classification and biogeography of panicoideae (poaceae) in the

Aliso: A Journal of Systematic and Evolutionary Botany

Volume 23 | Issue 1 Article 39

2007

Classification and Biogeography of Panicoideae(Poaceae) in the New WorldFernando O. ZuloagaInstituto de Botánica Darwinion, San Isidro, Argentina

Osvaldo MorroneInstituto de Botánica Darwinion, San Isidro, Argentina

Gerrit DavidseMissouri Botanical Garden, St. Louis

Susan J. PenningtonNational Museum of Natural History, Smithsonian Institution, Washington, D.C.

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Recommended CitationZuloaga, Fernando O.; Morrone, Osvaldo; Davidse, Gerrit; and Pennington, Susan J. (2007) "Classification and Biogeography ofPanicoideae (Poaceae) in the New World," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 23: Iss. 1, Article 39.Available at: http://scholarship.claremont.edu/aliso/vol23/iss1/39

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Aliso 23, pp. 503–529! 2007, Rancho Santa Ana Botanic Garden

CLASSIFICATION AND BIOGEOGRAPHY OF PANICOIDEAE (POACEAE) IN THE NEW WORLD

FERNANDO O. ZULOAGA,1,5 OSVALDO MORRONE,1,2 GERRIT DAVIDSE,3 AND SUSAN J. PENNINGTON4

1Instituto de Botanica Darwinion, Casilla de Correo 22, Labarden 200, San Isidro, B1642HYD, Argentina;2([email protected]); 3Missouri Botanical Garden, PO Box 299, St. Louis, Missouri 63166, USA

([email protected]); 4Department of Botany, National Museum of Natural History, Smithsonian Institution,Washington, D.C. 20013-7012, USA ([email protected])

5Corresponding author ([email protected])

ABSTRACT

Panicoideae (Poaceae) in the New World comprise 107 genera (86 native) and 1357 species (1248native). As circumscribed herein, Panicoideae include eight tribes: Andropogoneae, Arundinelleae,Centotheceae, Gynerieae, Isachneae, Paniceae, Steyermarkochloeae, and Thysanolaeneae. The twomajor tribes are Andropogoneae with 230 species (16.95% of all New World panicoids), and Paniceaewith 1082 species (79.73%). Andropogoneae are divided into nine subtribes (Andropogoninae, An-thistiriinae, Coicinae, Germainiinae, Ischaeminae, Rottboelliinae, Saccharinae, Sorghinae, and Trip-sacinae), while Paniceae are divided into seven subtribes (Arthropogoninae, Cenchrinae, Digitariinae,Melinidinae, Panicinae, Paspalinae, and Setariinae). Brazil is the center of diversity of New Worldpanicoids with 741 species (54.6% of all species). The distributions of endemic and restricted taxawere analyzed in terms of biogeographical regions and Kranz syndrome. Amphitropical disjuncts andwidespread species are also discussed.

Key words: Andropogoneae, biogeography, classification, endemism, Paniceae, Panicoideae, Poaceae,taxonomy.

INTRODUCTION

Subfamily Panicoideae (Poaceae) includes ca. 3270 spe-cies grouped in 206 genera (Grass Phylogeny WorkingGroup [GPWG] 2001). The subfamily is distributed world-wide, and is present on all continents except Antarctica.However, its members are dominant in tropical and subtrop-ical areas, but are less conspicuous in temperate or cold-temperate habitats.

The subfamily was described by Link (1827) and has beenrecognized as a monophyletic group because of its pairedflorets, the lower of which is staminate or sterile (Brown1810, 1814), and its distinctive simple starch grains (Tateoka1962; Kellogg and Campbell 1987; GPWG 2001).

In the past century several tribes were included in Pani-coideae on the basis of morphological characters. Pilger(1940) distinguished three tribes: Paniceae, with six sub-tribes (Anthephorinae; Boivinellinae; Lecomtellinae; Melin-idinae; Panicinae, which included Coelachne R. Br., Heter-anthoecia Stapf, and Isachne; and Trachyinae); Andropo-goneae, with six subtribes (Andropogoninae; Dimeriinae; Is-chaeminae; Rottboelliinae; Saccharinae; and Sorghinae); andMaydeae. Later, Pilger (1954) raised some of the subtribesto the tribal level, recognizing, along with Paniceae: Anthe-phoreae, with a single genus, Anthephora; Arthropogoneae,with Achlaena, Arthropogon, Reynaudia, and Snowdenia C.E. Hubb.; Boivinelleae, with Boivinella A. Camus, Cypho-chlaena Hack., and Perulifera A. Camus; Isachneae, withCoelachne, Heteranthoecia, Isachne, and Limnopoa C. E.Hubb.; Lecomtelleae, with Lecomtella A. Camus; Melini-deae, with Melinis, Rhynchelytrum Nees, and TricholaenaSchult.; and Trachyeae, with Trachys Pers. Pilger (1954) alsorecognized tribes Andropogoneae and Maydeae within sub-family Andropogonoideae.

Clayton and Renvoize (1986) followed Pilger (1940,1954) in part, recognizing seven tribes in Panicoideae. Thetwo largest tribes were Paniceae (with six subtribes, includ-ing Panicinae [! Setariinae]) and Andropogoneae (with 11subtribes), and the remaining tribes were small: Arundinel-leae, Eriachneae, Hubbardieae, Isachneae, and Steyermark-ochloeae. Clayton and Renvoize (1986) recognized Cento-thecoideae as an independent subfamily.

Watson and Dallwitz (1992) accepted six tribes in the sub-family: Andropogoneae, Arundinelleae, Isachneae, May-deae, Neurachneae, and Paniceae. These authors placed Cen-totheceae in Bambusoideae, Steyermarkochloeae and Eri-achneae in Arundinoideae, and Hubbardieae in Isachneae.

Six tribes were recognized by the GPWG (2001): Andro-pogoneae, Arundinelleae, Hubbardieae, Isachneae, Paniceae,and Steyermarkochloeae. They summarized the most impor-tant characters for the subfamily, including the presence ofproximal female-sterile florets and the transformation to theclassical NADP-ME C4 subtype. Other characters include thepresence of one female-fertile floret per spikelet, the gain ofa germination flap, and the loss of disarticulation above theglumes, a synapomorphy for Panicoideae excluding Dan-thoniopsis Stapf. The subfamily was characterized by theGPWG (2001) as follows: plants primarily herbaceous,mainly occurring in the tropics and subtropics, but also di-verse in temperate areas; spikelets bisexual or unisexual (ifthe latter, plants dioecious or monoecious), frequently paired,usually dorsally compressed and disarticulating below theglumes; lodicules two; stigmas two; caryopsis with the hilumusually short; endosperm hard, containing simple or lesscommonly compound starch grains; embryo usually withoutan epiblast, with a scutellar cleft present and with the me-socotyl internode elongated. The basic chromosome number

504 ALISOZuloaga, Morrone, Davidse, and Pennington

varies: x ! 5–9, 10 (12, 14). The photosynthetic pathwayvaries from C3 to C4, the latter including PEP-CK, NAD-ME and NADP-ME subtypes; there are also some C3/C4 in-termediates in Steinchisma. The Kranz syndrome is constantin some tribes, such as Andropogoneae, but is variable inothers, e.g., Arundinelleae and Paniceae (Soreng and Davis1998), which suggests that this character has evolved morethan once in Panicoideae (Clayton and Renvoize 1986; Gius-sani et al. 2001).

The monophyly of the subfamily has been supported byevery molecular phylogenetic study to date, including phy-logenies of both chloroplast and nuclear genes (Soreng andDavis 1998; Giussani et al. 2001; GPWG 2001). These re-sults suggest that the subfamily should include, within thePACCAD clade, all genera previously included within Cen-tothecoideae, and Gynerium (formerly in Arundinoideae).Soreng and Davis (1998) suggested that Centotheceae shouldbe assigned to Panicoideae, along with isolated genera pre-viously classified in Arundinoideae. These authors indicatedthat their conclusions are supported by previous molecularstudies (Barker et al. 1995, using rbcL sequences; Clark etal. 1995, using ndhF sequences), in which Centotheceae andThysanolaena (Arundinoideae) are embedded within Pani-coideae.

Duvall et al. (2007) have proposed that Isachne and Eri-achne R. Br. should be excluded from Panicoideae, becausethey are closely related to Micraira F. Muell. Also, Sanchez-Ken and Clark (2007) have provided new insights into re-lationships in the Panicoideae–Centothecoideae clade, usingthree data sets that also showed Gynerium as sister to Pan-icoideae.

The purpose of this work is to present a tentative classi-fication for Panicoideae, on the basis of New World taxa, todiscuss morphological characters, and to present biogeo-graphical trends for genera and species within the subfamily.

METHODS

As part of the ongoing Catalogue of New World Grassesproject (Judziewicz et al. 2000; Soreng et al. 2000 onwards;Peterson et al. 2001; Soreng et al. 2003; Zuloaga et al. 2003),the geographic distributions of all accepted taxa (native andintroduced) were recorded at the level of country with theCaribbean nations treated as a single unit. This paper rep-resents a summary of distribution patterns of the native spe-cies. All of the original data are available online at http://mobot.mobot.org/W3T/Search/nwgc.html (Jun 2005).

To analyze the distributions of endemic taxa, we followedthe classification of floristic regions of the world proposedby Takhtajan (1986).

CLASSIFICATION

In our classification of Panicoideae in the New World (Ta-ble 1; http://mobot.mobot.org/W3T/Search/nwgclass.html)we recognize eight tribes (Andropogoneae, Arundinelleae,Centotheceae, Gynerieae, Isachneae, Paniceae, Steyermark-ochloeae, and Thysanolaeneae), with nine subtribes in An-dropogoneae (Andropogoninae, Anthistiriinae, Coicinae,Germainiinae, Ischaeminae, Rottboelliinae, Saccharinae,Sorghinae, and Tripsacinae) and seven subtribes in Paniceae(Arthropogoninae, Cenchrinae, Digitariinae, Melinidinae,

Panicinae, Paspalinae, and Setariinae). This realignment isbased on previous morphological and molecular work (Clay-ton and Renvoize 1986; Watson and Dallwitz 1992; Gomez-Martinez and Culham 2000; Zuloaga et al. 2000; Giussaniet al. 2001; GPWG 2001; Sanchez-Ken and Clark 2001; Bar-ber et al. 2002; Aliscioni et al. 2003).

New World Panicoideae include a total of 107 genera and1357 species (with 21 genera and 109 species introduced)distributed in eight tribes, six (Arundinelleae, Centotheceae,Gynerieae, Isachneae, Steyermarkochloeae, and Thysanola-eneae) comprising a small number of genera and species,while Andropogoneae and Paniceae hold the majority of na-tive and introduced genera and species (Table 2).

Axonopus, Digitaria, Panicum, Paspalum, and Setaria arethe five largest genera of Panicoideae in the New World(Table 3), and together they account for 51.44% of all nativeNew World panicoids.

Andropogoneae

Recent phylogenetic studies have questioned the mono-phyly and subtribal classification of Andropogoneae (Ma-son-Gamer et al. 1998; Spangler et al. 1999; Kellogg 2000).Kellogg (2000) indicated that none of the accepted subtribesare monophyletic, and that further studies are necessary toobtain a more confident understanding of the phylogeny ofthe group. For current classification purposes, and acknowl-edging the necessity of future studies in the group, we followFilgueiras (2003), who adopted the classification of Claytonand Renvoize (1986) for the Catalogue of New WorldGrasses.

The tribe, the second largest in Panicoideae, includes taxawith spikelets paired, usually with one spikelet sessile andthe other pedicelled, alike or dissimilar, the sessile spikeletbeing bisexual and the pedicelled spikelet male or barren.The basic chromosome number is commonly x ! 5, occa-sionally 9 (Clayton and Renvoize 1986), and all taxa are C4,XyMS". Clayton and Renvoize (1986) included 85 generaand ca. 960 species in the tribe, which has a broad distri-bution throughout the tropics. Andropogoneae include 35genera and 230 species in the New World, grouped in ninesubtribes. Subtribes Andropogoninae (4 gen./81 spp.),Sorghinae (6/54), and Saccharinae (7/30) have the greatestnumber of genera and species in the tribe (Table 2). Endemicgenera are present in Anthistiriinae and Tripsacinae (Table 1).

Andropogon, Bothriochloa, and Schizachyrium have 49,25, and 23 species, respectively, while the remaining generaeach have 1–14 species. It is notable that 15 genera are in-troduced: Apluda, Arthraxon, Chrysopogon, Coix, Cymbo-pogon, Dichanthium, Eremochloa, Euclasta, Hyperthelia,Microstegium, Miscanthus, Pogonatherum, Polytrias, Rott-boellia, and Themeda. This accounts for the majority of in-troduced genera in Panicoideae. Three genera are endemic:Agenium (South America), Tripsacum (North America, Me-soamerica, and northern South America), and Zea (Mexicoand Central America).

Arundinelleae

Arundinelleae have a worldwide distribution, but mostmembers occur in the Old World tropics. The tribe includesfour genera in America usually found in savanna woodlands

VOLUME 23 505New World Panicoideae

Table 1. Classification of New World Panicoideae. Numbers in parentheses are the number of accepted species in the New World.* ! all species introduced. ** ! endemic.

Tribe AndropogoneaeSubtribe Andropogoninae

Andropogon L. (49)Arthraxon P. Beauv. (2)*Cymbopogon Spreng. (7)*

Loudetiopsis Conert (1)Tristachya Nees (6)

Tribe CentotheceaeCalderonella Soderstr. & H. F. Decker

(1)**

Sacciolepis Nash (6)Subtribe Paspalinae

Acostia Swallen (1)**Alloteropsis J. Presl (1)*Amphicarpum Kunth (2)**

Schizachyrium Nees (23)Subtribe Anthistiriinae

Agenium Nees (3)**Elymandra Stapf (1)Heteropogon Pers. (2)

Chasmanthium Link (5)**Orthoclada P. Beauv. (1)Pohlidium Davidse, Soderstr. & R. P.

