. 2:873. 1759. (See nomenchitural note No specimens were seen other than the below. holotype and its isotypes. Ct/nosurus scopariiis Lamarck, Encycl. Meth. 2:188. 1786. (HOLOTYPE: Not 48. CHLORIS RADIATA (Linnaeus) Swartz, positively identified, but there is a frag- Prodr. Veg. Indiam Occ. 26. 1788. (Based ment from P, in US!, labeled "Cynosurus on AgTOstis radiata Linnaeus.) Fig. 75, F-J. scoparius de St. Dominique Lam.") ) ) ) )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae) 111

Chloris scoparius (Lamarck) Desfontaines, described above or possibly another Caribbean Tabl. ficole Bot. Mus. Hist. Nat. 14. 1804. species, for example, C sagraeana. In the Lin-

( Based on Ct/nosurus scoparius Lamarck. naean herbarium there is a specimen of Pat-

Cliloris i^Iaucescens Steudcl, Ssn. PI. Glum. rick Browne's which Linnaeus might have seen; 1:206. 1854. (HOLOTYPE: "Guadeloupe. it is fragmented but seems to be more similar Duchaissing" P! Fragment in US! to C. sagraeana. No other information can be obtained either to substantiate or refute the Chloris tacncnsis Steudel in Lecliler Ber- name. While most of the evidence in this prob- berides Ainer. Australis 56. 1857. Nomen lem is either nonsubstantive, or even somewhat niulein, but a specimen "St. 1.569" is cited. negative, I do not wish to reject circumstantially This specimen, in P!, labeled C. tacnensis a name that has been used commonly for this in Steudel's script is C. radiata (Linnaeus) species for nearly two centuries. Until additional Swartz. information can be found, I propose continued Chloris biflora Kegel, Annotationes Bot. acceptance of Chloris radiata (Linnaeus) Index Seminum 28. 1863. Pro syn.. C. Swartz for the plant described above. radiata. Chloris radiata most closely resembles C. Chloris scoparius Kegel, Annotationes Bot. pycnothrix. Differences have been discussed Index Seminum 28. 1863. Pro syn.. C. under the latter species. Other Caribbean spe- radiata. cies are similar in spikelet and inflorescence Chloris pallescens Kegel, Annotationes Bot. features but have larger sterile florets (see Index Seminum 28. 1863. Pro syn., C. Table 13 for a further comparison. radiata. Chloris radiata is widely distributed through- Chloris crinita Salzman ex Doell in Martins out the eastern Caribbean, northern South Flora Brasiliensis 23; 64. 1878. Pro .syn., Africa, and Central America, where it is a C. radiata. Non Lagasca, 1805. common weed in waste areas, along paths, Gymnopogon radiata (Linnaeus) Parodi, and roadways, etc. (Fig. 76). Ph\sis 4:180. 1918. (Based on Agrostis Kepresentative specimens examined: BA- radiata Linnaeus. HAMA ISLANDS: Nassau, Hitchcock s.n., Nov Annual usually with dense fibrous roots, oc- 1890 (F, MO). BOLIVIA: Coripati, Yungas,

casionally rooting at lower nodes, 30 to 60 cm Bang 2.173 ( F, MO, NY, US). CAKOLINE tall; sheaths glabrous, occasionally pilose; ligule ISLANDS: Palau group, Koror Island, Fosberg a short-ciliate crown, sometimes with a few 32,327 (NY, US). COLOMBIA: Div. Antio- longer hairs intennixed; blades 10 to 30 cm quia: Fradonia, Archer 503 (US); Div. Cauca: long, up to 10 mm wide, usually pilose, occa- El Valle, Triana 345 (US); Div. Nariiio, be- sionally glabrous or scabrous, apex usualh' tween Dos Kios and La Union, Archer 3,394 acute; spikes five to fifteen, 5 to 8 cm long in (US); Div. Santander: Boca de Rosario, Kio one, two, or sometimes three ascending verti- Magdalena, Pennell 3,904 (MO, NY, US). cils; spikelets densely imbricate, ca 11 to 15 COSTA KICA: Port: Limon, Hitchcock 8,417, per cm of the scabrous rachis; glumes \ery 8,42S (US); Prov. San Jose: between San narrowly lanceolate, glabrous except for the Pedros Montes de Oca and Curridabat, Stand- scabrous midner\'e; first glume 0.7 to 1.6 mm ley 41.283 (US). CUBA: Prov. Pinar del Rio: long, 0.1 to 0.2 mm wide; second glume 2.0 Mariel, Ekman 12.939 (F, NY, US); Prov. to 2.7 mm long. 0.1 to 0.2 mm wide; fertile Habana: Guatao, Leon s.n., Oct. 1923 (US). lemma 2.8 to 3.3 mm long, 0.4 to 0.6 mm wide, DOMINICAN REPUBLIC: Prov. Pacificador: lanceolate to elliptic, glabrous, except upper Pimentel, Abbott 691 (US); Prov. Seibo: vi- margins short-ciliate, callus bearded, awn 6 to cinit\' of Hato Mayor, Allard 13,643 (US); Prov. 13 mm long; sterile floret one, 0.4 to 0.7 mm Santiago: vicinit)' of Santiago, Allard 14,553 );'^ long, ca 0.1 mm wide, greatK' reduced, borne on ( US Prov. Duarte: Valle de Cibao, Ekman a long rachilla joint, awn 3 to 5 mm long; car\- 13,259 (US); Prov. Puerto Plata: Puerto Plata, opsis 1.4 to 1.5 mm long, 0.3 to 0.4 mm wide, Ekman 14.448 (US). ECUADOR: Prov. Chim- trigonous; chromosome number 2n^40. borazo: Alausi, Hitchcock 20,695 (US); Prov. The correct name of this plant cannot be Esmeraldas: Rio Santiago, Acosta-Solis 19,710 established conclusively. Linnaeus' original de- (US); Prov. Manabi: Bahia, Acosta-Solis 12,002 scription refers to a plate and description in (F); Prov. Pichincha: between Santo Domingo

Sloane ("Sloane jam. t. 68. f. 3"). Neither the and Quininde, Acosta-Solis 13,289 (US). text nor the plate are clear enough for posi- FRENCH GUIANA: vicinity of Cayenne, tive identification; both could fit the species Broadway 44 (NY, US). GALAPAGOS IS- Bbigham Young University Science Bulletin 112

habit, x (B) spikelet partly dissected, Fig 75. ChloTis pycnothrix and C. radiata. (A-E) C. pycnothrix. (A) 1/5; group of sterile florets, showing vanation X 7.5- (C) group of fertile florets, showing variation, x 10; (D) rajiata. (F) habit, x 1/5; (G) spikelet. partly dissected, x 7.5; (H X 10- (E) caryopsis, x 10. ( F-J ) C. florets, showing variation, x 10; group of fertile spikelets, showing variation, x 7.5; (I) group of sterile (J) caryopsis, x 10. Biological Seiues, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 113

-^^"^^^^

.K^ «*> I *v / c 114 Brigham Young University Science Bulletin

Table 13. Comparison of Chloris arenaria, C. cuben^is, C. ekmanii, C. halophila, C. pycnothrix, C. radiata, and Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 115

C. sagraeana.

Fertile lemma Awn Sterile floret Spikelets per Length Width length Length Width cm of rachis (in mm) (in mm) (in mm) Margin pubescence (in mm) (in mm)

5-7 3.5-3.7 ca 0.5 20-24 appressed-ciliate; 1.6-1.9 0.2r0.3 hairs short

5-9 3.1-3.6 0.6-0.7 9-12 densely ciliate, at least some hairs more than 1 mm long

6-10 2.0-3.0 0.4-0.5 5.5-10 116 Brigham Young University Science Bulletin with the short-ciliate lemma margins of C. 637 (K); Golungo Alto, Welwitsch 2,985b (K). radiata, from parts of Venezuela and North ARGENTINA: Prov. Corrientes: Empedrado Africa, are referred to C. pycnothrix only be- Dap., Estancia "Las Tres Marias" Pedersen cause the preponderance of their characters 3,053 (US); Prov. Misioncs: Posades, Parodi belong to that species. Individual specimens 4,017 (US); Prov. Formosa: without precise from these areas may be difficult, if not im- location, Jorgensen 43,053 (US). BRAZIL: possible, to refer to one species or another on Prov. Bahia: 39 km NE of Bahia, A. Chase the basis of only a few key characters. These 8,016 (F, MO, NY, US); Prov. Maranhao: Caro- aberrant specimens must represent casual, ran- lina to San Antonio dc Balsas, Stcallen 3,951 dom variation in spikelet features, for these (US); Prov. Matto Grosso: Tres Lagoas, A. plants otherwise maintain the distinctive epi- Chase 10,759 (US); Prov. Parana: Guaratuba, dermal features of the species (see Table 13; Morro do Brejatuba, Hatschhach 4,469 (US); Fig. 1). Prov. Rio Grande do Sul: vicinity of Guaiba,

