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Vol. 67 September 1989 No.2 


CONTENTS 


URBATSCH, L.E.. New combinations in Ericameria (Asteraceae: 


Aste reae) SSHSSSSSSSSSSHSSSSSSSSSSSSSOSSSSSSSSSSSSHSHSSSSSHSSSSHOSSHSSSSSSSSHHHSSSSSSSHSSOSOOSS 1 09 


TURNER, B.L., New species and combinations from México in the 
Bartlettina complex of the genus Eupatorium (Asteracede) eccccsssseee 111 


: NESOM, G.L., A new combination in Stenotus (Compositae: Astereae) - 113 


. ~TURNER, B.L., Two new species of Ageratina (Asteraceae: Eupatorieae) 
from south-central Mexico SSCSHOSSSSSSSSSSSSSSSSSSSSSSSSSSSSSOSSHSSSSSSSSSSSHSSESSSSSES 11 5 


“ GALAN, A., Senecio bolivarianus in the Peruvian central Andes esscsse. 118 
_ TURNER, B.L., An overview of the Brickellia (Kuhnia) eupatorioides 
" (Asteraceae, Eupatorieae) complex SSCHSSSSSSSSSSSSSSSSSSSSSSSSOSSOSSSCSSSSSSSEOS Oe | 


_/ ESCOBAR, L.K., Passiflora chlorina a new species of subgenus Astrophea 
(Passifloraceae) from the cerrado vegetation Of Brazil esccccccscccseeeee 132 


_ REEDER, C.G. & J.R. REEDER, A further note on Agrostis elliottiana 
. (G ramineae) in Arizona PrrTTriiiitiiiiiitiiititiiiti tr) 134 
: -BUNTING, G:S., A reconsideration of Urospathella (Araceae) escccsseses 139 


Contents continued on inside cover. 


SS HIBRARY 


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Contents (cont.) 


NESOM, G.L., New species of Mexican Solidago (Compositae: 


Astereae COSOCOOOCOOOOOOOOSOOOOOOOOOOOHSSOS HOS OSOOSSOD SOOO OSSOS SOS OSOSSOOSSOSSSSESESESES 142 


, BRIDGES, E.L. & S.L. ORZELL, A new species of Carex (sect. Oligocarpae) 
from the Edwards Plateau of Texas COCCoCCOSOCO OOOO SOOO OOSEESOCOSESSEESESECESS 148 


NESOM, G.L., Solidago simplex (Compositae: Astereae), the correct name 


for Si glutinosa CO CCOOOOOOOOO OOD OEOSOOOOOOOOOOS HOODS OO OOOSSOSOOOOSOSSSSSESOOSSOSSSSSESESS 155 


NESOM, G.L., New species, new sections, and a taxonomic overview of 
American Pluchea (Compositae: Inuleae) cccccccccccccccccccoccccccccccsoeses 1 5G 


TURNER, B.L., Revisionary treatment of the genus Sinclairia, including 
Liabellum (Asteraceae, Liabeae) SCOOSSSSSSSSSSHSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSESEEE 168 


WARNOCK, M.J ap Book reviews COOCCOOOOOOOOOOOOOOOOOCOSOSOSOSOOOOOOOOOSOESESEEEESS 207 


Phytologia (September 1989) 67(2):109-110. 


NEW COMBINATIONS IN ERICAMERIA 
(ASTERACEAE: ASTEREAE) 


Lowell E. Urbatsch 
Department of Botany, Louisiana State University, Baton Rouge, Louisiana 
70803-1705 U.S.A. 


ABSTRACT 


Two new combinations are made in Ericameria. These are E. 
juarezensis and E. cooperi var. bajacalifornica. 


KEY WORDS: Systematics, Ericameria, México, Asteraceae. 


In preparation for the Asteraceae of México by B.L. Turner and G. Nesom, 
two new combinations need to be made in the genus Ericamerza. 


Ericameria juarezensis (Moran) Urbatsch, comb. nov. BASIONYM: Hap- 
lopappus juarezensis Moran. Trans. San Diego Soc. Nat. Hist. 15:154- 
155. 1969. TYPE: MEXICO. Baja California del Norte: 5 mi by road 
SE of Rancho las Filipinas, Sierra Juarez (near 31°47’N, 115°48’W), oc- 
casional on granitic hills, ca 1575 m, 17 Sep 1966, Moran 13556. (HOLO- 
TYPE: SD!; Isotypes: CAS!,DAV!,GH!,MICH!,NY!, RSA!,UC!,US!). 


Ericameria is sometimes accorded sectional status in the genus Haplopap- 
pus (Hall 1920; Keck 1959). That Ericameria warrants generic recognition 
has been reasonably well established (Clark 1979; Urbatsch & Wussow 1979) 
making the present combination necessary for this member of the E. cuneata 
(A. Gray) McClatchie species group. 


Ericameria cooperi (A. Gray) Hall var. bajacalifornica (Urbatsch & Wus- 
sow) Urbatsch, stat. nov. BASIONYM: Ericameria coopert (A. Gray) 
Hall ssp. bajacalifornica Urbatsch & Wussow. Brittonia 31:274. 1979. 
TYPE: MEXICO. Baja Califérnia del Norte: 2 mi N of Llano Colorado 
(31°39’N, 115°55’W), ca 1125 m, 20 May 1967, R. Moran 13874 (HOLO- 
TYPE: SD!; Isotypes: ARIZ!,CAS!,;COLO!,GH!,LL!,MICH!,RSA!,UC!, 
US!). 


The Asteraceae of México format specifies consistent treatment of infraspe- 
cific taxa at the varietal level, necessitating the present combination. 


109 


110 Payer O Ga A volume 67(2):109-110 September 1989 


ACKNOWLEDGMENTS 


I am grateful to Guy Nesom and B.L. Turner for calling this problem to 
my attention and for their willingness to review the paper itself. 


LITERATURE CITED 


Clark, W.D. 1979. The taxonomy of Hazardia (Compositae: Astereae). 
Madrono 26:105-127. 


Hall, H.M. 1928. The genus Haplopappus: a phylogenetic study in the Com- 
positae. Publ. Carnegie Inst. Wash. No. 389. 


Keck, D.D. 1959. in P.A. Munz A California Flora. University of California 
Press, Berkeley. 


Urbatsch, L.E. & J.R. Wussow. 1979. The taxonomic affinities of Haplopap- 
pus lineartfolius (Asteraceae: Astereae). Brittonia 31:265-275. 


Phytologia (September 1989) 67(2):111-112. 


NEW SPECIES AND COMBINATIONS FROM MEXICO IN THE 
BARTLETTINA COMPLEX OF THE GENUS EUPATORIUM (ASTERACEAE) 


B.L. Turner 
Department of Botany, University of Texas, Austin, Texas 78713 USA 


ABSTRACT 


A new species, Eupatorium calderonii, and 4 new combinations 
in the Bartlettina complex of Eupatorium (sensu lato) are proposed. 
All of the taxa requiring new names were previously treated within 
the genus Bartlettina by King & Robinson (1987). These include E. 
breedlovei (King & H. Robins.) B. Turner, E. cronquistii (King & 
H. Robins.) B. Turner, E. hintonii (King & H. Robins.) B. Turner 
and E. macdougallii (King & H. Robins.) B. Turner. 


KEY WORDS: Asteraceae, Eupatorieae, Eupatorium, Bartlettina, 
México. 


Preparation of a systematic treatment of the Bartlettzna group of Eupa- 
torium of the tribe Eupatorieae (Asteraceae) for México has necessitated the 
description of a new species and several name changes. King & Robinson 
(1987) have presented a brief account of their genus Bartlettzna in which they 
recognized 35 species. I have prepared a treatment of this taxon (Turner un- 
publ.) and note that the genus, as delimited by King & Robinson, appears 
to merge into Hebeclinium, Guayania, as well as other genera in South Amer- 
ica. Because of this I have opted to treat Bartlettzna within my broad view of 
Eupatorium. 


Eupatorium calderonii B. Turner, sp. nov. TYPE: MEXICO. Oaxaca: 
Mcpio. Comaltepec, Puerto Eligio, km 149 de la carretera Tuxtepec- 
Oaxaca, Sierra Juarez, 800 m, 26 Mar 1966, G. Martinez Calderon 770 
(HOLOTYPE: MEXU!). 


E. sordido Less. similis sed foliis anguste elliptici-ovatis petiolis 
0.5-1.0 cm longis, capitulis flosculis 20-30, et lobis corollarum dense 
puberulis differt. 


111 


112 Psy, Te LOG TA volume 67(2):111-112 September 1989 


Perennial suffruticose densely pubescent herbs or shrubs to 2 m high. 
Leaves lanceolate-elliptic, 10-15 cm long, 3.5-5.5 cm wide, tapering at both 
ends, widest at or near the middle, pinnately nerved, the margins minutely 
serrulate; petioles 0.5-1.0 cm long. Heads numerous, arranged in somewhat 
pyramidal corymbose panicles, the ultimate peduncles 1-3 mm long. Involucres 
7-8 mm high, campanulate, 3-4 seriate, the bracts gradate ovate to lanceolate, 
the inner bracts acute, hirsutullous. Receptacles convex, sclerose-alveolate, 
glabrous. Florets 20-30, the corollas purple, 4-5 mm long, glabrous except 
for the pubescent lobes, the latter ca 0.5 mm long. Achenes ca 2 mm long, 
sparsely hispid apically, the pappus of 35-40 white persistent bristles, 4-5 mm 
long. 

The species is closely related to the widespread E. sordidum Less. but has 
narrowly elliptic-ovate (vs broadly ovate to cordate) leaves with petioles 0.5- 
1.0 cm long (vs 2-10 cm); it also has fewer florets to a head (20-30 vs 50-200) 
and pubescent corolla lobes (vs glabrous or nearly so). 


Eupatorium breedlovei (King & H. Robins.) B. Turner, comb. nov. Based 
upon Bartlettina breedlove: King & H. Robins., Phytologia 28:286. 1974. 


Eupatorium cronquistii (King & H. Robins.) B. Turner, comb. nov. Based 
upon Bartlettina cronguistu King & H. Robins., Phytologia 38:108. 1977. 


Eupatorium hintonii (King & H. Robins.) B. Turner, comb. nov. Based 
upon Bartlettina hintoni King & H. Robins., Phytologia 31:62. 1975. 


Eupatorium macdougallii (King & H. Robins.) B. Turner, comb. nov. 
Based upon Bartletttna macdougallii King & H. Robins., Phytologia 
38:107. 1977. 


ACKNOWLEDGMENTS 


I am grateful to Dr. Guy Nesom for the Latin diagnosis and to both him 
and Dr. Carol Todzia for reviewing the manuscript. 


LITERATURE CITED 


King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae (Aster- 
aceae). Monographs Syst. Bot. 22:1-580. Missouri Bot. Garden, St. 
Louis. 


Phytologia (September 1989) 67(2):113-114. 


A NEW COMBINATION IN STENOTUS 
(COMPOSITAE: ASTEREAE) 


Guy L. Nesom 
Department of Botany, University of Texas, Austin, Texas 78713 USA 


ABSTRACT 
A new combination in Stenotus is presented. 


KEY WORDS: Stenotus, Haplopappus, Asteraceae, México, sys- 
tematics. 


The following combination is necessary in a taxonomic study of the Mexican 
taxa included by Hall (1928) as Haplopappus. 


Stenotus pulvinatus (R. Moran) Nesom, comb. nov. Based on Haplopappus 
pulvinatus R. Moran, Trans. San Diego Soc. Nat. Hist. 15:161. 1969. 


Stenotus pulvinatus, which is endemic to the Sierra San Pedro Martir of 
Baja California Norte, México, clearly belongs with the group of five other 
species circumscribed by Hall (1928) as Haplopappus sect. Stenotus (Nutt.) A. 
Gray. This relationship was noted by Moran (1969) in the original description 
of the species. In the separation of Haplopappus into component monophyletic 
groups, however, I believe that Stenotus Nutt. (1840) stands as a distinct 
genus. It is not congeneric with any other species of Mexican “Haplopappus.” 
The species of Stenotus range from Canada through the western United States 
to Baja California Norte and include S. acaulis Nutt. (the type), 5. armerioides 
Nutt., S. lanuginosus (A. Gray) E. Greene, S. macleani (Brandegee) Heller 
and S. stenophyllus (A. Gray) E. Greene. Stenotus pulvinatus is the only 
species of the genus with discoid heads, and in other features “it represents an 
extreme of reduction and compaction . . .” (Moran 1969, p. 163). 

Two of the six or seven species of Haplopappus sect. Tonestus (A. Nelson) 
H.M. Hall (the genus Tonestus A. Nelson, 1904) include names as Stenotus 
among their synonyms. The two groups are similar in their herbaceous habit, 
primarily monocephalous stems, 3 veined leaves, herbaceous, 3 veined phy]l- 
laries in 2-3 series and style appendages (disc flowers) with minute, short and 
dense collecting hairs. The species of Tonestus differ conspicuously from those 
of Stenotus in their leafy stems and heads immediately subtended by leaf-like 


113 


114 PHYTOL OGIA volume 67(2):113-114 September 1989 


bracts. A clear understanding of the relationship between Stenotus and Ton- 
estus has yet to be reached. Haplopappus alpinus L. Anderson & Goodrich, H. 
aberrans (A. Nelson) H.M. Hall and H. graniticus Tiehm & L. Shultz (of Ton- 
estus, the last species somewhat anomalous but best placed there related to 
the previous two) have several headed capitulescences and in that respect are 
at least superficially similar to species of sect. Macronema in habit. This fur- 
ther complicates the taxonomic solution and suggests that Macronema Nutt. 
(1840) may ultimately have priority as the generic name for all these species. 


ACKNOWLEDGMENTS 


I thank Drs. Billie Turner and Linda Escobar for their comments. 


LITERATURE CITED 


Hall, H.M. 1928. Sect. Stenotus, in The Genus Haplopappus—A phylogenetic 
study in the Compositae. Carnegie Inst. Washington, Publ. 389:161-171. 


Moran, R. 1969. Five new taxa of Haplopappus (Compositae) from Baja 
California, Mexico. Trans. San Diego Soc. Nat. Hist. 15:149-164. 


Phytologia (September 1989) 67(2):115-117. 


TWO NEW SPECIES OF AGERATINA 
(ASTERACEAE: EUPATORIEAE) FROM 
SOUTH-CENTRAL MEXICO 


B.L. Turner 
Department of Botany, University of Texas, Austin, Texas 78713 USA 


ABSTRACT 


Two new species of Ageratzna, A. chazaroana and A. perezii, 
belonging to the subgenus Ageratina, are described from the states of 
Veracruz and Morelos, México. 


KEY WORDS: Asteraceae, Eupatorieae, Ageratina, Mexico. 


Identification of material for a treatment of the Asteraceae of México 
(Turner & Nesom, in prep.) has necessitated description of the following new 
species of Ageratina. 


Ageratina chazaroana B. Turner, sp. nov. TYPE: MEXICO. Veracruz: 
Mcpio. Coatepec, ca Tierra Grande, between Mesa de Los Laureles and 
Cuesto del Pino, 30 Aug 1986, M. Chazaro B. & P.H. de Charazo 3993-a 
(HOLOTYPE: TEX!; Isotypes: TEX!,WIS!,XAL!). 


A. ciliatae (Less.) King & H. Robins. similis sed foliis sessilibus 
glabris et involucris glabris differt. 


Erect perennial herbs, 30-60 cm high, the stems moderately pubescent with 
crinkled multiseptate trichomes and arising from slender rhizomes which pre- 
sumably form colonies. Leaves opposite throughout, 3-5 cm long, 2-4 cm wide, 
sessile or nearly so; blades broadly ovate, 3-5 nervate from the base, sparsely 
pubescent along the veins, otherwise epunctate and glabrous. Heads 10-20 
in rather congested terminal or subterminal corymbs, the ultimate peduncles 
mostly 1-5 mm long. Involucres 4-5 mm high, campanulate, the bracts bise- 
riate, subequal, glabrous, except for the obtuse ciliate apices. Florets 15-25, 
corollas white, ca 3 mm long, the tube ca 1 mm long, the limb ampliate, ca 
2 mm long, the lobes, at least some of them, with a few well developed erect 
trichomes. Achenes ca 1.5 mm long, sparsely hispid along the ribs, the pappus 
of ca 20 delicate bristles, ca 3 mm long, in a single series. 


115 


116 PHY TOLO GIA volume 67(2):115-117 September 1989 


The species belongs to the subgenus Ageratina, having the habit and stem 
pubescence of A. ciliata, but differs markedly from that species in having 
sessile, nearly glabrous, coarsely serrate leaves. It is named for its major 
collector, Miguel Charazo, prolific plant collector working out of XAL. 


Ageratina perezii B. Turner, sp. nov. TYPE: MEXICO. Morelos: Mcpio. 
Tepoztlan, ca 8.5 km N Santo Domingo Ocotitlan, ca 2870 m, “bosque 
de pino-encino con zacatonal,” 11 Oct 1987, M.L. Espin, E. Cedillo, A. 
Villasenor & A. Vargas 259 (HOLOTYPE: TEX!; Isotype: CHAPA!). 


A. parayanae (Espinosa) B. Turner similis sed differt foliis bre- 
vipetiolatis deltatis et capitulis in pedunculis longioribus in capit- 
ulescentia corymbosa terminali disposita. 


Suffruticose perennial herb or shrub to 1.6 m high. Stems terete, striate, 
minutely pubescent with glandular, multiseptate trichomes, 0.5 mm long or 
less. Leaves opposite or subopposite below, those along the upper portions of 
the stem clearly alternate or subopposite, 5-12 cm long, 4-8 cm wide; peti- 
oles thick, 5-12 mm long; blades deltoid, 3 nervate from the base, sparsely 
pubescent above and below, the margins rather evenly crenulate. Heads nu- 
merous, arranged in rather strict, broad, flat topped, terminal cymes, the 
ultimate peduncles densely glandular pubescent, 1-2 cm long. Involucres 9-10 
mm high, the bracts 13-15, linear lanceolate, glandular pubescent, the apices 
acute. Florets 20-50; corollas white, 5-6 mm long, the lobes ca 0.5 mm long, 
decidedly pubescent with 2 or more multiseptate trichomes. Achenes linear, 
ca 4 mm long, appressed hispidulous, the pappus uniseriate, of 30-40 white 
bristles, mostly 5-6 mm long. 

According to label data, the species is an erect shrub to 1.6 m high, much 
branched from the base, the florets white and involucral bracts coffee red. It 
is said to be abundant in clearings of the forest. 

Ageratina perezti is superficially similar to A. cardiophylla (B.L. Robins.) 
King & H. Robins., but the latter is quite distinct, possessing mostly cordate 
leaves with much longer petioles, and belongs to the subgenus Neogreenella 
(sensu King & Robinson 1987), whilst the former belongs to the subgenus 
Ageratina, where it apparently relates to A. parayana (Espinosa) B. Turner. 
The latter taxon is readily distinguished by its longer petioles and smaller 
heads on shorter ultimate peduncles. 

It is a pleasure to name this taxon for José Garcia Pérez, long time herbar- 
ium assistant at CHAPA and dedicated worker on Mexican composites. 


Turner: New species of Ageratina ibe 


ACKNOWLEDGMENTS 


I am grateful to Dr. Guy Nesom for the Latin diagnoses and to him and 
Dr. Andrew McDonald for a critical review of the paper. 


LITERATURE CITED 


King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae (Aster- 
aceae). Monographs Syst. Bot. 22:1-580. Missouri Bot. Garden, St. 
Louis. 


Phytologia (September 1989) 67(2):118-120. 


SENECIO BOLIVARIANUS IN THE PERUVIAN 
CENTRAL ANDES 


Antonio Galan 
Departamento de Botanica, Facultad de Farmacia, Universidad Complutense, 
28040, Madrid ESPANA 


ABSTRACT 


A range extension and ecological characteristics are reported for 
Senecio bolivarianus. 


KEY WORDS: Asteraceae, Senecio, Peru, phytogeography. 


Senecio bolivarianus was described by Cuatrecasas (1969) based on indi- 
viduals coming from the regions of Huancayo and Urubamba, Peru. (TYPE: 
PERU. Huancayo: regién de San José de Acobambilla, NW slope Huacrav- 
ilca, 14800-15200 ft, cracks in granite, adpressed to crack, 27 Jul 1961, J.R. 
Lloyd & J.K. Marshall 257 (HOLOTYPE: K; Isotype: US]. Paratype: PERU. 
Cuzco: Urubamba, Pumahuanca, Cuyo, 4000-4400 m alt, en rocas humedas, 
13 Oct 1961, César Vargas 13702 [US]). During trips to central Peru, we have 
found a new location: PERU. Junin. Huancayo: Huaytapallana Peak, Cocha 
Grande Lagoon, NE964801; 4400 m, 19 Sep 1987, O. Tovar, M. Chanco & A. 
Galan 1294 (MAF 127799, USM). 

This new location is important because this species is an indicator of areas 
where glacial melt during the dry season causes a rupicolous condition with 
oozing water on vertical rocks. Such conditions are present at all three lo- 
calities. Populations of this plant appear as monospecific quilt-like mats and 
are found in fissures caused by glacier tongue erosion (phytosociological inven- 
tories: 50 m? area, 10% cover, SE orientation, index 2.2,3.3; location Huay- 
tapallana Peak). In the Puna region (sensu Rivas Martinez & Tovar 1983), 
the mid-minimum temperatures of the coldest month -m- (Rivas Martinez, 
Tovar & Galan 1988) decrease toward the south, thus allowing the formation 
of glaciers and probably accounting for the distribution of glacial melt plant 
communities in central and southern Peru. 