Ellis (1)**Zeugites P. Browne (10)**

Anthaenantia P. Beauv. (4)**Anthaenantiopsis Mez ex Pilg. (4)**Axonopus P. Beauv. (83)Canastra Morrone, Zuloaga, Davidse &

Filg. (1)**Hyparrhenia Andersson ex E. Fourn. (4)Hyperthelia Clayton (1)*Themeda Forssk. (2)*

Subtribe CoicinaeCoix L. (1)*

Tribe GynerieaeGynerium Willd. ex P. Beauv. (1)**

Tribe IsachneaeIsachne R. Br. (12)

Tribe Paniceae

Centrochloa Swallen (1)**Echinolaena Desv. (6)**Gerritea Zuloaga, Morrone & Killeen

(1)**Homolepis Chase (5)**

Subtribe GermainiinaeTrachypogon Nees (3)

Subtribe IschaeminaeApluda L. (1)*Ischaemum L. (6)

Subtribe ArthropogoninaeAltoparadisium Filg., Davidse, Zuloaga

& Morrone (2)**Achlaena Griseb. (1)**Arthropogon Nees (3)**

Hymenachne P. Beauv. (4)Ichnanthus P. Beauv. (31)Mesosetum Steud. (26)**Ophiochloa Filg., Davidse & Zuloaga

(1)**Subtribe Rottboelliinae

Elionurus Humb. & Bonpl. ex Willd. (4)Eremochloa Buse (1)*Hemarthria R. Br. (1)Mnesithea Kunth (12)

Reynaudia Kunth (1)**Subtribe Cenchrinae

Anthephora Schreb. (1)Cenchrus L. (15)Paratheria Griseb. (1)

Oplismenopsis Parodi (1)**Otachyrium Nees (8)**Panicum incertae sedis (64)Paspalum L. (285)Phanopyrum (Raf.) Nash (1)**

Rhytachne Desv. ex Ham. (3)Rottboellia L. f. (1)*

Subtribe SaccharinaeEriochrysis P. Beauv. (5)Imperata Cirillo (8)

Pennisetum Rich. (36)Subtribe Digitariinae

Digitaria Haller (89)Subtribe Melinidinae

Chaetium Nees (3)**

Plagiantha Renvoize (1)**Reimarochloa Hitchc. (3)**Spheneria Kuhlm. (1)**Steinchisma Raf. (7)**Streptostachys Desv. (4)**

Microstegium Nees (1)*Miscanthus Andersson (1)*Pogonatherum P. Beauv. (1)*Polytrias Hack. (1)*Saccharum L. (13)

Eriochloa Kunth (25)Megathyrsus (Pilg.) B. K. Simon &

S. W. L. Jacobs (1)*Melinis P. Beauv. (2)*Moorochloa Veldkamp (1)*

Tatianyx Zuloaga & Soderstr. (1)**Thrasya Kunth (22)**Thrasyopsis Parodi (2)**Triscenia Griseb. (1)**

Subtribe SetariinaeSubtribe Sorghinae

Bothriochloa Kuntze (25)Chrysopogon Trin. (3)*Dichanthium Willemet (4)*Euclasta Franch. (1)*

Scutachne Hitchc. & Chase (2)**Urochloa P. Beauv. (35)

Subtribe PanicinaeAcroceras Stapf (5)Cyrtococcum Stapf (1)*

Ixophorus Schltdl. (1)**Panicum incertae sedis (3)Paspalidium Stapf (6)Setaria P. Beauv. (60)Setariopsis Scribn. (2)**

Sorghastrum Nash (14)Sorghum Moench (7)

Subtribe TripsacinaeTripsacum L. (14)**Zea L. (5)**

Dichanthelium (Hitchc. & Chase)Gould (58)

Echinochloa P. Beauv. (20)Lasiacis (Griseb.) Hitchc. (15)**Oplismenus P. Beauv. (3)

Stenotaphrum Trin. (1)Tribe Steyermarkochloeae

Arundoclaytonia Davidse & R. P.Ellis (1)**

Steyermarkochloa Davidse & R. P.Tribe Arundinelleae

Arundinella Raddi (3)Loudetia Hochst. ex Steud. (1)

Panicum incertae sedis (50)Panicum L. (60)Pseudechinolaena Stapf (1)

Ellis (1)**Tribe Thysanolaeneae

Thysanolaena Nees (1)*

or flood plains: Arundinella, with three species from Mexicoand the Caribbean to Argentina; Tristachya, with six speciesfrom Mexico to southern Brazil and Paraguay; and two gen-era, Loudetia and Loudetiopsis, represented by single speciesin cerrados and open grasslands of Bolivia, Brazil, and Par-aguay. The tribe is characterized by spikelets often in triads,slightly laterally compressed, and the upper lemma awned,the awn often deciduous. The four New World genera havea basic chromosome number of x ! 6, 7, 10, 12, and 14,and all species are C4, XyMS".

Arundinelleae were placed in Panicoideae by several au-

thors, including Conert (1957), Clayton (1971), Renvoize(1982), Clayton and Renvoize (1986), and Watson and Dall-witz (1992). Clayton and Renvoize (1986) and Kellogg andWatson (1993) considered Arundinelleae to be phylogenet-ically basal to Andropogoneae. However, recent molecularevidence (Mason-Gamer et al. 1998; Spangler et al. 1999;Kellogg 2000) indicates the tribe is polyphyletic, and thatArundinella should be included in Andropogoneae. Also,Barker et al.’s (1995) rbcL data placed Tristachya sister toHyparrhenia within the Andropogoneae clade. In the presenttreatment Arundinelleae are still treated as a separate tribe


Table 2. Numerical summary of New World Panicoideae.

Tribe/subtribe GeneraNativespecies

Introducedspecies

Totalspecies

% of allspecies

AndropogoneaeAndropogoninaeAnthistiriinaeCoicinaeGerminiinae

354611

18370

803

4711

510

2308113

13

16.955.961.960.070.22

IschaeminaeRottboelliinaeSaccharinaeSorghinaeTripsacinae

26762

320213919

429

150

722305419

0.521.622.213.981.40

ArundinelleaeCentotheceaeGynerieaeIsachneaePaniceae

4511

58

1118

112

1021

0000

61

1118

112

1082

0.811.330.070.88

79.73ArthropogoninaeCenchrinaeDigitariinaeMelinidinaePanicinae

44179

7468647

205

073

2214

7538969

219

0.523.906.565.08

16.14PaspalinaeSetariinae

SteyermarkochloeaeThysanolaeneae

28521

5706020

213

01

57273

21

42.155.380.150.07

Total 107 1248 109 1357 100

Table 3. Numbers of species in the five largest genera occurringin the five New World countries most diverse in Panicoideae (seeTable 5).

Genus USA Mexico Venezuela Brazil Bolivia

AxonopusDigitariaPanicum s.l.PaspalumSetaria

329354023

1129677926

2716678316

5343

116167

28

2023367419

TotalTotal native species% of native species

130351

37.04

212464

45.69

209396

52.78

407741

54.93

172359

47.91

in its traditional circumscription until molecular phylogenet-ic studies that include all genera of the tribe can be con-ducted.

Centotheceae

The position of Centotheceae has been unstable in clas-sifications over the past four decades. The tribe was raisedto subfamily level by Soderstrom (1981), a decision fol-lowed by Clayton and Renvoize (1986), emphasizing thecentothecoid embryo and leaf anatomical characters, mainlythe palisade arrangement in the mesophyll. As previouslymentioned, Soderstrom (1981), Watson et al. (1985), andWatson and Dallwitz (1992) included the tribe within Bam-busoideae, while Barker et al. (1995), Clark et al. (1995),Soreng and Davis (1998), Gomez-Martınez and Culham(2000), and Sanchez-Ken and Clark (2003) pointed out,based on molecular characters, the relationship of Centothe-ceae with Panicoideae.

Centotheceae, as treated here, include five genera: Chas-manthium, with five North American species; Orthoclada,with one tropical American and one tropical African species;Zeugites, with ten species distributed from Mexico and theWest Indies to Bolivia; and Calderonella and Pohlidium,both monotypic, the former in Panama and Colombia andthe latter restricted to Panama. Centotheceae are distin-guished by spikelets one- to many-flowered, often com-pressed laterally; embryo with an epiblast, scutellar cleft,and mesocotyl internode present; mesophyll nonradiate, of-ten with an adaxial palisade layer; and a basic chromosomenumber of x ! 12. All species are C3.

Gynerieae

Tribe Gynerieae includes one Neotropical genus, Gyner-ium, known as ‘‘cana brava’’ or ‘‘giant reed,’’ and is distin-guished by distichous leaves, leathery blades with wide mid-ribs, dioecy, and plumose pistillate inflorescences. It has abasic chromosome number of x ! 11 and is C3. The mono-typic genus is distributed from Mexico and the West Indiesto northwestern Argentina, and it is found along river mar-gins or in swampy places. On the basis of morphology thegenus has been placed, together with other reedlike grasses(e.g., Arundo L., Phragmites Adans.), in Arundineae (Ren-voize 1981; Clayton and Renvoize 1986; Conert 1987; Ni-cora and Rugolo de Agrasar 1987). Hitchcock (1914) placedthe genus in Festuceae, Caro (1982) in Cortaderieae, andWatson and Dallwitz (1992) in Danthonieae. More recently,based on molecular studies, Hsiao et al. (1998, 1999) clas-sified Gynerium in Arundinoideae. On the contrary, Barker(1997), based on rbcL sequences, assigned the genus to thePanicoideae " Centothecoideae clade, as sister to Panicoi-deae or sister to Thysanolaena within Centothecoideae. TheGPWG (2001), in their combined study of morphologicaland molecular data, confirmed that Gynerium is locatedwithin the Panicoideae " Centothecoideae clade. Finally,Sanchez-Ken and Clark (2001) proposed the new tribe Gy-nerieae in the Panicoideae " Centothecoideae clade.

Isachneae

Isachneae are represented by a single genus, Isachne, and12 species in America, and there are several genera and near-ly 110 species in the Old World tropics. The tribe has longbeen considered part of Panicoideae (Brown 1810; Hackel1887; Pilger 1954; Clayton and Renvoize 1986; Watson andDallwitz 1992; GPWG 2001). It is characterized by biflow-ered spikelets, the lower floret bisexual and the upper floretbisexual or pistillate. Species of the tribe occur only in thetropics in forest interiors, forest edges, or open aquatic hab-itats. All species are nonKranz and have a basic chromosomenumber of x ! 10.

Recent molecular studies (Duvall et al. 2007; Sanchez-Ken and Clark 2007) suggest that Isachne is closely relatedto the Australian genus Micraira, and that both genera, plusothers, should be reassigned to a new subfamily in the PAC-CAD clade.


Paniceae

Classification of tribe Paniceae, the largest in Panicoideae,has been extensively discussed. Many classifications havebeen proposed. Pilger (1940), Hsu (1965), Butzin (1970),Brown (1977), and Clayton and Renvoize (1986) used dif-ferent morphological characters (e.g., spikelets arranged inopen and lax panicles, the spikelets borne on long or shortpedicels, vs. disposed in unilateral inflorescences branches),together with anatomical and karyological characters, fortheir classifications. Spikelet compression is usually dorsi-ventral, while the upper floret is indurate. Two main anatom-ical and physiological types are found, nonKranz and Kranz.The most common basic chromosome numbers are x ! 9and 10.

However, recent morphological and molecular studieshave questioned the monophyly of the tribe and also its sub-tribal classification; consequently we are still searching fora natural grouping of Paniceae (Gomez-Martınez and Cul-ham 2000; Zuloaga et al. 2000; Duvall et al. 2001; Giussaniet al. 2001; Aliscioni et al. 2003). Molecular studies (Go-mez-Martınez and Culham 2000; Giussani et al. 2001; Al-iscioni et al. 2003) have shown that the genera are initiallygrouped according to basic chromosome number, one cladex ! 10 and the other x ! 9, the former linked to Andropo-goneae. A morphological analysis (Zuloaga et al. 2000), inwhich neither Andropogoneae nor the basic chromosomenumber as a character were included, gave similar results forseveral groupings of genera in the tribe. These results sug-gest that Paniceae could even be split into two tribes.

Also, Panicum, one of the most controversial genera inthe tribe, has been scrutinized (Giussani et al. 2001; Alis-cioni et al. 2003), showing that the genus is not monophy-letic and recircumscription is needed. As a result, we treatPanicum in a strict sense, with species having open inflo-rescences, a basic chromosome number of x ! 9, C4 pho-tosynthesis of the NAD-ME subtype, and an indurated upperanthecium. Some of the other taxa previously positioned inPanicum have been placed in other genera, while other Pan-icum species or sections are in need of further study beforenomenclatural changes can be entertained. The latter areidentified as Panicum incertae sedis in Table 1. Several sub-tribes are here recognized. All genera of the x ! 10 cladeare grouped in subtribes Arthropogoninae and Paspalinae.The other subtribes of Paniceae are within the x ! 9 clade(Giussani et al. 2001) and include Cenchrinae, Digitariinae,Melinidinae, Panicinae, and Setariinae.

Arthropogoninae.—This subtribe is represented by four gen-era, each with one or a few species: Achlaena, Altoparadi-sium, Arthropogon, and Reynaudia. Achlaena and Reynaudiaare monotypic and restricted to Cuba. Altoparadisium andArthropogon have two and three species, respectively, in cer-rados of Bolivia and Brazil. The subtribe is characterized byspikelets laterally or dorsiventrally compressed, glumesawned, upper anthecium hyaline, and the lower palea re-duced or absent. There is a trend towards reduction of thenumber of spikelet bracts, from species of Arthropogon thatpossess both glumes and both florets to the extreme condi-tion in Altoparadisium where only the upper glume and low-er lemma may be present (A. chapadense). All species haveKranz anatomy of the MS, NADP-ME subtype. Based on

morphological characters, genera of the subtribe form aclade, although one species of Melinis is also included (Fil-gueiras et al. 2001). Recent molecular analyses (Giussani etal. 2001) grouped Altoparadisium and Arthropogon in astrongly supported clade within the x ! 10 clade, while Mel-inis appeared related to Moorochloa in the x ! 9 clade.

Cenchrinae.—Giussani et al. (2001), in a phylogenetic studyof Panicoideae based on the ndhF gene, found that Cenchrus,Paspalidium, Pennisetum, Setaria, and Stenotaphrum form arobust clade. These genera are characterized by the occur-rence of bristles (sometimes highly modified, as in Cen-chrus) in the inflorescences. The bristles may have very dif-ferent positions in the inflorescence. In some cases they arein a terminal position on the inflorescence or inflorescencebranches or they subtend individual spikelets. Clayton andRenvoize (1986) stated that bristles are not homologouswithin members of this group. However, molecular studies(Gomez-Martinez and Culham 2000; Giussani et al. 2001)suggest that the bristles have a common evolutionary his-tory. Also, morphological studies (Zuloaga et al. 2000) in-dicate that Cenchrus and related genera constitute a clade,and they share the presence of a point of articulation at thebase of first-order branches. Recently, the developmentalstudies of Doust and Kellogg (2002) provided strong evi-dence for a ‘‘bristle clade’’ and showed that small changesin the differentiation of primordial and axis ramification andelongation are responsible for most of the large differencesseen in mature inflorescences. Consequently, although wehere follow the circumscriptions of Cenchrinae and Setari-inae of Clayton and Renvoize (1986), it now seems verylikely that these subtribes should be merged into a singlesubtribe (Kellogg et al. 2003).