Chloris pycnothrix is widely distributed Beetle 1,498 (UC); Prov. Catarina; Campo Ere, through Africa and northern South America Fazenda Sao Vicente, Castellanos 24,742 (K). BURUNDI: environs de Karuzi, Vander Ben ( Fig. 77 ) . Collection data indicate that these plants are frequently found as weeds in dis- 2,075 (K). CAMEROONS: Bertoua, Bahlwin turbed areas. While usually described as an 13,954 (US); Wakwa, 10 km S of N'Gaoundere, annual, greenhouse specimens have persisted Breteler 488 (K); Nkambe Div.: Bamcnda for over two years before dying, suggesting Highlands, 6°35'N, 10''40'E, Hepper and Char- that this species might behave as a weak peter 1,938 (K). CENTRAL AFRICAN RE- rennial under favorable conditions. PUBLIC: 35 mi from Bambari, Kt/rle FAIO Representative specimens examined: AN- (K). ETHIOPIA: Bagemedcr Prov.: Lake Lana, Bahudur Gorge, Grahham 116/21 (K); GOLA: Huila Dtr.: Posta Zootechnica, Teixeira Harar Prov.: NE of Lake Alemaya, 15 km NW of Harar, Burger 1,079 (K). GALAPAGOS ISLANDS: Santa Cruz Island, Academy Bay, Schimpff 39 (NY, US). GHANA: Venchi, 7°49'N, 2°0TW, Rose-hmes 30,519 (K); Nzere- kore, Baldwin 9,709 (US), .9,725 (K). GUINEA: Mamou, Chevalier 34.887 (K). IVORY COAST: Pakobo Nord, Adjoushoun 384A (K). KENYA: S mi N of Tsavo Falls, 6,300 ft, Bar- ney 1,123 (K); Nairobi, Bogdan B44 (UC); Amboseli National Park, 50 mi E of Namanga, Bogdan AB 5,500 (K); S Lake Elmenteita,

6,000 ft, Clayton DC 27 (K); Mt. Suswa, Glover 3,785 (K). MALAGASY REPUBLIC: Mt. Belamboany, Perrier de la Bathie 14 (K). MALAWI: Shiri Highlands, Buchanan 55 (K); Mlanje, Burtt-Davy ^1962/29 (K); near Zomba, Cormack 165 (K). MARTINIQUE: roadside, Velez 3,447 (US). MAURITIUS: Curepipe. Vacoa, Vaughan A-29 (K). MOQAMBIQUE: Mocuba Dtr.: Namagoa, Faulkner 18 (K), 145 (K, US); 26 km W of Lourenyo Marcjues, Gomes e Sousa 3,247 (K). NIGERIA: Samam, Zaria, Clayton DC 1,332 (K); Bauchi Plateau, Lely P769 (MO); Toro, 20 mi E of Jos, Scmple 173 (US). PARAGUAY: Dep. Guaira: Villarica, Jorgensen 3,532, 3,533 (US); Dep. Misiones: Florida, Campal s.n., 15 Apr 1938 (US); Dep. San Pedro: Primavera, Woolston 6.40 .(K). PERU: Dep. Apurimac: Abancay, Stordt, without collection number and date (K). REPUB- LIC OF THE CONGO: Marungu, Dubois 1..363 Fig. 77. Distribution of Chloris pycnothrix. Insets: Yangambi, (A) Galapagos Islands. (B) Reunion and Mauritius. (K); Eala, Lebrun 454 (MO, US); Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 117

Louis 16,007 (K); Katanga, 25 km N of Luhum- nerves; first glume 1.5 to 3.0 mm long, 0.2 to bashi (Elizabethville), Schmitz 7,384 (K). RE- 0.3 mm wide; second glume 3.7 to 5.3 mm long, UNION: St. Denis, Rivels, without collection 0.3 to 0.4 mm wide; fertile lemmas 3.8 to 5.5 number and date (K). SOMALI REPUBLIC: mm long, 0.6 to 0.8 mm wide, narrowly lance- Hargeisa, 9°33'N, 44°E, GiUett 3.917 (K); olate or elliptic, apex acute to acuminate, Medishe, Glover and GiUihnd 941 (K, US). callus bearded, margins appressed-pilose, awn 9 SOUTHERN RHODESIA: Sali.sbuiy, Brain 440, to 26 mm long; sterile floret one, 1.5 to 2.5 mm 1,541 (MO); Stapleford Forest Reserve, Gilli- long, 0.2 to 0.4 mm wide, lanceolate to elliptic, land 237 (K); Kariba Gorge, Goldsmith 73/60 apex acute to acuminate, glabrous to somewhat (K); Urungvve National Reserve, Zwipani area, scabrous, awn 7 to 13 mm long; anthers ca 0.7 Goodier 513 (K); Victoria Falls, wet spray mm long; caryopsis 1.9 to 2.4 mm long, 0.4 to areas, Parker 4,202 (K); near Sinoia, Rodin 0.6 mm wide, ellipsoid, trigonous; chromosome 4,376 (F, MO, UC, US). SUDAN: Dai fur number 2n = 80. Prov.: Lambong, Marra, Wickens Gur Jebel Chloris halophila is similar to C radiata and 2,593 (K); Equatoria" Prov.: Yd, Harrison 488 C pijcnothrix in outward appearance but has a (K); Khartoum Prov.: Khartoum, Jackson 4,186 coarser inflorescence and larger spikelets ( com- (K); Northern Prov.: Wadi Haifa, Pettet 90 pare Fig. 75 and 78). Additional differences (K). TANZANIA: Kondoa Dtr.: near Sam- are given in Table 13. bala, Ritrtt 2,603 (K); Ngorongoro Crater, For many years the names Chloris beyrich- Burtt 4,338 (K); Musoma, Emson 241 (K); iana, C. radiata var. beijrichiana. and Gtjmno- Bukoba Dtr.: Ngono Bridge, Gillman 67 (K); pogon beijrichianaus were misapplied to this Kilimanjaro, Lyamungo, Greenwaij 3,133 (K); taxon, especially by Parodi. Parodi later (1945) Seronera, Serengeti, Greenwaij 9,874 ( ; Zan- K ) recognized these names as being properly ap- zibar, Hitchcock 24,445 (US). UGANDA: plied to C. radiata and provided a new name, Nabilatuk, near Emonagabem, Dtjson-Hudson C. halophila. for the plant described above. 55 (K); Entebbe, Hitchcock 24,874 (US); 7 mi Chloris halophila has been collected through W of Moroto Karamoja, Lan<^dak'-Broum 1,572 a great altitudinal range from ca 2,000 feet (K); Busoga, Nawendwa, Maitland 1,406 (K); elevation in Tucuman Province, Argentina, to Mbale, near Mt. Elgon, Snowden 1,090 A (K). over 11.000 feet in Bolivia (Fig. 71, circles). UNION OF SOUTH AFRICA: Cape Provinces, It occurs in grasslands, especially in saline East London, McCaUum-Wehster N94 (K); conditions. Natal, SW of Mapumulo, Umtavi Valley, Moll Representative specimens examined: AR- 1,843 (K); Transvaal, Kruger National Park, GENTINA: Prov. Catamarca: Dep. Ambato, de Winter and Codd 600 (US). VENEZUELA: Camino al Rodeo, Cristobal 406 (UC); Prov. E of La Guiana, Ctirran and Hainan 902 Cordoba: S of de Mar Chiquita, Burkart 20,457 (NY). ZAMBIA: Chilonga, Mt. Makula Re- (UC); Dep. Punilla La Cocha, Stuckert 20.191 search Station, Hinds 6 (K); Mpika Dtr.: (G); Prov. Tucuman: valle de Tafi, Bruch s.n., Luangwa Game Reserve, Mfuwe, Mitchell 2,945 1908 (NY); Dep. Burruyacu: Sal Morado, Dias (K); Kasama Dtr.: Mungwi, Robinson 4,080 799 (NY); Rio de Nio, Monetti 43,048 (US); (K); Mazabuka Dtr.: Mochipapa to Mabwin- Tapia, Rodriguez 182 (US); Leales, Chamar gombe Hills, White 7,523 (K). Pojo, Venturi 711 (K, US); Prov. Salta: Pasaje del Rio Juaramendo, Lorentz 344 (F); Rosario 50. CHLORIS HALOPHILA Parodi, Rev. Ar- de Lerma, Meyer 3,464 (NY); Dep. Agua Cal- gentina Agron. 12:45. Fig. 1. 1945. (ISO- iente: Cavelaria, Venturi 3,773 (MO, US). TYPE: "Argentina, Buenos Aires, Perga- BOLIVIA: Dep. Cochabamba: Cochabamba,