118 


New record for Senecio bolivarianus 119 


Senecio bolivarianus Cuatr. in Peru 


Cited locations 
New location 
In some weather bureaus 


Conococha =- Ancash (4020 m) - 4,6 
Pachachaca - Junf{n (4000 m) - 6,6 
Accnococha = Huancavelica (4520 m) - 7,4 
Angostura - Arequipa (4155 m) -11,1 
Imata - Arequipa (4436 m) -12,5 


Collacachi = Puno (3900 m) -14,5 


120 PHY TODOGLIA volume 67(2):118-120 September 1989 


LITERATURE CITED 


Cuatrecasas, J. 1969. Observaciones sobre Compositae. An. Esc. Nac. Cienc. 


Biol. Méx. 18:9-15. 


Rivas Martinez, S. & O. Tovar. 1983. Sintesis Biogeografica de los Andes. 
Collect. Bot. (Barcelona) 14:515-521. 


Rivas Martinez, S., O. Tovar & A. Galan. 1988. Pisos Bioclimdticos y Cul- 
tivos del Pert. ICI-INP, Madrid. 


Phytologia (September 1989) 67(2):121-131. 


AN OVERVIEW OF THE BRICKELLIA (KUHNIA) 
EUPATORIOIDES (ASTERACEAE, EUPATORIEAE) COMPLEX 


B.L. Turner 
Department of Botany, University of Texas, Austin, Texas, USA, 78713 


ABSTRACT 


Kuhnia eupatorioides is the generitype and principal species of the 
genus Kuhnia, which was retained by most workers up to the time 
of Shinners (1971), who reduced all of its members to synonymy un- 
der Brickellia. Subsequently, many workers have followed this submer- 
gence. Shinners recognized 7 species in the complex: B. schaffneri, B. 
leptophylla (with 2 varieties), B. chlorolepis, B. oretthales, B. mosieri, 
B. eupatorioides (with 4 varieties) and B. adenolepis. The distribution 
and morphological characters which distinguish these taxa are discussed 
and it is proposed that among those plants known to Shinners, only 
3 species are involved: B. oreithales (a Mexican, montane element of 
SW Chihuahua and adjacent Durango); B. adenolepis (a local endemic 
of Jalisco, México); and B. eupatorioides (a widespread species of the 
U.S.A. and adjacent Mexico). The latter species is believed to be com- 
posed of 6 regional varieties, 5 of which are contiguous, and all but one 
of which intergrade to some considerable extent with their allopatric 
consorts; these are: 1) var. eupatoriotdes of the eastern U.S.A., which 
grades into var. tezana to the southwest and into the more western var. 
corymbulosa to the west and northwest; 2) var. tezana (including var. 
ozarkana), which grades into the var. corymbulosa to the west and with 
var. eupatortoides to the east; 3) var. corymbulosa, which grades into 
the var. eupatoriotdes to the northeast and into the var. tezana to the 
southeast and into the more montane var. chlorolepis to the west and 
south; 4) var. gracillima, of south central Texas, which grades into the 
var. chlorolepis to the west; 5) var. floridana (of which B. mosier: is 
a synonym), which is disjunct in south Florida from the other taxa; 
and 6) var. chlorolepts which occurs in the more montane western por- 
tions of the U.S.A. extending southwards along the Central Plateau to 
south central México. The var. chlorolepis contains as synonyms the 
following taxa recognized by Shinners: B. chlorolepis, B. microphylla, 
B. schaffneri and B. leptophylla var. mezicana. A key to the various 
taxa is provided and maps showing their distributions are constructed. 


121 


122 PHY TOL O1G I A volume 67(2):121-131 September 1989 


The only new combinations needed to accommodate the above treat- 
ment are B. eupatorioides var. chlorolepis (Woot. & Standl.) B. 
Turner; B. eupatorioides var. gracillima (A. Gray) B. Turner; and 
B. eupatorioides var. floridana (R. Long) B. Turner. 


KEY WORDS: Asteraceae, Eupatorieae, Brickellia, Kuhnia, United 
States, México. 


The genus Kuhnia has been retained as distinct from Brickellia by most 
workers (Robinson 1917; Fernald 1950; Steyermark 1963; Correll & Johnston 
1970; Barkley 1986; etc.) up to the time of its submergence into the latter by 
Shinners (1971). Subsequent workers (e.g. McVaugh 1984; King & Robinson 
1987), including the present author, have agreed with this submergence, there 
being few, if any, characters by which to distinguish between them. 

Shinners (1946) rendered a definitive revision of the group, treating these as 
I have indicated in the above abstract. His was an excellent treatment, based 
upon the study of several thousand specimens and, to a limited extent, field 
work with most of the taxa in the United States. For a complete synonymy of 
the taxa recognized in the present paper, the interested reader is referred to 
the treatments of Shinners (1946; 1971). Where I differ with his nomenclature, 
this is discussed in the account that follows. 

Shinners was most familiar with the widespread Brickellia eupatorioides 
and treated this as comprising 4 regional varieties: an eastern var. eupato- 
rioides, a western var. corymbulosa, and the var. ozarkana and var. tezana, 
which together occupied a relatively narrow range between the varieties eu- 
patorioides and corymbulosa. Shinners was well aware that these several taxa 
intergraded and cited several intermediates. Unfortunately, he tended to iden- 
tify most of the plants he examined (by annotation or citation) as either this 
or that variety, without indication of intermediacy, tending to rely on only 
1 or 2 of his “key characters” for recognition purposes. This resulted in the 
occasional occurrence of “typical” elements outside of the purported range of 
his recognized varieties; though, for the most part, his arrangement is solid, 
(except for his ignoring, by annotation, the many intermediate plants he must 
have examined). 

The complexity of the Brickellia eupatorioides group is well illustrated in 
the treatment of this complex by Steyermark (1963), who mapped 3 varieties 
as occurring in Missouri. As shown by his dot map, var. eupatorzoides occurs 
throughout the state; var. corymbulosa occurs mostly in the northwestern por- 
tion of the state; and var. terana (as var. angustifolia) occurs in the central 
and southern portions of the state. In 2 widely separated counties of Missouri, 
he shows the occurrence of all 3 of these varieties in each of the counties. 
Indeed, the state of Missouri is an area, within the broader region of B. eupa- 
tortoides, where several of the varieties of B. eupatorioides come into contact 


Turner: Overview of Brickellia eupatoriotdes complex 123 


and presumably intergrade, hence the difficulty in assigning names, especially 
where individuals are labeled as opposed to populations. It is unlikely that 
this intergradation is due to present day hybridization between distinct taxa; 
rather, the intergradation is probably more the result of a sorting out of this 
or that combination of characters due to local or regional climatic or edaphic 
factors so that a mosaic of characters might occur in regions of contact. In any 
case, the only utilitarian way to treat such variation is to provide names for 
the populational units concerned as best one can, based upon the characters 
that, in combination, best exemplify those populations. 

In the overview that follows, I have treated var. terana and var. ozarkana, 
both recognized by Shinners (1946; 1971), as the same; he distinguished these 
largely by floret number (the former said to possess 18-33 florets per head vs 
10-14 florets) and little else. Because of this, I see no point in the recognition 
of 2 taxa. Possibly the var. terana (including var. ozarkana), with its filiform 
outer phyllaries, evolved in siiu out of the ancestral complex which gave rise 
to the var. corymbulosa to the northwest and var. eupatorzozdes to the east. 

Shinners recognized B. chlorolepis as a good species, differing from B. eu- 
patorioides by its sessile or subsessile, mostly narrower leaves, and reportedly, 
fewer heads in looser clusters on longer peduncles. Nevertheless, he clearly 
noted (1951, p. 125) that “intermediates with [B. e. var. corymbulosa| occur 
in eastern Colorado, northwestern Texas and adjacent New Mexico” which 
is painfully so, and because of this I reduce B. chlorolepis to varietal status 
under B. eupatorioides. The var. chlorolepis largely occupies the drier, more 
montane, portions of Colorado, New Mexico, Arizona and Texas, extending 
southwards along the Central Plateau of México to near México City. The 
var. corymbulosa occurs primarily in the great central grasslands of the U.S.A. 
from Canada to central Texas. 

Within México, Shinners (1946) proposed Kuhnza microphylla to accommo- 
date specimens from the southern Central Plateau which were said to be small 
(10-30 cm high) with relatively large involucres (8.8-11.2 mm high). He subse- 
quently resurrected the name Brickellia schaffneri for this (because of changes 
in the Code of Botanical Nomenclature) and the latter name was accepted by 
McVaugh (1984) in his treatment of Brickellia for the Flora Novo-Galiciana. 
McVaugh noted, however, that the ranges of B. chlorolepis and B. schaffnert 
(sensu Shinners) appear to coincide in the Central Plateau of México and that 
“it is very difficult to distinguish between these taxa.” Indeed, I cannot treat 
B. schaffneri as anything but a synonym of var. chlorolepis. 

Amongst the Mexican taxa recognized by Shinners (1946; 1971), this leaves 
only B. leptophylla var. mezicana, which is typified by material from near Cd. 
Durango, Durango; the only other specimens cited by Shinners were thought 
to be doubtful, perhaps belonging to B. chlorolepis, these being a Pringle 
collection from near Chihuahua City, Chihuahua and a Wright collection from 
New Mexico (U.S.A.). In my opinion, all of these are but leaf forms of var. 


124 PHY POLO GiA volume 67(2):121-131 September 1989 


chlorolepis. Certainly, they do not relate directly to B. leptophylla (=B. e. 
var. gracillima). The latter is typified by material from central Texas and is 
treated here as a localized varietal endemic, largely because it seems to be 
confined to stream bottoms along the Balcones Escarpment. 

The map (Figure 1) showing the general distribution of Brickellia eupato- 
rioides and its 6 varieties in the U.S.A. is based upon the numerous citations 
of Shinners (1946) and several hundred collections assembled since his study 
(TEX,LL). The map (Figure 1) showing the general distribution of B. eupa- 
torioides, and the dot maps of B. adenolepis and B. orezthales in México, are 
mostly based upon the numerous specimens of these taxa housed at LL and 


TEX. 
KEY TO TAXA WITHIN THE B. EUPATORIOIDES COMPLEX 


1. Mature heads mostly pendulous or nodding, in a raceme-like capitules- 
cence; w Chihuahua and Durango (México) ............ 2. B. oreithales 


1. Mature heads not pendulous, mostly erect or ascending in a terminal or 
subterminal, stiffly branching capitulescence ....................-.. (2) 


2. Involucral bracts, some or most of them, with glandular ciliate mar- 
gins; montane mesophytic habitats of S Zacatecas, Aguascalientes 
and Jalisco (Mexico) .2!%:..002.....J0. -aot eee 1. B. adenolepis 


2. Involucral bracts without glandular ciliate margins; widespread, 


central. Mesteaiyscies eee sae eee 3. Cab akorsisde. (3) 


eC | A ee an I ae (4) 
4. Heads with 9-13 florets; southernmost Florida ....... var. floridana 
4. Heads with 15-25 florets; south central Texas ....... var. gracillima 


5. Mid stem leaves predominantly sessile or subsessile, the blades mostly 2-9 
mm wide; ultimate peduncles mostly 1-10 cm long; montane regions of 
western U.S.A. (Colima, New Mexico, Arizona, W Texas) and Central 
Piatedu of México. ...20 525.41. sono eee var. chlorolepis 


5. Mid stem leaves mostly with petioles 1-10 mm long, the blades mostly 6-50 
mm wide; ultimate peduncles mostly 0.2-2.0 cm long; eastern temperate 
deciduous forests and grasslands of the central U.S.A. ............. (6) 


Overview of Brickellia eupatorioides complex 125 


Turner: 


‘senjeuen Bulmoys sepiouojyedne ‘g JO uONNquysip pezijei9ued “*} “B4 


sidajo1i0jyo 


a Buy) 19e16 
BuepIIO|} : 
euexe} 


{ 
SU Ranenes 


Te 8. 


seploiojedna 


A seas 
oS 


; fas 
CSE | {que vi iy ides coe i ED = AES 


4 


ave si tt ere AY Sava re ele Paks 
AY #1) ~~ SEETL a 
: Som 


BsojnqwAs09 


ad 


126 PHYTOL OG ILA volume 67(2):121-131 September 1989 


6. Middle and outer involucral bracts mostly with elongate, filiform, 
often contorted, apices, usually 1/2 as long or more than the inner 
bracts; central Texas northeastwards to S Illinois ..... var. terana 


6. Middle and outer involucral bracts acute or merely acuminate, 
mostly not contorted and usually 1/2 or less as long as inner 
DraCtS 2. a. eae e eo nee oon dn ieee wee er (7) 


7. Involucres mostly 7-11 mm high; florets mostly 6-15 per head; mostly 
sandy or silty soils in forested areas of the eastern 
WS .AS. ons sein so beer eiloone sa be ane oe var. eupatorioides 


7. Involucres mostly 9-15 mm high; florets mostly 15-35 per head; mostly 
calcareous or silty clay soils in grassy areas of the central and western 
1. Cae mM ee I er rorya ac var. corymbulosa 


1. Brickellia adenolepis (B.L. Robins.) Shinners, Sida 4:274. 1971. BA- 7 
SIONYM: Kuhnia adenolepis B.L. Robins. 


Distribution as shown in Figure 3 where it reportedly occurs in pine-oak 
forests from 1500 to 2300 m (McVaugh 1984). The species appears closely 
related to B. oreithales, both taxa possessing simple, mostly unbranched, stems 
and relatively naked capitulescences. McVaugh, who provided a thorough 
description, noted that the heads of B. adenolepis may droop prior to anthesis, 
but.at maturity these become stiffly erect on slender peduncles, much like those 
of B. eupatorioides. So far as known, the small, peculiar, brown, glandular 
hairs found along the margins of the involucral bracts are unique among the 
Kuhnia complex. 


2. Brickellia oretthales (B.L. Robins.) Shinners, Sida 4:274. 1971. BA- 
SIONYM: Kuhnia oreithales B.L. Robins. 


Brickellia extranea McVaugh. 


Kuhnia triplinervis S.F. Blake. 


Distribution as shown in Figure 3. Closely related to B. adenolepis but 
readily distinguished by several characters, as noted in the above key. Mc- 
Vaugh (1984) compared his B. eztranea with B. adenolepis but in my opinion 
it is synonymous with B. orezthales, differing from the latter only in its some- 
what shorter hairs along the pappus bristles. The type is from the southern- 
most portion of the distribution of B. oreithales and the length of the pappus 
hairs becomes somewhat longer northwards. Brickellia oreithales is occasion- 
ally mistaken for B. eupatorioides var. chlorolepis, with which it is partially 
sympatric; the latter however, occurs in drier sites at lower elevations. 


Turner: 


Overview of Brickellia eupatorioides complex 


127 


Fig. 2. Distribution of var chlorolepis_in Mexico. 


128 PHY TO LOG EA volume 67(2):121-131 September 1989 


‘peyeoipu! se “dds BTTSyOIq jo uonnquisiq ‘¢e *Bi4 


Turner: Overview of Brickellia eupatorioides complex 129 


4. Brickellia eupatorioides (L.) Shinners, Sida 4:274. 1971. var. eupatorzordes 


Distribution as shown in Figure 1. Shinners (1946) has given a fairly com- 
plete synonymy. I add here the recently proposed Brickellia rosmarinifolia 
(Vent.) W.A. Weber, Phytologia 53:187. 1985, at least as to intent (for which 


see the discussion under var. chlorolepis, below). 


4a. B. eupatorioides var. terana (Shinners) Shinners, Sida 4:274. 1971. Kuh- 
nia eupatorioides var. terana Shinners. 


Brickellia eupatorioides var. ozarkana (Shinners) Shinners. Kuhnia eupatorn- 
oides var. ozarkana Shinners. 


Distribution as shown in Figure 1. Its relationship to the other varieties is 
discussed in the above introduction. 


4b. B. eupatorioides var. corymbulosa (Torr. & Gray) Shinners, Sida 4:274. 
1971. 


Distribution as shown in Figure 1. Shinners (1946) has given a complete 
synonymy. 


4c. B. eupatorioides var. chlorolepis (Woot. & Standl.) B. Turner, comb. 
nov. Based upon Kuhnia chlorolepis Woot. & Standl., Contr. U.S. Natl. 
Herb. 16:177. 1913. Kuhnia rosmarinifolia var. chlorolepis (Woot. & 
Standl.) Blake (1940). Brickellia rosmarinifolia ssp. chlorolepis (Woot. 
& Standl.) W.A. Weber (1983). 


Clavigera corymbosa DC. (1836). Brickellia corymbosa(DC.) A. Gray (1852). 
Coleosanthus corymbosus (DC.) Kuntze (1891). 


Kuhnia schaffnerit A. Gray (1882). 


Kuhnia leptophylla var. mezicana Shinners (1946). Brickellia leptophylla var. 
mezicana (Shinners) Shinners (1971). 


Kuhnia microphylla Shinners (1946). 


Distribution as shown in figures 1 and 2. This is a relatively common 
roadside plant across most of the Central Plateau of México. As indicated in 
the above synonymy, if treated as a variety of B. eupatoriozdes, the earliest 
available name for that category is var. chlorolepis, Blake having proposed 
this in 1940. The automatic name, var. rosmarinifolia, cannot apply to this 
taxon, or any other taxon of the Brickellia eupatorioides complex, since it 
is apparently typified by an element that probably does not belong to the 
complex, perhaps not even to Brickellia itself (Shinners 1946 p. 143). 


130 PEYETOLrGCGCTA volume 67(2):121-131 September 1989 


I have included Brickellia corymbosa(DC.) A. Gray in synonymy here (not 
to be confused with B. eupatorioides var. corymbulosa) since, by its description, 
type locality (vicinity of Leén, Guanajuato, México, Mendez s.n., G-DC) and 
my examination of the excellent microfiche of the type (G-DC!), the plant 
concerned appears to belong to the var. chlorolepis. Nevertheless, Robinson 
(1918), McVaugh (1984) and King & Robinson (1987) maintain the species. 
McVaugh, however, keyed it next to his B. schaffneri (=B. eupatorioides var. 
chlorolepis of the present treatment), distinguishing between these largely by 
the length of the lateral hairs along the pappus bristles, a weak and variable 
character for specific recognition. 


4d. B. eupatorioides var. floridana (R.W. Long) B. Turner, comb. nov. 
Based upon Kuhnia eupatorioides var. floridana, R.W. Long, Rhodora 
72:39. 1970. 


Kuhnia mosieri Small, Man. S.E. Fl. 1329,1508. 1933. Brickelha moszera 
(Small) Shinners, Sida 4:274. 1971. 


Distribution as shown in Figure 1. Long, correctly I think, reduced this 
taxon to varietal rank, placing it under the generic name Kuhnia. Unfortu- 
nately, he provided a new varietal name based upon a newly designated type. 
If treated at the varietal level, this is the correct name to be used, hence the 
new combination within Brickellza as adopted here. 


4e. B. eupatorioides var. gracillima (A. Gray) B. Turner, comb. nov. 
Based upon Kuhnia eupatorioides var. gracillima A. Gray, Boston J. 
Nat. Hist. 6:218. 1850. Kuhnza rosmarinifolia var. gracillima (A. Gray) 
Blank (1907). 


Kuhnia leptophylla Scheele (1848). Brickellia leptophylla (Scheele) Shinners 
(1971). 


Distribution as shown in Figure 1. If treated at the varietal level, this 
taxon must take the name var. gracillima, in spite of the earlier specific name, 
K. leptophylla. 

The var. gracillima is clearly closely related to, and probably derived from, 
the var. chlorolepis. Gray, as noted above, treated it as a variety of Kuhnia 
eupatorioides, but Shinners (1946), maintained it as a species, largely because 
he thought it to be confined to a unique habitat, gravelly stream beds along the 
Balcones Escarpment of central Texas. I have observed quite large, relatively 
uniform, populations of the taxon in such habitats, especially along the upper 
reaches of Barton Creek in Travis County. In this area, B. eupatoriozdes var. 
corymbulosa may be found on the drier calcareous hillsides, usually in grassy 
sites of juniper-oak woodlands. I have yet to find the taxa growing together 


Turner: Overview of Brickellia eupatorioides complex 131 


or with any indication that they might be forming hybrids. Nevertheless, var. 
gracillima appears to intergrade westward into the var. chlorolepis and I can 
see little justification in its recognition at the species level, since there is little 
other than leaf width to distinguish between the two. 


ACKNOWLEDGMENTS 


I am grateful to Dr. Guy Nesom and Dr. Carol Todzia for a review and 
helpful comments on the present manuscript. 


LITERATURE CITED 


Barkley, T. 1986. Kuhnia, in Flora of the Great Plains, Univ. of Kansas 
Press, Lawrence, Kansas, p. 966. 


Correll, D. & M.C. Johnston. 1970. Kuhnia, in Manual of the Vascular 
Plants of Teras. Contr. Texas Res. Found., Bot. Studies 6:1547-1549. 


Fernald, M.L. 1950. Kuhnia, in Gray’s Manual of Botany. Dioscorides Press, 
Portland, Oregon, p. 1370. 


King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae (Aster- 
aceae). Monographs Syst. Bot. 22:1-580. Missouri Bot. Garden, St. 
Louis. 


McVaugh, R. 1984. Brickellia, in Flora Novo-Galiciana 12:153-187. 
Shinners, L.H. 1946. Revision of the genus Kuhnia L. Wrightia 1:122-144. 
Shinners, L.H.. 1971. Kuhnia L. transferred to Brickellia. Sida 4:274. 


Steyermark, J 1963. Kuhnza, in Flora of Missouri, lowa State Univ. Press, 
Ames, Iowa, p. 1468. 


Phytologia (September 1989) 67(2):132-133. 