Cenchrinae include four genera, none endemic, in theAmericas. Three genera have an involucre of bristles fallingwith each spikelet: Anthephora, with one species; Cenchrus,with 15 species; and Pennisetum with 36 species, some in-troduced. The monotypic Paratheria has a single bristle fall-ing with each spikelet. All genera are Kranz, of the MSsubtype (! NADP-ME).

Digitariinae.—Digitariinae contain a single genus, Digitaria,with 89 species (three introduced from the Old World) thatoccur in tropical and warm-temperate regions from the USAto Argentina and Chile. Common in open places, roadsides,and streamsides, several species are weeds and others arecultivated in pastures. The genus is distinguished by spike-lets with a reduced lower glume, upper anthecium cartilag-inous, and upper lemma margin thin and not inrolled. Allspecies are C4, of the NADP-ME subtype.

Melinidinae.—All genera in this subtribe are Kranz, of thePEP-CK subtype. Most of the genera have an indurate andtransversely rugose upper anthecium, and spikelets arrangedin unilateral branches and with short pedicels. Melinidinaeinclude seven genera in America: Eriochloa (25 spp.) andUrochloa (35 spp.) mostly occur in tropical and subtropicalregions; Brachiaria (Trin.) Griseb. (! Moorochloa) (1 sp.),Megathyrsus (1 sp.), and Melinis (2 spp.) are introduced; andChaetium and Scutachne are endemic. Chaetium has threespecies ranging from Mesoamerica and Cuba to SouthAmerica, while Scutachne has two species in Cuba.


Panicinae.—This subtribe is characterized by inflorescenceslax and open, spikelets with or without a lower flower, thelower palea always developed, and the upper anthecium in-durate. Widespread genera (number of New World speciesin parentheses) within the subtribe are: Acroceras (5), Di-chanthelium (58), Echinochloa (20), Oplismenus (3), Pani-cum s.s. (60), Pseudechinolaena (1), and Sacciolepis (6).Lasiacis is an endemic genus with 15 species, while themonotypic Cyrtococcum is introduced. There are also sev-eral incertae sedis sections of Panicum, and two ungroupedincertae sedis species of the genus, in Panicinae, includingsect. Monticolae Stapf with three widespread species; sect.Parvifolia Hitchc. & Chase ex Pilg. with ca. 32 species fromMesoamerica to Argentina; sect. Verrucosa Hitchc. & Chaseex C. C. Hsu with two North American species; and sects.Cordovensia Parodi and Parviglumia Hitchc. & Chase exPilg., occurring from Mexico to Argentina, with five and sixspecies, respectively. Section Cordovensia, previously in-cluded in Dichanthelium, is actually related to Acroceras,Lasiacis, and Pseudechinolaena (Aliscioni et al. 2003). Twophotosynthetic pathways are present in the subtribe: non-Kranz (in Acroceras, Cyrtococcum, Dichanthelium, Lasiacis,Oplismenus, Pseudechinolaena, and incertae sedis species ofPanicum) and Kranz (Echinochloa, of the NADP-ME sub-type, and Panicum s.s., of the NAD-ME subtype).

Paspalinae.—As in Arthropogoninae, in this subtribe thereis also a trend towards reduction of the number of bracts inthe spikelet, with some genera (e.g., Axonopus and Paspa-lum) having species whose spikelets have only a lower lem-ma and an upper floret. Also, taxa of Paspalinae have spike-lets usually arranged in unilateral inflorescence branches andare either nonKranz or Kranz of the MS, NAPD-ME sub-type. It is worth emphasizing that 23 genera, of 28 total, areendemic in the New World and only one genus, Alloteropsis,is introduced in the Americas from the Old World. Paspalumand Axonopus are the largest genera, with 285 and 83 totalspecies, respectively, the New World representatives ofwhich are distributed from the USA to Argentina, Uruguay,and Chile; these genera have only a few native species inthe Old World. Mesosetum includes 26 species from Me-soamerica to Brazil and Argentina. Thrasya has 22 species(see discussion of the status of this genus in Denham andZuloaga 2007), and Ichnanthus has 31 species in the NewWorld tropics, with one species, I. pallens (Sw.) Munro exBenth., having a pantropical distribution. A total of 11 en-demic genera are monotypic. Of these, nine occur in SouthAmerica (Acostia, Canastra, Centrochloa, Gerritea,Ophiochloa, Oplismenopsis, Plagiantha, Spheneria, and Ta-tianyx); Triscenia is found in Cuba; and Phanopyrum occursin the southeastern USA. Genera confined to South Americaand having 2–4 species are: Anthaenantiopsis (4 spp.),Streptostachys (4 spp.), and Thrasyopsis (2 spp. restricted tocerrados of central Brazil). All other genera have 2–8 spe-cies: Amphicarpum (2 North American spp.), Anthaenantia(4 spp.), Echinolaena (6 spp.), Homolepis (5 spp.), Hymen-achne (4 spp.), Otachyrium (8 spp., one widespread and allconfined to South America), Reimarochloa (3 spp.), andSteinchisma (7 spp.—two widespread, one in Cuba, one inMexico, and three in South America).

Several sections of Panicum also are included in this sub-

tribe as incertae sedis. Their definite placement at the genericlevel awaits further phylogenetic and morphological study.These sections include sect. Laxa Hitchc. & Chase ex Pilg.,excluding the type species (P. laxum Sw.), with nine Amer-ican species related to Hymenachne, Plagiantha, and Stein-chisma; sect. Lorea Zuloaga, with 27 South American spe-cies found in the Guiana Highlands and in the Brazilian pla-teau; sect. Obtusa Pilg., with a single North American spe-cies related to Paspalum; sect. Prionitia Zuloaga, with oneCuban and another South American species; sect. StoloniferaHitchc. & Chase ex Pilg., with 13 species distributed fromMexico to Argentina and linked to Echinolaena and Ichnan-thus; sect. Tuerckheimiana (Hitchc.) Zuloaga, with a singlespecies in Guatemala and Mexico; as well as sects. Agro-stoidea Hitchc. & Chase ex C. C. Hsu and Tenera (Hitchc.& Chase) Pilg., with two and four species, respectively, dis-tributed from North America and the West Indies to SouthAmerica; and sects. Megista Pilg. and Valida Zuloaga &Morrone, both monotypic, Megista with a widespread spe-cies and Valida with P. validum, confined to southeasternSouth America.

Setariinae.—This subtribe, represented by five genera in theNew World, is characterized by the presence of bristles thatpersist at the spikelet base after the spikelets have fallen.Setaria has nearly 140 species worldwide, mostly in thetropics and subtropics of Africa, Asia, and the New World(60 spp.). Paspalidium, as circumscribed in this treatment,has six species with unilateral, distant branches on the inflo-rescences; Veldkamp (1994) and Webster (1995) transferredspecies of this genus to Setaria. Setariopsis, with two spe-cies, and the monotypic Ixophorus, are endemic to the NewWorld, while Stenotaphrum is pantropical. The subtribe hasKranz anatomy, of the MS (! NADP-ME) subtype.

Steyermarkochloeae

Davidse and Ellis (1984), when describing the new genusSteyermarkochloa, created the new tribe Steyermarkoch-loeae for it, and largely on the basis of anatomical charactersplaced it within subfamily Arundinoideae. They compared itwith Arundo, Gynerium, Phragmites, and Thysanolaena, allgenera with anomalous anatomical characters, but becauseof numerous morphological differences considered the genusto be a peripheral and best placed in its own specializedtribe. Later, Davidse and Ellis (1987) placed the newly de-scribed genus Arundoclaytonia in Steyermarkochloeae, stillincluded in Arundinoideae. Watson and Dallwitz (1992) fol-lowed Davidse and Ellis (1984), but expressed doubt re-garding the position of Steyermarkochloeae, indicating apossible relationship with Bambusoideae. Clayton and Ren-voize (1986) classified Steyermarkochloeae in subfamilyPanicoideae, pointing out ‘‘an obvious resemblance of Stey-ermarkochloa to Hymenachne,’’ which was clearly only avery superficial one. Sanchez-Ken and Clark (2007) haveindicated, based in part on a partial ndhF sequence, that theposition of Arundoclaytonia is unstable in the PACCADclade. We retain, for the time being, Steyermarkochloeaewithin Panicoideae by virtue of their unisexual, dorsallycompressed spikelets that disarticulate below the glumes andfall as a unit. Both genera are nonKranz, while chromosomenumbers are unknown.


Table 4. Number of endemic species and genera per continent/region and percentage of total native Panicoideae.

Species % Genera %

North AmericaMesoamerica ! CaribbeanSouth America

165115577

12.168.50

42.52

36

18

2.85.7

17.0

Steyermarkochloeae include two endemic and monotypicSouth American genera which share similar unisexual spike-lets, multiflowered male spikelets, two stamens per flower,and lack lodicules. Arundoclaytonia is found in white sandsavannas of the Amazon region. The genus is characterizedby its caespitose habit; basal culm internodes numerous, pro-liferating, and lignified; cauline leaves with spiral phyllo-taxy; inflorescences compact, aggregated into a false panicle;spikelets mostly unisexual; male spikelets 3–13-flowered,bearing two stamens per flower; female spikelets three-flow-ered, with only the middle floret fertile, the palea convoluteand many-nerved; lodicules absent; stigmas and stamens ter-minally exserted; and the caryopsis fusiform with an elliptic-punctate hilum. Steyermarkochloa, the other genus, grows inthe Amazonian llanos of Colombia, Venezuela, and adjacentBrazil, in seasonally flooded savannas. The genus has di-morphic culms and leaves, with solid sheaths and cylindricaland flattened blades, a unique foliar morphology withingrasses (Davidse and Ellis 1984). Spikelets are mostly uni-sexual, three-flowered, and the terminal floret is always ru-dimentary. Male and the few bisexual spikelets usually havetwo fully developed florets.

Thysanolaeneae

Tribe Thysanolaeneae has one monotypic, introduced, andornamental Asiatic genus, Thysanolaena, present in the Ca-ribbean, USA, and El Salvador and possessing dimorphicflowers and spikelets with a rachilla extension, the spikeletfalling as a single unit with the attached pedicel. The biflow-ered spikelets falling entire at maturity are characters linkingthis tribe to Panicoideae. The genus has a basic chromosomenumber of x " 11 or 12 (Watson and Dallwitz 1992) and isC3. Hilu and Wright (1982) noted that Thysanolaena hasbeen included in five different tribes, Arundineae, Arundi-nelleae, Paniceae, Thysanolaeneae, and Tristegineae. How-ever, their phenetic analysis provided strong evidence for theindependence of the tribe and they mentioned that it is re-lated to Centothecoideae, a hypothesis confirmed in the sub-sequent molecular studies of Barker et al. (1995), Barker(1997), and Clark et al. (1995). Clayton and Renvoize (1986)also recognized the tribe but placed it in Arundinoideae.

BIOGEOGRAPHY

Subfamily Panicoideae is best represented in tropical andsubtropical regions of the world. Hartley (1958a, b) pointedout that Paniceae and Andropogoneae may have originatedin tropical areas of the Old World, most likely in easternAfrica and Madagascar. Centers of diversity in eastern Af-rica, India/China, and Australia have been discussed by Har-tley (1958a, b) and Cross (1980).

Of the 206 genera and 3270 species included in Panicoi-deae (GPWG 2001), 44 of the 86 native genera and 1248species are restricted to the New World (Tables 1, 2), witha total of 911 species confined to one or two of the subre-gions here considered (Appendix 1). Nearly endemic to theNew World are the very species-rich genera Axonopus (withone Old World vs. 83 New World spp.) and Paspalum (withsix Old World vs. 285 New World spp.). Of the restrictedgenera and near-endemics Axonopus and Paspalum, the fol-lowing are more or less widespread: Anthaenantia, Axono-

pus, Chaetium, Echinolaena, Gynerium, Homolepis, Ixopho-rus, Lasiacis, Mesosetum, Paspalum, Reimarochloa, Setar-iopsis, Steinchisma, Thrasya, Tripsacum, and Zeugites;22.55% of the restricted species are widespread from Northto South America. Nevertheless, many of the genera are notequally distributed throughout the American continents.They tend to be more concentrated in one or more of thebiogeographical areas discussed below.

Distribution in Relation to Continents and Political Units

For this comparison the New World was divided intoNorth America, South America, Mesoamerica, and the Ca-ribbean, the last two categories combined in Table 4. Threegenera are confined to North America: Chasmanthium (Cen-totheceae), with four species in the eastern and southeasternUSA and one species in northern Mexico, and two generaof Paniceae, Amphicarpum and Phanopyrum, with two andone species, respectively, in the southeastern USA. A totalof 165 species of Panicoideae are endemic to North Amer-ica. It is noteworthy that most of the endemic genera ofAndropogoneae and Centotheceae are limited in their distri-bution to North America and Mesoamerica, with only a fewspecies reaching northern South America (e.g., species ofCalderonella, Tripsacum, and Zeugites). Two genera are en-demic in Mesoamerica, Pohlidium (Centotheceae) in Panamaand Zea (Andropogoneae), while four genera are confined tothe Caribbean, specifically Cuba (Achlaena, Reynaudia, Scu-tachne, and Triscenia). A total of 115 species are endemicin Mesoamerica and the Caribbean.

A total of 18 genera are present only in South America:Agenium (Andropogoneae); Arundoclaytonia andSteyermarkochloa (Steyermarkochloeae); and Acostia, Alto-paradisium, Anthaenantiopsis, Arthropogon, Canastra, Cen-trochloa, Gerritea, Ophiochloa, Oplismenopsis, Otachyrium,Plagiantha, Spheneria, Streptostachys, Tatianyx, and Thras-yopsis (Paniceae). Some genera of Paniceae are primarilySouth American, but have one or a few species in Mesoam-erica (e.g., Chaetium, Echinolaena, and Thrasya).

As can be observed in Table 5, Brazil, Mexico, Venezuela,the Caribbean region, and Bolivia have the greatest numberof species in Panicoideae, with more than 50% of the speciesin Brazil and 26–34% in the other countries. Also, Brazil,Mexico, USA, the Caribbean, and Venezuela account for thegreatest number of endemic species. The distributions of thefive largest genera in the subfamily are provided in Table 3.

Distribution in Relation to Biogeographical Areas

For this analysis we used 15 of the subdivisions (regionsand provinces) of Takhtajan’s (1986) biogeographical clas-sification of the world (Appendix 1). Species designated asendemic are those limited in their distribution to one of the


Table 5. Numerical summary of New World Panicoideae speciesfor 23 countries and the Caribbean region.