mino . . . Parodi 2769" US! ) Fig. 78. Cutter 7,381 (US); Mizque, Eyerdam 24,987 Perennial 15 to 50 cm tall, sometimes up (F, G, K, MO, UC); Rio Tapacari. Kuntz s.n., to 80 cm, weakly stoloniferous; sheaths glabrous, 1892 (NY); Dep. La Paz: vicinity of La Paz, sometimes sparsely pilose above; ligule ciliate ca 10,000 ft. Bang 82 (BM, G, MO, NY, US). or a short, membranous crown; blades glabrous ECUADOR: Prov. Carchi: Pistu a la PalesHna, or scabrous, sometimes becoming sparsely pilose Acosta-Solis 16.557 (US); Prov. Loja: between on upper surface near the base; spikes four to Las Lomas and Loja, Hitchcock 21,409 (US); thirteen, 3.5 to 12.0 cm long (usually 7 to 8 Prov. Pichincha: descenso del Quincha a Guayl- cm), arranged in several verticils; spikelets ap- labamba, Acosta-Solis 10,329 (F, US); Caja- pressed, somewhat distant, ca four to seven marca Prov.: Cajamarca, 2,600 m, Anderson 630 per cm of the scabrous rachis; glumes narrowly (UC, US). PERU: Dep. Cuzco: Cuzco, Hitch- lanceolate, glabrous except for the scabrous mid- cock 22,492 (US); 5 km S of Ollantaytambe, University Science Bulletin 118 Brigham Young

dissected, x 7.5; (C-^H) fertile florets, show- Fig. 78. Chloris hahphila. (A) habit, x 1/5; (B) spikelet, partly caryopsis, x 7.5. ing variation, x 5; (I-N) sterile florets, showing variation, x 7.5; (O) )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae) 119

Hitchcock 22,552 (K, NY, US); Dep. Hauncave- glume, less than 1.5 mm long, and the sterile lica, Izcuchaca, entre Cucnca y La Mejorada, floret are both smaller than any other species in 3,100-3,150 m, Tovar 2,448 (US); Prov. Huan- this complex. Occasional specimens have gla- cavo: Mantero, N of Huancavo, 3,300 m, An- brous rather than hain' leaves, but this character derson 683 (UC, US). does not correlate with either geographic origin of the specimen or with other features. 51. CHLORIS EKMANII Hitchcock, U.S. Dept. This species is apparently widely distributed

Agrie. Misc. Publ. 243:129. 19.36. (HOLO- in Cuba, Jamaica, and Hispaniola ( Fig. 80, cir- TYPE: "Bal)iijal, Jagiicy Grande, prov. Ma- cles) in a variety of ecological sites, such as

tanzas, Cuba . . . Ekman . . . 16594." US! along railways, roadways, and rivers, shrubby Fig. 79, A-F. savannas, pastures, and pinclands. It is often collected in bright red clay soils, though other Perennial 30 to 45 cm tall, occa.sionally up to collections have been made on rocky or even 60 cm tall, tufted to short-stoloniferous; culm.s serpentine soils. slender, wir\', somewhat geniculate at the base; Representative specimens examined: CUBA: sheaths compressed-keeled; blades up to 10 cm Prov. Camagiiey: Camagiiey to Santayana, Brit- long, usually less than 1 mm wide, sparsely to ton 2,428 (NY); San Serapio, N of Camaguey densely pilose on both sides, occasionally gla- City, Leon 16,927 (US); Prov. Havana: near brous; spikes three to six, 1.5 to 5.0 cm long, slen- Almandares River, Leon 287 (US); San Miguel, der, flexuous, erect to appressed when young, at Eknuin 10,912 (US); La Pitanill, San Miguel de maturity spreading horizontally or becoming Casanova, Leon 12,454, 19,517 (US); Guanaba- somewhat reflexed; spikelets imbricate, ap- coa, Ekman 13,645 (F, NY, US), Hitchcock pressed when young, somewhat divergent at 23,299 (K, US), 23,240 (US); Jata Hills, Ekman maturits', ca 6 to 10 per cm of the scabrous 856 (US); Isla de Pifios, Nueva Gerona, along rachis; glumes lanceolate, glabrous except for Rio Los Casas, Ekman 11,682 (NY); Prov. Las the scabrous midnerve; first glume 1.1 to 2.0 mm Villas ( Santa Clara ) : Trinidad, road to San Juan long, 0.1 to 0.3 mm wide; second glume 2.3 to de Letran, Ekman 13,975 ( US ) ; Gara, Leon 3.6 mm long, 0.1 to 0.3 mm wide; fertile lemma 1,480 (NY); vicinity of Sancti Spiritus, Serguis 2 to 3 mm long, 0.4 to 0.5 mm wide, narrowh 2,569 (US); Pitajones to Ciegos de Ponciano, lanceolate to elhptic, callus bearded, sparsely to Shafer 12,225 (NY, US); Prov. Matanzas: Ja- densely appresscd-ciliate on upper margins, giiey Grande, Ekman 16,954 (F); Prov. Oriente: otherwise glabrous, awn 5.5 to 10.0 mm long; Guamo, Hitchcock 23,412, 23,427 (US); Hol- sterile floret one, 0.4 to 1.1 mm long, less than guin, Eknmn 3,229 (F); Prov. Pinar del Rio: El 0.2 mm wide, usually included in the fertile Sabalo, Finca Sabanalamar, Killip 32,294 (US); lemma, awn 3.3 to 6.0 mm long; carj'opsis ca Loma Pelada de Buenovista, Cavajabos, Leon 1.4 mm long, ca 0.3 mm wide, ellipsoid, trigo- 13,226 (US); Bahia Honda, Wilson 9,421 (NY). nous. DOMINICAN REPUBLIC: Prov. Montecristi: Chloris arenaria, C. cul)ensis, C. ekmanii, C. Guayabin, Abbot 916 (US); Santiago Prov.: racliata, and C. sagraeana form a complex of Licev al Medo, near Peiia, Jiminez 3,910 (US). species in the Caribbean region, the area to JAMAICA: Cornwall Co., Montego Bay, Hitch- which all except C. racliata are restricted. While cock 9,695 (US); Middlesex Co.: Lititz, Man- most specimens can be named with reasonable chester, Harris 12,434 (F, K, NY, US), Harris confidence, some either have intermediate fea- 12,697 (K, NY, US); Inverness, Lower Clar- tures or else recombine characteristics of two endon, Harris 12,727 (K, NY, US); between species. These are often found in, but are not Ewarton and Linstead, Hitchcock 652 ( F, K, restricted to, areas of sympatr)'. Field studies MO, NY, UC, US); New Forest, Southern Man- are needed, especially in Cuba; and precise def- chester, Hitchcock 9,823. 9,847 (US). initions of the taxa invoh'cd and evaluation of their biological significance must await further 52. CHLORIS ARENARIA Hitchcock & Ek- infomiation. The treatment presented here will man ex Hitchcock, U.S. Dept. Agric. Misc. enable one to name most specimens. A further Publ. 243:131. 1936. (HOLOTYPE: "Cuba, discussion of variation patterns is presented un- La Grifa, prov. Pinar del Rio. Ekman der the species most closely concerned. Table 13 11253." US! ISOTYPES: F! K!) Fig. 79, presents a comparison of the characteristics of G-K. these species.

Chloris ekmanii is most similar to C. sagrae- Chloris eleusinoides Grisebach var. vestita

arm but is a more delicate plant with wiry stems Greenman ex Combs, Trans. Acad. Sci. St. and, usually, very narrow leaves. The first Louis 7:477, pi. 39. 1897. (Fragment of 120 Bricham Young University Science Bulletin

habit, x ligiilc region, x 5; (C,D) Fig. 79. Chloris ekmanii and C. arenaria. (A-F), , C._ ckmatiii._ (A),.-, -1/3;, (B). , „ „ spikelets, partly dissected, x 7.5; (E) sterile florets, showingshowinj variation, x 10; (F) earyopsis, x 10. (G-L) florets, show- C. arenarUi. (G) habit, x 1/4; (H,I) ligule region, x 5; (J) spikelet, x 7.5; (K, L) sterile ing variation, x 15. . ) . )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 121

TTPE: "Chloris sagraeana nob. no. v. fol. ." obtusa Cuba. . . P! Fragment of type:

US! ) Fig. 81, I-T. Chloris morales-coelloi Leon ex Britton, Bull. Torrey Bot. Club 53:4.58. 1926. (ISO- ". TYPE: . . Cajobabo . . . not far from

mouth of the Jojo River, Oriente . . . Leon

. . . 12.320." US!