PASSIFLORA CHLORINA A NEW SPECIES OF SUBGENUS ASTROPHEA 
(PASSIFLORACEAE) FROM THE CERRADO VEGETATION OF BRAZIL 


Linda K. Escobar 
Departamento de Biologia, Universidad de Antioquia, Apartado Aéreo 1226, 
Medellin, COLOMBIA” 


ABSTRACT 


During the course of a taxonomic revision of Passiflora subgenus 
Astrophea (Passifloraceae) a new species was found from the cerrado 
vegetation of the state of Minas Gerais, Brazil. Passiflora chlorina 
most closely resembles P. sclerophylla which is known from collections 
made on the sandstone tepuis of Venezuela and Guyana. 


KEY WORDS: Passifloraceae, Passiflora, systematics, Neotropics. 


Passiflora chlorina L. Escobar, sp. nov. TYPE: BRAZIL. Minas Gerais: 
ca 17 km SW of the Minas Gerais-Bahia state border along Highway 
BR-116, 30 Mar 1976, Davidse et al. 11605 (HOLOTYPE: MO!). 


P. sclerophyllae Harms similis sed nectariis foliaribus infra lam- 
inam in paginis adaxialibus vel lateralibus petiolorum, stipitibus 
florum brevioribus, et forma filamentorum coronae differt. 


Lianas with concentric rings of secondary wood and stout tendrils. Plants 
pubescent with straight to curved yellow-green trichomes ca 0.1 mm long, the 
adaxial surface of leaves and inner flower parts glabrous. Young stems with 
smooth, dark red-black, flaking bark. Leaf blades obovate to elliptic, 3.8-5.7 
cm long, 2.3-3.5 cm wide, rounded and retuse at apex, mostly acute at base, 
entire at margins, stiffly coriaceous, lustrous on adaxial surface, with scattered 
trichomes on mid vein, drying reddish brown; lateral veins 5-7, with expanded 
fine veins forming an incrassate reticulum on abaxial surface; petioles 8.5-11.0 
mm long, 1.1-1.2 mm diam, with 2, paired, subspherical, bordered nectaries 
1.5-2.0 mm diam below blade on adaxial or lateral surfaces; stipules triangular, 
ca 3 mm long, ca 1.2 mm wide at base, deciduous. 

“Present address: Plant Resources Center, Department of Botany, Univer- 
sity of Texas, Austin, Texas 78713 USA. 


132 


Escobar: New Passiflora from Brazilian cerrado 133 


Peduncles stout, solitary in leaf axil, 2.8-7.0 mm long, 1.1-1.5 mm diam; 
bracts triangular, ca 1 mm long, ca 0.5 mm wide, reddish. Flowers narrowly 
campanulate, ca 3 cm long, 2.0-2.5 cm diam, white; flower stipe stout, 4-5 mm 
long, 1.1-1.2 mm diam; hypanthium campanulate, 8-9 mm long, 3.2-3.5 mm 
wide at base, ca 6 mm wide at apex; sepals oblong, 18-20 mm long, 4.3-5.0 
mm wide, subcoriaceous; petals subequal to sepals, membranaceous; corona 
in 2 series, at apex of hypanthium, the outer ca 9 mm long, ca 1 mm wide 
at base, abruptly widening ca 2.5 mm from apex to 1.5 mm, then tapering to 
blunt apex; the second series linear, ca 3 mm long, ca 0.2 mm wide, bent to 
ca 45 degree angle in upper 1/3; operculum erect, arising ca 3 mm from base 
of hypanthium, ca 3 mm long, composed of fleshy, papillose filaments; ovary 
oblong, ca 3.5 mm long, ca 2 mm wide, 12 ribbed, densely pubescent, with 
straight, tan trichomes; styles ca 2.3 mm long, ca 0.6 mm diam, pubescent ca 
2/3 their length from base. Fruits unknown. 

Passtflora chlorina is only known from the type collection in cerrado vege- 
tation of Brazil, on the border of the States of Minas Gerais and Bahia, at 950 
meters elevation. It is most similar to P. sclerophylla Harms, by virtue of the 
stiffly coriaceous leaves with thickened fine veins of the abaxial surface forming 
an incrassate reticulum. It is distinguished from P. sclerophylla by the place- 
ment of the foliar nectaries below the blade on the adaxial or lateral surfaces 
of the petioles, the shorter flower stipe and the shape of the coronal filaments. 
The specific epithet refers to the yellow green color of the pubescence, which 
is especially noteworthy on the flowers. 


ACKNOWLEDGMENTS 


I am indebted to Dr. Guy Nesom for criticism of the manuscript and for 
the Latin diagnosis. 


Phytologia (September 1989) 67(2):134-138. 


A FURTHER NOTE ON AGROSTIS ELLIOTTIANA 
(GRAMINEAE) IN ARIZONA 


Charlotte G. Reeder & John R. Reeder 
Herbarium, University of Arizona, Tucson, Arizona 85721 USA 


ABSTRACT 


It has been discovered that Agrostis elliottiana, thought to be new 
to Arizona and New Mexico in 1986, was actually collected by C.G. 
Pringle near Tucson, Pima County, AZ, more than 100 years ago. The 
only voucher specimen is apparently at VT. Included also is information 
about other species collected in the area at about the same time, with 
notes on habitat changes. 


KEY WORDS: Agrostis elliottzana, Arizona, Pringle. 


Three years ago we reported the occurrence of Agrostis elliottzana Schultes 
in Arizona and New Mexico (Reeder & Reeder 1986); these were considered to 
be first records. In the spring of 1985 and again in 1986 we had encountered 
this delicate annual grass in Pima County, Arizona, along Ash Creek which 
flows near the border with Cochise County on the eastern side of the Rincon 
Mountains. While we were studying the collections, we received a specimen 
of the same species from R.D. Worthington (UTEP), who had found it on the 
westernmost edge of the Peloncillo Mountains in Hidalgo County, New Mexico, 
near the Arizona border. Like our specimens, his had been found growing along 
the edge of a stream. These records extended the known distribution for A. 
elhottiana westward from central Texas. 

We believed that a thorough search of the literature had been made be- 
fore we published our account. Neither Hitchcock (1951), Gould (1951), nor 
Swallen (1960) gave any clue that this grass had ever been found in Arizona. 
At the time we recorded our findings, we had studied Agrostis exigua Thurber, 
an annual California species, and concluded that the two are conspecific; A. 
elliottiana Schultes is the older binomial. 

Recently, while researching an unrelated matter, we were alerted to a 
startling bit of information in a series of papers by F. Lamson Scribner (1882- 
1883). Among the 85 species mentioned, some 16 had been collected in Cal- 
ifornia, while the remainder had come from Arizona. Most of the Arizona 
collections were from the area around Tucson, particularly Fort Lowell. Oth- 
ers were from the Santa Cruz Valley, Santa Rita Mts., Santa Catalina Mts., 


134 


Reeder & Reeder: Further note on Agrostis elliottiana 135 


one from Pantano and another from near Yuma. To our surprise, we noted 
Agrostis arachnoides Elliott (=A. elliottiana] in the list, as having come from 
“near Fort Lowell”!! ARIZ had no such collection, and it is clear US had 
none. It seemed unlikely that this was a misidentification; Scribner, who had 
studied the grasses in Tennessee and the southeastern United States, would 
certainly have known this species. There seemed a chance that the Pringle 
Herbarium (VT) might have the specimen we were seeking. We made inquiry, 
and through the courtesy of Dr. David S. Barrington, we shortly received the 
loan of an herbarium sheet with a fine example of this delicate awned an- 
nual species of Agrostis. The label data are: “FLORA OF THE PACIFIC 
SLOPE, Arizona, Agrostis arachnoides Ell., near Camp Lowell, April, 1881. 
C.G. Pringle.” (Above the label someone had written in pencil: “A. Elliotvana 
[sic!] Schultes.” This left no doubt that the species, indeed, had been collected 
in Arizona over 100 years before our gatherings! 

At present the locality “near Fort Lowell” seems a most unlikely habitat 
for this little grass which is to be found growing along the sandy margins of 
running streams. Fort Lowell is now surrounded by the city of Tucson, and the 
washes nearby are dry except for short periods during heavy summer rains. 
One must look back in time to understand this distribution. 

Pringle made his first trip to the Pacific Slope in the autumn of 1880, the 
year the railroad arrived in Arizona. No doubt he utilized this newly available 
means of transportation. During this time, he had three objectives: 1) to 
explore the forests of the area for the U.S. Census Department; 2) to collect 
woods for the Jesup Collection of the American Museum of Natural History; 
and 3) to make general collections for Dr. Gray’s studies. At that time, Asa 
Gray was working on The Synoptical Flora of North America and felt the 
need for material from the arid Southwest. Pringle was permitted to make 
duplicate specimens for his own herbarium and for sets which were to be sold 
to help defray the expenses of the trip. He worked at these tasks through 1884 
(Brainerd 1911). 

It was in 1881 that Pringle collected rather extensively around Tucson, 
probably staying at Fort Lowell. Camp Lowell or Fort Lowell, as it was var- 
iously called, was first established in 1862 by California Volunteers at a site 
which is now in the center of downtown Tucson. It remained active for two 
years when in September, 1864, it was abandoned, but was reoccupied in May, 
1865. In 1866 the site was made a permanent post and given the name Camp 
Lowell. It developed that the original site was not a good location. The need 
for an adequate water supply for the post, as well as hay for the horses, re- 
sulted in the Camp’s being moved in 1873 to a spot on which the ruins of 
the once lively fort now stand. The new location was at the edge of the then 
running Rillito stream, where there was an abundant growth of grass nearby. 
Soon it became a thriving community with a public school, church, two weekly 
papers and various places of entertainment for the soldiers. It remained an 


136 PHYTO LPO"GTA volume 67(2):134-138 September 1989 


important Fort until the 1880’s when the Apaches were finally subdued. In 
1886 it was abandoned, and the area left to become ruins. Within recent 
years, however, efforts have been made to restore portions of the old Fort; the 
immediate surroundings now constitute Fort Lowell Park (Barnes 1960). 

The list of grasses presented in the above mentioned list by Scribner in- 
cluded 14 species which had come from Camp Lowell. In addition to Agrostis 
arachnovdes, there were the following which are commonly found in moist soil: 
Agrostis ezarata Trin. “of two forms,” one from the banks of the Rillito in May; 
the other with a more slender panicle from the Santa Rita Mts. in July. (It is 
of interest that Hitchcock in 1905 p. 38, confirms these two forms of Agrostis 
ezarata.) Also, Panicum sanguinale L. |= Digitaria sanguinalis (L.) Scop.]; Poa 
annua L. var. stricta Vasey, from the banks of the Rillito (= Poa bigelovii Vasey 
& Scribner). The Pringle gathering of 1881 is the type collection for this latter 
taxon, one of Arizona’s early spring blooming grasses. Other species collected 
by Pringle in 1881 from the Santa Cruz Valley and Camp Lowell area appear 
to be grasses of the foothills and/or mesas in that vicinity: e.g. (with the 
modern binomials): Digitaria insularis (L.) Mez ex Ekman; Hilarza belangerz 
(Steud.) Nash; Aristida adscensionis L.; Aristida hamulosa Henr.; Sporobolus 
contractus, A.S. Hitchcock and several species of Bouteloua: B. aristidoides 
L., B. chondrosioides (H.B.K.) Benth. ex S. Wats., B. barbata Lag. (It is note- 
worthy that in 1901 Scribner & Merrill described Bouteloua micrantha based 
on Griffiths 1556 from Fort Lowell. This is now regarded as a synonym of B. 
barbata.) Pappophorum apertum Munro ex Scribner, which was said to have 
come from the “Mesas near Camp Lowell, Arizona” was also described from 
a Pringle collection of the 1881 set. This is now considered to be a synonym 
of Pappophorum vaginatum Buckley. Another name based on a Pringle collec- 
tion of 1881 is Sporobolus cryptandrus (Torrey) A. Gray var. strictus Scribner. 
The author indicated that it was from “Banks of the Rillita [sic!] near Camp 
Lowell. June.” This is Sporobolus contractus A.S. Hitchcock, which is based 
on the same type. 

J.J- Thornber (1909) discussed the vegetation groups of the Desert Labo- 
ratory Domain, which included such areas as I. Tumamoc Hill [location of the 
Laboratory buildings]; II. Mesa-like Mountain Slopes; III. Santa Cruz Flood- 
plain; and IV. Santa Cruz River and Irrigation Ditches. Among the numerous 
grass species listed for the Santa Cruz flood-plain, there is no mention of any 
Agrostis species. In the category of Santa Cruz River and Irrigation Ditches, 
he does list Agrostis verticillata Vill. [=Polypogon viridis (Gouan) Breistr.] 
and Paspalum distichum L., both species to be found in moist places. In 
the ARIZ Herbarium there are numerous specimens collected by Thornber 
from Fort Lowell during the early 1900’s, but we found no Agrostis elliottiana. 
Both Thornber and David Griffiths were actively collecting plants, especially 
grasses, in 1901-1903 in southern Arizona. Most of the Thornber collections 
and many of the Griffiths numbers are in ARIZ. Two collections of Agrostis 


Lard 


Reeder & Reeder: Further note on Agrostis elliottiana 137 


scabra Willd., which came from Fort Lowell are: Thornber s.n., June 1, 1903 
and Thornber 491, May 25, 1903. This perennial species was associated with 
the annual, inconspicuous, ephemeral Agrostis elliottiana where we found it in 
1985 and 1986. This would tend to suggest that in the early 1900’s there was 
some water in the Rillito Stream, which joins the Santa Cruz River not far 
from the Fort Lowell site. 

Agrostis elliottiana is not a newcomer to Arizona. It was here in 1881, and 
presently is to be found in at least one locality in the State. It is not likely 
that it will again be found in the Fort Lowell area. In our experience, this 
little grass is confined to riparian habitats — Fort Lowell scarcely qualifies for 
that designation today. 

The fact that Pringle found Agrostzs ellottiana in 1881 demonstrates again 
that he was a keen plant observer. Although neither Griffiths nor Thornber 
collected the species, this does not necessarily mean that it was there only in 
the year Pringle found it. We emphasize again that it is a delicate, incon- 
spicuous plant, which is frequently (perhaps always) found in association with 
Agrostis scabra. It might easily be mistaken for a young plant of that species. 


ACKNOWLEDGMENTS 


Our thanks to David S. Barrington (VT) for the loan of the Pringle speci- 
men, and to Charles T. Mason, Jr. for use of the Herbarium facilities at ARIZ. 
Richard S. Felger and Laurence J. Toolin reviewed the manuscript and offered 
valuable suggestions. 


PERTINENT LITERATURE 


Barnes, W.C. 1960. Arizona Place Names. 1960. Revised and Enlarged by 
Byrd H. Granger. Tucson: Univ. Arizona Press. 


Brainerd, Ezra. 1911. Cyrus Guernsey Pringle. Rhodora 13(155):225-232. 


Gould, F.W. 1951. Grasses of the Southwestern United States. Tucson: Univ. 
Arizona Press (Reprinted 1973). 


Hitchcock, A.S. 1905. North American Species of Agrostis. Bull. U.S. Dept. 
Agric. Bur. Plant Industry 68. 


1951. Manual of the Grasses of the United States. U.S. Dept. 
Agric. Misc. Publ. 200. (Ed. 2, revised by Agnes Chase.) 


138 PRY TOLOGTA volume 67(2):134-138 September 1989 


Reeder, C.G. & J.R. Reeder. 1986. Agrostis Elliottiana (Gramineae) new to 
Arizona and New Mexico. Phytologia 60(6):453-458. 


Scribner, F.L. 1882-1883. A list of grasses collected by Mr. C.G. Pringle in 
Arizona and California during the summer of 1881, with descriptions 
of those species not already described in American publications. Bull. 
Torrey Bot. Club 9:74-77,86-89,103-105,145-149; 10:29-32. 


Swallen, J.R. 1960. Gramineae, pp. 70-145. Jn: Kearney, T.H., R.H. Pee- 
bles, & Collaborators. Arizona Flora. Ed. 2 + Suppl. Berkeley: Univ. 
California Press. 


Thornber, J.J. 1909. Vegetation groups of the Desert Laboratory Domain. 
Publ. Carnegie Inst. Wash. 113:103-112. 


Phytologia (September 1989) 67(2):139-141. 


A RECONSIDERATION OF UROSPATHELLA 
(ARACEAE) 


George S. Bunting 
101 Linden Avenue, Pocomoke City, Maryland 21851 USA 


ABSTRACT 


The relationships of Urospathella are discussed; anatomical data of 
the seed are presented to support its separation from Urospatha and 
information justifying its exclusion from Cyrtosperma is evaluated. 


KEY WORDS: Urospathella, Araceae, Neotropics, anatomy. 


The genus Urospathella Bunting has a single species, Urospathella wurdacku 
(Bunting) Bunting, a small plant known to occur only in white sand savannas 
of Amazonian Venezuela. It differs from Urospatha sagittifolia (Rudge) Schott 
by its slender habit, oblong or elliptic leaf blades lacking posterior lobes and a 
unilocular ovary with two sub-basal ovules. The lack of precise data concern- 
ing the presence of endosperm in the seed has caused speculation about its 
correct generic placement. Originally described in the genus Cryptosperma, 
it has subsequently been transferred to Urospatha (Hay 1988) and then to 
Urospathella (Bunting 1988). 

Recent study of a seed of Urospathella wurdacku, by Dr. James C. French at 
Rutgers University, revealed that the seed “most definitely contains endosperm 
[with the] large plump embryo surrounded by a layer of cells filled with lipid 
droplets and small irregular crystalline particles.” He “ did not determine 
if they were starch, but their birefringence was not typical of starch grains. 
Outside the endosperm was the thick seed coat.” Dr. French has “no doubt 
about the presence of endosperm in U. {Urospathella| wurdacki. The other 
species. . .sent [Urospatha sagittifolia], reputed to lack endosperm, differed in 
lacking this lipid-rich layer of cells.” 

The albuminous nature of the seeds of Urospathella wurdacku obliges a 
reconsideration of the genus Urospathella, since the seeds were originally de- 
scribed as apparently without endosperm. With this character negated, the 
slender habit of this species and its leaf form remain the chief morphological 
features to distinguish it from Cyrtosperma (cf. Hay 1988). Even the sparsely 
and finely verruculose condition of the petiole suggests the spiny condition well 
developed in Cyrtosperma. However, the petiole of Urospathella wurdackz does 


139 


140 PAY FO LOG TA volume 67(2):139-141 September 1989 


not appear to be geniculate as in the latter genus. Moreover, the geographic 
isolation of this species in north-central South America reinforces its segrega- 
tion from Cyrtosperma, now considered to be an entirely Asian genus. 

Hay (1988 and in personal correspondence) indicated that the Neotropical 
species previously placed in Cyrtosperma must, on the basis of their exal- 
buminous seeds or aspect, be excluded from this genus. These include the 
following: 


“Cyrtosperma americanum Engl. in Martius = Anaphyllopsis americana 


(Engl.) A. Hay, ined. 
Cyrtosperma spruceanum (Schott) Engl. = Dracontium sp. 


Cyrtosperma wurdacku Bunting = Urospatha wurdackiw (Bunting) A. Hay, 
comb. nov.” 


The last transfer was formally made, but because this species has seeds with 
endosperm, its inclusion in Urospathais untenable. Further, Hay’s comments 
about the comparable unilocular condition of the ovary of this species and that 
of Urospatha savannarum Steyerm. (Publ. Field Mus. Nat. Hist. Bot. Ser. 
28:102. 1951) seem tenuous. The type specimen of this latter name cannot be 
located for restudy to verify the unilocular condition of its ovary with a single 
ovule arising from the base at one side, as described by Steyermark. It is my 
experience that the removal of a pistil from a spadix of Urospatha, done even 
with great care, often results in all or part of the ovules, together with the very 
short partition, remaining fast to the axis of the spadix and being pulled out of 
the ovary rather than remaining in situin the latter. Until its ovary characters 
can be verified, it appears unwise to consider Urospatha savannarum in these 
discussions, since the drawing accompanying its original description depicts a 
plant very similar in both foliage and inflorescence to a young plant of the 
widespread Urospatha sagittifolia (Rudge) Schott sensu lato. 


NOMENCLATURE SYNOPSIS 


Urospathella wurdackii (Bunting) Bunting, Phytologia 65:391. 1988. Cyr- 
tosperma wurdacku Bunting, Acta Bot. Venez. 10:285. 1975. Urospatha 
wurdacku (Bunting) A. Hay, Blumea 33:457. 1988. 


ACKNOWLEDGMENTS 


I am very grateful to Dr. James C. French, of Rutgers University, for de- 
termination of the endosperm content of seeds of pertinent material and to 
Dr. D.H. Nicolson and Dr. T. Croat for their reviews of the manuscript. 


Bunting: Reconsideration of Urospathella 141 


LITERATURE CITED 


Bunting, G.S. 1988. Urospathella, new genus of Venezuelan Araceae. Phy- 
tologia 65:391-392. 


Hay, A. 1988. Cyrtosperma (Araceae) and its Old World allies. Blumea 
33:427-469. 


Phytologia (September 1989) 67(2):142-147. 


NEW SPECIES OF MEXICAN SOLIDAGO 
(COMPOSITAE: ASTEREAE) 


Guy L. Nesom 
Department of Botany, University of Texas, Austin, Texas 78713 USA 


ABSTRACT 


Three new species and a new variety of Solidago are described 
from Nuevo Leon and Coahuila in NE México. Solidago ericame- 
rioides and S. gypsophila are apparently obligate gypsophiles with 
highly restricted distributions in Nuevo Leon and Coahuila, respectively. 
Solidago hintoniorum is known from a number of localities in the 
mountains of southeastern Coahuila, central Nuevo Leon and adjacent 
Tamaulipas. Solidago wrightii var. orientalis is known from a sin- 
gle collection from the Sierra La Marta of Coahuila and is the only 
collection of this species from the eastern sierra of México. 


KEY WORDS: Solidago, Asteraceae, México, floristics. 