Total

%of all

species Endemic NativeNative

(%)Intro-duced

Intro-duced(%)

CanadaUSAMexicoCaribbeanBelize

43351464370194

3.225.934.227.314.3

06266560

35281423316179

2.822.533.925.314.3

870415415

18.619.9

8.814.6

7.7GuatemalaEl SalvadorHondurasNicaraguaCosta Rica

274175289227266

20.212.921.316.719.6

20112

256158258203231

20.512.720.716.318.5

1817312435

6.69.7

10.710.613.2

PanamaColombiaVenezuelaGuyanaSuriname

238327396236194

17.524.129.217.414.3

29

3421

208308373210171

16.724.729.916.813.7

3019232623

12.65.85.8

11.011.9

French GuianaEcuadorPeruBrazilBolivia

197231251741359

14.517.018.554.626.5

112

721311

176206228688331

14.116.518.355.126.5

2125235328

10.710.8

9.27.27.8

ChileParaguayArgentinaUruguay

40303346172

2.922.325.512.7

3381

31290311158

2.523.224.912.7

9133514

22.54.3

10.18.1

Total 1357 100 497 1248 100 109 8.0

major American continental masses: Caribbean, Mesoamer-ica, North America, or South America. Restricted species arethose that occur in only two of the continental masses.

Holarctic Kingdom

North American Atlantic Region (NAAR).—A total of 72species are endemic to this region, which stretches acrossNorth America from the Atlantic Ocean to the Rocky Moun-tains, and from the Gulf of Mexico to southern Canada, andis characterized by deciduous forests in the east and grass-lands in the west. Endemic genera include Amphicarpum (2spp.) and Phanopyrum (1 sp.) in the southeastern USA. Inaddition, there are 31 and 21 endemic species, respectively,of subtribes Panicinae and Paspalinae. This region shares 17restricted species in 13 genera with the West Indian Provinceof the Caribbean Region.

Madrean Region (MR).—This area includes most of thesouthwestern USA (Arizona, California, Nevada, New Mex-ico, Utah, and parts of adjacent states) and northern andcentral Mexico, and is predominantly associated with dryhabitats. There are 93 endemic species in the region (whichaccounts for 10% of all endemic species) and no endemicgenera. This is the center of diversification of genera in Cen-totheceae (e.g., Zeugites), Zea (Tripsacinae), and the isolatedgenus Ixophorus (Setariinae).

Neotropical Kingdom

Caribbean Region (CAR).—This region includes tropicallowland plains in southern North America (tropical Florida),

southern Mexico, almost all of Central America, the islandsin the Caribbean Sea, and the Galapagos Islands. There aresix endemic genera and 120 endemic species among Pani-coideae. These are analyzed below within each province.

Central American Province (CAP).—A total of 61 speciesare confined to this province, which extends from southernMexico to Panama and neighboring areas of Colombia. To-pographically diverse, various kinds of forests dominate thisregion. Endemic genera are found in Centotheceae: Calde-ronella and Pohlidium, each with a single species in Panama,the former extending to Colombia. Other noteworthy endem-ic taxa found in this province belong to Tripsacinae (Trip-sacum spp.) and Centotheceae (Zeugites spp.). Sixteen re-stricted species are in common with the adjacent MadreanRegion.

West Indian Province (WIP).—This province includestropical areas of southern Florida, and the Greater and LesserAntilles. Four genera are limited to Cuba: Achlaena and Rey-naudia (Arthropogoninae), Scutachne (Melinidinae), andTriscenia (Paspalinae). Fifty-six Cuban species are endemic,nearly 50% belonging to Isachne and Paspalum. The prov-ince has 17 restricted species in common with the NorthAmerican Atlantic Region. In contrast, there are very fewspecies in common with the Central American Province andother biogeographical regions of northern South America.

Galapageian Province (GP).—Five species are endemicto the Galapagos Islands, two in Paspalum, and one each inCenchrus, Pennisetum,and Urochloa. No genera are endemic.

Andean Region (AR).—This region extends along the An-dean backbone of South America from Venezuela to north-ern Chile and Argentina. Highlands dominate, with mountainsummits of ca. 5000–6000 m. A total of 65 endemic speciesare present at medium or high altitudes. Two are in mono-typic genera (both Paspalinae) and have very limited, localdistributions: Acostia gracilis in Ecuador and Gerritea pseu-dopetiolata near La Paz, Bolivia. Paspalum and Pennisetumaccount for more than 56% of the endemic species in theregion, with 26 and 10, respectively.

Region of the Guiana Highlands (GUR).—This region oc-cupies the Guiana Highlands of northern South America, in-cluding southern Venezuela, western Guyana, western Co-lombia, and portions of northern Brazil. The vegetation com-prises tropical forests and savannas. A total of 55 speciesare endemic, most of them representatives of Axonopus,Panicum, and Paspalum. There are no endemic genera. Onlyseven restricted species are in common with the AmazonianRegion and 15 with cerrado areas of the Brazilian Region.

Amazonian Region (AMR).—This region includes the low-lands of the Amazon Basin, the coasts of Venezuela and itsadjacent islands, Trinidad and Tobago, the greater part ofGuyana, all of Suriname, all of French Guiana, and parts ofnortheastern Brazil. Tropical forests are characteristic. Thetwo genera in Steyermarkochloeae, Arundoclaytonia andSteyermarkochloa, are endemic, along with 36 species in 11other genera. Only four species are shared with the WestIndian Province, while 13 are in common with central areasof the Brazilian Region, including the species in the mono-typic genus Spheneria.


Brazilian Region (BR).—This region includes the BrazilianHighlands and the Caatinga, Paranaense, Atlantic, and GranChaco provinces, embracing most of Brazil, eastern Para-guay, Bolivia, and northern Argentina. Dominant vegetationtypes are savannas with xerophytic shrubs and woodlands(cerrados; Eiten 1972), and tropical or subtropical forests.The region has the greatest number of endemic species (376)and genera (11). The endemic genera are Agenium in An-dropogoneae, and Altoparadisium, Anthaenantiopsis, Arthro-pogon, Canastra, Centrochloa, Ophiochloa, Oplismenopsis,Plagiantha, Tatianyx, and Thrasyopsis in Paniceae. Ota-chyrium, Spheneria, and Streptostachys are best representedin the region, but O. versicolor (Doll) Henrard is a wide-spread species and Spheneria kegelii, and Streptostachys as-perifolia extend into the Amazonian Region. Subtribes Pan-icinae and Paspalinae are dominant in the region, with 50and 223 species, respectively.

Caatinga Province (CP).—The typical vegetation of thisprovince includes thorny shrubs, cactus parklands, and dif-ferent types of savannas. Five total species in Axonopus,Dichanthelium, Panicum, and Paspalum are endemic. Noendemic or restricted genera are present.

Uplands of Central Brazil Province (UPL).—The flora ofthis province, in which the predominant vegetation consistsof open savannas with a dry season, is characterized by highlevels of endemism. Within Panicoideae there are 196 en-demic species (or 21% of the total number of endemic spe-cies) and seven endemic genera in Paniceae: Altoparadisium,Arthropogon, Canastra, Centrochloa, Ophiochloa, Plagian-tha, and Tatianyx. Subtribe Arthropogoninae is well repre-sented with two genera and five species, but it is curious thatthe other two (monotypic) genera of the subtribe are restrict-ed to Cuba in the West Indian Province.

Chacoan Province (CHA).—This province includes theGran Chaco and Yungas. It is predominantly characterizedby arid tropical forests and woodlands. There are 23 endemicspecies and another 31 restricted species only shared withthe Parana Province. Subtribe Setariinae is well representedin the province. No endemic or restricted genera are present.

Atlantic Province (ATLP).—This province includes thesoutheastern coast of Brazil and its islands, and is charac-terized by evergreen tropical rain forests. A total of 19 spe-cies, belonging to seven genera (Acroceras, Dichanthelium,Digitaria, Ichnanthus, Isachne, Panicum, and Paspalum),are endemic. Three restricted species are shared with theParana Province and six with the Uplands of Central BrazilProvince. No genera are endemic to the province.

Parana Province (PAR).—Fifty species and Oplismenop-sis (Paniceae) are endemic to this province. The provincecovers areas of southern Brazil, eastern Paraguay, and north-eastern Argentina, in which the characteristic vegetationconsists of subtropical forests dominated by the genus Ar-aucaria Juss., together with tall-grass grasslands. A total of22 restricted species are in common with cerrados of centralBrazil, as well as Anthaenantiopsis and Thrasyopsis.

Amphitropical Disjuncts

A total of 31 species are disjunct between the MadreanRegion or the Caribbean Region and different regions of

South America. Seventeen species occur both in the Mad-rean Region and in South America in the Chacoan Provinceor Andean Region. Another 14 species are present in theCentral American Province and also in several areas ofSouth America (i.e., the Andean, Brazilian, and Amazonianregions). Bothriochloa, Digitaria, Panicum, and Paspalumare the genera with the majority of such amphitropical dis-junct species.

Distribution of Endemic Species in Relation to the KranzSyndrome

There are several interesting geographic patterns of Kranz(C4) and nonKranz (C3) taxa. The North American AtlanticRegion has 28 nonKranz species (38.89% of the total num-ber of endemic species) and 42 Kranz species (58.33%), ofwhich just two are of the PS subtype. Many of the nonKranzspecies belong to Dichanthelium, which is best representedin more temperate areas. On the other hand, the MadreanRegion has more than 90% Kranz species. Of these, it isnoteworthy that 27% are of the PS subtype, a subtype un-common elsewhere in the New World. Possessing this sub-type are 12 Panicum species, seven Urochloa species, andthree Eriochloa species.

The percentage of Kranz species is also high in the Ca-ribbean Region, with 97 species (79.5%). The GalapageianProvince has five Kranz species and no endemic C3 species.The West Indies Province has 48 Kranz species (44 MS, 4PS). The Central American Province has 44 Kranz species,with just one species of the PS subtype. Only seven non-Kranz species and one C3/C4 intermediate species occur inthe West Indies Province, while there are 17 nonKranz spe-cies in the Central American Province. A similar pattern ispresent in the Amazonian Region, where there are 27 Kranzspecies (26 MS, 1 PS) and seven nonKranz species. Theproportion of Kranz to nonKranz species is more balancedin the Guiana Highlands, with 20 nonKranz species and 35Kranz species, all of the MS subtype.

The distribution of Kranz species traditionally has beenthought to be correlated with warm climates and low alti-tudes. However, in the Andean Region we found 61 species(92%) to be C4, 54 of the MS subtype and just seven of thePS subtype. Kranz species in genera such as Paspalum reach4500 m elevation in this region.

Curiously, the percentage of C4 species is lower than ex-pected in the warm, low elevations of the Brazilian Region,with 284 Kranz species (76%). However, when consideringthe photosynthetic pathway by province within the region,16 species (84%) are nonKranz in the Atlantic Province,where tropical rainforests dominate. The number of Kranzspecies increases in the Caatinga and Parana provinces, withmore than 80% Kranz species. In the Chacoan Province allendemics are C4, with several representatives of the Kranzgenus Setaria. Finally, all species shared by the Parana andChacoan provinces, 31, are Kranz, while of the restrictedspecies shared by the Uplands, Chacoan, and Parana prov-inces, 39 (88%) are Kranz and three nonKranz. The UplandsProvince of Central Brazil has 71% C4 species, with Pas-palum (56 spp.), Axonopus (16 spp.), and Mesosetum (13spp.) most diverse. Within this province, Panicum is the


richest among the genera possessing C3 species, with 33 C3

species and just two C4 species.This study indicates that there are at least three centers of

richness of C4 taxa—the southeastern USA and Mexico, theAndean Region, and southern South America within theChacoan and Parana provinces. Our results for South Amer-ica disagree with previous studies of the distribution of C3

and C4 grasses (Ellis et al. 1980; Hattersley 1983; Cabido etal. 1997) for other parts of the world. These authors all in-dicated that the number of C4 grasses is greater as temper-ature and rainfall increase, which is correlated with relativelylow elevations. In our analysis the percentage of nonKranzspecies increases from west to east and decreases from northto south, with lower values in the Andean Region and thesouthern portion of the Brazilian Region.

ACKNOWLEDGMENTS

This work was supported by ANPCyT (Agencia Nacionalde Promocion Cientıfica y Tecnica, Argentina) grant 11739and CONICET grant 2131. Collections were supported bythe National Geographic Society through grants 6825-00 and7425-03. We thank Paul Peterson, J. Travis Columbus, andtwo anonymous reviewers for helpful comments on the man-uscript.

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Appendix 1. Endemic and restricted species of New World Panicoideae in relation to Kranz syndrome. Endemics in this analysis arethose species that occur in only one of the following four continental masses: Caribbean (CB), Mesoamerica (MA), North America (NA),and South America (SA). Restricted species are those that occur in two adjoining continental masses. Floristic regions (Takhatajan 1986):AMR ! Amazonian Region, AR ! Andean Region, ATLP ! Atlantic Province, BR ! Brazilian Region, CAP ! Central AmericanProvince, CAR ! Caribbean Region, CHA ! Chacoan Province, CP ! Caatinga Province, GP ! Galapageian Province, GUR ! GuianaHighlands Region, MR ! Madrean Region, NAAR ! North American Atlantic Region, PAR ! Parana Province, UPL ! Uplands ofCentral Brazil Province, WIP ! West Indian Province.

Taxon Continent Country Floristic region/provinceKranz

syndrome

Tribe Andropogoneae

Subtribe Andropogoninae

Andropogon aequatoriensis Hitchc.A. arctatus Chapm.A. arenarius Hack.