Perennial 15 to 80 cm tall, cespitose, erect or ascending from a geniculate base; sheaths glabrous, ligule short-ciliate; blades up to 15 cm long, 1.3 to 3.0 mm wide, glabrous; spikes three to six, .3.5 to 15.0 cm long (usually ca 4 to 9 cm long), becoming widely divergent or somewhat Fig. 80. Distribution of Chloris cubensis (squares), reflexed at maturity'; glumes narrowly lanceolate, C. chmanii (circles), and C. sagraeana (triangles). glabrous except for the scabrous midrib; spike- Inset A: Puerto Rico and the Lesser Antilles. lets appressed, ca four to nine per cm of the scabrous rachis; first glume 1.6 to 2.2 mm long,

type: "Rio Damuji, Rodas Cuba . . . Combs 0.2 to 0.4 mm wide; second glume 2.4 to 3.5 mm 631" US! Origin of fragment unknown. long, 0.5 to 0.7 mm wide; fertile lemma 2.6 to 3.5 Perennial 60 to 70 cm tall; culms cespitose, mm long, 0.5 to 0.7 mm wide, elliptic, callus erect to somewhat decumbent at base; sheaths bearded, with hairs less than 0.8 mm long, mar- compressed-keeled, puberulent. becoming dense- gins strongly appressed-pilose, the hairs usually ly so near the apex; ligule a dense tuft of less than 0.5 mm long, keels glabrous, occasion- hairs; lower leaves 5 to 10 cm long, 1.5 to 3.0 ally very sparsely pilose, awn 7 to 13 mm long; mm wide, becoming shorter upward on the sterile floret one (occasionally two), 0.7 to 1.6 culm; blades scabrous to densely pilose above, mm long, 0.2 to 0.5 mm wide, cvlindrical, gla- denseh' pilose below; spikes five to six, 4 to 6 brous, apex acute, awn 3.3 to 9.0 mm long; cary- cm long, digitate, ascending when young, be- opsis 1.6 to 1.7 mm long, 0.4 to 0.5 mm wide, coming horizontal or somewhat reflexed at ma- ellipsoid, trigonous. turit)'; spikelets rather denselv imbricate, ap- Chloris sagraeana is the most variable species pressed, ca five to seven per cm of the scabrous within the complex that includes C. arenaria, C. rachis; glumes narrowly lanceolate, acuminate, cubensis, C. ekmanii, and C. radiata. This vari- glabrous except for the scabrous midnerve; first ation is especially noticeable with respect to glume 2.2 to 2.5 mm long, 0.2 to 0.3 mm wide; the size of the fertile and sterile lemmas. Fre- second glume 3.3 to 3.5 mm long, 0.2 to 0.4 mm quently specimens with exceptionally long lem- wide; fertile lemma 3.5 to 3.7 mm long, ca 0.5 mas will produce two sterile florets; and on oc- mm wide, lanceolate to elliptic, callus bearded, casion, the lowennost "sterile" floret will pro- otherwise glabrous except for the short, ap- duce a caryopsis. though it otherwise resembles pressed-ciliate margins, awn 20 to 24 mm long; the upper sterile floret in nK)rphology. This vari- sterile floret one, 1.6 to 1.9 mm long, 0.2 to 0.3 ation in stmcture mav or mav not be correlated mm wide, narrowly c\lindrical, glabrous, awn with size. A specimen with two sterile florets, 15 to 20 mm long; caryopsis not seen. coincidentally very robust, was named Chloris Chloris arenaria mav' be separated from other morales-coelloi by Leon (in Britton, 1926). species in this complex by its densely villous Chloris sagraeana may be separated from C. leaf blades and sheaths and bv its long awns on ekmanii by its more robust stature and broader the lemmas ( 20 to 24 mm ) leaves and from C. cubensis in having shorter, Only one of the two specimens studied had appressed hairs on the lemma margins (Table collection infonnation. This plant ( Eknuin 13). 11,253) was collected in pineland on white sand. This species has been collected from a vari- Specimens examined: CUBA: without pre- ety of habitats, including pinelands, serpentine cise location, Wright 3,819 (K, NY, US). outcrops, coral rocks, and pastures ( Fig. 80,

triangles ) 53. CHLORIS SAGRAEANA Achille Richard Representative specimens examined: BAHA- in Sagra, Hist. Fisica Politica Natural Isla MA ISLANDS: Fortune Island, Hitchcock s.n., Cuba XI. Botanica 315. 1850. (HOLO- Nov 1890 (F, MO); Inagua Island, Hitchcock 122 Brigham Young UNivEHsrri- Science Bulletin

Fig. 81. Chloris ciihensis and C. sagraeana. (A-H) C. ciibensis. (A) habit, .\ 1/3; (B) ligule region, x 3; (C) spikelet, partly dissected, x7.5; (D) florets, x 7.5; (E-G) sterile florets, showing variation, x 10; (H) caryop-

( ) ligule x spikelet, sis, X 10. (I-T) C. sagraeana. I,J habit, x 1/3; (K) region, 3; (L) partly dissected, x 7.5; (M) spikelet with two sterile florets, partly dissected, x 7.5; (N) spikelet from type, partly dissected, .\ 7.5;

(0,P) additional florets, showing variation, x 7. .5; ( Q-S ) sterile florets, showing variation, x 10; (T) caryopsis, X 10. )

Biological Series, Vol. 19, No. Taxonomy of the Genu,s Chloris (Gramineae) 123

s.n., 3 Dec 1S90 ( F, MO, NY); Nassau, Britton long, 0.2 to 0.3 mm wide; second glume 2.9 to and Brace 596 (F, NY), Curtiss 81 (US). BAR- 3.9 mm long, 0.2 to 0.4 mm wide; fertile" lemma BADOS: Allen s.n., Jan 1942 (K, NY). BARBU- 3.1 to 3.6 mm long, 0.6 to 0.7 mm wide, lanceo- DA: S of Codrington, Danfortit 3 (US). CUBA: late, callus long-bearded, upper 2 3 of margins Prov. Havana: vicinit\' of Havana, Ansouin 779 long-ciliate with hairs up to 1 mm long, keel and (US); between Morro and Cojimar, Ekmun 389 sides sparseh' pilose, lemma awn 9 to 12 mm (US); Guanabacoa, Lomas de los Jatos, Ekman long; sterile floret one, 1.4 to 1.9 mm long, 0.4 to 10,902a (US); Near Marianas, Leon 1,965 (US); 0.6 mm wide, cylindrical, apex obtuse, awn 6 to near Luzano, Leon 4,807 (US); Santa Fe, Leon 9 mm long; caryopsis ca 1.3 mm long, ca 0.3 to 13,096 (US); Cuabal de Jesus Maria Minas, 0.4 mm wide, ellipsoid.

Leon and Hioram 4,783, 4,787 (US); Prov. Ma- Chloris cubensis is similar to C. saeraeana, tanzas: Babiyal, Ekman 16,954 (US); Prov. Pi- from which it may be separated by its generally nar del Rio: La Grifa, Ekman 11,252 (US); larger size, less divergent mature spikes, and by Mariel, coral reef E of town, Ekman 12.837 the long hairs on the lemma margins and, usu- (US); Prov. Oriente: pinelands, crest of Sierra ally, on the lemma keel. (Fig. 81; Table 13). (US). Nipe, Leonard and Leonard 3,183 DO- Specimens have been collected on roadsides Prov. Montecriste: MINICAN REPUBLIC: and open dr)- areas (Fig. SO, squares). Valle de Cibao, Villa Vasquez, Ekman 13,128 Representative specimens examined: AN- (K, US); Prov. de Santiago: Mao, Abbott s.n., TIGUA: Nonsuch Bav, Facey Creek, Box 556 23 Feb 1921 (US). GUADELOUPE: without (US). BARBADOS: 'without precise location, precise localit)', Diiss 4,110 (F, NY, US); Les Hitchcock 16.499 (US). CUBA: Prov. Cama- Pointes, Quesial 1,740 (US). HAITI: Pignon to gii(>y: Minas, Ekman 15,368 ( F, NY, US); Prov. San Rafael, Cook et al. 138 (US); Port-au- Havana: Loma de Tierra, Ekman 13,469 (K, Prince, Ekman 2.107 ( K, US ) ; Tortue Island l)c- US); Vibora, Leon 859 (US); Calvarco, Leon tween La Vallee and Ravine-Rosetiere, Ekman 860 (US); Prov. Las Villas: Loma de Belen, 4,058 (K, US); Corail, Dutreuil, Ekman H Leon 15,627 (US); Prov. Oriente: Holguin, Ek- 10,748 (K. US); Grande Cayemite, Pointe man 1,014 (F, K, MO, NY, UC, US), Ekman Macon, Eyerdam 336 ( F, NY, US); vicinit\' of 15,715 (F, NY, US). HAITI: Tortue Island, vi- Petionville, Leonard 4,984 ( NY, US ) ; vicinit)- of cinitv of Basse Terre, Leonard and Leonard 12,- Gros Morne, Dep. Artibonite, Leonard 9,839 5.:5,S'(US). JAMAICA: Hanover Park, Bailev's (US); Port-de-Paix, Leonard and Leonard Point, Adams 7,964 ( BM ) ; Cockpit, Vere, Har- 11.051 (US); Baie des Moustiques, Leonard and ris 12.462 (F, MO, NY, US); Spanish Lower Leonard 12,067 (US); vicinitv of Jean Rabcl, Road, Harris 12,476 (K). PUERTO RICO: Rio Leoiuird and Leonard 12,552 (K, MO, NY, US); Piednxs, Alberts 273 (US). vicinitv of Mole St. Nicolas to Bombardopolis, Leonard and Leonard 13,225 (UC, US). 55. CHLORIS MOSSAMBICENSIS Schumann, JAMAICA: Cornwall Co., Savanna-la-Mar, Notizbl. Bot. Gart. Berlin 1:104. 1895. (ISO- Hitchcock 9,890 (F, MO, NY). Middlesex Co.,

TYPE: "Mozambik, Cabareino Grande . . . Ashlev Hall Savannah, Lower Clarendon, Harris No. 88. leg. Prelado." BM!) Fig. 82. 12,738 (F, NY, US).