In the preparation of a taxonomic treatment of Mexican Solidago, three 
previously undescribed taxa warrant recognition as species and another as a 
variety. Two of the species apparently are obligate gypsophiles and narrow 
endemics. The third species is relatively more widespread and known from a 
number of collections. 


Solidago gypsophila Nesom, sp. nov. TYPE: MEXICO. Coahuila: Mpio. 
Guatro Cienegas, ca 9 air mi SW of Cuatro Cienegas, ca 2 mi N of Poza 
Becerra, in a fen-like area, with Flaveria, Baccharis, Scirpus, Anemopsis, 
Sporobolus, Distichlis, 709 m, 10 Oct 1976, J. Henrickson 15576 with M. 
Dillon (HOLOTYPE: LL!). 


S. canadensi L. similis sed paginis infernis foliorum dense et 
grosse hispidulis laminis profunde recessis et nervaturis valde pro- 
trudentibus differt. 


Herbaceous perennials 1.5-2.0 m tall, base not seen. Stems 5-7 mm thick 
at mid stem, yellowish, densely villous-pilose with white, crinkly hairs. Leaves 
erect-ascending, crowded and overlapping with internodes mostly 1 cm long, 3 
veined, densely and evenly hispidulous-hirtellous on both surfaces, with deeply 


142 


Nesom: New taxa of Solidago from México 143 


recessed lamina and strongly protruding venation, oblanceolate-obovate, 8-12 
mm wide, 35-75 mm long, attenuate to a petioliform base 1-2 mm long, the up- 
per ovate to obovate, slightly reduced in size. Heads campanulate on bracteate 
ultimate pedicels 1-2 mm long, secund and crowded on lateral branches with 
narrowly elliptic bracteal leaves, the whole capitulescence a pyramid 30 cm 
broad at base and 35 cm high; receptacles deeply foveolate; phyllaries yel- 
low, glabrous, with a prominently raised, resinous midvein, in 3-4 strongly 
graduated series, the inner 2.5-3.0 mm long. Ray flowers 7-10, the corollas 
3.0-3.5 mm long, glabrous, the ligules 1.5-2.0 mm long, 0.5 mm wide, api- 
cally toothed or cleft. Disc flowers 4-5, the corollas 3.5 mm long, the lobes 
triangular-lanceolate, ca 1.5 mm long, the throat 1 mm long, tube 1 mm long; 
style appendages triangular. Achenes sparsely strigose, mature size not seen; 
pappus a single series of 26-32 barbellate bristles 2.5-3.0 mm long. 

Additional collections examined: MEXICO. Coahuila: Mpio. Cuatro Ciene- 
gas, Cuatro Cienegas Basin, small laguna between Lagunas Churince and 
Grande, 13 Aug 1967 [just into flower], Cole, Minckley & Pinkava 3790 (ASU); 
Dos Quatos, 21 Aug 1967 [immature], Cole, Minckley & Pinkava 4336 (ASU). 

Solidago gypsophila is similar and obviously related to S. canadensis var. 
canescens A. Gray in its villous stem pubescence, crowded leaves, large, tri- 
angular capitulescence and the small heads with few disc flowers. The new 
taxon could be added to the numerous varieties of S. canadensis, but it dif- 
fers strongly from it and all other species in its leaves with deeply recessed 
lamina and strongly protruding venation and in its dense, even length, stiffly 
hispid-hirtellous foliar pubescence. 

The closest known locality for Solidago canadensis var. canescens is about 
150 miles north of Cuatro Cienegas. The widespread and commonly collected 
S. velutina DC. is the only other species of the genus that occurs close to the 
type locality. Typical plants of S. velutina have been collected from several 
localities in the nearby Sierra de la Madera and Sierra de la Gavia and from 
within the Cuatro Cienegas basin itself (“summit - Anteojo Complex,” Meyer 
s.n., ASU), but the latter site may be a limestone outcropping rather than 
gypsum. Solidago gypsophila almost certainly is an obligate gypsophile. 


Solidago ericamerioides Nesom, sp. nov. TYPE: MEXICO. Nuevo Leén: 
Mpio. Galeana, 10 km NE of Pocitos, gypsum cliff over water, 1850. 
m, 26 Aug 1984, Hinton et al. 18763 (HOLOTYPE: TEX!; Isotype: 
ASU!,NY!). 


S. missouriensi Nutt. similis sed differt statura brevioribus (6- 
15 cm alto), caulibus pubescentibus pilis crassis brevis, foliis bre- 
vioribus (8-20 mm longis) elliptici-oblanceolatis, capitulescentiis 
corymbosis capitulis paucioribus, phyllariis 3 nervatis, flosculis radii 
paucioribus, et acheniis longioribus. 


144 PAY 7 OE O GIsA volume 67(2):142-147 September 1989 


Suffrutescent perennials 6-17 cm tall, producing basal offsets or stolons 
that thicken and lignify. Stems moderately to densely pubescent with thick, 
short, upturned hairs. Leaves apparently fleshy when fresh, wrinkling upon 
drying, entire, elliptic-oblanceolate or oblanceolate-obovate, 8-20 mm long, 2-3 
mm wide, not at all reduced in size upwards but the internodes lengthening to 
2-4 mm, margins scabrous-ciliate, the lower surface sparsely and microscop- 
ically strigose. Heads campanulate on bracteate pedicels 3-10 mm long, in 
compact corymbs 2.0-3.5 mm broad of 8-20 heads each; phyllaries in 3-4 se- 
ries, oblong-oblanceolate, whitish indurated except for the prominent, orange 
resinous midvein, the middle series 3 veined, the inner series 4.5-5.0 mm long 
with minutely ciliolate margins, the outermost 1/2-1/3 as long as the rest. Ray 
flowers 6-10, yellow, the corollas 4-5 mm long, the ligules 2-3 mm long, 1 mm 
wide, 3-4 veined, with 3 shallow apical teeth, the tube glabrous.’ Disc flowers 
6-13, the corollas ca 3.5 mm long, tubular-funnelform, the lobes lanceolate, 
1.0 mm long, throat 1.5 mm long, tube 1 mm long; style appendages deltate- 
lanceolate. Achenes fusiform-cylindric, 2.0-2.3 mm long, densely strigose, with 
5-7 raised, orange resinous nerves; pappus a single series of 29-34 barbellate 
bristles mostly 2-3 mm long. 

Solidago ericamerioides may be closely related to S. mzssouriensis Nutt., 
although they are very different in a number of features. Forms of S. mis- 
souriensts occur in the same gypseous area as 5. ericameriozdes around Galeana, 
Nuevo Leon, but even in a highly gypseous substrate, plants of S. missourien- 
sis maintain a general morphology relatively typical of the species in its main 
range in the United States. The two species (in México) are distinguished by 
the following couplet. 


1. Plants 6-17 cm tall with thick-hairy stems; leaves 8-20 mm long, elliptic- 
oblanceolate; capitulescence corymbose, distinctly flat topped, few head- 
ed; phyllaries of nearly equal length, the middle 3 veined; ray flowers 
6-10; achenes .2.0-2.3 mm long <2... 06... gov atau S. ericamertoides 


1. Plants 50-100 cm tall with glabrous to glabrate stems; leaves 40-90(-110) 
mm long, linear-lanceolate; capitulescence cylindric to oval or pyramidal 
with numerous heads; phyllaries strongly graduated in length, the middle 
1 veined; ray flowers 13-18; achenes 1.0-1.5 mm 
One odtaa ce Se de ote ce tice cr ees ck ans. es eee S. missourzensis 


Typical plants of Solidago missouriensis have been collected on gypsum 
very near the type locality of S. ericamerioides (Hinton et al. 18780, TEX). 
This sheet appears to be a mixed collection, bearing six typical plants of S. 
missourtensis and one small plant that has the habit and capitulescence of S. 
ericamerioides, but longer leaves and strongly graduated phyllaries. I believe 
this individual may represent a hybrid or introgressant. 


Nesom: New taxa of Solidago from México 145 


Collections of the following undescribed species have been accumulating 
at TEX-LL for some time, identified as Solidago muellert Standley, which 
however, has proved to be a synonym of S. missourtensis Nutt. 


Solidago hintoniorum Nesom, sp. nov. TYPE: MEXICO. Nuevo Leén: 
Mpio. Galeana, Cerro El Potosi, clearing in pine forest, 3150 m, 15 Sep 
1969, Hinton et al. 17264 (HOLOTYPE: TEX!). 


S. petiolari Ait. similis sed caulibus et foliis sparsim villosis, 
phyllariis eglandulosis glabris 3 nervatis marginibus hyalinis, et 
acheniis strigosis differt. 


Rhizomatous perennials. Stems 0.5-1.0 m tall, sparsely hairy with spread- 
ing crisped hairs mostly 0.5-1.0 mm long, eglandular. Leaves entire to serrate 
with 3-6(-9) teeth, obovate to obovate-oblanceolate, the basal often persistent, 
basally attenuate to a broad petiolar region, lower to midcauline 5-9 cm long, 
10-25(-35) mm wide, little reduced upwards but becoming sessile, eglandular, 
margins spreading ciliate, the major veins usually sparsely villous, the lamina 
glabrous to sparsely villous. Heads 5-6 mm wide, sessile to subsessile on ul- 
timate peduncles 1-2 mm long, in tightly cylindrical panicles mostly 6-15 cm 
long, the capitulescence sometimes branched and producing lateral ascend- 
ing cylinders below the primary one; phyllaries oblong with erect, rounded to 
obtuse apices, with a prominent, green apical patch with numerous visible sto- 
mates, glabrous or with a few, minute hairs near the apex, strongly graduated 
in 3-4 series with the inner 5-6 mm long, midvein thick and orange resinous, 
often flanked by 1-2 pairs of smaller, lateral veins on the middle and inner, 
margins with broad, hyaline flanges, sometimes slightly lacerate-ciliate near 
the apex. Ray flowers 5-10, the corollas 5.5-6.0 mm long, ligules 2.5-3.5 mm 
long, 0.8-1.3 mm wide. Disc flowers (6-)8-11, the corollas 5.0-5.5 mm long. 
Achenes moderately strigose, cylindric, mature size not observed; pappus of 
25-36 barbellate bristles. 

Additional collections examined: MEXICO. Coahuila: Mpio. Arteaga: 
Sierra La Marta, with Pinus, Ceanothus, Quercus and Garrya, 24 Oct 1981, 
Poole 2449 (TEX); Sierra Los Camargos, bushy hillside, 2920 m, 19 Jul 1980, 
Hinton et al. 17884 (TEX); Sierra Los Camargos, 2870 m, 4 Aug 1980, Hinton 
et al. 17925 (ASU); Sierra La Viga, N side, with Pznus, Pseudotsuga, Abies 
and Picea, 2700-3000 m, 24 Oct 1984, McDonald & Gomez 1179 (TEX); Sierra 
La Viga, SE side, Pinus culminicola and P. hartwegzz, 3300 m, 25 Oct 1984, 
McDonald & Gomez 1216 (TEX); Sierra de Arteaga, Las Vigas, canon de 
la Carbonera, Pinus, Pseudotsuga, Abies, Quercus and Ceanothus, 2100-2600 
m, 15 Sep 1988, Villarreal et al. 4590 (TEX). Nuevo Leon: Mpio. Galeana: 
Sierra La Marta, rocky mountainside, 2870 m, 4 Aug 1980, Hznton et al. 
17925 (TEX); Sierra La Marta, rocky mountain, 2800 m, 6 Sep 1981, Hinton 
et al. 18335 (TEX); El Carrizo, oak woods, 1900 m, 16 Oct 1983, Hinton et al. 


146 PHYTOL O GIA volume 67(2):142-147 September 1989 


18613 (TEX,WAT); Dulces Nombres, open rocky places, 2000 m, 12 Aug 1948, 
Meyer & Rogers 2955 (MO); W of Galeana, 7.5 mi SW of Dieciocho de Marzo 
along the road to the top of Cerro Potosi, pine and fir forest with shrubby 
oaks, 12 Oct 1984, Sundberg 3114 (TEX). Tamaulipas: 3-5 km S of Huisachal, 
limestone slopes, ca 6000 ft, 27 Jun 1949, Stanford et al. 2096 (SMU,TEX). 

The Tamaulipan collections have strongly reduced upper cauline leaves but 
are otherwise typical of Solidago hintoniorum. They are perhaps hybrids be- 
tween S. hintoniorum and S. simplez. Plants of another collection (Meyer 
& Rogers 2955) may also be hybrids between the same two species. I have 
tentatively identified them as S. simpler, however, because they have the re- 
duced habit, purplish stems, and weakly 3 nerved leaves of that species, but 
in vestiture they are much more similar to S. hintoniorum. The long ciliate 
petioles of S. hintontorum are distinctive from both of the putative hybrids. 
Both S. simpler and S. hintoniorum have a cylindrical capitulescence, mostly 
1 nerved, obovate (lower) leaves and glabrous, oblong phyllaries with rounded 
apices. 

Solidago hintoniorum is similar to S. petiolaris and S. wrighti in its 1 
nerved leaves, which are relatively unreduced upwards and its large, prominent 
ray flowers. With S. petiolaris it shares a cylindric capitulescence and with 
S. wrightw strigose achenes. It is different from both species in its glabrous 
phyllaries and completely non glandular, sparsely villous vestiture. 

Solidago wrightt and S. petiolaris appear to be more similar to each other 
than either is to S. hintoniorum, although the three clearly form a closely re- 
lated group. In fact, the first two are so similar they perhaps should be treated 
as a western and eastern phase of a single species. They are distinguished by 
capitular features and achene vestiture, as in the key below, but intergradient 
population systems occur in southwest Texas, where their geographic ranges 
meet. A review of the taxonomy of the whole group is in progress (Nesom, in 
prep.). 

The following key identifies the Mexican taxa of Solidago that have 1 nerved 
leaves, not or only slightly reduced in size upwards, and heads in corymbs or 
cylindrical panicles, not at all secund. The geographic ranges are only for 
México. 


1. Phyllaries (1-)3(-5) veined, glabrous, eglandular; capitulescence cylindri- 
cal; stems and leaves sparsely villous with spreading crisped hairs 0.5- 
1.0 mm long; southeast Coahuila to central Nuevo Leon and adjacent 
Wamamlipas’ 7). 005 ssc sas nt oces coed ete s bs aise en S. hintoni 


1. Phyllaries 1 veined, hairy, glandular, or both; capitulescence cylindrical 
to corymbose; stems and leaves stipitate glandular or merely hairy, but 
if the latter the hairs mostly 0.1-0.2 mm high ..................... (2) 


2. Capitulescence a terminal corymb usually as wide as long or an open, 


Nesom: New taxa of Solidago from México 147 


paniculate cluster of similar corymbs; achenes obviously strigose; 
Chihuahua to central Durango and southwest to east central 
Caahoile oe5530: - 32 2ST a. SS S. wrightw 


2. Capitulescence a tightly cylindric panicle; achenes glabrous or nearly 
so; N Coahuila and central Nuevo Leon .............. S. petiolaris 


A collection from mountains of Coahuila east of Saltillo is related to Sol- 
tdago wrighti and deserves recognition at the varietal level. 


Solidago wrightii var. orientalis Nesom, var. nov. TYPE: MEXICO. 
Coahuila: Mpio. Arteaga, Sierra La Marta, pine and spruce woods, 
3100 m, 6 Sep 1981, Hinton et al. 18333 (HOLOTYPE: TEX!; Isotype: 
NY! TEX!). 


Varietatibus aliis S. wright A. Gray similis sed marginibus 
foliorum serrat-crenatis dentibus 5-10(-14) imprimis differt. 


Perennials, base not seen. Stems 0.5-0.6 m tall, puberulous with a mixture 
of stipitate glands and short, spreading, upturned hairs ca 0.2 mm high. Leaves 
thick, elliptic to elliptic obovate, 35-75 mm long, 15-25 mm wide, serrate with 
5-10(-14) pairs of teeth, the lamina hirtellous and stipitate glandular. Heads 
sessile to subsessile on ultimate peduncles 1-3 mm long, in compact, terminal 
corymbs; phyllaries thick, oblong to lanceolate oblong, often with spreading- 
reflexing apices, with the inner often strongly convex, graduated in 3-4 series, 
the innermost 5-6 mm long. Ray flowers 4-6. Disc flowers 9-12. Achenes 
sparsely strigose. 

The plants of this collection have a corymbose capitulescence and densely 
stipitate glandular upper stems, leaves and phyllaries. Leaves of plants from 
elsewhere in the range of Solidago wrightw are entire but they very rarely 
produce 1 or 2 pairs of small teeth. The prominently toothed leaves of var. 
ortentalis may reflect genetic input from S. petzolaris, which is known from 
several areas in the eastern Sierra and has similarly toothed leaves in the area 
where S. wrightti occurs. The plants of var. orientalis are the only ones of 
the species known from the eastern Sierra Madre, but S. wrightw has been 
collected from the Sierra de Jimulco of southwestern Coahuila, and the species 
may have migrated eastward across Coahuila. In qualification, however, the 
number of ray flowers of var. orientalis (4-6) is more similar to that of plants of 
S. wright in New Mexico (6-11) than in the western Sierra of México (9-14). 


ACKNOWLEDGMENTS 


I thank Billie Turner and T. P. Ramamoorthy for reviews of this manuscript 
and ASU, MO and SMU for loans of specimens. 


Phytologia (September 1989) 67(2):148-154. 


A NEW SPECIES OF CAREX (SECT. OLIGOCARPAE) 
FROM THE EDWARDS PLATEAU OF TEXAS 


Edwin L. Bridges 
The University of Texas Herbarium, Austin, Texas 78713 USA 
and 
Steve L. Orzell 
Texas Natural Heritage Program, Texas Parks and Wildlife Department 
Austin, Texas 78744 USA 


ABSTRACT 


Carex edwardsiana, a new species of sect. Oligocarpae, is de- 
scribed and its diagnostic features, habitat and relation to other en- 
demics of the Balcones Escarpment of the Edwards Plateau of Texas 
are discussed. This species has previously been considered to fall within 
the variability known for C. oligocarpa, but it is morphologically dis- 
tinct and geographically isolated from this more northern and eastern 
species. 


KEY WORDS: Carez, Cyperaceae, systematics, Texas, Edwards 
Plateau. 


In the process of an intensive field study of the distribution and ecology 
of the Cyperaceae of Texas, we have noted for several years that the central 
Texas entity identified as Carez oligocarpa Schkuhr. does not closely resemble 
specimens of that species from northern and eastern states. The discussion of 
this entity (as C. oligocarpa) by Dr. Marshall Johnston noted some of these dif- 
ferences, and he stated that it should perhaps not be referred to C. oligocarpa 
(Correll & Johnston 1970). However, this species had rarely been collected in 
Texas before 1970, and the consistency of its differences was not known. After 
making field observations at several sites and examining a series of collections 
made after the Texas flora was published, we are convinced that it represents 
a distinct, previously undescribed species. 


Carex edwardsiana Bridges & Orzell sp. nov. (Figure 1). TYPE: UNITED 
STATES. Texas: Travis County, mesic colluvial limestone slope forests 
with rock outcrops in valley of Barton Creek, ca 2 mi upstream from Bar- 
ton Springs in Zilker Park, and 2.5 mi upstream from Colorado River, 


148 


Bridges & Orzell: New Carer from the Edwards Plateau 149 


f hia 


ih 
| trail 


Figure 1. Carex edwardsiana Bridges & Orzell (from type). 
a. Habit; b. Lower pistillate scale; c. Perigynium; 
d. Fruit with attached style and stigmatic branches. 


150 PAM TO EF OiGbA volume 67(2):148-154 September 1989 


ca 0.5 air mi NE of Loop 360 bridge over Barton Creek, Oak Hill 7.5’ 
Quad, 30° 14’ 55” N, 97° 47’ 47” W, 17 Apr 1989, Orzell & Bridges 9280 
(HOLOTYPE: TEX!; Isotypes: GH!,MICH!,MO!,NCU!,NY!,SMU!,US!). 


Planta perennis, 25-35 cm longa, laxe vel dense cespitosa, vel 
breviter crasse rhizomatosa, foliis principalibus culmo longioribus. 
Cataphylla infra folia oblonga vel ovata, ca 5-40 mm longa, multi- 
costata, distaliter carinata, acuta, brunneola vel fuscopurpurea. 
Pseudoculmi laterales steriles foliosissimi. Folia principalia ap- 
proximata; vaginae foliorum brevissimae, pallide albovirides, saepe 
rubropunctatae, laeves, ventraliter subscariosae; ligula erecta squa- 
miformis, angusta, ca 1 mm lata, hippocrepica; laminae patulae, 
compressae, anguste lineares, 20-40(-65) cm longae, 2-4 mm latae, 
margine scabridulae, pagina valde nervosa, glabra. Spicae 2-5, lin- 
eares vel anguste oblongae, remotae, ad basin plantae confertae. 
Spica ultima mascula, anguste lineari-ellipsoidea, 1.8-2.5 cm longa, 
2-3 mm crassa, straminea, sessilis vel pedunculo 8 cm usque longo 
elevata, ultimam spicam femineam multo superans vel ei brevior. 
Bractea spicae masculae sine vagina, 5-50 mm longa; glumae mas- 
culae 4-5 mm longae, zona costali angusta, viridi, matrice scariosa, 
lata, alba; antherae 2.5-3 mm longae. Pedunculi spicarum fem- 
inearum crassi, filiformes vel lineares, scaberuli, 4-20 mm longi, 
infimi longissimi, pro parte maxima in vaginis bracteis inclusi; 
bracteae spicae feminae foliaceae, infimae longissimae, vaginis 4-15 
mm longis, laminis erectis, spicas et culmos multo superantibus, 
saepe foliis longissimis aequilongis. Spicae laterales omnino fem- 
ineae, (3-)7-9 florae, 7-20 mm longae, 4-7 mm crassae, floribus ap- 
proximatis, rhachidi recta vel leviter fractiflexa; glumae femineae 
ovatae vel lanceolatae, cuspide aut mucrone incluso (2-)4-5(-11) 
mm longae, 1-1.5 mm latae, zona costali viridi, 3-costata, de latere 
stariosae, albidae, ad apicem contractae vel retusae, apice omnes 
valde cuspidatae, cuspide valde scabro, in ultimis glumis abbrevi- 
_ato. Perigynia obovoidea vel ellipsoidea, 4.5-5.0 mm longa, 2.0-2.2 
mm lata, obscure vel acute trigona, faciebus in medio planis vel 
leviter concavis, subtiliter impressinerviis, viridibus vel brunneolis, 
rostro brevi (0.5 mm), recto vel leviter ad angulum 30° excurvato. 
Achenia sessilia aut stipite ad 0.1 mm longo insidenti, obovoidea, 
quasi sine rostro, 2.5-3.0 mm longa, 1.5-2.0 mm lata, trigona, arcte 
inclusa, minute papillosa, cum styli basi tumida articulata. 