SANASA

EcuadorUSA

ARNAARBR–CHA " PAR

C4–MSC4–MSC4–MS

A. bourgeaei Hack.A. brachystachys Chapm.A. cabanisii Hack.

MANANA

USAUSA

CAR–CAPNAARNAAR


A. campestris Trin.A. coloratus Hack.A. cordatus SwallenA. crassus SohnsA. crispifolius Guala & Filg.

SASASASASA

BrazilArgentinaBoliviaVenezuelaBrazil

BR–UPLBR–CHA " PARBR–CHA " PARGURBR–UPL

C4–MSC4–MSC4–MSC4–MSC4–MS

A. crucianus RenvoizeA. diuturnus SohnsA. durifolius RendleA. exaratus R. Br.A. floridanus Scribn.

SASASASANA

VenezuelaBrazil

USA

BR–UPLGURBR–UPLBR–CHA " PARNAAR


A. gerardii VitmanA. glaucescens KunthA. glaucophyllus Roseng., B. R. Arrill. & Izag.

NA/MASASA

NAAR " CAR–CAPARBR–CHA " PAR


A. glaziovii Hack.A. indetonsus SohnsA. ingrates Hack.

SASASA

BrazilBrazil

BR–UPLAMRBR–UPL


A. insolitus SohnsA. lehmannii Pilg.A. liebmannii Hack.A. lindmanii Hack.A. longiberbis Hack.

SASANASA

CB/NA

ColombiaMexico

AMRARMRBR–CHA " PARNAAR " CAR–WIP


A. macrothrix Trin.A. multiflorus RenvoizeA. perdignus SohnsA. pohlianus Hack.

SASASASA

BoliviaVenezuelaBrazil

BR–UPL " CHA " PARBR–UPLAMRBR–UPL

C4–MSC4–MSC4–MSC4–MS

A. pringlei Scribn. & Merr.A. scabriglumis SwallenA. ternaries Michx.A. tracyi Nash

NASA

NA/MANA

MexicoEcuador

USA

MRARNAAR " MRNAAR


A. urbanianus Hitchc.A. vetus SohnsSchizachyrium beckii KilleenS. cubense (Hack.) Nash

CBSASACB

VenezuelaBolivia

CAR–WIPAMRBR–UPLCAR–WIP


S. gaumeri NashS. gracile (Spreng.) NashS. gracilipes (Hack.) A. CamusS. maritimum (Chapm.) NashS. mexicanum (Hitchc.) A. Camus

NACB/NA

SANANA

Mexico

USAMexico

MRNAAR " CAR–WIPBR–UPL " CHA " PARNAARMR


S. muelleri NashS. multinervosum NashS. niveum (Swallen) GouldS. parvifolium (Hitchc.) Borhidi & CatasusS. praematurum (Fernald) C. F. Reed

NACBNACBNA

Mexico

USA

USA

MRCAR–WIPNAARCAR–WIPNAAR


S. reedii (Hitchc. & Ekman) Borhidi & CatasusS. rhizomatum (Swallen) GouldS. scabriflorum (Rupr. ex Hack.) A. CamusS. scoparium (Michx.) Nash

CBNASANA

USACAR–WIPNAARBR–CHA " PARMR



Appendix 1. Continued.


syndrome

S. sericatum (Swallen) GouldS. spicatum (Spreng.) Herter

NASA

USA NAARBR–CHA ! PAR

C4–MSC4–MS

Subtribe Anthistiriinae

Agenium leptocladum (Hack.) ClaytonA. majus Pilg.A. villosum (Nees) Pilg.Elymandra lithophila (Trin.) Clayton

SASASASA Brazil

BR–UPL ! CHA ! PARBR–UPL ! CHA ! PARBR–UPL ! CHA ! PARBR–UPL


Subtribe Germaniniinae

Trachypogon macroglossus Trin.T. vestitus Andersson

SAMA/SA

Brazil BR–UPLCAR–CAP ! AMR

C4–MSC4–MS

Subtribe Ischaeminae

Ischaemum arenosum SohnsI. guianense Kunth ex Hack.I. minus J. Presl

SASASA

Venezuela GURAMR ! GURBR–CHA ! PAR


Subtribe Rottboelliinae

Mnesithea balansae (Hack.) de Koning & Sosef SA BR–CHA ! PAR C4–MSM. cylindrica (Michx.) de Koning & SosefM. impressa (Griseb.) de Koning & SosefM. parodiana (Henrard) de Koning & SosefM. rugosa (Nutt.) de Koning & SosefM. selloana (Hack.) de Koning & Sosef

NACBSA

CB/NASA

USA NAARCAR–WIPBR–CHA ! PARNAAR ! CAR–WIPBR–CHA ! PAR


M. tessellata (Steud.) de Koning & SosefM. tuberculosa (Nash) de Koning & SosefRhytachne gonzalezii DavidseR. guianensis (Hitchc.) Clayton

NACB/NA

SASA

USA NAARNAARAMRAMR ! BR–UPL


Subtribe Saccharinae

Imperata brevifolia VaseyI. condensata Steud.I. parodii AcevedoI. tenuis Hack.Saccharum alopecuroides (L.) Nutt.

NASASASANA

Chile

USA

MRARARBR–UPL ! CHA ! PARNAAR


S. angustifolium (Nees) Trin.S. baldwinii Spreng.S. brevibarbe (Michx.) Pes.S. coarctatum (Fernald) R. D. WebsterS. giganteum (Walter) Pers.

SANANANANA

USAUSAUSA

BR–CHA ! PARNAARNAARNAARNAAR


Subtribe Sorghinae

Bothriochloa eurylemma M. Marchi & Longhi-WagnerB. hirtifolia (J. Presl) HenrardB. hybrida (Gould) GouldB. imperatoides (Hack.) Herter

SANA/MA

NASA

BR–CHA ! PARMR ! CAR–CAPMRBR–CHA ! PAR


B. meridionalis M. Marchi & Longhi-WagnerB. palmeri (Hack.) Pilg.B. reevesii (Gould) GouldB. velutina M. Marchi & Longhi-WagnerB. wrightii (Hack.) Henrard

SANANASANA

Brazil

BR–CHA ! PARMRMRBR–CHA ! PARMR


Chrysopogon pauciflorus (Champ.) Benth. ex VaseySorghastrum balansae (Hack.) DavilaS. brunneum SwallenS. chasae Swallen

CB/NASA

NA/MASA

Paraguay

Brazil

NAAR ! CAR–WIPBR–CHA ! PARMR ! CAR–CAPBR–UPL


S. contractum (Hack.) Kuhlm. & A. KuhnS. elliotti (C. Mohr) NashS. minarum (Nees) Hitchc.S. nudipes Nash

SANASANA

BrazilUSA

Mexico

BR–UPLNAARBR–UPL ! CHA ! PARMR


S. pohlianum DavilaS. scaberrimum (Nees) HerterS. secundum (Elliott) NashS. viride SwallenSorghum trichocladum (Rupr. ex Hack.) Kuntze

NASANASAMA

MexicoBrazilUSA

MRBR–UPLNAARBR–UPL ! CHA ! PARCAR–CAP





syndrome

Subtribe Tripsacinae

Tripsacum australe H. C. Cutler & E. S. AndersonT. bravum J. R. GrayT. cundinamarcae de Wet & TimothyT. intermedium de Wet & J. R. HarlanT. jalapense de Wet & Brink

SANASAMAMA

MexicoColombia

ARMRARCAR–CAPCAR–CAP


T. lanceolatum Rupr. ex E. Fourn.T. maizar Hern.-Xol. & RandolphT. manisuroides de Wet & J. R. Harlan

NA/MAMAMA

MR ! CAR–CAPCAR–CAPCAR–CAP


T. peruvianum de Wet & TimothyT. pilosum Scribn. & Merr.T. zopilotense Hern.-Xol. & Randolph

SAMAMA

ARCAR–CAPCAR–CAP


Zea diploperennis H. H. Iltis, Doebley & R. GuzmanZ. nicaraguensis H. H. Iltis & B. F. BenzZ. perennis (Hitchc.) Reeves & Mangelsd.

NAMANA

MexicoNicaraguaMexico

MRCAR–CAPMR


Tribe Arundinelleae

Loudetia flammida (Trin.) C. E. HubbLoudetiopsis chrysothrix (Nees) ConertTristachya angustifolia Hitchc.

SASANA Mexico

BR–UPLBR–UPLMR


T. avenacea (J. Presl) Scribn. & Merr.T. contrerasii R. GuzmanT. laxa Scribn. & Merr.T. leiostachya NeesT. papilosa R. Guzman

CANANASANA

MexicoMexico

Mexico

CAR–CAPMRMRBR–UPLMR


Tribe Centotheceae

Calderonella sylvatica Soderstr. & H. F. DeckerChasmanthium curvifolium (Valdes-Reyna, Morden &

S. L. Hatch) Wipff & S. D. JonesC. latifolium (Michx.) H. O. YatesC. laxum (L.) H. O. Yates

MA/SANA

NANA

Mexico

USAUSA

CAR–CAP ! AMRMR

NAARNAAR

C3

C3

C3

C3

C. nitidum (Baldwin) H. O. YatesC. ornithorhynchum NeesPohlidium petiolatum Davidse, Soderstr. & R. P. Ellis

NANAMA

USAUSAPanama

NAARNAARCAR–CAP

C3

C3

C3

Zeugites capillaris (Hitchc.) SwallenZ. hackelii SwallenZ. hintonii HartleyZ. latifolia (E. Fourn.) Hemsl.Z. munroana Hemsl.

NANANANAMA

MexicoMexicoMexicoMexico

MRMRMRMRCAR–CAP

C3

C3

C3

C3

C3

Z. panamensis SwallenZ. pittieri Hack.Z. sagittata HartleyZ. smilacifolia Scribn.

MAMANANA

Panama

MexicoMexico

CAR–CAPCAR–CAPMRMR

C3

C3

C3

C3

Tribe Isachneae

Isachne angustifolia NashI. disperma (Lam.) DollI. goiasensis RenvoizeI. leersioides Griseb.I. ligulata Swallen

CBCBSACBSA

Brazil

CAR–WIPCAR–WIPBR–UPLCAR–WIPCAR ! AMR

C3

C3

C3

C3

C3

I. pubescens SwallenI. pygmaea Griseb.I. rigidifolia (Sw.) Trin.I. salzmannii (Trin. ex Steud.) Renvoize

MACBCBSA Brazil

CAR–CAPCAR–WIPCAR–WIPBR–ATLP

C3

C3

C3

C3

Tribe Paniceae

Subtribe Arthropogoninae

Achlaena piptostachya Griseb.Altoparadisium chapadense Filg., Davidse, Zuloaga

& MorroneA. scabrum (Pilg. & Kuhlm.) Filg., Davidse, Zuloaga

& Morrone

CBSA

SA

BrazilCAR–WIPBR–UPL

BR–UPL

C4–MSC4–MS

C4–MS




syndrome

Arthropogon filifolius Filg.A. villosus NeesA. xerachne EkmanReynaudia filiformis (Spreng. ex Schult.) Kunth

SASASACB

Brazil

Brazil

BR–UPLBR–UPLBR–UPLCAR–WIP


Subtribe Cenchrinae

Cenchrus distichophyllus Griseb.C. gracillimus NashC. multiflorus J. PreslC. palmeri VaseyC. platyacanthus Andersson

CBCB/NA

MANASA

MexicoEcuador

CAR–WIPNAAR ! CAR–WIPCAR–CAPMRCAR–GP


Pennisetum advena Wipff & VeldkampP. annuum MezP. chilense (E. Desv.) B. D. Jackson ex R. E. Fr.P. complanatum (Nees) Hemsl.

NASASAMA

USAPeru

NAARARARCAR–CAP


P. crinitum (Kunth) Spreng.P. distachyon (E. Fourn.) Rupr. ex ChaseP. dominguense (Spreng. ex Schult.) Spreng.

NAMACB

Mexico MRCAR–CAPCAR–WIP


P. durum BealP. frutescens LeekeP. intectum ChaseP. montanum (Griseb.) Hack.P. occidentale Chase

NASASASASA

Mexico MRBR–CHA ! PARARARAR


P. pauperum Nees ex Steud.P. peruvianum Trin.P. prolificum ChaseP. rigidum (Griseb.) Hack.P. rupestre Chase

SASANASASA

Ecuador

MexicoArgentina

CAR–GPARMRARAR


P. sagittatum HenrardP. tempisquense R. W. PohlP. tristachyum (Kunth) Spreng.P. weberbaueri Mez

SAMASASA

Costa RicaARCAR–CAPAR ! BR–UPLAR


Subtribe Digitariinae

Digitaria aequatoriensis (Hitchc.) Henrard SA AR C4–MSD. aequiglumis (Hack. & Arechav.) ParodiD. arenicola (Swallen) BeetleD. atra LucesD. badia (Scribn. & Merr.) FernaldD. balansae HenrardD. bonplandii Henrard

SANASA

NA/MASASA

USAVenezuela

BR–CHA ! PARNAARAMRMR ! CAR–CAPBR–CHA ! PARBR–UPL

C4–MSC4–MSC4–MSC4–MSC4–MSC4–MS

D. breedlovei R. W. Pohl & DavidseD. catamarcensis RugoloD. cayoensis SwallenD. chacoensis (Parodi) HenrardD. chaseae Henrard

MASAMASASA

Mexico

MexicoArgentinaBrazil

CAR–CAPARCAR–CAPBR–CHABR–UPL


D. clavitricha R. W. PohlD. cognata (Schult.) Pilg.D. connivens (Trin.) HenrardD. corynotricha (Hack.) Henrard

MANASASA

PanamaUSABrazilBrazil

CAR–CAPNAAR ! MRBR–CHA ! PARBR–UPL


D. costaricensis R. W. PohlD. curtigluma Hitchc.D. dioica Killeen & RugoloD. distans (Chase) FernaldD. eggersii (Hack.) Henrard

MANA/MA

SANACB

Mexico

CAR–CAPMR ! CAR–CAPAMR ! BR–UPLMRCAR–WIP


D. ekmanii Hitchc.D. enodis (Hack. ex Arechav.) ParodiD. eriostachya NeesD. floridana Hitchc.D. fragilis (Steud.) Luces

CBSASANASA

Uruguay

USA

CAR–WIPBR–CHA ! PARBR–CHA ! PARNAARAMR ! BR–UPL


D. gardnerii HenrardD. gracillima (Scribn.) FernaldD. hitchcockii (Chase) Stuck.D. hololeuca Henrard

SANANASA

BrazilUSAUSABrazil

BR–UPLNAARMRBR–UPL





syndrome

D. induta SwallenD. lanuginosa (Nees) HenrardD. leiantha (Hack.) Parodi

SASASA

Brazil BR–UPLBR–UPL ! CHA ! PARBR–UPL ! CHA ! PAR


D. leucites (Trin.) HenrardD. matogrossensis (Pilg.) HenrardD. multiflora SwallenD. myriostachya (Hack.) HenrardD. neesiana Henrard

MASAMASASA

BrazilBrazil

CAR–CAPBR–UPL ! CHA ! PARCAR–CAPBR–CHA ! PARBR–UPL


D. pampinosa HenrardD. panicea (Sw.) Urb.D. paniculata Soderstr. ex McVaughD. paraguayensis HenrardD. parva Swallen

SACBNASASA

Brazil

Mexico

Brazil

BR–UPLCAR–WIPMRBR–UPL ! CHA ! PARAMR


D. patens (Swallen) HenrardD. pauciflora Hitchc.D. phaeothrix (Trin.) ParodiD. pinetorum Hitchc.

NANASACB

USAMRNAARBR–UPL ! CHA ! PARCAR–WIP


D. pittieri (Hack.) HenrardD. pubiflora (Vasey) WipffD. purpurea Swallen

MANASA Brazil

CAR–CAPMRBR–CHA ! PAR


D. sabulicola HenrardD. sejuncta (Hack. ex Pilg.) HenrardD. serotina (Walter) Michx.D. simpsonii (Vasey) FernaldD. singularis Mez

SASA

CB/NACB/NA

SA

BrazilBrazil

BR–ATLPBR–UPLNAAR ! CAR–WIPNAAR ! CAR–WIPBR–CHA ! PAR


D. swalleniana HenrardD. tenuis (Nees) HenrardD. texana Hitchc.D. venezuelae HenrardD. villiculmis Henrard

SASANASASA Venezuela

BR–CHA ! PARGUR ! BR–UPLMRGURAMR


Subtribe Melinidinae

Chaetium bromoides (J. Presl) Benth. ex Hemsl.C. cubanum (C. Wright) Hitchc.C. festucoides NeesEriochloa acuminata (J. Presl) KunthE. boliviensis Renvoize

NACBSANASA

Mexico MRCAR–WIPAMRNAAR ! MRBR–UPL

C4–PSC4–MSC4–MSC4–PSC4–PS

E. contracta Hitchc.E. eggersii Hitchc.E. grandiflora (Trin.) Benth.E. lemmonii Vasey & Scribn.E. michauxii (Poir.) Hitchc.