Tetrapogon mossambicensis ( Schumann 54. CHLORIS CUBENSIS Hitchcock & Ek- Chippcndahl ex B.S. Fisher in Meredith, man c\ Hitchcock, U.S. Dept. Agric. Misc. Grasses and Pastures of South Africa 198. " 1955. (Based on Chloris mossambicensis Publ. 243:131. 1936. (HOLOTYPE: . . . Schumann.) Cuba, prov. Oriente, Gamboa . . . Ekman

. . . 14960." US!) Fig. 81, A-H. Perennial, tufted to aggressively stolonifer- Perennial usualh' 50 to 60 cm tall, occasion- ous, 20 to 100 cm tall (usually 40 to 80 cm); ally up to 90 cm, tufted to stoloniferous; sheaths sheaths glabrous to scabrous; ligule a short-cili- glabrous, ligule a short-ciliate or naked crown ate crov.'n, often with a few long hairs inter- up to 0.5 mm tall; leaves 10 to 15 cm long, ca spersed, especially near the margins; blades up 1..5 mm wide, often folded, scabrous to sparseh' to 30 cm long, ca 8 mm wide, obtuse to acumi- pilose above, glabrous below; spikes three to nate; spikes three to six (often four or five), 3 seven, 5.5 to 8.5 cm long, radiate, ascending; to 10 cm long, divaricate to horizontally spread- spikelets appressed, imbricate, ca five to nine ing; spikelets sparsely inserted, imbricate, diver- per cm of the scabrous rachis; glumes narrowly gent, ca eight or nine per cm of the scabrous lanceolate, membranous, glabrous except for tlie rachis; glumes narrowly lanceolate, thin, mem- scabrous midnerve; first glume 1.7 to 2.4 mm branous, glabrous except for the scabrous mid- 124 Brigham Young University Science Bulletin

Fig. 82. ChloTis mossambicensis. (A) habit, x 1/5; (B) spikelet, partly dissected, x 5; (C) sterile florets, x

7.5; (D) partly dissected "sterile" florets, showing upper sterile floret (u.s.f. ), palea of lower "sterile" floret (pa.), lemma of lower "sterile" floret (1.), and flower, x 20; (E-J) florets, showing variation, x 5; (K) caryopsis, x 7.5. nerve; first glume 1.6 to 2.6 mm long, ca 0.2 mm pilose, awn 2 to 9 mm long; caryopsis 1.2 to 1.3 wide; second glume 3.0 to 4.4 mm long, 0.3 to mm long, 0.3 to 0.5 mm wide, ovoid to obovoid, 0.4 mm wide; fertile lemma 3.0 tO' 4.7 mm long, rounded to trigonous. 0.8 to 1.2 mm wide, elliptic to ovate, apex acute Tliis species has been treated variously as a to obtuse, callus densely bearded, margins ap- member of Chloris or of Tetrapogon. As brought pressed-pilose below, becoming ciliate near the out in i-arlier discussion, the two genera are very apex with hairs ca 1 mm long, keel appressed- closely related and must be somewhat arbitrari- pilose, internerves glabrous, awn 3.5 to 12.0 mm ly separated. Essentially, Tetrapogon regularly long; sterile floret usually one (occasionally has more than one basal fertile floret, vyhile two), 1.5 to 3.3 mm long, 0.7 to 1.5 mm wide, Chloris has just one. A sampling of nineteen narrowly to broadh' triangular, usually barren, specimens, from different locations in southern occasionally fertile and setting seed, apex trun- Africa, showed seventeen with a single fertile cate, usually glabrous, very occasionally sparsely floret and two \\ath two fertile florets. Because Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 125

dcntes, 6-18 cm longae, dilute virides. Spiculae appressae, secus rhachin scabram ± 9-10 per utnimque cm dispositae. Glumae lanceolatae tenues membranaceae, praeter carinam scabram glabrae. Cluma prima 1.7-2.2 mm longa, 0.2 -0..3 mm lata. Gluma secunda 2.5-3.2 mm longa. :+: 0.5 mm lata. Lemma fertile ellipticum, apiee acutum, 2.5-2.8 mm longum, 0..8-0.9 mm latum, callo angusto barbato, secus margines carinamque dense appressepilosum; arista 1-2 mm longa. Flosculi steriles 2, inferior obovoideus obtusus vel acutus, 1.5-1.7 mm longus, 0.6- 0.7 mm latus, praeter carinam sparsim appresse- pilosam, glabcr, arista 0.7 to 1.6 mm longa, superior ob()\oideus apicc ± tmncatus 1.0-1.5 mm longus, 0.5-0.8 mm latus, versus apicem scaber ceterum glaber, arista rb 0.5 mm longa. Fig. 83. Distribution of Chloris mossambicensis (south- Caryopsis ignota. em Africa). Lower stems and leaves unknown. Leaf sheaths glabrous, ligule a densely scabrous or of this and an overall rc,sein1)laco to tlie prepond- short-eiliate, tnincate crown; upper leaf blades erance of Chloris species, I am treating this 10 to 18 cm long, 5 to 16 wide, scabrous species as a member of the genus CJdoris. mm above, glabrous below, acuminate; spikes seven Chloris mossambicensis is wideK' distributed to nine, 6 to 18 cm long, ascending, pale green; in soutliern Africa (Fig. 83), where it lias l)een spikelets appressed, ca nine or ten per cm of collected in a variety of habitats. A more de- the scabrous rachis; glumes lanceolate tailed and expanded distribution map may be to narrowly ovate, thin and mtMubranous, glabrous ex- found in Chippendall ( 1955). cept for the scabrous Representative specimens examined; KEN- midnervc; first glume 1.7 to 2.2 mm long, 0.2 to 0.3 wide; second YA: 14 Falls, Athi River, near Doinyo Sapuk, mm glume 2.5 to 3.2 mm long, ca 0.5 nmi wide; fertile lem- Bogdan AB 1,590 (K); Nairobi to Mombasa ma 2.5 to 2.8 mm long, 0.8 to 0.9 wide, el- Road, near Samburu Station, Bogdan AB 3,616 mm liptic, callus nanow, bearded, margins (K); Mariakani, Bogdan AB 5,429 (K); Kwale and keel densel\- appressed-pilose, apex acute, Dtr., near Tani, bc-twetn Samburu and MacKin- awn 1 to 2 mm long; sterile florets two; lower sterile floret non Road, Drummond and llensleij 4,189 (K). 1.5 to 1.7 mm long, 0.6 to 0.7 wide, obo- MOgAMBIQUE; Sul do Save, Maputo River, mm \oid, keel lightly appressed-pilose, entre Changalane e Catuane, Mijre and Carval- otherwise glabrous, apex acute to obtuse, hao 1,440 (K); between Moamba and Boane, awn 0.7 to 1.6 mm long, upper sterile floret I.O 1.5 Schwcickcrdt 1,919A (K); Louren90 Marques. to mm long, 0.5 to 0.8 mm wide, broadlv obovoid, Rio Limpopo, entre Mianga e Guija, Torre 6,614 glabrous below, scabrous at the tmncate to rounded apex, (K). SWAZILAND: 11 mi W of Stegi, Codd awn ca 0.5 long; anthers ca 0.4 long; and Dyer 2,863 (K). TANZANIA: Tanga Prov.. mm mm car\opsis unknown. Mkwaja, Pangani, banks of Bohomaia River, van Renshurg 516 (K). UNION OF SOUTH AFRI- HOLOTYPE: Burma, roadside between CA: Natal: 8 mi N of Mkuze on Candover Road, Meiktila and Pvawbye. On calcareous stiff clay. Edwards 3,257 (K); Zululand, lower Umfolozi V. Them Lain 311.' 23 Oct 1947. K! region. Ward 2,442 (K); Transvaal: 16.5 mi SE Chloris biirmcnsis is, \\ithout question, dis- of Pretoria Kop, Acocks 16,713 (K); Barberton tinct from all other species in Chloris. The com- Dtr., near Krokodipoort siding, Liehenherg 2,- bination of awned sterile florets in pairs and the 406 (K); Kruger National Park, 4 mi S of Sku- densely appressed-pubescent, elliptic fertile kuza, de WinU'r and Codd 546 (K). florets (Fig. 84) separates this taxon from all other species. The onl\- specimen available—the 56. CHLORIS BURMENSIS sp. nov. Fig. 84. holotype— is unfortunately without any basal Culmi inferiores cum foliis igiioti. Foliorum parts; so infonnation on height, habit, or longev- vaginae glabrae; ligulae breves ± truncatae, ity is not available. Superficially, C. hurmensis scabrae vel brcviterciliatae; laminae 10-18 cm resembles some species of Tetrapogon, but de- longae, 5-16 mm latae, acuminatae, inferne tails of its spikelet structure place it definitely in glabrae, superne scabrae. Spicae 7-9, adscen- Chloris. 126 Brigham Young University Science Bulletin

K VL VM

Fig. 84. Chloris burmensis. (A) inflorescence and upper culm, x 1/3; (B) ventral view of portion of rachis, X 3; (C) dorsal view of portion of rachis, x 3; (D) spikelet, partly dissected, x 10; (E) fertile floret with one half removed, showing palea and flower, x 15; (F) palea, ventral view, x 15; (G-I) lower sterile florets, showing variation, x 10; (J-M) upper sterile floret, showing variation, x 10.