Perennial, 25-35 cm long, loosely to densely cespitose, or with stout, short 
rhizomes, the principal leaves longer than the culms. Cataphylls below leaves 
oblong to ovate, ca 5-40 mm long, multicostate, distally carinate, acute, brown 


Bridges & Orzell: New Carez from the Edwards Plateau 151 


or reddish brown. Pseudoculms lateral, sterile, very leafy. Principal leaves 
approximate, sheaths short, pale greenish white, often with scattered reddish 
dots, smooth, ventrally subscarious; ligule an erect narrow scale, ca 1 mm wide, 
horseshoe shaped; blades spreading, flattened, narrowly linear, 20-40(-65) cm 
long, 2-4 mm wide, marginally scabridulous, surface strongly nerved, smooth. 
Spikes 2-5, oblong-linear, borne remotely on culms but densest at base of plant. 
Terminal spike male, narrowly ellipsoid-linear, 1.8-2.5 cm long, 2-3 mm thick, 
stramineous, sessile or on a peduncle to 8 cm long, from shorter than to much 
overtopping the upper female spike. Male spike bract sheathless, 5-50 mm long, 
male scales 4-5 mm long, white hyaline to stramineous with green midnerve; 
anthers 2.5-3 mm long. Peduncles of female spikes stout, filiform to linear, 
scaberulous, 4-20 mm long, the lowest longest, mostly included in the bract 
sheaths; female spike bracts as leaves, the lowest longest, with sheaths 4-15 
mm long, the blades erect, much exceeding the spikes and culms, often as long 
as the longest leaves. Lateral spikes all female, (3-)7-9 flowered, 7-20 mm long, 
4-7 mm thick, flowers approximate, rachis straight to slightly zigzag; female 
scales ovate to lanceolate, including cusp or mucro (2-)4-5(-11) mm long, 1-1.5 
mm wide, the costal zone green, 3-ribbed, the sides scarious, white, tapered to 
retuse at apex, the tips of all scales strongly cuspidate, cusps strongly scabrous, 
with cusp length reduced in upper scales. Perigynia obovoid-ellipsoid, 4.5-5.0 
mm long, 2.0-2.2 mm wide, obscurely to sharply trigonous, the faces at middle 
flat to slightly concave, finely impressed nerved, greenish to brownish, the beak 
short (0.5 mm), straight to slightly excurved, to a 30 degree angle. Achene 
sessile or with a stipe to 0.1 mm long, obovoid, essentially beakless, 2.5-3.0 
mm long, 1.5-2.0 mm wide, trigonous, tightly included, minutely papillose, 
jointed with the swollen style base. 


Additional collections examined (Paratypes): UNITED STATES. Texas: 
Kendall Co., mesic, wooded, N facing rocky slope on S bank of Guadalupe 
River, from W boundary of Guadalupe River State Park to at least 500 ft E; 
Anhalt 7.5’ Quad, 29° 52’ 22” N, 98° 29’ 48” W, 18 Apr 1989, W.R. Carr 
9557 (SMU,TEX). Medina Co., mesic, rocky, wooded, N facing slope on S 
bank of Bandera Creek, ca 500-1000 ft E of W boundary of Hill Country State 
Natural Area; Twin Hollow 7.5’ Quad, 29° 37’ 12” N, 99° 12’ 17” W, 20 Apr 
1989, W.R. Carr 9595 (SMU,TEX). Travis Co.: same as type locality, 7 Apr 
1986, Bridges 86-175 (SMU,TEX); 25 Apr 1987, Bridges 87-140 (TEX); 30 
Mar 1989, Orzell & Bridges 8809 (GH,MICH,MO,NCU,NY,TAES,TEX); 30 
May 1989, Orzell & Bridges 10145 (SMU,TEX); seasonally mesic upper N 
facing slope forest at head of ravine on N side of Mountain Trail, ca 0.4 mi 
N of Comanche Trail, ca 2.5 air mi NW of int FM 2222 and FM 620 at Four 
Points, between Cypress Creek Arm and main pool of Lake Travis, Mansfield 
Dam 7.5’ Quad, 30° 25’ 07” N, 97° 53’ 16” W, 8 Apr 1989, Orzell & Bridges 
9062 (MICH,MO,NCU,SMU,TEX); wooded slopes in ravine, E end of 200 ft 
limestone cliff on S bank of Bull Creek, SE of Loop 360, 0.5 mi NE of Lakewood 


152 PHYO LOG PA volume 67(2):148-154 September 1989 


Drive; Jollyville 7.5’ Quad, 30° 22’ 38” N, 97° 46’ 22” W, 9 Apr 1983, W.R. 
Carr 4654 (SMU); rocky bank of ephemeral creek in cedar woods ca 2700 ft 
NE of summit of Cat Mtn., Austin West 7.5’ Quad, 30° 21’ 24” N, 97° 46’ 38” 
W, 1 May 1983, W.R. Carr 4794 (SMU,TEX); in stream bottom, in shade 
of walnuts and hackberries, between limestone ridges ca 0.75 mi NNW of int 
Loop 360 and Lakewood Drive, Jollyville 7.5’ Quad, 30° 22’ 57” N, 97° 47’ 
28” W, 8 Apr 1984, W.R. Carr 6009 (SMU,TEX); in cedar/Texas oak woods 
on steep slope in limestone ravine, ca 1200 ft W of Spicewood Springs Rd, ca 
0.7 mi S of Loop 360, Jollyville 7.5’ Quad, 30° 22’ 33” N, 97° 45’ 58” W, 14 
Apr 1984, W.R. Carr 6010 (SMU,TEX); along banks of ephemeral stream in 
small cedar-wooded ravine, ca 2500 ft NW of int Loop 360 and N branch of 
Spicewood Springs Rd, Jollyville 7.5’ Quad; 30° 23’ 15” N, 97° 46’ 40” W, 22 
Apr 1984, W.R. Carr 6042 (SMU). 

Carez edwardsianais locally abundant in moist, rich, black, clayey to loamy 
soil rich in humus, mostly in protected slope and ravine forests in the Eastern 
and Southern Balcones Escarpment sections of the Edwards Plateau of Texas. 
These forests are similar to the north slope deciduous forests described by 
Van Auken (1988) and the canyon flora of Palmer (1920). The rich, calcare- 
ous, seasonally moist soils support many species at their western and southern 
range limits, as well as providing habitat for numerous local and regional en- 
demics. Canopy trees associated with C. edwardsiana include Quercus buck- 
leyi, Q. durandii, Q. muhlenbergii, Juniperus ashet, Frazinus tezensis, Celtis 
reticulata, Carya illinoiensis and Prunus serotina. Shrub and woody vine as- 
sociates include Aesculus pavia, Garrya lindhetmeri, Lindera benzoin, Rhus 
radicans, Callicarpa americana, Ungnadia speciosa, Parthenocissus quinque- 
folia, Morus rubra, Cornus drummondii and Juglans microcarpa. The herb 
layer at some sites includes the local endemics {primarily the PC1 endemics of 
Amos & Rowell (1988)] Anemone edwardsiana, Commelinantia anomala, Eu- 
phorbia roemeriana, Onosmodium helleri and Ruellia drummondiana, and the 
regionally endemic Carez planostachys, Desmodium psilophyllum, Dichanthe- 
lium pedicellatum, Hedeoma acinoides, Mirabilis indhermeri, Nolina terana, 
Polytaenia tezana, Salvia roemeriana and Tetragonotheca terana. More wide 
ranging herb associates, generally at their western limit, include Aristolochia 
serpentaria, Aquilegia canadensis, Arisaema dracontium, Aster teranus, Chas- 
manthium latifolium, Geum canadense, Parietaria obtusa, Senecio obovatus 
and Verbesina virginica. Carex edwardsiana is closest morphologically to C. 
oligocarpa, to which it has previously been referred. Both have the trigonous, 
basally tapering, apically constricted or beaked perigynia with numerous finely 
impressed nerves characteristic of section Oligocarpae. In general aspect, C. ed- 
wardsiana differs from C. oligocarpa in having shorter culms with more basally 
clustered spikelets and leaves usually much exceeding the culms. It is not as 
densely cespitose as C. oligocarpa and forms clumps from 10-20 cm in diam- 
eter by means of short, stout rhizomes. It tends to have more perigynia per 


Bridges & Orzell: New Carez from the Edwards Plateau 153 


spikelet, with these more closely clustered on the spikelet axis. The perigynia 
and achenes are larger than those reported for C. oligocarpa. The achene lacks 
an apiculus, and is not as long stiped as C. oligocarpa. C. oligocarpa also has 
reddish zones on the staminate scales, while those of C. edwardszana are all 
white hyaline. The following key can serve to distinguish these two species. 


Plants densely cespitose, to 50 cm tall, culms about the length of the longest 
leaves, spikelets mostly in the upper half of the culms; perigynia aver- 
aging 4 per spikelet (-8 in some cases), overlapping each other less than 
half their length; perigynia 3.5-4 mm long, achene 2.0-2.3 mm long, dis- 
tinctly stipitate and apiculate; scales of the staminate spike with white 
to reddish brown with green midnerve ...... Carez oligocarpa Schkuhr. 


Plants loosely to densely cespitose, often forming clumps to 10-20 cm in 
diameter by short, stout rhizomes, 25-35 cm tall, longest leaves much 
exceeding the culms, spikelets mostly in the lower half of the culms, 
densest in the lowest 5 cm of the culm; perigynia averaging 7 per spikelet 
(fewer on upper spikelets or on poorly developed culms), overlapping for 
half of their length; perigynia 4.5-5.0 mm long, achene 2.5-3.0 mm long, 
essentially sessile and beakless; scales of staminate spike white hyaline 
with green midnerve ............. Carer edwardsiana Bridges & Orzell 


The nearest approaches of C. oligocarpa to the range of C. edwardsiana 
appear to be in Dallas and Cooke counties in north central Texas, and perhaps 
in western Louisiana or extreme eastern Texas. Both of these regions are about 
320 km from the nearest locations of C. edwardsiana. Although restricted to 
specific, isolated microhabitats, Carer edwardsiana seems to be frequent within 
this habitat and probably occurs in numerous localities in the Eastern and 
Southern Balcones Escarpment Sections of the Edwards Plateau. It may occur 
in all the counties between Travis and Medina and perhaps extends slightly 
farther to the north and west. Its total distribution is expected to follow the 
pattern of the other Balcones Escarpment endemics or near endemics (Amos & 
Rowell 1988), which are a conspicuous component of the vernal flora of these 
geographically isolated mesic ravines. 


ACKNOWLEDGMENTS 


Special thanks to William R. Carr of Austin, Texas, who gave us access 
to his collections of this species and provided ecological data from several 
localities. We wish to acknowledge Clemente F. Guzman III of Austin for 
preparing the illustration. Thanks to reviewers Guy Nesom and Marshall 
Johnston for useful comments and to Rupert Barneby for editing the Latin 


154 PHYTOLOGTA volume 67(2):148-154 September 1989 


description. Publication costs were provided by the Plant Resources Center 
at the University of Texas at Austin. 


LITERATURE CITED 


Amos, B.B. & C.M. Rowell, Jr. 1988. Floristic geography of woody and 
endemic plants. pp. 26-42 7n B.B. Amos & F.R. Gehlbach, eds., Edwards 
Plateau Vegetation. Baylor University Press, Waco. 


Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of 
Tezas. Texas Research Foundation, Renner. 


Palmer, E.J. 1920. The canyon flora of the Edwards Plateau of Texas. J. 
Arnold Arb. 1:233-239. 


Van Auken, O.W. 1988. Woody vegetation of the southeastern escarpment 
and plateau. pp. 43-55 7n B.B. Amos & F:R. Gehlbach, eds., Edwards 
Plateau Vegetation. Baylor University Press, Waco. 


Phytologia (September 1989) 67(2):155-157. 


SOLIDAGO SIMPLEX (COMPOSITAE: ASTEREAE), 
THE CORRECT NAME FOR S. GLUTINOSA 


Guy L. Nesom 
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A 


ABSTRACT 


Solidago simplez Kunth, typified by plants from south central México, 
is an earlier name for a widespread species of the United States and 
Canada that has been variously called S. decumbens, S. glutinosa, S. 
spathulata var. nana and S. spathulata var. neomezicanus. The recog- 
nition of a “tall” and “short” form of S. simplez in the western United 
States is difficult and may reflect the existence of independently derived 
ecotypes rather than separate lineages. 


KEY WORDS: Solidago, Asteraceae, taxonomy, México, North 


America. 


A population system of Solidago in the eastern Sierra Madre of México is 
conspecific with a widespread taxon of the United States and Canada. The 
Mexican plants are S. stmplez Kunth, which was described from plants from 
south central México, probably in the vicinity of México City, and they have 
been collected in the eastern Sierra Madre in the states of México, Hidalgo, 
Guanajuato, Tamaulipas and Nuevo Leon. Those from north of México have 
been variously called S. spathulata, S. decumbens and S. glutinosa. Rzedowski 
(1985) was the first to call attention to the probable synonymy of S. sim- 
plez with these taxa. Ringius (1986) and Ringius & Semple (1987) mapped 
the Mexican populations but apparently were unaware they had been named 
early in the 19th century. The latter workers essentially followed an ear- 
her study by Cronquist (1947) by treating the western (diploid) and eastern 
(tetraploid) segments of this species complex (in the U.S. and Canada) as sub- 
species of S. gluttnosa and recognizing varieties from within both subspecies. 
Solidago spathulata DC., a population system from the coastal strand of Ore- 
gon and California, was segregated by Ringius (1986) as a species distinct from 
widespread S. glutinosa, and | agree with this. 

Since the earlier name for S. glutinosa and its infraspecific taxa is S. sim- 
plex, a number of new combinations will be necessary for taxa north of México, 
but with the exception of the caveat offered below, I defer to Dr. Ringius in 
applying a taxonomic scheme to the variation pattern he has studied in detail. 


155 


156 PAR YS TOLL. OCG TA volume 67(2):155-157 September 1989 


My view of the taxonomy of the western segment of Solidago simplez is 
presented below. The synonymy is representative but not complete. 


Solidago simpler Kunth, Nov. Gen. Sp. 4 [folioj:81. 1818; 4 [quarto]:103. 
1820. TYPE: MEXICO. Santa Rosa (“Sanctae Rosae Mexicanorum”), 
September, Humboldt & Bonpland s.n. (HOLOTYPE: P, fiche!). 


S. glutinosa Nutt., Trans. Amer. Philos. Soc. 2. 7:332. 1840. S. spathulata 
subsp. glutinosa (Nutt.) Keck, Aliso, 4:104. 1958. [tall form]. 


S. multiradiata var. neomezicana A. Gray, Proc. Amer. Acad. Arts 17:191. 
1882. S. neomezicana (A. Gray) Woot. & Standl., Contr. U.S. Natl. 
Herb. 16:182. 1913. S. spathulata var. neomezicana (A. Gray) Crongq., 
Vasc. Pl. Pacif. Northw. 5:311. 1955. {tall form]. 


S. humilis var. nana A. Gray, Syn. Fl. N. Amer. 1(2):148. 1884. S. glutinosa 
var. nana (A. Gray) Cronq., Rhodora 49:76. 1947. S. spathulata var. 
nana (A. Gray) Cronq., Vasc. Pl. Pacif. Northw. 5:311. 1955. S. purshu 
var. nana (A. Gray) Farw., Amer. Midl. Nat. 12:72. 1930. [alpine, short 
form]. 


S. bonplandi O. Kuntze, Rev. Gen. Pl. 1:315. 1891. 
S. decumbens E. Greene, Pittonia 3:161. 1897. [tall form]. 


S. oreophila Rydb., Mem. N.Y. Bot. Gard. 1:387. 1900. S. decumbens var. 
oreophila (Rydb.) Fernald, Rhodora 38:202. 1936. [tall form}. 


Two varieties have been distinguished among the plants of Solidago simpler 
in western North America (var. nana and var. neomezicana of S. spathulata, or 
var. nana and var. glutinosa of S. glutinosa). The alpine plants are generally 
shorter with a more compact, fewer-headed capitulescence and are restricted in 
distribution and habitat. The taller plants are found from British Columbia to 
central México in a much wider range of habitats. In my opinion, it is difficult 
to provide more than an arbitrary separation of an alpine variety (var. “nana”) 
of short stature from a taller one of lower elevations (var. “glutinosa”). Both 
forms, with intermediates, have sometimes been collected from the same site 
and mounted on the same sheet. The taller plants may have a compact or 
more elongated capitulescence. Cronquist’s observation (1955, p. 311) that 
the varietal names in this group refer to “intergradient but fairly well marked 
ecotypes” appears to be true, and I suggest that in the western United States 
the alpine forms may be independently derived at various localities rather 
than constituting a single lineage. If so, such forms should not receive formal 
taxonomic recognition. 

The Mexican plants have noticeably less pubescent achenes than plants 
widespread in the western United States and a greater tendency to produce 


Nesom: Solidago stmplez correct name for S. glutinosa 157 


scattered cilia along the petiole margins. The differences, however, are only of 
degree and do not call for separate taxonomic status. Comments on a putative 
hybrid between S. simpler and a species endemic to eastern México are found 
in a separate paper (Nesom 1989). 


ACKNOWLEDGMENTS 


I appreciate the helpful comments of B.L. Turner and Marshall Enquist. 


LITERATURE CITED 


Cronquist, A. 1947. Notes on the Compositae of northeastern United States. 
IV. Solidago. Rhodora 47:69-79. 


1955. Part 5: Compositae. Jn Hitchcock, C.L. et al. Vascular 
Plants of the Pacific Northwest. Univ. of Washington Press, Seattle. 


Nesom, G.L. 1989. New species of Mexican Solidago (Compositae: Astereae). 
Phytologia 67(2):142-147. 


Ringius, G.S. 1986. A biosystematic study of the Solidago spathulata DC. - 
S. glutanosa Nutt. complex (Compositae: Astereae). Ph.D. dissertation, 
Univ. Waterloo, Waterloo, Canada. 


Ringius G.S. & J.S. Semple. 1987. Cytogeography of the Solidago spathulata 
- glutinosa complex (Compositae: Astereae). Canad. J. Bot. 65:2458- 
2462. 


Rzedowski, J. 1985. Flora Fanerogamica del Valle de Mézico. Vol. 2, Di- 
cotyledoneae. Instituto de Ecologia, México, D.F. 


Phytologia (September 1989) 67(2):158-167. 


NEW SPECIES, NEW SECTIONS, AND A TAXONOMIC OVERVIEW 
OF AMERICAN PLUCHEA (COMPOSITAE: INULEAE) 


Guy L. Nesom 
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A 


ABSTRACT 


Two new species of Pluchea, both related to P. foetida and P. rosea, 
are proposed from México. Pluchea yucatanensis sp. nov. is de- 
scribed from Belize and the Yucatan peninsula of México and its occur- 
rence is noted as an adventive in the United States. Pluchea mexicana 
comb. nov. is endemic to alkaline plains of northeastern San Luis Po- 
tosi. The two newly proposed species and others with sessile, clasping 
and mostly oblong leaves are segregated as Pluchea sect. Amplecti- 
folium sect. nov., and a key is provided to the six American species 
of that group. In addition, the identity of sect: Pluchea is clarified and 
another section is proposed, sect. Pterocaulis sect. nov., with plants 
characterized by winged stems from decurrent leaves. Pluchea sericea 
is maintained within the genus as sect. Phalacroline A. Gray, possibly 
closely related to Pluchea lanceolata. 


KEY WORDS: Asteraceae, Pluchea, México, floristics, taxonomy. 


Pluchea comprises about 40 species. More than half of these are in the 
New World, with the remainder in Africa, Asia and Australia. Robinson & 
Cuatrecasas (1973) reviewed the generic boundaries among Pluchea, Tessaria 
and Berthelotia, but apart from this and the early studies of de Candolle and 
Asa Gray, there has been no attempt to delimit the infrageneric structure of 
Pluchea. In the the course of a floristic study of Mexican Pluchea, it quickly 
became clear that for the most part, the American species could be arranged 
in morphologically distinct groups, and in the present paper formal status is 
proposed for these as sections. Old World species have been associated with 
these groups in some cases, but a more complete view of the genus still remains 
to be provided. Nevertheless, I believe the preliminary structure proposed here 
will prove to be helpful. Descriptions are first presented for two new Latin 
American species. 

Godfrey (1952) segregated Pluchea rosea Godfrey from P. foetzda (L.) DC. 
Both are species primarily of the southeastern United States and the West 


158 


Nesom: Overview of North American Pluchea 159 


Indies. He also described P. rosea var. mezicana Godfrey, a series of popula- 
tions restricted to México and disjunct from the main range of P. rosea. In 
reviewing the taxonomy of Mexican Pluchea, I find that two separate elements 
are represented among the plants cited by Godfrey as P. rosea var. mezicana, 
and I recognize one of them as a species distinct from P. rosea. 