NASASANANA USA

MRARBR–UPL ! CHA ! PARMRNAAR

C4–PSC4–PSC4–PSC4–PSC4–PS

E. montevidensis Griseb.E. nana ArriagaE. pacifica MezE. peruviana MezE. sericea (Scheele) Munro ex Vasey

SASASASANA

ArgentinaBR–CHA ! PARBR–CHAARARMR


E. setosa (A. Rich.) Hitchc.E. tridentata (Trin.) Kuhlm.E. weberbaueri Mez

CBSASA

CAR–CAP ! WIPBR–CHA ! PARAR

C4–PSC4–PSC4–PS

Scutachne amphistemon (C. Wright) Hitchc. & ChaseS. dura (C. Wright) Hitchc. & ChaseUrochloa acuminata (Renvoize) Morrone & Zuloaga

CBCBSA Brazil

CAR–WIPCAR–WIPBR–UPL


U. albicoma (Swallen & Garcıa-Barr.) Morrone& Zuloaga

SA Colombia AR C4–PS

U. arizonica (Scribn. & Merr.) Morrone & Zuloaga NA MR C4–PSU. ciliatissima (Buckley) R. D. Webster NA MR C4–PSU. decidua Morrone & Zuloaga SA Brazil BR–UPL C4–PSU. discifera (E. Fourn.) Morrone & Zuloaga NA Mexico MR C4–PSU. lorentziana (Mez) Morrone & ZuloagaU. megastachya (Nees ex Trin.) Morrone & ZuloagaU. meziana (Hitchc.) Morrone & ZuloagaU. multiculma (Andersson) Morrone & Zuloaga

SASANASA

BrazilMexicoEcuador

BR–CHABR–UPLMRCAR–GP

C4–PSC4–PSC4–PSC4–PS




syndrome

U. oblita (Swallen) Morrone & ZuloagaU. ophryodes (Chase) Morrone & ZuloagaU. paucispicata (Morong) Morrone & ZuloagaU. texana (Buckley) R. D. WebsterU. venosa (Swallen) Morrone & Zuloaga

SANASANANA

Mexico

Mexico

ARMRBR–CHAMRMR


Subtribe Panicinae

Acroceras chaseae Zuloaga & MorroneA. excavatum (Henrard) Zuloaga & MorroneA. fluminense (Hack.) Zuloaga & Morrone

SASASA

Brazil

Brazil

BR–ATLPBR–ATLP ! PARBR–ATLP

C3

C3

C3

Dichanthelium adenorachis (Zuloaga & Morrone)Zuloaga

D. aequivaginatum (Swallen) ZuloagaD. assurgens (Renvoize) ZuloagaD. boreale (Nash) FreckmannD. boscii (Poir.) Gould & C. A. Clark

SA

SASANANA

Brazil BR–UPL

AMR ! BR–UPLBR–UPLNAARNAAR

C3

C3

C3

C3

C3

D. cabrerae (Zuloaga & Morrone) ZuloagaD. caparoense (Zuloaga & Morrone) ZuloagaD. clandestinum (L.) GouldD. commutatum (Schult.) Gould

SASANA

CB/NA

BrazilBrazil

BR–UPLBR–UPLNAARNAAR ! CAR–WIP

C3

C3

C3

C3

D. congestum (Renvoize) ZuloagaD. cucaense (Zuloaga & Morrone) ZuloagaD. cumbucana (Renvoize) Zuloaga

SASASA

BrazilBrazilBrazil

BR–UPLBR–ATLPBR–UPL

C3

C3

C3

D. davidsei (Zuloaga & Morrone) ZuloagaD. depauperatum (Muhl.) GouldD. ensifolium (Baldwin ex Elliot) GouldD. heliophilum (Chase ex Zuloaga & Morrone) ZuloagaD. itatiaiae (Swallen) Zuloaga

SANA

CB/NASASA

BrazilBrazil

AMRNAARNAAR ! CAR–WIPBR–UPLBR–ATLP

C3

C3

C3

C3

C3

D. latifolium (L.) HarvillD. laxiflorum (Lam.) GouldD. leibergii (Vasey) FreckmannD. linearifolium (Scribn.) Gould

NACB/NA

NANA

NAARNAAR ! CAR–WIPNAARNAAR

C3

C3

C3

C3

D. macrospermum GouldD. malacophyllum (Nash) GouldD. nodatum (Hitchc. & Chase) GouldD. nudicaule (Vasey) B. F. Hansen & Wunderlin

NANANANA

MexicoUSA

USA

MRNAARNAARNAAR

C3

C3

C3

C3

D. oligosanthes (Schult.) GouldD. ovale (Elliott) Gould & C. A. ClarkD. pedicellatum (Vasey) GouldD. perlongum (Nash) FreckmannD. peristypum (Zuloaga & Morrone) Zuloaga

NANA

NA/MANASA

USA

USABrazil

NAARNAARNAARNAARBR–ATLP

C3

C3

C3

C3

C3

D. petropolitanum (Zuloaga & Morrone) ZuloagaD. polyanthes (Schult.) Mohlenbr.D. portoricense (Desv. ex Ham.) B. F. Hansen

& Wunderlin

SANA

CB/NA

BrazilUSA

BR–ATLPNAARNAAR ! CAR–WIP

C3

C3

C3

D. ravenelii (Scribn. & Merr.) GouldD. sabulorum (Lam.) Gould & C. A. ClarkD. scabriusculum (Elliott) Gould & C. A. Clark

NASANA

USA

USA

NAARBR–ATLP ! PARNAAR

C3

C3

C3

D. sciurotis (Trin.) DavidseD. scoparium (Lam.) GouldD. sendulskyii (Zuloaga & Morrone) ZuloagaD. stigmosum (Trin.) ZuloagaD. stipiflorum (Renvoize) Zuloaga

SACB/NA

SASASA

Brazil

BrazilBrazilBrazil

BR–CPNAAR ! CAR–WIPBR–UPLBR–ATLP ! UPLBR–UPL

C3

C3

C3

C3

C3

D. superatum (Hack.) ZuloagaD. surrectum (Chase ex Zuloaga & Morrone) ZuloagaD. telmatum (Swallen) ZuloagaD. willcoxianum (Vasey) FreckmannD. xanthophysum (A. Gray) Freckmann

SASASANANA

BrazilBrazilVenezuelaUSA

BR–ATLP ! UPLBR–UPLGURNAARNAAR

C3

C3

C3

C3

C3

D. yadkinense (Ashe) Mohlenbr.Echinochloa chacoensis P. W. Michael ex RenvoizeE. helodes (Hack.) Parodi

NASASA

USA NAARBR–CHABR–CHA

C3

C4–MSC4–MS

E. holciformis (Kunth) ChaseE. jaliscana McVaughE. muricata (P. Beauv.) Fernald

NA/MANANA

MexicoMR ! CAR–CAPMRNAAR ! MR





syndrome

E. oplismenoides (E. Fourn.) Hitchc.E. paludigena Wiegand

NANA

MexicoUSA

MRNAAR

C4–MSC4–MS

Lasiacis anomala Hitchc.L. linearis SwallenL. rugellii (Griseb.) Hitchc.Panicum aquarum Zuloaga & MorroneP. arctum Swallen

CB/SAMA

CB/MASASA

AMR ! CAR–WIPCAR–CAPCAR–CAP ! WIPAMRGUR

C3

C3

C3

C4–PSC3

P. auricomum Nees ex Trin.P. aztecanum Zuloaga & MorroneP. bartletii Swallen

SANAMA

BrazilMexico

BR–UPLMRCAR–CAP

C3

C4–PSC3

P. bergii Arechav.P. brachyanthum Steud.P. caaguazuense Henrard

SANASA

USABR–CHA ! PARNAARBR–UPL ! PAR

C4–PSC3

C3

P. campestre Nees ex Trin.P. capillare L.P. capillarioides VaseyP. cayoense SwallenP. cervicatum Chase

SANANAMASA

Brazil BR–UPLMRMRCAR–CAPGUR ! BR–UPL

C4–PSC4–PSC4–PSC3

C4–PSP. chasiae Roseng., B. R. Arill. & Izag.P. chloroleucum Griseb.P. deciduum SwallenP. decolorans KunthP. diffusum Sw.

SASASANACB

ArgentinaVenezuelaMexico

BR–PARARGURMRCAR–WIP

C4–PSC4–PSC3

C4–PSC4–PS

P. ephemeroides Zuloaga & MorroneP. exiguum MezP. flexile (Gatt.) Scribn.

SASANA

Brazil BR–UPLBR–UPL ! PARNAAR ! MR


P. fonticolum SwallenP. furvuum SwallenP. glabripes Doll

SAMASA

VenezuelaGuatemala

GURCAR–CAPBR–PAR

C3

C4–PSC4–PS

P. granuliferum KunthP. guatemalense SwallenP. hallii VaseyP. havardii VaseyP. hillmanii Chase

SAMANANANA USA

GURCAR–CAPMRMRMR

C3

C3


P. ichunense SwallenP. incumbens SwallenP. lacustre Hitchc. & EkmanP. lepidulum Hitchc. & ChaseP. ligulare Nees ex Trin.

SAMACBNASA

Venezuela

Mexico

GURCAR–CAPCAR–WIPMRGUR ! BR–UPL

C3

C3


P. longipedicellatum SwallenP. longissimum (Mez) HenrardP. machrisiana Swallen

SASASA

BrazilParaguayBrazil

BR–ATLPBR–PARBR–UPL

C3

C4–PSC3

P. magnispicula Zuloaga, Morrone & VallsP. micranthum KunthP. missionum Ekman

SASASA

Brazil BR–UPLGURBR–PAR

C4–PSC3

C3

P. mohavense ReederP. mucronulatum MezP. mystasipum Zuloaga & MorroneP. nervosum Lam.P. noterophilum Renvoize

NASASASASA

USABrazilBrazil

Brazil

MRBR–CPBR–UPLGUR ! BR–UPLBR–UPL

C4–PSC4–PSC4–PSC3

C3

P. nutabundum Zuloaga & MorroneP. olyroides KunthP. orinocanum LucesP. pampinosum Hitchc. & ChaseP. pandum Swallen

SASASANASA

Brazil

Venezuela

BR–UPLBR–UPL ! PARGURMRGUR

C3

C4–PSC3

C4–PSC3

P. parcum Hitchc. & ChaseP. parviglume Hack.P. pedersenii ZuloagaP. peladoense Henrard

NA/MAMASASA

MR ! CAR–CAPCAR–CAPBR–PARBR–UPL ! CHA ! PAR

C4–PSC3

C4–PSC4–PS

P. penicillatum Nees ex Trin.P. petilum SwallenP. petrense Swallen

SASASA

BrazilColombiaVenezuela

BR–ATLPGURGUR

C3

C3

C3




syndrome

P. philadelphicum Bernh. ex Trin.P. pseudisachne MezP. quadriglume (Doll) Hitchc.P. racemosum (P. Beauv.) Spreng.P. rhizogonum Hack.

NASASASASA

USABrazil

NAARBR–UPLBR–UPL ! PARBR–ATLPBR–PAR

C4–PSC3

C4–PSC4–PSC3

P. rivale SwallenP. rupestre Trin.P. subulatum Spreng.P. tamaulipense F. R. Waller & MordenP. tepuianum Davidse & Zuloaga

SASASANASA

BrazilBrazilMexicoVenezuela

GURBR–UPLBR–UPLMRGUR

C3

C3

C3

C4–PSC3

P. vaseyanum Scribn. ex BealP. verrucosum Muhl.P. wettsteinii Hack.P. yavitaense SwallenSacciolepis otachyrioides Judz.

NANASASASA

MexicoUSABrazilVenezuela

MRNAARBR–UPLGURGUR ! AMR

C4–PSC3

C3

C3

C3

Subtribe Paspalinae

Acostia gracilis SwallenAmphicarphum muhlenbergianum (Schult.) Hitchc.A. purshii KunthAnthaenanthia rufa (Elliott) Schult.A. villaregalis (McVaugh & R. Guzman) Espejo

& Lopez-Ferrari

SANANANANA

EcuadorUSAUSAUSAMexico

ARNAARNAARNAARMR


A. villosa (Michx.) P. Beauv.Anthaenantiopsis fiebrigii (Mez) ParodiA. perforata (Nees) ParodiA. rojasiana ParodiA. trachystachya (Nees) Mez ex Pilg.

NASASASASA

USA NAARBR–UPL ! PARBR–UPLBR–UPL ! PARBR–UPL


Axonopus amapaensis G. A. BlackA. andinus G. A. BlackA. apricus G. A. BlackA. arcuatus (Mez) G. A. Black

SASASASA

BrazilBoliviaBrazilBrazil

AMRARBR–UPLBR–UPL


A. argentinus ParodiA. arsenei SwallenA. boliviensis RenvoizeA. brasiliensis (Spreng.) Kuhlm.A. canescens (Nees ex Trin.) Pilg.

SANASASASA

MexicoBolivia

BR–UPL ! CHA ! PARMRBR–UPLBR–UPL ! PARAMR ! BR–UPL


A. carajasensis BastosA. casiquiarensis DavidseA. caulescens (Mez) HenrardA. chaseae G. A. BlackA. chimantensis Davidse

SASASASASA

Brazil

BrazilVenezuela

GURGURGURBR–UPLGUR


A. chrysostachyus (Lag.) ChaseA. ciliatifolius SwallenA. comans (Trin. ex Doll) HenrardA. comatus (Mez) Swallen

SAMASASA Brazil

BR–ATLP ! UPLCAR–CAPBR–UPLBR–UPL


A. complanatus (Nees) DedeccaA. cuatrecasasii G. A. BlackA. debilis G. A. Black

SASACB

Brazil BR–CPAMRCAR–WIP


A. deludens ChaseA. elegantulus (J. Presl) Hitchc.A. eminens (Nees) G. A. BlackA. equitans Hitchc. & ChaseA. fastigiatus (Nees ex Trin.) Kuhlm.