UNIDENTIFIED, REJECTED, OR REASSIGNED NAMES

Tlie following alphabetical list includes all Chloris acuminata Trinius, Species Gram. names used in Chloris that are presently uniden- Icon. 3:/)/. 305. 1831. =Eustachi/s disticho- tifiable, not located, or assigned to other genera. phtjlla (Lagasca) Nees. Chloris acicularis Lindley ex Mitchell, Jour. Chloris affinis Caro and Sanchez, Kurtziana Exped. Interior Trop. Australia 33. 1848. Basis 6:224. 1971. Fig. 2. Specimens of this plant of Enteropogon acicularis (Lindley) Lazarides. were not available in time to be considered Chloris acicularis var. queenslandiae Domin, in this publication. On the basis of the original Biblioth. Bot. 85:368. 1915. = Enteropogon description and accompanying illustration, it acicularis (Lindley) Lazarides, on basis of the would seem to be most closely related to description. Chloris pijcnothrix Trin. Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 127

Chloris agowensis Hochstetter ex Kegel, An- 1971. Specimens of this plant were not avail- notationes Bot. Index Seminum 26. 1862. able in time to be considered in this publica- Nomen nudum, synonym of Chloris spathacea tion. On the basis of the original description Hochstetter. it would seem to be part of the variation pat- Chloris anoplia (Nees) Foumier ex Hem- tern of Chloris castilloniaija Lillo and Parodi. sley, Biol. Cent.-Amer. Bot. 3:558. 1885. =?, Chloris cenchriformis (Achille Richard) possibly Leptocldoa sp. Based on Leptochloa Baillon ex Schweinfurth and Ascherson, Beitr. anoplia Nees. Flora Aethiopiens 310. 1867. Based on Lepi- Chloris argentina (Hackel) Lillo and Parodi, dopironia cenchriformis Achille Richard. =Tet- Physis 4:180. 1918. Based on Chloris clis- rapogon cenchriformis (Achille Richard) Pilger. tichophijlhi var. argentina Hackel which — Eu- Chloris cheesmanii Hackel ex Cheesman, stacht/s retiisa (Lagasca) Kunth. Trans. Linnean Soc. Bot. 6:305. 1903. =En- Chloris argentinensis Parodi, Rev. Fac. Ag- teropogon unispiceus (F. Mueller) Clayton on ron. and Vet. Univ. Nac. Buenos Aires 2:28.3. basis of description. 1919. Error for Chloris argentina? Chloris ciliata Swartz var. brachyathera Chloris aristida Salzmann ex Steudel, Syn. Hackel ex Kurtz, Bol. Acad. Ciencias Cordoba PI. Glum. 1:218. 1854. Nomen nudum, as s\n- (Rep. Argentina) 16:257. 1900. Without de-

onyin of Gijmnopogon foliosus Nees. scription. As described bv Parodi ( Rev. Fac. Chloris hahiensis Steudel, S\n. Pi. Glum. .\gron. and Vet. Univ. Nac. Buenos Aires 2:271. 1:208. 1854. =Eitstachi)s caribaea (Sprongcl) 1919) = Chloris ciliata Swartz. Herter. Chloris compressa De Candollc, Cat. Horti

Chloris hahiensis f. glabrescens Hackel, Re- VIonspeliensis 94. 1813. =? Specimens P!, pert. Sp. Nov. 8:46. 1910. =Eustac}it/s cari- labeled as this species, of uncertain origin, are baea (Spri'iigel) Herter. C. radiuta Swartz. A photo, in US!, of a speci- Chloris hoivinii A. Camus, Bull. Soc. Bot. men in Montpellier labeled as C compressa, France 79:845. 1933. Basis of Daknopholis perhaps in de Candolle's script, is also C hoivinii (A. Camus) Clayton. radiata. The protologue might applv equally to Chloris brachi/athcra (Hackel) Herter, Re- C. radiata or C. virgata Swartz. vista Sudamer. Bot. 6:146. 1940. Based on Chloris confertifolia Trinius in Sprengel, Chloris ciliata Swartz var. brachi/athera Hackel Neue Entdeckungen 2:74. 1821. =Eustaclnjs in Kurtz, which see. disticiu^plnjlla (Lagasca) Nees. Chloris brevipila Rosengurtt and IzaDactyloctenium sp., on basis of

2. 1968. ^Eustachtjs sp., on basis of descrip- fragment of type ( US! ) from CN. tion and illustration. Chloris curtipendula Michaux, Flora Bor. Chloris calvesccns Hackel in Chodat and Amer. 1:59. 1803. Basis of Bouteloua curti- Hassler, Bull. Herb. Boissier II. 4:279. 1904. pendula (Michaux) Torrev in Emory. = Eustaclii/s sp., on basis of description. Chloris cynodoidea Lagasca, Elenchus Plan- Chloris campulodes Trinius ex Steudel, Noin. tarum 5. 1816. Nomen nudem. Bot. ed. 2. 1:353. 1840. Nomen nudum. Chloris cynodon (Linnaeus) Trinius, Gram. Chloris capensis (Houttyn) Thellung, Re- Unifloris 229. 1824. Based on Panicum dactylon pert. Sp. Nov. 10:289. 1912. Based on Andro- Linnaeus —Cynodon dactylon (Linnaeus) Per-

pogon capensis Houttyn, which is based on soon.

Andropogon muticum Linnaeus, a superfluous Chloris delicatula Clarke in Hooker, f. Flora

name. Andropogon muticum Linnaeus is uniden- British India 7:290. 1896. Basis of Gymnopogon tifiable. delicatula (Clarke) Bor. Chloris capensis var. hahiensis (Steudel) Chloris dichanthioides Everist, Queensland Parodi, Revista Argentina Agron. 20:26. 1953. — Agric. Jour. 49:4.32. /)/. 155, 19.38. Basis of Eustachys caribaea ( Sprengel ) Herter. Austrochloris dichanthioides (Everist) Lazar- Chloris capensis var. glabrescens (Hackel) ides. Parodi, Revista Argentina Agron. 20:20. 1953. Chloris digitaria Humboldt, Bonpland, = E«.sfflt/i(/s caribaea (Sprengel) Herter. Kunth, Nov. Gen. and Sp. PI. 1:168. 1816. Chloris card)aea Sprengel, Syst. Veg. 1:295. = Leptochloa sp. on basis of description. 1825. Basis of Eustachys caribaea (Sprengel) Chloris digitata (Roxburgh) Steudel, Syn. Herter. PI. Glum. 1:207. 1854. Based on Melica digi- Chloris castilloniaiui Lillo and Parodi var. tata Roxburgh which = Enteropogon dolicho- pubescens Caro and Sanchez, Kurtziana 6:230. stachya (Lagasca) Lazarides. 128 Bricham Young University Science Bulletin

Chloris (listacluja Bojer, Hortus Mauritianus fasciculata Schrader in Schultes. Based on An- 371. 1837. = Chloris filiformis (Vahl) Poiret? dropogon fa.yciculatum Linnaeus =Microstegium

An unnamed Bojer specimen from Mauritius sp. on basis of holotype ( LINN! ).