Pluchea mexicana (Godfrey) Nesom, comb. et stat. nov. Based on Pluchea 
rosea Godfrey var. mezicana Godfrey, J. Elisha Mitchell Sci. Soc. 68:269. 
1952. TYPE: MEXICO. San Luis Potosi: alkaline plains, Hacienda de 
Angostura, 5 Aug 1891, C.G. Pringle 3813 (HOLOTYPE: GH; Isotypes: 
C,F!,MU!.NY,PH,US!). 


Erect, herbaceous perennials ca 4-6 dm tall. Stems with a mixture of sessile 
and stipitate glands and long, thick, multicellular, non glandular trichomes. 
Leaves alternate, lanceolate- oblanceolate-oblong, erect-ascending, 25-45 mm 
long, 7-18 mm wide, with minutely serrulate margins, sessile, clasping, slightly 
auriculate, coarsely pubescent with a dense mixture of thick, long and short, 
gland tipped hairs and thick, viscid hairs with colored crosswalls, sessile resin 
glands absent or mixed with the stipitate ones. Heads 6-7 mm wide, sessile, 
immediately subtended by bracteal or reduced leaves. Phyllaries lanceolate, in 
3-4 series of nearly equal length, the innermost 5-6 mm long, rose-purple with 
sharply delimited, light colored margins, sclerified and mostly glabrous on the 
lower half, stipitate glandular and viscid pubescent with thick trichomes on 
the upper half. Pistillate flowers numerous, the corollas filiform, pink to pur- 
plish, 4.4-4.9 mm long, minutely 3 lobed and glandular at the apex, the style 
branches barely if at all exserted; hermaphroditic flowers ca 25-30, the corol- 
las 5 mm long, gradually ampliate, the lobes triangular-lanceolate, glandular, 
purplish. Achenes cylindrical, 6 nerved, very sparsely strigose and glandular, 
mature size not seen; pappus a single series of 18(-20) minutely barbellate 
bristles 4 mm long. 

Additional collections examined. MEXICO. San Luis Potosi. Media Luna, 
near Rio Verde, Palmer 75 (CM,F,MO,US); Minas de San Rafael, plains be- 
tween Tiburcio and Angostura, June 1911, Purpus s.n. (F,US); and July 1911, 
Purpus s.n. (F,MO,US). These three collections were cited by Godfrey (1952) 
as P. rosea var. mezicana. 

Pluchea mezicana is endemic to the gypseous, alkaline plains (ca 800-1000 
m) north of Rio Verde, San Luis Potosi and flowers June to July. This area 
is common to several other narrow endemics of Compositae (i.e., Geissolepis 
suaedaefoha B. Robinson, Stephanodorza tomentella (B. Robinson) E. Greene, 
Viguiera potosina S.F. Blake, Pinaropappus multicaulis Brandegee and Cir- 
sium ezcelsius (B. Robinson) Petrak. Attempts in September 1988 to relocate 
populations of the Pluchea and Cirsium were unsuccessful, although the other 
species noted above were found. 


160 PHYTOLOGIA volume 67(2):158-167 September 1989 


Pluchea mezicana differs from P. rosea in its vestiture of stipitate glandular 
hairs, sometimes mixed with sessile glands and thick, viscid but non glandular 
hairs. Pluchea rosea has only sessile glands and a puberulous vestiture of non 
glandular hairs that are apically attenuate and usually drawn out into long 
filiform extensions. Additionally, the phyllaries of P. mezicana are mostly 
of even length, the basal portions are strongly sclerified, they are rosy with 
sharply delimited, white margins and only the outermost are densely hairy. 
The inner are glandular but otherwise sparsely hairy to glabrous. Godfrey 
(1952, p. 269) also observed some of these differences and noted that the 
vestiture of the plants from San Luis Potosi is “. . . quite unlike anything 
found in [P. rosea] elsewhere.” I also call attention to the disjunct geographical 
position of the plants from San Luis Potosi and their very different habitat; 
P. rosea is known only from coastal or near coastal sites, and in México, 
it is known from scattered localities (cited below). I believe the distinctive 
morphological features of these inland plants are of enough significance to 
warrant separating P. mezicana at the specific rank, although it seems clear 
that the two taxa are closely related. 

Pluchea rosea was noted by Godfrey (1952) to occur in the southeastern 
United States, West Indies and México. With the separation of P. mezicana, 
only one of the Mexican collections he cited as P. rosea remains correctly iden- 
tified as that species (Quintana Roo), but I have studied additional collections 
of P. rosea from Central America, which are included among the following: 
BELIZE. Between London and Rancho along the old Northern Highway to 
Maskall, wet margin of Eleocharis marsh, 50 m, 20 Mar 1987, Davidse & 
Brant 32896 (US). 

HONDURAS. Depto. Gracias a Dios, W of Brus Laguna, cut over forest, 
sea level, 23 Apr 1971, Nelson & Herndndez s.n. (MO). 

MEXICO. Quintana Roo, in savanna, Lake Chichancanab (Laguna Chan- 
kabnab), 28-29 Jul 1932, Steere 2407 (F,US). 

NICARAGUA. Dept. Zelaya: ca 11.9 km SW of Bismuna Tara, along river 
in savanna, 19 Apr 1978, Stevens 7782 (F-2 sheets); vicinity of Awastara, 
savanna, 2 Jul 1980, Stevens 17759 (F). 

Another taxon distinctive in morphology but apparently unnamed is known 
from several collections primarily from the Yucatan peninsula of Belize and 
México. 


Pluchea yucatanensis Nesom, sp. nov. TYPE: MEXICO. Campeche: In 
savanna, Champoton, 7-15 Jul 1932, Steere 1844 (HOLOTYPE: LL!; 
Isotype: US!). 


P. mezicanae habitu et morphologia foliorum sed absentia tri- 
chomatum nonglandiferorum differt. Phyllaria interioria lineari- 
lanceolata, 5-6 mm longa, extima ovati-lanceolata longitudine in- 
terioria aequantia. 


Nesom: Overview of North American Pluchea 161 


Perennials 2-6 dm tall, single stemmed from the base and unbranched until 
the capitulescence, the stems, leaves and phyllaries glandular but lacking other 
pubescence. Leaves leathery and slightly succulent, with sessile to slightly 
stipitate glands, alternate, mostly oblong-obovate with obtuse, mucronulate 
apices, subclasping and slightly auriculate, ascending to erect, 3-5 cm long, (6- 
)15-19 mm wide, slightly reduced in size immediately below the capitulescence, 
the margins serrulate with (6-)8-11 pairs of small teeth or mucros. Heads 8-14 
in small, terminal, corymboid clusters, on ultimate pedicels 2-5 mm long, each 
immediately subtended by a lanceolate bracteal leaf, turbinate to campanulate 
turbinate, 4-5 mm wide; phyllaries densely and prominently stipitate glandular 
near the apices, stipitate to sessile glandular below, without other pubescence, 
in 3-4 slightly graduated series, the innermost linear lanceolate, 5-6 mm long, 
the outer ovate lanceolate, equally as long or nearly so, or sometimes only 
ca half as long. Pistillate flowers numerous, the corollas pink to lavender, 
filiform, 3.0-3.3 mm long, minutely 3 lobed and glandular at the apex, the 
style branches exserted 0.3-1.0 mm; hermaphroditic flowers 20-22, the corollas 
pink to cream, 4.5-5.0 mm long, gradually ampliate, the lobes linear lanceolate, 
0.9-1.0 mm long, glandular. Achenes cylindrical, 0.8-0.9 mm long, 6-8 nerved, 
sparsely strigose, eglandular; pappus of 18-20, minutely barbellate bristles 4.3- 
4.8 mm long. 

Additional collections examined: BELIZE. 9.8 mi S of Belize, mangrove 
swamp, 5 Jun 1973, Croat 23817(LL,MO,US); 1 mi W of jct with road to Fer- 
guson Bank, ca 8 mi E of Hattiesville, along Hector Creek Rd, low, intermit- 
tently wet area, many temporary ponds, sedges, palmetto, and broad-leaved 
scrub, 16 Aug 1971, Sorensen 7072 (F,MO,US); mile 13, Western Highway, 
growing in pools in sandy, waste ground, 30 Aug 1980, Whztefoord 2268 (MO). 

UNITED STATES. Mississippi: Hancock Co., sandy, low area along Jordan 
River S of Kiln, edge of mixed woods with Serenoa, Sabal, Nyssa, Tarodium, 
30 May 1967, Jones 12656 (TEX). 

Pluchea yucatanensis appears to be restricted essentially to the Yucatan 
peninsula of México and Belize, where it occurs at low elevations near the coast 
and flowers June to August. One collection of P. yucatanensis has been made 
from a near coastal locality in Mississippi of the southern United States, where 
it probably is adventive. It is similar to P. roseain habit and leaf morphology 
but the whole plant without any type of pubescence except primarily sessile 
glands. The leaves are leathery and slightly succulent and are restricted at 
flowering to the upper half of the strictly erect, unbranched stems. 


PLUCHEA SECTION AMPLECTIFOLIUM 


Both taxa newly proposed here as species, as well as P. oblongzfolia of east- 
ern South America and Pluchea rosea, P. foetidaand P. longifolia of North and 
Central America, have erect to ascending, lanceolate to oblanceolate-oblong 


162 PRY TOPOGTA volume 67(2):158-167 September 1989 


leaves with serrulate margins and sessile, clasping, slightly auriculate bases. 
The plants are herbaceous perennials and produce strictly erect, mostly un- 
branched stems. The inner phyllaries are not strongly differentiated from the 
outer. The species vary among themselves in the production of glands and eg- 
landular trichomes on the achenes. Both pistillate and hermaphroditic corollas 
have glandular lobes. In all of these species, the uppermost bracteal leaves in 
the capitulescence are reduced in size and petiolate, pointing to a close re- 
lationship with sect. Pluchea, which has petioled leaves. These taxa form a 
closely related cluster of species that I recognize as a distinctive section within 
the genus. The African species Pluchea dioscoridis (L.) DC. has similar leaf 
morphology and may also belong here, but it has eglandular corolla lobes and 
glandular anther appendages. A key to the American species of the section 
follows. 


Pluchea sect. Amplectifolium Nesom, sect. nov. TYPE SPECIES: Pluchea | 
foetida (L.) DC. 


Folia erecti-ascendentia oblanceolati-oblonga marginibus ser- 
tatis et basibus sessilibus amplectentibus parum auriculatis. 


ARTIFICIAL KEY TO THE AMERICAN SPECIES OF SECT. 
AMPLECTIFOLIUM 


1. Leaves mostly 8-20 cm long, 3-7 cm wide; heads campanulate-cylindric, 
the innermost phyllaries 9-11 mm long, middle phyllaries 2.5-3.0 mm 
wide; southern Florida -... ..):/55 fiise side's ecto eae P. longifolia Nash 


1. Leaves mostly 3-10 cm long, 1-3 cm wide; heads turbinate-campanulate, 
the innermost phyllaries 5-8 mm long, middle phyllaries 1.0-1.5 mm wide 

(2) 
2. Plants stipitate to sessile glandular, without other pubescence; Yu- 
catan peninsula of México and Belize, adventive in Mississippi, 
United States) s.cso.t. cet. co sees os Vee ee P. yucatanensis Nesom 


2. Plants glandular with sessile, resin glands or gland tipped hairs, also 
Hairy foes ie cece dene nen obec ec eseue st eee (3) 


3. Plants with gland tipped hairs, sessile resin glands lacking or mixed 
with numerous stipitate ones; phyllaries in several series of nearly equal 
length, sclerified and mostly glabrous on the proximal half, viscid pubes- 
cent with thick trichomes and stipitate glandular on the distal half; east 
central San Luis Potosi, México ........ P. mexicana (Godfrey) Nesom 


Nesom: Overview of North American Pluchea 163 


3. Plants with sessile, translucent, yellowish, resin glands, stipitate glands 
absent or rare; phyllaries in several strongly graduated or even series, 
relatively evenly thick-herbaceous, moderately to densely puberulent or 
appressed villous from base to tip with fine hairs .................. (4) 


4. Heads basally rounded with a prominent impression at the peduncle 
insertion; phyllaries with 3-7, prominent, thin, longitudinal nerves, 
the outer and inner phyllaries of relatively even length, glandular 
and cobwebby ciliate margined but mostly without other vestiture; 
COASLAISD CASI Gs a coishens mmeila e Siete wien Hier arene oe P. oblongifolia DC. 


4. Heads basally rounded and impressed to obtuse; phyllaries 1 nerved, 
equal to prominently graduated in length, at least the outermost 
glandular ‘and ipuberulent, villous: sje): ss: anes! 0% sicreyayoistornis) ear si (5) 


5. Heads 6-12 mm wide, basally rounded to impressed; phyllaries thinly 
arachnoid pubescent, with sessile glands, the inner 5-8 mm high, the 
outer ovate, much shorter than the inner 2-3 series; phyllaries and corol- 
las creamy white; southeastern United States to Texas, also Hispanola 
and Veracruz, México, where probably adventive .. P. foetida (L.) DC. 


5. Heads 5-9 mm wide, basally obtuse to barely acute; phyllaries usually 
arachnoid and commonly also with dense, thick, viscid hairs, the in- 
ner 4-6 mm high, the outer ovate-acuminate to ovate-lanceolate, half to 
equally as long as the innermost; corollas and often the phyllaries pink 
to purplish; SE United States to Texas, West Indies, Gulf and scattered 
localities along the Caribbean coast from Campeche, México, to 
Ramage Pare cere eee. eect ne ine ea Geren are P. rosea Godfrey 


Pluchea roseais very similar to P. foetida and many apparent intermediates 
are found in the southeastern United States. The notes by Cronquist (1980) 
are particularly helpful in distinguishing the two. The plants of P. rosea cited 
from Nicaragua may be influenced by genes from P. yucatanensts, because the 
heads are somewhat narrower and less pubescent and the phyllaries narrower 
and more equal in length than in plants typical of the southeastern U.S. 

I have seen only one collection of Pluchea foetida from anywhere in Central 
America or México: Veracruz, 3 km antes de llegar a las Choapas, 13 May 
1980, Calzada 5964 (TEX). It is typical in morphology of conspecific plants of 
the southeastern United States. 


PLUCHEA SECT. PLUCHEA 
Pluchea Cass., Bull. Sci. Soc. Philom. Paris 1817:31. 1817. TYPE SPECIES: 


Conyza marilandica Michx. [=P. purpurascens (Sw.) DC. = P. odorata 
(L.) Cass., sensu Gillis, 1977]. 


164 PHYIEOLOGTA volume 67(2):158-167 September 1989 


Stylimnus Rafin., J. Phys. Chim. Hist. Nat. 89:100. 1819. TYPE SPECIES: 
Conyza marilandica Michx. Pluchea sect. Stylimnus (Rafin.) DC., Prodr. 
5:540. 1836. Other synonyms as noted by Godfrey (1952). 


The species of sect. Pluchea are herbaceous to suffrutescent perennials char- 
acterized by petioled, mostly elliptic lanceolate leaves with attenuate to obtuse 
bases. The inner phyllaries are but little differentiated from the outer, and the 
achenes are sparsely hairy and variably glandular on the nerves and faces. 
Both pistillate and hermaphroditic corollas have glandular lobes. 

Species included: Pluchea camphorata (L.) DC., P. odorata (L.) Cass., P. 
fastigiata Griseb., P. symphytifolia (Miller) DC. and perhaps others. Pluchea 
indica Less., an Asian species, almost certainly belongs here, but it has more 
strongly differentiated inner phyllaries and glandular anther appendages. As 
noted by Cooperrider & Galang (1965), on Pacific islands it forms sterile hy- 
brids with P. symphytzfolia. 


PLUCHEA SECT. PTEROCAULIS 


Another group of primarily American species shows strong morphological 
unity and warrants recognition as a section, which is proposed here. 


Pluchea sect. Pterocaulis Nesom, sect. nov. TYPE SPECIES: Pluchea 
salictfolia (Miller) S.F. Blake 


Caules alati extensionibus foliorum lineari-lanceolatorum ser- 
ratorum longi-decurrentium. 


Species included: Pluchea lariflora Hook. & Arn., P. macrocephala DC., 
P. microcephala Godfrey, P. parvifolia(A. Gray) Godfrey, P. sagittalis (Lam.) 
Cabr. (including P. suaveolens (Vell.) O. Kuntze and P. salicifolia (including 
P. subdecurrens Cass. and P. adnata (Willd.) Mohr). The African species P. 
ovalis Pers.) DC. appears to belong here as well. 

The species of Pluchea sect. Pterocaulis are suffrutescent perennials char- 
acterized by stems completely winged by the basal extensions of the linear- 
lanceolate, serrate, long decurrent leaves. Both pistillate and hermaphroditic 
corollas have glandular lobes. The achenes vary among species in their pro- 
duction of hairs and glands. The species limits among these taxa are in need 
of taxonomic study. 


PLUCHEA SECT. PHALACROLINE 


Robinson & Cuatrecasas (1973) presented a seminal discussion of generic 
limits among the New and Old World species centered around Pluchea, includ- 
ing Tessaria and Berthelotia, although I believe the groupings proposed in the 


Nesom: Overview of North American Pluchea 165 


present paper will also prove to be useful as a starting point for understanding 
the phylogeny. In my opinion, Robinson & Cuatrecasas were correct in recog- 
nizing the polyphyletic aspect of Tessaria sensu Cabrera (1939) but incorrect 
in limiting the genus to only T. integrifoha Ruiz & Pavon, which clearly is 
congeneric with T. absinthioides (Hook. & Arn.) Cabrera. The two are set 
apart from the rest of Pluchea by their combination of alveolate and palea- 
ceous receptacles, spreading reflexed inner phyllaries, extremely peculiar style 
nectary floral morphology, thickened margins of the hermaphroditic flowers, 
pappus bristles basally united into a thick cylindrical cup and glabrous achenes. 
Tessaria integrifolia is the more advanced of the two species in its typically 
solitary (but variable in number), hermaphroditic flowers with deeply cleft 
corolla lobes. The taxonomy accepted by Robinson & Cuatrecasas (1973) was 
summarized by their recognition of two genera among the plucheoid species: 
1) a heterogeneous Pluchea, including Berthelotia and 2) Tessarza. 

Pluchea sericea (Nutt.) Coville of the southwestern United States and 
northwestern México has recently been treated as Tessaria (Correll & John- 
ston, 1970), but it clearly does not belong with that genus, as it lacks the 
distinctive features of floral and pappus morphology. As suggested by Ryd- 
berg in the publication of his combination to Berthelotza, it may be related to 
the Afro-Asian species P. (Berthelotia) lanceolata (DC.) Hiern. (the type of 
Berthelotza), with which it shares pappus bristles with expanded apices and a 
sericeous vestiture. Gray also allied it with Berthelotia (see below). If treated 
as a monotypic genus, Aven Nelson’s name is already available. Until more 
detailed and incisive techniques are applied to the study of its relationships, 
I maintain it as Pluchea and recognize its isolated position among American 
members of the genus, but possible relatedness to P. lanceolata, by treating it 
as a member of section Phalacroline. 


Pluchea sect. Phalacroline A. Gray, Pl. Wright. 1:102. 1849. TYPE SPECIES: 
Tessaria borealis A. Gray (=Pluchea sericea). Gray discussed a possible 
relationship between the type species and Berthelotza lanceolata. 


Pluchea sect. Berthelotia (DC.) A. Gray, Proc. Amer. Acad. Arts 17:212. 
1882. Berthelotta DC., Prodr. 5:375. 1836. TYPE SPECIES: B. lanceo- 
lata DC. (=Pluchea lanceolata (DC.) Hiern.). Gray cited both B. lance- 


olata and Tessaria borealis as members. 


Eremohylema A. Nelson, Univ. Wyoming Pub. Bot. 1:54. 1924. TYPE 
SPECIES: E. sericea (Nutt.) A. Nelson (=Pluchea sericea). 


Because the nomenclature of Pluchea sericea has been somewhat confused, 
a summary is presented here. 


Pluchea sericea (Nutt.) Coville, Contr. U.S. Natl. Herb. 4:128. 1893. Based 
on Polypappus sericeus Nutt., J. Acad. Nat. Sci. Philad. ser. 2, 1:178. 


166 PHY TOLOGIA volume 67(2):158-167 September 1989 


1847. TYPE: UNITED STATES. “Rocky Mountains of Upper Cali- 
fornia,” Gambell s.n. Berthelotia sericea (Nutt.) Rydb., Bull. Torrey 
Bot. Club 33:154. 1906. Eremohylema sericea (Nutt.) A. Nelson, Univ. 
Wyoming Pub. Bot. 1:54. 1924. Tessaria sericea (Nutt.) Shinners, Sida 
3:122. 1967. 


Tessaria borealis Torrey & A. Gray ex A. Gray, Pl. Fendl. 75. 1849. Gray 
cited collections (SYNTYPES) by Fremont, Coulter and Emory. Tes- 
sarta borealis A. Gray, nom. superfl., Pl. Wright. 1:102. 1852. Pluchea 
borealis (Torrey & A. Gray ex A. Gray) A. Gray, Proc. Amer. Acad. Arts 
17:212. 1882. 


Tessaria borealis was attributed several times by Gray (1849; 1882) to 
Torrey & Gray (ex A. Gray in Pl. Fendl.). Shortly after his study of the Fendler 
collections, Gray republished the name with a type and Latin description as 
T. borealis A. Gray (1852), but this must be regarded as superfluous. In the 
same paper, he noted that a still earlier publication of the same name was 
mistakenly attributed to DeCandolle by Torrey (in Emory, Notes Military 
Reconn. 143. 1848) and that it had been intended as an enumeration, not a 
description. Nuttall’s original name, Polypappus sericeus (1847), was given as 
a synonym in both of Gray’s reports (1849; 1852). 