NASASA

CB/SASA

Mexico

Brazil

MRARAMR ! BR–UPLAMR ! CAR–WIPBR–UPL


A. flabelliformis SwallenA. furcatus (Flugge) Hitchc.A. grandifolius RenvoizeA. herzogii (Hack.) Hitchc.A. hirsutus G. A. Black

SACB/NA

SASASA

Brazil

GUR ! AMRNAAR ! CAR–WIPBR–UPLBR–UPLBR–UPL


A. iridifolius (Poepp.) G. A. BlackA. jeanyae DavidseA. jesuiticus (Araujo) VallsA. junciformis G. A. Black

SAMASASA

BrazilBrazil

AR ! GURCAR–CAPBR–PARBR–UPL





syndrome

A. laxiflorus (Trin.) ChaseA. laxus LucesA. longispicus (Doll) Kuhlm.

SASASA

VenezuelaGUR ! AMRAMRAMR


A. marginatus (Trin.) ChaseA. mathewsii (Mez) Hitchc.A. mexicanus G. A. BlackA. micay Garcıa-Barr.A. monticola G. A. Black

SASANA

MA/SASA

Mexico

Brazil

BR–UPLARMRAR ! CAR–CAPBR–UPL


A. morronei Giraldo-CanasA. obtusifolius (Raddi) ChaseA. oiapocensis G. A. BlackA. passourae G. A. BlackA. pellitus (Nees ex Trin.) Hitchc. & Chase

SASASASASA

Colombia

BrazilFrench GuianaBrazil

AMRBR–UPL ! PARGURAMRBR–UPL


A. pennelii G. A. BlackA. piccae Giraldo-CanasA. polydactylus (Steud.) Dedecca

SASASA Brazil

GURGURBR–CP


A. polystachyus G. A. BlackA. pressus (Nees ex Steud.) ParodiA. pubivaginatus Henrard

SASASA

Brazil BR–UPL ! PARBR–UPLAMR ! BR–CP


A. ramboi G. A. BlackA. ramosus SwallenA. rosei (Scribn. & Merr.) ChaseA. rosengurtii G. A. BlackA. rupestris Davidse

SASANASASA

Brazil

Mexico

Brazil

BR–PARGURMRBR–PARBR–UPL


A. senescens (Doll) HenrardA. siccus (Nees) Kuhlm.A. steyermarkii SwallenA. succulentus G. A. BlackA. suffultiformis G. A. Black

SASASASASA

VenezuelaParaguayVenezuela

AMRBR–PARGURBR–PARGUR


A. suffultus (Mikan ex Trin.) ParodiA. sulcatus G. A. BlackA. surinamensis (Hochst. ex Steud.) Henrard

SASASA

BrazilBR–PARAMRGUR ! AMR


A. triglochinoides (Mez) DedeccaA. uninodis (Hack.) G. A. BlackA. villosus Swallen

SASASA

Brazil

Venezuela

AMRBR–UPL ! PARGUR


A. volcanicus R. W. PohlA. yutajensis G. A. BlackA. zuloagae Giraldo-CanasCanastra lanceolata (Filg.) Morrone, Zuloaga, Davidse

& Filg.Centrochloa singularis Swallen

MASASASA

SA

VenezuelaColombiaBrazil

Brazil

CAR–CAPGURGURBR–UPL

BR–UPL

C4–MSC4–MSC4–MSC3

C4–MSEchinolaena ecuadoriana Filg.E. inflexa (Poir.) ChaseE. minarum (Nees) Pilg.E. oplismenoides (Munro ex Doll) StieberE. standleyi (Hitchc.) Stieber

SASASASAMA

Ecuador ARBR–UPLBR–UPL ! PARBR–UPL ! ARCAR–CAP

C3

C3

C3

C3

C3

Gerritea pseudopetiolata Zuloaga, Morrone & KilleenHomolepis longispicula (Doll) ChaseH. villaricensis (Mez) Zuloaga & Soderstr.

SASASA

BoliviaBrazil

ARBR–UPLBR–PAR

C3

C3

C3

Hymenachne grumosa (Nees) ZuloagaH. pernambucense (Spreng. ex Mez) ZuloagaIchnanthus annuus Killeen & KirpesI. bambusiflorus (Trin.) DollI. breviscrobs Doll

SASASASASA

Brazil

BR–PARBR–ATLP ! PARBR–UPL ! PARBR–UPL ! ATLPAMR

C3

C3

C3

C3

C3

I. camporum SwallenI. ephemeroblepharis G. A. Black & Froes ex G. A.

Black & PiresI. glaber (Raddi) Hitchc.I. grandifolius (Doll) Zuloaga & Soderstr.

SASA

SASA

Brazil

BrazilBrazil

BR–UPLGUR

BR–ATLPBR–ATLP

C3

C3

C3

C3

I. hirtus (Raddi) ChaseI. hoffmannseggii (Roem. & Schult.) DollI. lanceolatus Scribn. & J. G. Sm.

SASAMA

BR–ATLPAMRCAR–CAP

C3

C3

C3




syndrome

I. leptophyllus DollI. longiglumis MezI. mayarensis (C. Wright) Hitchc.I. mollis EkmanI. procurrens (Nees ex Trin.) Swallen

SASACBSASA

BrazilBrazil

Brazil

AMRBR–UPLCAR–WIPBR–UPLBR–UPL ! PAR

C3

C3

C3

C3

C3

I. riedelii (Trin.) DollI. tarijianus K. E. RogersI. tarumanensis G. A. Black & FroesI. tectus SwallenI. zehntneri Mez

SASASASASA

Brazil

BrazilVenezuelaBrazil

BR–ATLPARAMRGURBR–UPL

C3

C3

C3

C3

C3

Mesosetum agropyroides MezM. alatum Filg.M. annuum SwallenM. ansatum (Trin.) Kuhlm.

SASASASA

BrazilBrazilBrazil

BR–UPLBR–UPLBR–UPLBR–UPL


M. arenarium SwallenM. bifarium (Hack.) ChaseM. blakei Swallen

SASAMA

BrazilBrazil

BR–UPLBR–UPLCAR–CAP


M. chlorostachyum (Doll) ChaseM. comatum SwallenM. elytrochaetum (Hack.) SwallenM. exaratum (Trin.) Chase

SASASASA

ArgentinaBrazilBrazil

AMRBR–PARAMRBR–UPL


M. ferrugineum (Trin.) ChaseM. gibbosum (Renvoize) Filg.M. longiaristatum Filg.M. pappophorum (Nees) Kuhlm.M. penicillatum Mez

SASASASASA

BrazilBrazilBrazil

Brazil

BR–UPLBR–UPLBR–UPLBR–UPL ! AMRBR–UPL


M. pittieri Hitchc.M. rottboellioides (Kunth) Hitchc.M. sclerochloa (Trin.) Hitchc.

MASASA Brazil

CAR–CAPGUR ! AMRBR–UPL


M. stoloniferum SwallenM. wrightii Hitchc.Ophiochloa hydrolithica Filg., Davidse & Zuloaga

MACBSA Brazil

CAR–CAPCAR–WIPBR–UPL


Oplismenopsis najada (Hack. & Arechav.) ParodiOtachyrium aquaticum Send. & Soderstr.O. boliviensis RenvoizeO. grandiflorum Send. & Soderstr.O. piligerum Send. & Soderstr.

SASASASASA

BrazilBolivia

Brazil

BR–PARBR–UPLBR–UPLGUR ! BR–UPLBR–UPL

C3

C3

C3

C3

C3

O. pterygodium (Trin.) Pilg.O. seminudum Hack. ex Send. & Soderstr.O. succisum (Swallen) Send. & Soderstr.Panicum acicularifolium Renvoize & Zuloaga

SASASASA

BrazilBrazil

Brazil

BR–UPLBR–UPLBR–UPL ! AMRBR–UPL

C3

C3

C3

C3

P. anceps Michx.P. andreanum MezP. animarum Renvoize

NASASA

USA

Brazil

NAARARBR–UPL

C4–MSC3

C3

P. aristellum DollP. bahiense RenvoizeP. beyeri Hitchc. & EkmanP. biglandulare Scribn. & J. G. Sm.P. brachystachyum Steud.

SASACBMASA

BrazilBrazil

Brazil

BR–UPLBR–UPLCAR–WIPCAR–CAPBR–UPL

C3

C3

C4–MSC3

C3

P. bresolinii L. B. Sm. & Wassh.P. chapadense SwallenP. chnoodes Trin.P. cipoense Renvoize & Send.P. condensatum Bertol.

SASASASASA

BrazilBrazil

BrazilBrazil

BR–PARBR–UPLGUR ! BR–UPLBR–UPLBR–ATLP

C3

C3

C3

C3

C3

P. crateriferum SohnsP. durifolium Renvoize & ZuloagaP. eligulatum N. E. Br.P. euprepes Renvoize

NASASASA

MexicoBrazil

Brazil

MRBR–UPLGUR ! BR–UPLBR–UPL

C3

C3

C3

C3

P. fontanale SwallenP. glaziovii Hack.P. hemitomon Schult.

SASA

CB/NA

VenezuelaBrazil

GURBR–UPLNAAR ! CAR–WIP

C3

C3

C3




syndrome

P. irregulare SwallenP. jauanum DavidseP. lagostachyum Renvoize & ZuloagaP. latissimum Mikan ex Trin.P. leptachne Doll

MA/SASASASASA

VenezuelaBrazilBrazilBrazil

CAR–CAP ! ARGURBR–UPLBR–ATLPBR–ATLP

C3

C3

C3

C3

C3

P. longum Hitchc. & ChaseP. loreum Trin.P. lutzii SwallenP. marauense Renvoize & ZuloagaP. molinioides Trin.

NASASASASA

MexicoBrazilBrazilBrazilBrazil

MRBR–UPLBR–UPLBR–UPLBR–UPL

C3

C3

C3

C3

C3

P. obtusum KunthP. petersonii Hitchc. & EkmanP. piauiense SwallenP. poliophyllum Renvoize & Zuloaga

NACBSASA

BrazilBrazil

MRCAR–WIPBR–UPLBR–UPL

C4–MSC4–MSC3

C3

P. prionitis NeesP. restingae Renvoize & ZuloagaP. rude Nees

SASASA

BrazilBrazil

BR–PARBR–UPLBR–ATLP ! UPL

C4–MSC3

C3

P. sacciolepoides Renvoize & ZuloagaP. scabridum DollP. sipapoense SwallenP. soderstromii Zuloaga & Send.P. stagnatile Hitchc. & Chase

SASASASA

NA/MA

Brazil

VenezuelaBrazil

BR–UPLAMRGURBR–UPLMR ! CAR–CAP

C3

C3

C3

C3

C3

P. stevensianum Hitchc. & ChaseP. steyermarkii SwallenP. subtiramulosum Renvoize & ZuloagaP. teretifolium Hack.P. tijucae Renvoize

CB/SASASASASA

VenezuelaBrazilBrazilBrazil

AMR ! CAR–WIPGURBR–UPLBR–UPLBR–UPL

C3

C3

C3

C3

C3

P. trinii KunthP. tuerckheimii Hack.P. vaginiviscosum Renvoize & ZuloagaP. validum Mez

SAMASASA

Brazil

Brazil

BR–UPLCAR–CAPBR–UPLBR–PAR

C3

C4–MSC3

C4–MSPaspalum acutifolium LeonP. acutum ChaseP. adoperiens (E. Fourn.) ChaseP. affine Steud.P. alaini Leon

CBMAMAMACB

CAR–WIPCAR–CAPCAR–CAPCAR–CAPCAR–WIP


P. albidulum HenrardP. almum ChaseP. alterniflorum A. Rich.P. altsonii ChaseP. ammodes Trin.

SASACBSASA

GURBR–CHACAR–WIPGURGUR ! BR–UPL


P. amphicarpum EkmanP. andersonii MezP. apiculatum DollP. approximatum Doll

CBSASASA

PeruCAR–WIPARGUR ! AMRBR–UPL


P. arsenei ChaseP. arundinellum MezP. aspidiotes Trin.

NASASA

Mexico MRBR–PARGUR ! BR–UPL


P. atabapense Davidse & ZuloagaP. azuayense SohnsP. bakeri Hack.P. barbinode Hack.P. barclayii Chase

SASA

CB/SASASA

VenezuelaEcuador

BrazilPeru

GURARAMR ! CAR–WIPBR–UPLAR


P. bertonii Hack.P. biaristatum Filg. & DavidseP. bifidifolium Soderstr.P. bifidum (Bertol.) NashP. bonplandianum Flugge

SASASANASA

BrazilGuyanaUSA

BR–PARBR–UPLGURNAARAR


P. botteri (E. Fourn.) Chase NA/MA MR ! CAR–CAP C4–MSP. brachytrychum Hack.P. breve Chase

SACB

Brazil BR–UPLCAR–WIP

C4–MSC4–MS

P. buchtienii Hack.P. burchelli Munro ex Oliv.

SASA

BoliviaBrazil

ARBR–UPL

C4–MSC4–MS




syndrome

P. burmanii Filg., Morrone & Zuloaga SA Brazil BR–UPL C4–MSP. cachimboense Davidse, Morrone & ZuloagaP. campinarum Filg. & DavidseP. capillifolium NashP. centrale ChaseP. ceresia (Kuntze) Chase

SASACBMASA

BrazilBrazil

BR–UPLAMRCAR–WIPCAR–CAPAR


P. chacoense ParodiP. chaffanjonii MauryP. chaseanum ParodiP. cinerascens (Doll) A. G. Burm. & C. N. BastosP. circulare Nash