(in G!) is C. filiformis. The protologue is in- Chloris floccifolia ( Forskal ) Pcjiret in complete. Lamarck, Encycl. Method. Bot. Suppl. 2:238. Chloris distichophijlla Lagasca, Gen. Sp. 1811. Based on Cynosurus floccifolius Forskal Nov. 4. 1816. Basis of Eustachys distichophijlla = Eleusine floccifolia (Forskal) Sprengel. (Lagasca) Nees. Chloris floridona (Chapman) Wood, Amer. Chloris distichophijlla var. acuminata (Trin- Bot. and Florist Pt. 2. 407. 1871. = Eustachys ius) Hackel, Bull. Herb. Boissier II. 4:279. floridana Chapman. 1903. = Eustachys distichophijlla (Lagasca) Chloris foliosa Willdenovv, Sp. PI. 4:924. Nees. 1806. Basis of Gymnopogon foliosus (Will- Chloris distichophijlla var. argentina Hackel denovv) Nees. Fragment of t\pe, from B, in in Stuckcrt, Anales Mus. Nac. Hist. Nat. Buenos US! Aires 11:113. 1904. = Eustachys disticlwphylla Chloris foliosa Lagasca, Gen. Sp. PI. Nov. 5. (Lagasca) Nees. 1816. Non Chloris foliosa Willdenovv. Uniden- Chloris distichophijlla var. f^emdna Hackel tified. A fragment, in US!, of a Lagasca speci- in Stuckert, Anales Mus. Nac. Hist. Nat. Buenos men dated 1820. from M, is Gymnojwgon sp. Aires 11:113. 1904. = Eustachys distichophijlla Chloris geminata Hochstetter, Flora 38:205. (Lagasca) Nees. 1855. =Tetrapogon sp. on basis of description. Chloris dolichostachya Lagasca, Gen. Sp. Chloris glaiica (Chapman) Wood, Amer. Nov. 5. 816. Basis of Enteropogon dolicho- Bf>t. and Florist Pt. 2. 407. 1871. = Eustachys stachya (Lagasca) Lazarides. glauca Chapman. Chloris duhia Humboldt, Bonpland, Kunth, Chloris glahrescens (Hackel) Rosengurtt and Nov. Gen. and Sp. PI. 1:169. 1816; /;/. 694. Izaguirre, Bol. Soc. Argentina Bot. 12:123. 1825. Basis of Leptochloa duhia (Humboldt, 1968. Based on Chloris hahicnsis var. glabres- Bonpland, Kunth) Net>s. cens Hackel = Eustachys caribaea (Sprengel) Chloris durandiana Schultes, Mantissa 2:.341. Herter. 1824. Based on Chloris gracilis Durand, which Chloris gracilis Durand, Chloridis Speciebus may equal Chloris radiata (Linnaeus) Swartz. 10. 1808. =? Tvpe not seen, description may See also gracilis Durand in this list. C be of Chloris radiata (Linnaeus) Swartz. Non Chloris dusenii Ekman, Arkiv for Botanik Chloris gracilis Humboldt, Bonpland, Kunth, 10:26. 1911. = Eustachys uligino.sus (Hackel) 1816. Herter; isotype in US! Chloris gracilis Humboldt, Bonpland, Kunth, Chloris elegans (Kunth) Roberty, Petite Nov. Gen. Sp. PI. 1:168. 1816. Non Chloris Flore Ouest.-Afrique 387. 1954. Basionym not gracilis Durand. = Leptochloa sp. on basis of cited. description. Chloris elegans (Kunth) Roberty, Bull. Inst. Chloris gryllus (Linnaeus) Honckeny, Svn. Fran9aLse Afrique Noire Ser. A. 17:51. 1955. PI. Gennaniae 1:437. 1792. =:Chry.wpogon Based on Ctenium elegans Kunth. gryllus (Linnaeus) Trinius on basis of descrip- Chloris eleusinoides Grisebach, Flora British tion. Based on Andropogon gryllus Linnaeus. West Indian Islands 539. 1864. Unidentified. Chloris "uineensis Schumacher, Beskr. Guin- Type not seen, but description suggests Chloris Planter 75. 1827. =Dactijlocteniuin sp. sagraeana A. Rich. eiske photograph type ( in US! ) in the Chloris emarginata Beauvois, Ess. Agrost. 79, on basis of of Isert Thonning Herbarium. 158. 1812. Nomen midem. and Chloris ecpdtans Trinius, Sp. Gram. Icon. 3. Chloris hlspida Durand, Chloridis Speciebus pi. 309. 1831. =Eustachijs paspaloides (Vahl) 17. 1808. =Bouteloua sp. on basis of Lanza and Mattei. description. Chloris falcata Swartz, Ges. Naturf. Freunde Chloris imherhis Desfontaines, Cat. PI. Pa-

Berlin Neuc Schrift 3:160. /;/. 1. Fig. 1. 1801. risiensis ed. 3. 15. 1829. Nomen nudem. = Harpechloa capensis Kunth. Chloris incomplcta Roth ex Roemer and Chloris fasciculata Schrader in Schultes, Schultes, Syst. Veg. 2:607. 1817. = Entero- Mantissa 2;.3.39. 1824. =Eustachys disticho- pogon dolichostachya (Lagasca) Lazarides on phijlla (Lagasca) Nees. basis of photograph of tvpe in K! Chloris fasciculata (Linnaeus) Thellung, Chloris lepidopironia Desveaux ex Regel, Repert. Sp. Nov. 10:289. 1912. Non Chloris Annotationes Bot. Index Seminum 26. 1862. )

Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 129

Nomen mulcm. lierbarium name published as a pallida Willdenow. =Schoenfeldia pallida synonym. Edgeworth, on basis of description. Chloris longiharbis Michaux ex Beauvois, Chloris panicea Willdenow, Sp. PI. 4:923. Ess. Agrost. 79, 158. 1812. Nomen luidcm. 1806. ^Enteropogon dolichostachya (Lagasca) Chloris nmcrantha Desveaux, Opuscules 73. Lazarides. Type not seen, but description 1831. =Tetrapogon sp. on basis of description. strongly suggests this species. Chloris macrantha Jaubcrt and Spach, lllustr. Chloris parva G. Mimeur, Bull. Mus. Hist.

PI. Orient. 4:42. /;/. 328. 1851. Non Chloris Nat. (Paris) 22:128. 1950. =Chloris prieiirii macrantha Desveaux. —Tetrapogon sp. on basis Kunth? Mimeur cites a specimen sent in 1880, of description. Talmy 86, from Senegal and deposited in P. Chloris macrostachija Hochstetter ex Achille This cannot be located, but another specimen Richard, Tent. Florae Abxssinicae 2:408. 1850. sent by Talmy, also from Senegal, but in 1882, Basis of Enteropogon macrostachyus (Hoch- is Chloris prictirii. The original description like- stetter ex Achille Richard) Munro ex Bentham. wise strongly suggests this species. Chloris maritimiis (Kunth) Trinius, Gram. Chloris parvispictila Caro and Sanchez, Unifloris 235. 1824. Based on Ct/nodon mari- Kuitziana 6:227. 1971. Fig. 3. Specimens of timus Kunth. =- Cijnodon sp. this plant were not available in time to be Chloris mearnsii Merrill, Philippine Jour. considered in this publication. On the basis Sci. 3:220. 1908. = Enteropogon sp. on basis of the original description and accompanying of description. illustration it would seem to lie most closely Chloris monostacJiya Michaux, Flora Bor. related to Chloris dandyana. Anier. 1:59. 1S03. —Ctenium sp. on basis of Chloris paspaloidcs Hochstetter, Flora 38:206. description. 1885. =Etistachys paspaloidcs (Vahl) Lanza Chloris monostachijo (Vahl) Poiret in and Mattei. Lamarck, Enc\cl. Method. Bot. Suppl. 2:238. Chloris paiili-diicis Schweinfurth, Beitr. 1811. Non Chloris monostachi/a Michaux. Basis Flora Aethiopiensis 298. 1867. Nomen inidtim. of Enteropogon monostachi/os (\'ahl) K. Schu- Chloris paijtensis Steudel, S\ii. PI. Glum. mann ex Engl. ( Based on Ct/nosurus niono- 1:207. 1854. =Cynodon dactijlon on basis of stachijus X'ahl. fragment of type (in US!) from Urville Chloris moorci F. Mueller, Linnaea 25:444. Herbarium. 1852. Unidentified. A specimen at K!, collected Chloris pedicellata Steudel, Flora 33:232. and labeled as C. moorei by von Mueller, 1850. Nomen ntidttm, as s\ nonvm of Schellingia though not the type, is Enteropogon acicuhiris tenera Steudel.

( Lindlev ) Lazarides. Chloris pendtda Salzmann ex Steudel, Syn. Chloris mucronata Michaux, Flora Bor. PI. Glum. 1:208, 1854. Nomen nudum, as Amer. 1:59. 1803. =Dactijloctenium sp. on svnonym of Chloris hahiensis Steudel. basis of description. Chloris penicellata (Vahl) Persoon, Syn. PI. Chloris multiradiata Hochstetter \'ar. ragazzii 1:87. 1805. =Tetrapogon sp.? on basis of de- Pirotta, Anales 1st. Reale Bot. Roma 6:157. scription of Cynostirtis penicillatus Vahl, upon 1896. Unidentific>d, possibh' a \'ariant of Chloris which it is based. virgata Swartz. Chloris penicillata Jan. ex Trinius, Gram. Chloris neglecta Nash, Bull. Tone\' Bot. Club Uniflores 231. 1824. Non Chloris penicillata 22:423. 1895. Basis of Eustachijs neglecta (Vahl) Persoon. Nomen ntidtim. as synonym (Nash) Nash. of Chloris compressa De Candolle. Chloris ohttisifolia Trinius, Gram. Unifloris Chloris perrieri A. Camus, Bull. Soc. Bot. 233. 1824. Unidentified, description suggests France 96:93. 1949. =Daknopholis hoivinii