Some shrubby species of Pluchea in South America do not belong with any 
of the groups discussed here. P. dodoneaefolia (Hook. & Arn.) H. Robins. 
& Cuatr., P. zamalloae (Cabrera) H. Robins. & Cuatr. and P. fiebrigu H. 
Robins. & Cuatr. were noted by Robinson & Cuatrecasas (1973, p. 280) to 
form a “macroscopically evident group,” and P. chingoyo DC. of Peru appears 
to be morphologically isolated, although its cordate, short petiolate leaves may 
point to a relationship with the species of sect. Amplectifolium. 

The only other named section of Pluchea of which I am aware is sect. 
Hebephora DC. (Prodr. 5:453. 1836). In it De Candolle included two Asian 
species, P. hirsuta (L.) Less. and P. scabrida DC., which have fimbrillate- 
hirsute receptacles. Judging from the descriptions, these may be related to 
sect. Pluchea. 


ACKNOWLEDGMENTS 


I thank Carol Todzia and Billie Turner for their reviews and comments and 
F, MO, US and CM for loans of specimens. 


Nesom: Overview of North American Pluchea 167 


LITERATURE CITED 


Cabrera, A.L. 1939. Las especies Argentinas del genero “Tessaria.” Lilloa 
4:181-189. 


Cooperrider, T.S. & M.M. Galang. 1965. A Pluchea hybrid from the Pacific. 
Amer. J. Bot. 52:1020-1026. 


Correll, D.S. & M.C. Johnston. 1970. Manual of the Vascular Plants of 


Tezas. Texas Research Foundation, Renner, Texas. 


Cronquist, A. 1980. Vascular Flora of the Southeastern United States. Vol- 
ume I, The Asteraceae. Univ. North Carolina Press, Chapel Hill. 


Gillis, W.T. 1977. Pluchea revisited. Taxon 26:587-591. 


Godfrey, R.K. 1952. Pluchea, section Stylimnus, in North America. J. Elisha 
Mitchell Sci. Soc. 68:238-271. 


Robinson, H. & J. Cuatrecasas. 1973. The generic limits of Pluchea and 
Tessaria. Phytologia 27:277-285. 


Phytologia (September 1989) 67(2):168-206. 


REVISIONARY TREATMENT OF THE GENUS SJNCLAIRIA, INCLUDING 
LIABELLUM (ASTERACEAE, LIABEAE) 


B. L. Turner 
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A 


ABSTRACT 


A revisionary study of the genus Sinclatria (including Liabellum) 
is rendered. It is composed of 23 species, all confined to México and 
Central America, except for S. polyantha which extends into Colombia 
in South America. Two subgenera are recognized: Stnclairta with 15 
species and Megaliabum with 8 species. The latter subgenus contains 
2 sections: Megaliabum with 3 species and Liabellum with 5 species. 
One new species, S. hintoniorum B. Turner, belonging to the section 
Liabellum, is described and illustrated, and a new variety of S. hy- 
poleuca, S. h. var. minor B. Turner, is proposed. Keys to the species 
and distribution maps are provided and a diagram showing hypothetical 
relationships is presented. 


KEY WORDS: Asteraceae, Liabeae, Sznclairia, Liabellum, México, 
Central America. 


INTRODUCTION 


Sinclatria is a wholly American genus of 23 species largely confined to 
tropical and subtropical regions of México and Central America. A single 
species (5. polyantha) just barely extends into Colombia. Prior to Rydberg’s 
(1927) treatment, most of the species were placed in the large genus Liabum. 
Rydberg treated all of the species known to him, placing these in four genera: 
Sinclairta, Sinclairiopsis, Megaliabum and Liabellum. 

Robinson (1983) provided an excellent overview of this complex, pointing 
out its relationship to Liabum and yet other genera of the tribe Liabeae. He 
placed the 29 species known to him in 2 genera, Sinclairia (25 spp) and Liabel- 
lum (4 spp); the former included Rydberg’s genera, Sinclairza, Sinclarriopsis 
and Megaliabum; the latter included all of the species which Rydberg relegated 
to Liabellum. The present treatment places all of the species recognized by 
Rydberg (1927) and Robinson (1983) in the genus Sinclairia. | recognize two 
subgenera within the genus: Sinclairza and Megaliabum. The former contains 


168 


Turner: Revision of Sinclatria including Liabellum 169 


Rydberg’s Sinclazria and Sinclairiopsis, the latter contains his Megaliabum and 
Liabellum (both of which I treat as sectional categories). My account differs 
from that of Robinson (1983) in that Lzabellum and Megaliabum are treated 
as sections of Sinclairia, whereas Robinson positioned Megaliabum within his 
concept of Sinclairia, retaining Liabellum in the sense of Rydberg. The re- 
lationships of these several categories are shown in my Figure 1. Overall, I 
recognize but 23 species in the genus Sznclairia, 15 in subgenus Sznclairia and 
8 in the subgenus Megaliabum. 


CHROMOSOME NUMBERS 


Chromosome counts are known for relatively few species of Sinclairia and 
these were summarized by Robinson et al. (1985). They gave the base chromo- 
some number of Sinclairia as z = 16, but this is largely conjectural because the 
few counts reported are poorly documented. The numbers of only 3 species of 
Sinclairia, all belonging to the subgenus Sinclazrza, have been reported. These 
are: 

S. discolor n = 17 or 18 pairs 
S. hypochlora n = 15 or 16 pairs 
S. sublobata n = 15, 16 or 17 pairs. 

Apparently the difficulty in establishing an accurate count for the genus re- 
lates to the presence of an exceptionally large chromosome “pair” at diakinesis 
in meiotic material. This “pair” might be a bivalent, tetravalent or perhaps a 
hexavalent. Most likely it is either a bivalent or tetravalent, making the base 
number z = 15 or 16. 


GENERIC RELATIONSHIPS 


Robinson (1983) has provided an admirable study of the generic relation- 
ships of the tribe Liabeae. He positioned Sinclairia (as conceived here) in 
the subtribe Liabinae, aligning it to the South American genera Microhabum 
and Austroliabum (cf. his figure 1, showing a suggested evolutionary scheme 
for the tribe). I am not able to improve upon his suggestions; my superficial 
examination of the genera concerned suggests the alignment is sound. 


SPECIES RELATIONSHIPS 


Sinclairia, with 23 species, nearly all of which are confined to México or 
Central America, appears to be a monophyletic group. Within this, three evo- 
lutionary lines can be recognized. I have recognized these as sectional groups: 
1.) sect. Sinclazria with 15 species, these comprising subgenus Sinclazria; 
2.) sect. Megaliabum with 3 species; and 3.) sect. Liabellum with 5 species. 
Sections Megaliabum and Liabellum have been positioned in the subgenus Me- 
gahabum. Rydberg (1927) recognized these several sections as distinct genera 


170 PHY LOD OiGdA volume 67(2):168-206 September 1989 


S. discolor (1) 
S. broomeae (2) 
S. glabra (3) 
S. hypochlora (4) 
S. vagans (7) 
S. polyantha (8) 
see itis Sap 4 S&S. andromachioides (10) 
qiotvtes with small S. deppeana ( 11) 
to moderate-sized S.tajumulcensis (1 2) 
heads 
Subgen. 
Sinclairia (sect. Sinclairia) 
clambering shrubs 
or hemiapiniphyies: i ['S Veutiobetalts) 
headed capitulescence | 9. liebmannii (13) 
S. similis (14) 
S. deamii (9) 
SINCLAIRIA 
S. sericolepis (6) 
S. klattii (15) 
S. andrieuxii (16) 
S. moorei (17) 
ubgen. 
egaliabum 
(sect. Megaliabum) 
—e ne ae 
onaibe walltoepe Hemis S. pringlei (18) oe} 
(sect. Liabellum S. palmer (20) 
tuberous herbs S. gentryi (21) 
with very large heads S. cervina (22) 
S. angustissima (23) 


Fig. 1. Hypothetical relationships among species of Sinclairia (numbers in parenthesis refer 
to arrangement in text). 


Turner: Revision of Sinclairia including Liabellum 171 


(plus the monotypic genus Sznclazriopsis, based upon S. klattwz, which I po- 
sition in the sect. Sinclairia). Robinson (1983), however, included the sect. 
Megaliabum in his concept of Sinclazria, but retained Lzabellum as distinct. I 
tend to believe that Megaliabum stands, morphologically, somewhere between 
the largely clambering or hemiepiphytic sect. Sznclazrza and the largely suffru- 
ticose or herbaceous sect. Liabellum. In total attributes the section Megaliabum 
appears closest to the section Liabellum and, therefore, I have included both of 
these in the subgenus Megaliabum. Their treatment as three subgenera within 
Sinclairia might be equally defensible but I have emphasized habit, capitules- 
cence, involucral structure and biogeography in positioning Megaliabum with 
Liabellum. Robinson (1983) positioned the former within Sznclairza largely 
because of its habit (shrubby) and lack of tuber-like roots, Liabellum being 
strictly herbaceous with tuber-like roots. He also emphasized the mostly ses- 
sile leaves of the latter (vs petiolate in Sinclairia). My view of the relation- 
ships of the various species within these several sections is shown in Figure 1. 
Whether the newly described, S. hintoniorum has tuber-like roots or not is 
not known, but it appears to link the sect. Megaliabum to sect. Liabellum by 
yet other features (e.g. head size, achenial characters and suffruticose habit). 

Clearly, the several sections recognized here are in much need of critical 
study, especially chromosomal and comparative DNA investigations. Until 
then, much that has been written about their relationships, one to the other, 
will remain speculative. 


TAXONOMIC TREATMENT 


Sinclairia Hook. & Arn., Bot. Beech. Voy. 433. 1841. TYPE SPECIES: 
Sinclairia discolor Hook. & Arn. 


Liabellum Rydb., N. Amer. Fl. 34:294. 1927. (not Liabellum Cabrera, 1954). 
Megalhabum Rydb., N. Amer FI. 34:293. 1927. 
Sinclairiopsis Rydb., N. Amer. Fl. 34:292. 1927. 


Herbaceous or suffruticose perennials with tuber-like roots, 0.5-2.0 m high, 
soft wooded shrubs or small trees to 7 m high, clambering vines to 30 m high 
or hemiepiphytes (sprouting in trees but soon producing large aerial roots 
which make their way to the ground and become anchored). Leaves opposite, 
petiolate or not, usually simple, but often deeply lobed, 3 nervate from below, 
the undersurfaces usually densely white tomentose or arachnoid, rarely not. 
Heads turbicampanulate to campanulate, radiate or eradiate, very large and 
borne 1-15 in stiffly erect cymes (subgenus Megaliabum) or relatively small 
and numerous borne in divaricately branched, corymbose panicle or rarely in 
drooping subfasciculate corymbs. Involucres subturbinate to campanulate, 3- 
6 seriate, imbricate or subimbricate, gradate. Receptacle epaleate, plane to 


172 PHY POLO GTA volume 67(2):168-206 September 1989 


somewhat convex, glabrous or rarely alveolate with pubescent ridges. Ray 
florets, when present, pistillate, fertile, the ligules yellow or rarely orange. 
Disk florets few to numerous, yellow, creamy white or orange, glabrous or 
sparsely pubescent without, the tube shorter than the poorly developed limbs, 
the lobes usually linear, 2-4 times as long as wide, but shorter than the throats. 
Anthers mostly yellow, the appendages lanceolate and acute to ovate rounded, 
tailed at the base, the filaments glabrous throughout. Style branches rather 
filiform, hispidulous like the upper shafts, merely recurving at maturity, the 
apices rounded. Achenes obpyramidal to prismatic, 8-10 ribbed, glabrous to 
densely pubescent, the carpopodium a sclerose annulus; pappus 1-4 seriate, the 
inner series of 30 or more, white or tawny, persistent, bristles, the outer series 
of much shorter, readily deciduous, squamellae, these often differentiated as 
whitened scales. Base chromosome numbers, z = 15, 16, 17 or 18. 


KEY TO SPECIES 


1. Involucres small (mostly 0.4-1.5 cm high; 0.5-1.0 cm wide), borne numerous 
in divaricately branched corymbose panicles or in drooping subfasciculate 
CYMER gee nee sok ane oe ev is Ree. oe ee Subgenus I. Sznclairia 


1. Involucres large (mostly 1-3 cm high, 1-4 cm wide), borne 1-7 in stiffly 
erect terminal cymes, the peduncles mostly (1-)2-8 cm 
Hon ee Sistas oss! amend Saldana eee Subgenus II. Megaliabum 


I. SUBGENUS SINCLAIRIA 


1. Heads arranged in drooping subfasciculate cymes, the ultimate peduncles 
mostly 2-5 cm long. ..... .jc + «csi. 2 aac don eyo 15. S. klatte 


1. Heads arranged in divaricately branched corymbose panicles ......... (2) 


2. Leaves glabrous or green on both surfaces, not at all 


bicolored ’y). 2 oes de ene cence ces sone oer (3) 
2. Leaves bicolored, the lower surfaces dirty white or white beneath 
with a matted or arachnoid tomentum ....................45. (4) 


3. Involucres mostly 4-6 mm high; eastern Chiapas and 
Guatemalar si: ih. owt. 's ies >. ede: 108 ee 4. S. hypochlora 


3. Involucres mostly 7-10 mm high; central and northwestern 
IPOS) 5 ssw apes dhaySTasoin Soke: Spe oy a aratwin hs apensloy oxens Seapaae ea 3. S. glabra 


4. Leaf blades, at least some of them, with distinct basal lobes or 
flanges; involucral bracts with rather rigid acute tips; Sinaloa to 
Oaxaca along the Pacific'slopes .....:..........-. 22 oon (5) 


Turner: Revision of Stnclairia including Liabellum 173 


4. Leaf blades not noticeably lobed, if sublobate then the involucral 
bracts broadly acute; obtuse or-rounded’'. .25202..0.00 0.5 a. .20s (6) 


5. Involucres 8-9 mm high; florets 15-20 per head; peduncles with glandular 
GMCS) fcr.) yt ets Lebo aout. eae, 2 ee 14. S. samilis 


5. Involucres 5-8 mm high; florets 9-12 per head; peduncles without glandular 
PGE he Bache eee Boas Sreetritacs contenu ad diacetate. oR: 13. S. hebmanni 


6. Achenes evenly and moderately to densely pubescent 
PO MINOE pein tat Mae ceria ia a a aide. Me ois chagaa nine eis cae See IES (7) 


6. Achenes glabrous or nearly so, or else the upper portion moder- 
ately pubescent but the lower portion glabrous or nearly so, rarely 


somewhat sparsely atomiferous glandular throughout ........ (Viz) 
favomucresl2-)o mm: high: (6 c..0 foes PN Pe eee eae ee ce (8) 
Meplnvohseres:A-O(- 12) mim hight wz! nas. wt. hkes en ea (9) 


8. Involucral bracts appressed, the apices acute; 
Girabemipla secretes aac o sh alicia 12. S. tajamulcensis 


8. Involucral bracts loosely appressed apically, the apices obtuse or 
rounded: Veracruz) Mexico 456028. 2248 st SS 11. S. deppeana 


9. Leaves densely brown villous beneath, not white tomentose; Veracruz, 
UNS RAC se tetas Seek SOG ie ees aA ti ns Sean rs ee crate 10. S. andromachioides 


9. Leaves densely white tomentose or white arachnoid beneath ........ (10) 


10. Branches of capitulescence with spreading, subglandular or glan- 
dular, trichomes; these often intermixed with a low tomentum; ray 
florets: absentywoseep eahnisenn alt peels sc. shee 9. S. deami 


10. Branches of capitulescence tomentulose, arachnoid or subarachnoid, 
if spreading trichomes present then ray florets present or else the 
achenes PlabrOUs OF NEATLY SO. <n co fin = een wie cee es eae (11) 


11. Inner involucral bracts glabrous or nearly so, the apices clearly rounded 
or broadly obtuse, without terminal apiculations; 
PACLESDLEAGH LE Nan Ae ccm aee Pie eek see 8. S. polyantha 


11. Inner involucral bracts clearly pubescent (glabrate with age), the apices 
mostly acute and often with minute terminal apiculations; southwestern 
Guatemala and-adjacent Chiapas)... 0c) anced dso <oase 7. S. vagans 


12. Involucres broadly campanulate, 8-12 mm wide at midsection, the 
bracts 30-40, closely and stiffly appressed; 
WEL MerU ZS Rr Ahan aes ae a kU ee cee ene 6. S. sericolepis 


174 PHY TOL OIGTA volume 67(2):168-206 September 1989 


12. Involucres turbocampanulate, 4-7 mm wide at midsection, the 
bracts 10-25, loosely or closely appressed ........--+-++-+-+-+- (13) 


13. Leaves, some of them, trullate or to some extent lobate or with basal 
flanges (rarely with merely large teeth); Guatemala to 
Nicaragua: «sis. 28.412 10. oae8 oct eee 5. S. sublobata 


13. Leaves ovate to elliptic, the margins unlobed and without basal flanges 
or teeth; Pacific slopes of México from Sinaloa to Oaxaca, or Gulf slopes 
of México and interior portions of Central America ............... (14) 


14. Ray florets absent; disk corollas orange or orange yellow; Pacific 
slopes of México, Sinaloa to Guerrero ................ 3. S. glabra 


14. Ray florets present (rarely not); disk and ray corollas 
yellow 2... eee cee nbn e oo neisle biomes + nce ate leet eae (15) 


15. Involucres 6-7 mm high, the inner bracts strictly appressed with acute 
or narrowly obtuse apices; disk florets 15-25; Pacific slopes of México, 
Jalisco to GuerrerO 020.20.) cee seen nec sone nan 2. S. broomeae 


15. Involucres 8-15 mm high, the inner bracts usually somewhat loose and 
scarious above, the apices broadly obtuse or rounded; disk florets 10-15; 
Gulf slopes of México and Chiapas, southwards to 
Panama 220 ee Pe ee on eer 1. S. discolor 


II. SUBGENUS MEGALIABUM 


1. Suffruticose branched herbs or soft wooded shrubs 1-4 m high, the stems 
arising (so far as known) from woody crowns, not forming tuber-like 
roots; leaves clearly petiolate, the petioles not winged, 


Sect. Megabiabrrn. 0-0. - «200.010 oie» 0:4 0s 0100 < bin ents ciate gee ee (2) 


1. Perennial, rarely suffruticose, herbs 0.3-1.0(-2.0) m high with simple stems, 
the root system with prominent tuber-like structures (not known in S. 
hintoniorum); leaves sessile or the petiole broadly winged to the base, 
Sect. Leabellurm crv .ass seeci. vbs ioe eee ee (4) 


2. Ray florets absent; petioles mostly 0.2-0.8 cm long; Jalisco, 
Méxice)\:).2/s8o0.2. PP Re 18. S. pringler 
2. Ray florets present; petioles mostly 2-5 cm long; Guerrero, México 


to Guatemala .......0esceecssee ne ones 6 oon oe (3) 


3. Ray florets 11-13; disk florets 25-35; outer involucral bracts mostly 2.5-3.0 
mm wide; Guerrero, México ..... WOE acted ose eee 17. S. mooretz 


Turner: Revision of Sinclairia including Liabellum 175 


3. Ray florets 21-34; disk florets 50+; outer involucral bracts mostly 1-2 mm 
wide; Oaxaca and Chiapas, Mexico and adjacent 
SPADE CO arene Ae Rene, Pe Oe PEST 16. S. andrieuri 


4. Leaves divided to the base, appearing whorled, the divisions narrowly 
linear: toonnGy Wide eh tes Sots dos Oak cates 23. S. angustissima 


4. Leaves entire or variously divided, but not as above, the divisions 
MIDst sd AU TAM WAGE wie. date n os wrasse Re nls cape aes aeRO (5) 


5. Leaves ovate elliptic, unlobed, confined to the very 
CERI nae cas NS colette ws ciate ts aveins «kr = larn eh ee WHOS REE 21. S. gentry 


5. Leaves obovate or variously parted, clearly cauline .................. (6) 


6. Suffruticose herbs 1-2 m high; achenes 5-6 mm long; pappus rusty 
FEU MIECIICO: SHALE. saciinn aici Gripe 4.5 srmodyieree aoe 19. S. hintontorum 


6. Non suffruticose herbs 0.3-9.0 m high; achenes 3-4 mm long; pappus 
tawny white or yellowish white; Chihuahua to Michoacan ..... (7) 


7. Leaves scattered along the entire length of the stem; peduncles without 
glandular hairs; outer pappus undifferentiated or nearly 
Ne = ee ae eS cid Se eyetecs tas as dads eae oon age eas aro 22. S. cervina 


7. Leaves mostly confined to the lower third of the stem; peduncles with 
glandular hairs; outer pappus of white scales ............ 20. S. palmeri 


I. SUBGENUS Sinclairia (species 1-15) 


1. Sinclairia discolor Hook. & Arn., Bot. Beech. Voy. 433. 1841. Lzabum 
discolor (Hook. & Arn.) Benth. & Hook. ex Hemsl., Biol. Centr. Amer. 
Bot. 2:232. 1881. TYPE: NICARAGUA. Realejo, Sinclair s.n. (HOLO- 
TYPE: K; Phototype: GH!). 


Liabum platylepis Sch.-Bip. ex Klatt, Leopoldina 23:146. 1887. Snclavria 
platylepts (Sch.-Bip.) Rydb., N. Amer. Fl. 34:296. 1927. TYPE: 
MEXICO. Veracruz: Mirador, w/o date, C. Sartorius s.n. (“in Linden! 
n. 1236”) (LECTOTYPE: GH!; fragment of lectotype: GH!). 