SASASASANA

Venezuela

USA

BR–CHAGURBR–CHAGUR ! BR–UPLNAAR


P. clandestinum SwallenP. commune LilloP. compressifolium SwallenP. conduplicatum Canto-Dorow, Valls

& Longhi-Wagner

SASASASA

Brazil

Brazil

BR–UPLARBR–UPL ! PARBR–PAR


P. cordaense SwallenP. cordatum Hack.P. costaricense MezP. costellatum SwallenP. crinitum Chase

SASAMASANA

Brazil

BrazilMexico

BR–UPLGUR ! BR–UPLCAR–CAPBR–UPLMR


P. crispatum Hack.P. crispulum SwallenP. cromyorrhizon Trin. ex DollP. crustarium SwallenP. culiacanum Vasey

SASASASA

NA/MA

Brazil

Brazil

BR–UPLBR–UPLBR–PARBR–UPLMR ! CAR–CAP


P. curasavicum ChaseP. cymbiforme E. Fourn.P. dasytrichium Dusen ex SwallenP. dedeccae Quarın

CBNA/MA

SASA

Brazil

CAR–WIPMR ! CAR–CAPBR–UPLBR–UPL ! PAR


P. delicatum SwallenP. difforme J. Le ConteP. dilatatum Poir.

SANASA

USAGUR ! BR–UPLNAARBR–PAR


P. dispar ChaseP. dissectum (L.) L.P. distachyon Poir. ex Trin.P. distortum ChaseP. divergens Doll

CBCB/NA

CBCBSA Brazil

CAR–WIPNAAR ! CAR–WIPCAR–WIPCAR–WIPBR–UPL


P. durifolium MezP. edmondii LeonP. ekmanianum HenrardP. ellipticum DollP. equitans Mez

SACBSASASA

Brazil

BR–PARCAR–WIPARBR–UPLBR–PAR


P. erectum ChaseP. erianthoides Lindm.P. eucomum Nees ex Trin.P. exaltatum J. Presl

NASASASA

Mexico

Brazil

MRBR–PARBR–UPLBR–PAR


P. expansum DollP. falcatum Nees ex Steud.P. filifolium Nees ex Steud.

SASASA

Brazil

Brazil

BR–UPLBR–UPL ! PARBR–UPL


P. filiforme Sw.P. flaccidum NeesP. flavum J. PreslP. floridanum Michx.P. formosum Swallen

CBSASANASA

USABrazil

CAR–WIPBR–UPLARNAARBR–UPL


P. galapageium ChaseP. geminiflorum Steud.P. gemmosum Chase ex RenvoizeP. giganteum Baldwin ex VaseyP. glabrinode (Hack.) Morrone & Zuloaga

SASASANASA

Ecuador

BrazilUSA

CAR–GPAMR ! BR–UPLBR–UPLNAARBR–PAR

C4–MSC4–MS

C4–MSC4–MS

P. glaucescens Hack.P. goyasense Davidse, Morrone & ZuloagaP. guaricense Swallen

SASASA

BrazilVenezuela

BR–UPL ! CHA ! PARBR–UPLAMR





syndrome

P. guenoarum Arechav.P. guttatum Trin.

SASA Brazil

BR–UPL ! CHA ! PARBR–UPL

C4–MSC4–MS

P. haenkeanum J. PreslP. hartwegianum E. Fourn.P. hatschbachii Zuloaga & MorroneP. haumanii ParodiP. hintonii Chase

SANASASANA

PeruMexicoBrazil

Mexico

ARMRBR–UPLBR–PARMR


P. hirtum KunthP. hispidum SwallenP. hitchcockii ChaseP. imbricatum Filg.P. inconstans Chase

SAMA

MA/SASASA

ColombiaGuatemala

Brazil

ARCAR–CAPCAR–CAP ! ARBR–UPLAR


P. indecorum MezP. insulare Ekman ex ChaseP. ionanthum ChaseP. jaliscanum Chase

SACBSA

NA/MA

BR–PARCAR–WIPBR–PARMR ! CAR–CAP


P. jesuiticum ParodiP. jimenezii ChaseP. killipii (Hitchc.) Zuloaga and Soderstr.

SAMASA

Costa RicaPeru

BR–PARCAR–CAPAR


P. lachneum Nees ex Steud.P. lacustre Chase ex SwallenP. laeve Michx.P. lanciflorum Trin.P. latipes Swallen

SASANASASA

BrazilUSA

Brazil

BR–UPLAMRNAARGUR ! BR–UPLBR–UPL


P. lenticulare KunthP. lentiferum Lam.P. lentifinosum J. PreslP. leptachne ChaseP. lilloi Hack.

SANA

NA/MANASA

VenezuelaUSA

Mexico

AMRNAARMR ! CAR–CAPMRBR–PAR


P. limbatum HenrardP. lindenianum A. Rich.P. loefgrenii Ekman

SACBSA

BR–PARCAR–WIPBR–CP


P. longiaristatum Davidse & Filg.P. longicuspe NashP. longipilum Nash

SANANA

BrazilMexicoUSA

BR–UPLMRNAAR


P. longum ChaseP. macranthecium ParodiP. macrophyllum KunthP. madorense RenvoizeP. malmeanum Ekman

NASASASASA

Mexico

Brazil

MRBR–UPLARBR–UPLBR–UPL


P. mandiocanum Trin.P. marmoratum Kuhlm.P. mayanum ChaseP. minarum Hack.P. modestum Mez

SASAMASASA

BrazilMexicoBrazil

BR–UPL ! PARBR–UPLCAR–CAPBR–UPLBR–PAR


P. monostachyum VaseyP. morichalense Davidse, Zuloaga & Filg.P. multinervium A. G. Burm.

NASASA

NAARGUR ! AMRGUR ! BR–UPL


P. mutabile ChaseP. nanum C. Wright ex Griseb.P. nelsonii Chase

NACBNA

Mexico

Mexico

MRCAR–WIPMR


P. nesiotes ChaseP. nicorae ParodiP. niquelandiae Filg.P. nudatum LucesP. nummularium Chase ex Sendulsky & A. G. Burm.

CBSASASASA

Brazil

Brazil

CAR–WIPBR–PARBR–UPLAMR ! BR–UPLBR–ATLP ! UPL


P. oteroi SwallenP. ovale Nees ex Steud.P. pallens SwallenP. pallidum KunthP. palmeri Chase

SASASASANA

Brazil

Mexico

BR–UPLBR–PARBR–UPLARMR


P. paludosum SwallenP. palustre MezP. pauciciliatum (Parodi) Herter

SASASA

Brazil BR–UPLBR–PARBR–PAR





syndrome

P. paucifolium SwallenP. peckii F. T. Hubb.P. penicillatum Hook. f.

SAMASA

BR–PARCAR–CAPAR


P. petilum ChaseP. petrense A. G. Burm.P. petrosum SwallenP. phyllorachis Hack.P. pictum Ekman

SASASASA

MA/SA

BrazilColombiaBrazil

GURBR–UPLGURBR–UPLCAR–CAP ! AMR


P. pilgerianum ChaseP. pissinum SwallenP. planum Hack.P. pontanalis SwallenP. praecox Walter

SASASASANA

Brazil

BrazilUSA

ARBR–UPLBR–PARBR–UPLNAAR


P. procurrens QuarınP. proximum MezP. quadrifarium Lam.P. quarinii Morrone & Zuloaga

SASASASA

BR–PARBR–UPL ! PARBR–PARBR–PAR


P. racemosum Lam.P. ramboi I. L. BarretoP. rectum Nees

SASASA

BrazilBrazil

ARBR–PARBR–UPL


P. redondense SwallenP. reduncum Nees ex Steud.P. redundans ChaseP. regnellii MezP. remotum J. Remy

SASASASASA

BrazilBrazilEcuador

BR–UPLBR–UPLCAR–GPBR–UPL ! PARAR


P. reptatum Hitchc. & ChaseP. restingense RenvoizeP. reticulinerve RenvoizeP. riedellii MezP. riparium Nees

CBSASASASA

BrazilBoliviaBrazil

CAR–WIPBR–UPLBR–UPLBR–UPLAMR


P. rocanum LeonP. rottboellioides C. WrightP. rufum Nees ex Steud.P. rugulosum Morrone & Zuloaga

CBCBSASA Ecuador

CAR–WIPCAR–WIPBR–PARAR


P. rupestre Trin.P. rupium RenvoizeP. saugetii Chase

CBSACB

BrazilCAR–WIPBR–UPLCAR–WIP


P. scalare Trin.P. schultesii SwallenP. scutatum Nees ex Trin.P. simplex MorongP. sodiroanum Hack.

SASASASASA

BrazilColombiaBrazil

BR–UPLGURBR–UPL ! CPBR–CHAAR


P. soukupii CarbonoP. squamulatum E. Fourn.P. strigosum Doll ex ChaseP. subciliatum ChaseP. subfalcatum (Doll) Tutin

SANA/MA

SAMA/SA

SA

Peru

Brazil

Venezuela

ARMR ! CAR–CAPBR–ATLPCAR–CAP ! AMRAMR


P. subsesquiglume DollP. telmatum SwallenP. tilletii Davidse & ZuloagaP. tinctum Chase

SASASA

NA/MA

Brazil BR–UPLBR–UPLGURMR ! CAR–CAP

C4–MSNA


P. tolucensis R. GuzmanP. trianae Pilg.P. trichophyllum Henrard

NASASA

Mexico MRARBR–UPL


P. trichostomum Hack.P. trinii SwallenP. tuberosum MezP. turriforme R. W. PohlP. umbrosum Trin.

SASASAMASA

Brazil

Peru

BR–UPLARARCAR–CAPBR–UPL ! PAR


P. urbanianum Ekman ex Hitchc.P. urvillei Steud.P. usteri Hack.

CBSASA

CAR–WIPBR–UPL ! PARBR–PAR





syndrome

P. uyucense R. W. PohlP. validum Swallen

MASA

HondurasBrazil

CAR–CAPBR–UPL

C4–MSC4–MS

P. variabile (E. Fourn.) NashP. venezuelae (Chase) A. G. Burm.P. virletii E. Fourn.

NA/MASANA

VenezuelaMexico

MR ! CAR–CAPGURMR


P. volcanensis Zuloaga, Morrone & DenhamP. zuloagae Davidse & Filg.Phanopyrum gymnocarpon (Elliott) Nash

SASANA

BrazilUSA

ARBR–UPLNAAR

C4–MSC4–MSC3

Plagiantha tenella RenvoizeReimarochloa aberrans (Doll) ChaseR. oligostachya (Munro ex Benth.) Hitchc.Spheneria kegelii (Mull. Hal.) Pilg.Steinchisma cuprea (Hitchc. & Chase) W. V. Br.

SASA

CB/NASANA

Brazil

Mexico

BR–UPLAMRNAAR ! CAR–WIPBR–UPL ! AMRMR

C3

C4–MSC4–MSC3/C4

S. decipiens (Nees ex Trin.) W. V. Br.S. exiguiflora (Griseb.) W. V. Br.S. spathellosa (Doll) RenvoizeS. stenophylla (Hack.) Zuloaga & Morrone

SACBSASA

GUR ! BR–CHA ! UPL ! PARCAR–WIPBR–PARBR–UPL ! AMR

C3/C4

C3/C4

C3/C4

C3/C4

Streptostachys asperifolia Desv.S. macrantha (Trin.) Zuloaga & MorroneS. ramosa Zuloaga & Soderstr.

CB/SASASA Brazil

AMR ! BR–UPL ! CAR–WIPBR–UPLBR–UPL


S. rigidifolia Filg., Morrone & ZuloagaTatianyx arnacites (Trin.) Zuloaga & Soderstr.Thrasya achlysophila Soderstr.T. auricoma A. G. Burm.T. axillaris (Swallen) A. G. Burm. ex Judz.

SASASASASA

BrazilBrazilGuyanaBrazil

BR–UPLBR–UPLGURBR–UPLGUR


T. crucensis KilleenT. glaziovii A. G. Burm.T. granitica A. G. Burm.T. hitchcockii Chase

SASASA

MA/SA

BoliviaBrazilSuriname

BR–UPLBR–UPLGURCAR–CAP ! AMR


T. longiligulata Bastos & A. G. Burm.T. mosquitensis Davidse & A. G. Burm.T. oreophila A. G. Burm.

SAMASA

Brazil

Brazil

BR–UPLCAR–CAPBR–UPL

NANANA

T. parvula A. G. Burm.T. paspaloides KunthT. scandens (Trin.) Soderstr. & A. G. Burm.T. schumannii (Pilg.) Pilg.T. seminuda A. G. Burm.

SACB/SA

SASASA

Brazil

BrazilBrazil

AMRCAR–WIP ! AMRGURBR–UPLGUR ! BR–UPL


T. setosa SwallenT. stricta A. G. Burm.T. thrasyoides (Trin.) ChaseThrasyopsis juergensii (Hack.) Soderstr. & A. G. Burm.

SASASASA

Venezuela

Brazil

GURGURBR–UPLBR–PAR


T. repanda (Nees ex Trin.) ParodiTriscenia ovina Griseb.

SACB

Brazil BR–UPL ! PARCAR–WIP

C4–MSC3

Subtribe Setariinae

Panicum plenum Hitchc. & ChasePaspalidium chapmanii (Vasey) R. W. Pohl

NACB/NA

MRCAR–CAP ! WIP ! NAAR

C4–MSC4–MS

Setaria alonsoi Pensiero & AntonS. arizonica RomingerS. barbinodis R. A. W. Herrm.S. cernua KunthS. cordobensis R. A. W. Herrm.

SANASASASA

Argentina

BoliviaEcuador

BR–CHAMRARARAR


S. corrugata (Elliott) Schult.S. distantiflora (A. Rich.) Pilg.S. fiebrigii R. A. W. Herrm.S. globulifera (Steud.) Griseb.S. hunzikeri Anton

CB/NACBSASASA

NAAR ! CAR–WIPCAR–WIPBR–CHABR–CHA ! PARBR–CHA


S. lachnea (Nees) KunthS. latifolia (Scribn.) R. A. W. Herrm.S. leonis (Ekman) LeonS. longipila E. Fourn.

SANACBMA

MexicoBR–CHAMRCAR–WIPCAR–CAP


S. mendocina Phil.S. oblongata (Griseb.) ParodiS. palmeri Henrard

SASANA

Argentina

Mexico

BR–CHABR–CHAMR





syndrome

S. pampeana Parodi ex NicoraS. paraguayensis PensieroS. parodii NicoraS. paucifolia (Morong) Lindm.S. pflanzii Pensiero

SASASASASA

BR–CHABR–CHABR–CHABR–UPL ! CHABR–CHA


S. pradana (Leon) LeonS. reverchonii (Vasey) Pilg.S. rosengurtii NicoraS. scabrifolia (Nees) KunthS. scheelei (Steud.) Hitchc.

CBNASASANA

CAR–WIPMRBR–CHABR–UPL ! CHAMR


S. stolonifera BoldriniS. submacrostachya LucesS. texana EmeryS. utowaneae (Scribn.) Pilg.

SASANA

CB/SA

VenezuelaBR–CHAGURMRCAR–WIP ! AMR


S. vaginata Spreng.S. variifolia (Swallen) DavidseS. villosissima (Scribn. & Merr.) K. Schum.

SAMANA

BR–CHACAR–CAPMR


Setariopsis latiglumis (Vasey) Scribn. NA Mexico MR C4–MS

Tribe Steyermarkochloeae

Arundoclaytonia dissimilis Davidse & R. P. EllisSteyermarkochloa unifolia Davidse & R. P. Ellis

SASA

Brazil AMRAMR

C3

C3

classification and biogeography of panicoideae (poaceae) in the

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