Eustaclu/s tener ( Presl ) A. Camus. (A. Camus) Clayton on basis of holot\pe in P! Chloris ohttisifolia Balansa, Jour, de Bot. Chloris peregrina Durand, Chlorides Specie- (Morot) 4:166. 1890. Non Chloris ohttisifolia bus 11. 1808. Unidentified. Type not seen, Trinius 1824. =Etistachys tener (Presl) A. description suggests Chloris inflata Link. Camus. Chloris petraea Swartz, Prodr. Veg. Ind. Occ. Chloris pallida Willdenow, Sp. PI. 4:293. 25. 1788. Basis of Eustachys petraea (Swartz) 1806. Based on Andropogon provinciale Retz. = Desfontaines. ?. The letter name unidentified, though descrip- Chloris piperita Michaux ex Steudel, Nom. tion is not of a Chloris. Bot. ed. 2. 1:353. 1840. Nomen ntidum, as

Chloris pallida ( Edgeworth ) Hooker, f.. synonym of Campulosus rnonostachytis Beauvois. Flora British India 7:289. 1896. Non Chloris Chloris poaeformis Humboldt, Bonpland, 130 Brigham Young University Science Bijlletin

Kunth, Nov. Gen. and Sp. PL 1:169. 1816. Chloris retusa Lagasca, Elenchus Plantarum = LeptochIoa sp. on basis of description. 5. 1816. = Eustachys retusa (Lagasca) Kunth Chloris poltjdacttjla Swartz var. hreviaristata on basis of fragment and photograph of type Hackel in Kurtz, Bol. Acad. Ciencias, Cordoba in US! (Rep. Argentina) 16:257. 1899. Unidentified. Chloris rhachitricha Steudel, Syn. PI. Glum. Type specimen not seen, but description sug- 1:205. 1854. =Eulalia ? on basis of type speci- gests Chloris ciliata Swartz. men in P!

Chloris pohjdactijla Swartz f. major Kegel, Chloris ridleyi Hackel, Oesterr. Bot. Zeitschr. Annotationes Bot. Index Seminum 25. 1862. 52:237. 1902. = Eustachys tener (Presl) A. Unidentified. Type not seen. Description incon- Camus. Isotype at K! clusive. CIdoris roxhurghii Edgeworth, Jour. Asiatic

Chloris polydactijla Swartz f. media Kegel, Soc. Bengal 21:183. 1851. Based on Melica Annotationes Bot. Index Seminum. 25. 1862. digitata Roxburgh which = Enteropogon doli- Nomen nudum. chostachi/a (Lagasca) Lazarides.

Chloris pohjdactijla Swartz f. minor Kegel, Chloris rufescens Lagasca, Var. Ciencias Lit. Annotationes Bot. Index Seminum 25. 1862. and Artes (Madrid) 4:143. 1805. Type not Based on Chloris elegans Humboldt, Bonpland, located. Original description too meager for Kunth, which =Chloris vir^ata Swartz. Prob- recognition of plant. ably misapplied to Chloris pohjdactijla Swartz. Chloris savatieri Baillon, Bull. Soc. Linn. Chloris polijstachia Lagasca, Elenchus Plan- (Paris) 2:1070. 1893. Nomen nudum, given as tarum 5. 1816. Nomen nudum. an herbarium name in a discussion of Tetra- Chloris procumhens Durand, Chlorides Spe- pogon sp. ciebus 16. 1808. =Bouteloua sp. on basis of Chloris scariosa Beauvois, Ess. Agrost. 79, description. 158. 1812. Non Chloris scariosa F. Mueller.

Chloris prostrata ( Willdenow ) Poiret in Nomen nudum. Lamarck, Encycl. Method. Bot. Suppl. 2:239. Chloris secundus (Pursh) A. Eaton, Manual 1811. Based on Dacttjloctenium prostratum Bot. North Amer. ed. 5. 173. 1829. Based on Willdenow. = Dacttjloctenium sp. Ctpwsurus secundus Pursh. =Bouteloua sp. on Chloris pubescens Peyritsch in Wawra and basis of description. Peyritsch, Akad. Wiss. Wien Sitzungsber. Math. Clitoris septentrionalis C. Mueller, Bot. -Naturwiss. Klasse 38:385. 1860. Non C. Zeitung (BerHn) 19:340. 1861. =Eu.stachys pubescens Lagasca, 1805. —Eustachys sp. on petraea (Swartz) Desveaux on basis of descrip- basis of description. tion. Chloris pulchra Schumacher, Beskr. Guinei- Chloris simplex Schumacher, Beskr. Guinei- ske Planter 76. 1827. =Ctenium sp. Photo- ske Planter 74. 1827. = Enteropogon sp. on graph of type, in US!, from Isert and Thonning basis of photograph (in K! ) of type speci- Herbarium. men in Isert and Thonning Herbarium. Chloris queenslandiac Domin, Biblioth. Bot. Chloris spathacea Hochstetter ex Steudel, 85:368. 1915. Nomen nudum, as synonym of Syn. PI. Glum. 1:204. 1854. =Tetrapogon Chloris acicularis var. queenslandiac Domin. cenchriformis on basis of isotype at K! Chloris radiata Heyne ex Both, Nov. PI. Chloris spathacea Baillon, Bull. Soc. Linn. Indiae Orientalis 61. 1821. Non Chloris radiata (Paris) 2:1070. 1893. Non Chloris spathacea Swartz. Nomen nudum, as synonym of Chloris Hochstetter, 1854. =Tetrapogon sp. on basis incompleta Roth, which = Enteropogon dolicho- of tvpe in P! stachtja (Lagasca) Lazarides. Chloris subaequigluma Rendle, Cat. African Chloris radiata Swartz var. beijrichiana Plants collected by Dr. F. Welwitsch 2:222. (Hackel) Kurtz in Rio and Achaval, Geog. Prov. 1899. Basis of Bracteola .subaequigluma Cordoba 1:331. 1904. Attributed incorrectly to (Rendle) C. E. Hubbard. Hackel by Kurtz, presumably based on Chloris Chloris swartziana Doell in Martins, Flora betjrichiana Kunth, but basionym not cited. Brasiliensis 23:68. 1878. —Eustachys petraea Chloris ramosissima A. Camus, Not. Syst. (Swaitz) Desveaux, on basis of description. Mus. Hist. Nat. Paris 12:155. 1946. =Dakno- Chloris swartzii C. Mueller, Bot. Zeitung pholis boivinii ( Camus ) Clayton on basis of (Berlin) 19:341. 1861. = Eustachys petraea holotype in P! (Swartz) Desveaux. Based on Chloris petraea Chloris repens Steudel, Nom. Bot. ed. 2. Swartz. 1:353. 1840. Nomen nudum, as synonym of Chloris tenella Koen ex Roxburgh. Hortus Eleusine indica (Linnaeus) Gaertner. Bengalensis 82. 1814. Nomen nudum. Chloris )

Biological Series, Vol. 19, No. 2 T.\xonomy of the Genus Chloris (Gramineae) 131 tenella Roxburgh, Flora Indica ed. Carey, 1:330. (Hackel) Herter. 1820. =Tetrapogon sp. on basis of description. Chloris unispicea F. Mueller, Fragm. Chloris tenera (Prcsl) Scribncr, Ann. Rcpt. Phytogr. Australiae 7:118. 1870. Basis of En-

Missouri Bot. Card. 10:41. /)/. 40. 1899. teropogon unispicea (Mueller) Clayton.

= Eustachys tener (Prcsl) A. Camus. Chloris villosa ( Desfontaines ) Persoon, Syn. Chloris temiior Poiret ex Schultes, Mantissa PI. 1:87. 1805. Based on Tetrapogon villosits 2:321. 1824. Nomen nii(him, as svnon\'ni of Desfontaines. = Tetrapogon sp. Leptcliloa cynosuroicles Rocmcr and Schultes. Chloris villosa var. sinaica Decaisne, Ann. Chloris tetrameris Trinius, Gram. L'nitlorcs Sci. Nat. Bot. (Paris) II. 2:12. 1834. = Tetra- 235. 1824. =Enterof)ogon. doliclwstadnju pogon sp. on basis of description. (Lagasca) Lazarides on basis of description. Chloris virgattts Nees ex Steudel, Syn. PI. Chloris transiens Pilger, Bot. Jahrb. 51:418. Glum. 1:213. 1854. Nan Chloris virgata 1914. Basis of Schocnefehlia transiens (Pilger) Swartz. Unidentified. Description ma\- be of Chiovenda. Leptochloa sp. Chloris triangiihita Hochstctter ex Achille Leptochloris greggii Munro ex Merrill, U.S. Richard, Tent. Florae Ab\ssinicae 2:409. 1850. Dept. Agric. Div. Agrost. Circ. 32:7. 1901. Not = Tetrapogon sp. on basis of description. located. Chloris uliginosa Hackel, Repert. Sp. Nov. Trichloris latifolia Vasey, U.S. Dept. Agric. 7:320. 1909. Basis of Eustachijs nligiiwsus Spec. Rept. 63:32. 1883. Not located.

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