Clambering vines or hemiepiphytes. Leaves thin, strongly discolored, the 
undersurfaces densely white tomentose. Heads numerous in terminal, con- 
gested, mostly rounded, corymbose panicles. Involucres mostly 8-15 mm high, 
the inner bracts mostly scarious and obtuse or rounded at the apices, usually 
loose and recurved with age. Ray florets 5-8, the ligules 3-8 mm long. Disk 
florets 10-15, the corollas ca 8 mm long, the tube ca 4 mm long, the lobes 
glabrous or nearly so. Achenes 1.5-2.0 mm long, glabrous or a few hispid hairs 


176 PHYTOLOGIA volume 67(2):168-206 September 1989 


near the apex, the inner pappus tawny or tawny white, 6-9 mm long, the outer 
pappus of well defined scales ca 1 mm long; chromosome number, n = 17 or 
18 pairs. 

DISTRIBUTION (Figure 2): Southern México (Gulf slopes of Veracruz 
and Chiapas) to Panama, montane rain forests, 1200-2400 m; Dec-Mar. 

REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: Mcpio. Siltepec, 
ridge above Siltepec, 2000-2400 m, 1 Feb 1982, Breedlove 58311 (TEX); San 
Luis, 1300 m, 24 Jan 1945, Matuda 5294 (LL); Mcpio. Tuxtla Gutierrez, 19 
km from Tuxtla Gutierrez on road to canyon El Sumidero, 1210 m, 11 Mar 
1978, Garcia 578 (F,TEX). Veracruz: Mcpio. Los Reyes, ca 1600 m, 22 Mar 
1976, Torres W320 (F,XAL). 

GUATEMALA. Alta Verapaz: Taitic, 1600 m, Tuerckhetm 2116 (GH,LL). 
Cortes: between Banaderos and Cusnco, 1400 m, 26 Mar 1963, Molina 11456 


(LL). GUATEMALA: Guatemala, 1465 m, 3 Feb 1907, Kellerman 6277(F,GH). 


Izabal: ca Lago Izabal, 500-600 m, 11 May 1966, Jones 3366 (F). Quezalte- 
nango: above Mujulia, ca 1800 m, 1 Feb 1941, Standley 85651 (F). San Marcos: 
ca Aldea Fraternidad, 1800-2400 m, 10 Dec 1963, Willams 26240 (F). 

EL SALVADOR. San Salvador: Volcan de San Salvador, 1410-1800 m, 2 
Feb 1946, Carlson 442 (F). Santa Ana: Cerro Monte Cristo, 6000-6500 m, 17 
Jan 1959, Allen 7147 (F,GH,LL). Sonsonate: Cerro El Pilon, 1800 m, 23 Feb 
1968, Molina 21610 (F). 

BELIZE. Toledo: Soloman Camp, 80-420 m, 5-12 Mar 1987, Davidse 32036 
(LL,MO); high ridge, Edwards Road, beyond Columbia, 20 Mar 1948, Gentle 
6479 (F,LL). 

HONDURAS. Cortes: Montana San Idalfonso, 1500 m, 27 Mar 1963, 
Molina 11591 (F). Morazan: mountains above San Juancito, 1800 m, 20 Feb 
1948, Willams 1378 (GH); 24 Mar 1951, Williams 17457 (GH). Ocotepeque: 
ca El Portillo, 2000 m, 23 Jan 1976, Molina 31386 (F). 

NICARAGUA. Granada: Volcan Mombacho, 9 Mar 1922, Greenman 5792 
(GH). Matagalpa: El Ocotal, 1300 m, 9 Mar 1967, Molina 20431 (F). 

COSTA RICA. San José: Cerro Zurqui, 1800-2000 m, 19 Feb 1978, Almeda 
$685 (F); La Palma, 5 Feb 1922, Greenman 5499 (GH). 

PANAMA. Goofy Lake to ca 8 mi S of Goofy, 26 Nov 1966, Dwyer 7052 
(GH). 

A widespread species superficially similar to S. polyantha but the involu- 
cral bracts loose at maturity and the achenes glabrous or merely sparsely 
pubescent, especially just below the pappus. 


2. Sinclairia broomeae H. Robins., Phytologia 33:287. 1976. TYPE: MEXICO. 


Guerrero: “along the Milpillas-Atoyac road via Puerto del Gallo, about 
47 miles southwest of México Highway 95, 16 miles southwest of Filo de 
Caballo and 10.4 miles southwest of Carrazal del Bravo, 12.4 miles north- 
east of Yerba Santa,” 17 Oct 1975, J.L. Reveal et al. 4267(HOLOTYPE: 


Turner: 


Revision of Sinclairza including Liabellum 


e Sinclairia discolor 


larg 


Fig.’ 2. Distribution of Sinclairia discolor. 


178 PHY F0u,0 GilA volume 67(2):168-206 September 1989 


US; Isotype: TEX!; Photoisotype: GH!). 


Clambering shrubs and probably hemiepiphytic, occurring up to 40 m high 
in trees; leaves 12-16 cm long, 6-12 cm wide, thin, bicolored; petioles mostly 
3-6 cm long; blades broadly ovate to subdeltoid, white tomentose beneath, the 
nerves brown, the margins minutely serrulate. Heads numerous in corymbose 
panicles, the ultimate peduncles 5-15 mm long. Involucres 6-7 mm high, tur- 
bocampanulate, the bracts appressed with mostly acute apices. Ray florets 5, 
the ligules 5-8 mm long, yellow. Disk florets 15-25, the corollas yellow, 8-10 
mm long, the lobes ca 2 mm long. Achenes sparsely to densely pubescent 
above, glabrous below, 1.5-2.0 mm long, the inner pappus of tawny bristles ca 
8 mm long, the outer pappus scales 1.2 mm long. 

DISTRIBUTION (Figure 4): Jalisco (Sierra de Manantlan) and Guerrero, 
pine-oak cloud forests, 1800-2100; Oct-Dec. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Jalisco: Sierra Man- 
antlan, Playa Las Joyas, 1900-1950 m, 21 Dec 1984, Judziewicz & Cochrane 
4798 (TEX,WIS) Sierra de Manantlan, 3 km S of Rincon de Manantlan, going 
to Las Joyas (crossing no. 4), 1850-2100 m, 11 Oct 1982, Iltzs et al. 28866 
(TEX,WIS); Sierra Manantlan (19° 37’, 104° 15’), 1980 m, 12 Nov 1985, 
Vasquez 3764 (WIS); Las Joyas, Autlan, 1980 m, 12 Nov 1985, Vasquez 3726 
(WIS). Guerrero: Distr. Mina, Cerro Azul, 2550 m, 4 Dec 1939, Hinton et al. 
14951 (GH). 

The species is closely related to S. discolor but is readily distinguished by its 
smaller involucres with appressed acute bracts. Recent collections have been 
obtained from the Sierra de Manantlan, as indicated in the above citations. 


3. Sinclairia glabra (Hemsl.) Rydb., N. Amer. Fl. 34:297. 1927. Li- 
abum glabrum Hemsl., Biol. Centr. Amer., Bot. 2:232. 1881. TYPE: 
MEXICO. Morelos: Cuernavaca, Iturbide, 1865, Bourgeau 1401 (HOLO- 
TYPE: K; Isotypes: P,F!,GH!; Photoisotypes: F!,LL!). 


Shrubs or small trees to 6 m high. Leaves 10-22 cm long, 3-8 cm wide; 
petioles 3-6 cm long; blades mostly ovate, with acute bases, the undersurfaces 
glabrous or nearly so. The margins serrulate. Heads discoid, numerous in 
rather congested terminal corymbose panicles, 2-3 times as long as wide, the 
capitulescence usually without leafy inclusions. Involucres campanulate, 7-9 
mm high, the bracts appressed, glabrous with ciliate margins, the inner series 
obtuse or rounded at the apices. Ray florets absent. Disk florets 14-20 per 
head, glabrous, 8-10 mm long. Achenes 1.5-2.0 mm long, glabrous or sparsely 
pubescent above, rarely a few glandular hairs, the pappus of numerous tawny 
bristles 6-8 mm long, the outer scales ca 0.5 mm long. 

I recognize 3 allopatric regional varieties under this taxon, as keyed be- 
low. The above description applies to the var. glabra; all of the varieties 


179 


Revision of Sinclairia including Liabellum 


Turner: 


snyeqojqns “Ss peyeje: euj) s 


BUBSUDSP “SG pue Soplooewoipue S ‘(2 Aq peyeoipu! 
BIJEUBA S}! pue BIQETD ELIEPOUIS yO UO!NQuIsIp |e19UeD 


‘e Bid 


180 PHYEOLOGHEA volume 67(2):168-206 September 1989 


intergrade in regions of contact. Williams (1976) expanded S. glabra so as to 
include S. sublobatus, a species of Central America and southernmost México 
which has usually bicolored lobed leaves, strictly yellow corollas and heads ar- 
ranged in both axillary and terminal corymbs, the entire capitulescence being 
leafy within at anthesis. Sinclairia glabra has orange or orange yellow, mostly 
longer, corollas, unlobed leaves and a strictly terminal capitulescence (without 
leafy inclusions). 


Key to varieties (see Figure 3 for distributions). 


1. Leaves not bicolored, glabrous beneath; corollas mostly 8-10 mm long; E 
Michoacan, México, Morelos and NE Guerrero ............. var. glabra 


1. Leaves bicolored, variously white tomentose or arachnoid pubescent be- 
neath; corollas 6-15 mm long; Sinaloa to central Michoacan and NW 
Guerrero (nig bie Fan nee garde ode tees acters & ce che ae (2) 


2. Involucres mostly 7-8 mm high; corollas 9-14 mm long; pappus 
bristles yellow-white or pale tawny; Sinaloa to Colima and northern 
Michoacan . 2.0 24.s jas. n eines cs hee 2 <o.0 aoe var. hypoleuca 


2. Involucres mostly 5-6 mm high; corollas 6-8 mm long; pappus bristles 
rusty tawny; S Michoacan and NW Guerrero .......... var. minor 


S. glabra var. glabra. For nomenclature and synonymy, see above. 


DISTRIBUTION: E Michoacan, México, Morelos and NE Guerrero, trop- 
ical deciduous and oak forests, 300-1700 m; Oct-Dec. As described above; in- 
tergrades between var. glabra and var. hypoleuca occur, as noted by McVaugh 
(1984). 

REPRESENTATIVE SPECIMENS: MEXICO. México: Distr. Temascal- 
tepec, Vigas, 17 Nov 1932, Hinton 2627(GH). Morelos: barrancas near Cuer- 
navaca, 5000 ft, 7 Nov 1895, Pringle 6182 (F,GH). Michoacan: Zitacuaro, 24 
Oct 1938, Hinton 18401 (GH). Guerrero: Distr. Aldama, Temisco, 310 m, 12 
Nov 1937, Mexia 8797 (F,GH,LL). 


S. glabra var. hypoleuca (Greenm.) B. Turner, comb. nov. Based on: L1- 
abum glabrum var. hypoleucum Greenm., Proc. Amer. Acad. Arts 32:294. 
1987. Sinclairia hypoleuca (Greenm.) Rydb., N. Amer. Fl. 34:297. 1927. 
TYPE: MEXICO. Jalisco: canyons near Guadalajara, 8 Dec 1988, C.G. 
Pringle 2169 (LECTOTYPE: GH!, as selected by McVaugh in Flora 
Novo-Galiciana 12:578. 1984). 


181 


Revision of Sinclairia including Liabellum 


Turner: 


eT 
BIO|YOOAAY 


RESON 


PUB SITUS S ‘STASOSTSS F ‘CIOTOOUAY“S ‘SESWOSIYS Jo uo 


aeauloolg 
a 


SSUSO MUMS 


! | ! 
ANquisip jereued 


*y “Bid 


182 PB ¥ G0, 0) GLA volume 67(2):168-206 September 1989 


DISTRIBUTION: S Sinaloa, Nayarit, Jalisco, Colima and N Michoacan, 
tropical deciduous or pine-oak forests, 1000-2000 m, Oct-Dec. 

My circumscription of this taxon is essentially the same as that of McVaugh 
(1984), who also recognized its probable intergradation with var. glabra. 

REPRESENTATIVE SPECIMENS: MEXICO. Sinaloa: 9 mi E of Chupo- 
deras, 2600 ft, 22 Nov 1969, Breckon 990(LL). Nayarit: 9 mi N of Compostela, 
1000-1200 m, 13 Nov 1959, McVaugh & Koelz 557 (LL). Jalisco: 15 mi N of 
Talpa, 20 Oct 1983, Ayers 277(TEX). Colima: 18 road mi NE of Colima, 3600 
ft, 30 Oct 1962, Cronquist (TEX). Michoacan: 5 mi N of Cotiga, 6000-6200 ft, 
5-9 Oct 1961, King & Soderstrom 4685 (TEX). 


S. glabra var. minor B. Turner, var. nov. TYPE: MEXICO. Michoacan: 
Distr. Coalcoman, Aguila, 17 Nov 1941, G.B. Hinton et al. 16161(HOLO- 
TYPE: LE!). 


S. hypoleucae var. hypoleucae sed involucris minoribus (5-6 mm 
altis vs 7-8 mm) et corollis brevioribus (6-8 mm longis vs 9-14 mm) 
differt. 


DISTRIBUTION: S Michoacan and adjacent E Guerrero, Pacific slopes in 
tropical deciduous forests, 600-1400 m; Oct-Dec. 

The taxon differs from the var. glabra in possessing smaller involucres with 
mostly narrowly obtuse or acute involucral bracts, heads with fewer florets, 
leaves tomentose beneath and pappus bristles rusty tawny. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Michoacan: Distr. 
Coalcoman, Puerto Las Cruces, 1340 m, 28 Oct 1938, Hinton 12467 (GH,LL); 
S. Naranjillo, 1360 m, 26 Nov 1938, Hinton 12678 (GH,LL). Guerrero: Distr. 
Montes de Ocas., San Antonio, 16 Dec 1937, Hinton et al. 11682 (GH,LL). 


4. Sinclairia hypochlora (S.F. Blake) Rydb., N. Amer. Fl. 34:301. 1927. 
Liabum hypochlorum §.F. Blake, Contr. Gray Herb., n.s. 53:27. 1918. 
FYPE: GUATEMALA. Retalhuleu: San Felipe, wet places, 13 Jan 1917, 
E.W.D. Holway 708 (HOLOTYPE: GH!). 


Clambering shrubs or vines to 15 m high, probably also hemiepiphytic 
upon occasion. Leaves not bicolored, 10-25 cm long, 6-10 cm wide; petioles 
2-5 cm long; blades ovate to elliptic ovate, glabrous on both surfaces, or nearly 
so, minutely glandular punctate, the margins minutely denticulate. Heads ra- 
diate, numerous in broad terminal and subterminal corymbose panicles, the 
ultimate peduncles mostly 2-5 mm long. Involucres 4-5 mm high, the bracts 
pubescent 2-3 seriate, rather stifly appressed and acute at the apices. Recep- 
tacles with markedly pubescent alveolae. Ray florets 5-8, the ligules 1-3 mm 
long, 0.5-1.0 mm wide, yellow. Disk florets 5-7, the corollas yellow, 7-8 mm 
long, glabrous. Achenes ca 2.5 mm long, pubescent throughout, the pappus 


Turner: Revision of Sinclairia including Liabellum 183 


of numerous yellowish bristles 6-7 mm long, the outer scales ca 0.5 mm long; 
chromosome number, n = 15 or 16 pairs. 

DISTRIBUTION (Figure 4): Guatemala and adjacent Chiapas, montane 
cloud forests, 500-900 m; Nov-Jan. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Chiapas: Mt. Ovan- 
do, ca Escuintla, 14 Nov 1945, Matuda 6194 (LL); Jalapa, Triunfo, 6 Nov 1948, 
Matuda 18461 (F). 

GUATEMALA. Quezaltenango: Colomba, 3000 ft, 29 Dec 1934, Skutch 
9031 (AA,F,GH,LL). Retalhuleu: San Felipe, 12 Jan 1917, Holway s.n. (GH). 
Schitepequez: Volcan Zunil, 2 Feb 1940, Steyermark 35429 (F). 

This species would appear to be but a populational form of S. vagans, or 
vice versa, but differs, in addition to its glabrous foliage, by having smaller 
rays and very pubescent alveolate ridges on the receptacle and corollas lobes 
nearly glabrous. In addition, 5. vagans appears to occur mostly at higher 
elevations (2000-2500 m vs 500-1000 m). 

In addition to the type itself, Holway (cited above without number) appar- 
ently collected a rayless form of this plant with weakly arachnoid tomentose 
leaves and peculiar glandular achenes. This is perhaps a hybrid or hybrid 
derivative with yet another species, possibly S. sublobatus, which has rayless 
heads and occurs with or near S. hypochlora. 


5. Sinclairia sublobata (B.L. Robins.) Rydb., N. Amer. Fl. 34:297. 1927. 
Liabum sublobatum B.L. Robins., Proc. Amer. Acad. Arts 32:294. 1897. 
TYPE: GUATEMALA. Solola: San Lucas Toliman, 5100 ft, 2 Feb 1915, 
E.W.D. Holway 179 (HOLOTYPE: GH!). 


Sinclairia brachypus Rydb., N. Amer. Fl. 34:299. 1927. TYPE: GUATEMA- 
LA. Chiquimula: Volcan Ipala, 1700 m, Jan 1907, H. Pitter 1886 (HOLO- 
TYPE: US; Isotype: F'). 


Slender erect shrubs or small trees to 12 m high. Leaves 10-20 cm long, 5- 
18 cm wide, probably deciduous shortly after flowering; petioles mostly 3-6 cm 
long; blades trullate to broadly ovate or somewhat subcordate, usually thick- 
ened with age, bicolored (a dense tomentum beneath but rarely only weakly 
arachnoid and not totally bicolored), 3 nervate, the basal portion to some 
extent with acute lobes or flanges (sometimes merely large teeth). Heads dis- 
coid, numerous in both terminal and axillary somewhat divaricate corymbs, 
the entire capitulescence therefore a leafy arrangement, the ultimate peduncles 
mostly 1-8 mm long. Involucres mostly 7-9 mm high, the bracts mostly ap- 
pressed through maturity, the inner bracts with apices obtuse or rounded. Ray 
florets absent. Disk florets 8-10, the corollas yellow, glabrous, 7-9 mm long, the 
tube ca 3 mm long, grading into the throat, the lobes ca 2 mm long. Achenes 
3-4 mm long, sparsely pubescent above, mostly glabrous below, the pappus 
of numerous, yellow white or tawny, bristles 6-7 mm long, the outer series of 


184 PHYTOLOGIA volume 67(2):168-206 September 1989 


short scales ca 1 mm long, these well differentiated; chromosome number, n = 
15 or 16 pairs. 

DISTRIBUTION (Figure 5): Southernmost México, Guatemala, Honduras, 
E] Salvador and Nicaragua, mostly Pacific slopes along barrancas on dry slopes 
and ridges, 200-1800 m; Dec-Mar. 

REPRESENTATIVE SPECIMENS: MEXICO. Oaxaca: Mcpio. Sta. Maria 
Chimalapa, 4 km NE of Santa Maria, 170 m, 7 Mar 1984, Wendt 4300(CHAPA, 
TEX); 4 km N of Sta. Santa Maria, 300 m, 26 Feb 1925, Herndndez 916 
(CHAPA, TEX). Chiapas: Mcpio. Tenejapa, pokolum, 520 m, 18 Mar 1985, 
Breedlove 9485 (LL); Mcpio. Vajalon, Rancho Carmen, 25 Mar 1984, Ton 7471 
(MEXU, TEX). 

GUATEMALA. Alta Verapaz: between San Cristobal and Chixoy, 1200- 
1300 m, 19 Feb 1942, Steyermark 43896(F,GH). Chimaltenango: ca Sibaja, 
ca 1050 m, 6 Jan 1939, Standley 62256 (F,GH). Guatemala: ca Guatemala, 
1465 m, 1 Jan 1907, Kellerman 6298 (F). Retalhuleu: ca Retalhuleu, 240 m, 
Standley 88799 (F,GH). Sacatepequez: ca Santa Maria, 5500 ft, 28 Jan 1937, 
Hunnewell 14909 (GH). SANTA ROSA: Carrizal, 5000 ft, Jan 1893, Heyde & 
Luz 4225 (GH). 

HONDURAS. Morazan: Choluteca: ca San Marcos, 12 Jan 1949, Standley 
15717 (F). Cuesta La Moroloa, 1300 m, 15 Feb 1951, Molina 3896 (F,GH). 
Sonsonate: Cerro Verde, 4 Feb 1959, Allen 7223 (F). 

EL SALVADOR. Ahuachapin: ca Apaneca, 24 Jan 1947, Standley 3002(F). 
Chalatenango: between San Ignacio and Citala, 1500-3000 ft, 19 Nov 1958, 
Allen 7108 (F,GH). San Salvador: ca Tonacatepeque, 30 Dec 1921, Standley 
19530 (GH). Sonsonate: ca Sonsonate, 220-300 m, 18 Mar 1922, Standley 
22265 (GH). 

NICARAGUA. Esteli: El Zacaton, 1400 m, 11 Feb 1984, Moreno 22687 
(F). Managua: Sierra de Managua, Garnier 21 (F). Matagalpa: 5-10 km W of 
Matagalpa, 600-700 m, 3 Jan 1963, Wilhams 23820 (F,LL). Ocotepeque: 15.1 
m E of Santa Fe, 1460 m, 28 Jan 1987, Croat 63834 (TEX). 

Because of its lobed leaves, capitulescence and floral details, this species 
appears related to S. liebmannii of México. The latter, however, has smaller 
leaves, smaller involucres with acute involucral bracts and creamy white corol- 
las (vs bright yellow). Occasional plants of S. sublobata possess weakly bi- 
colored leaves (the tomentum beneath being mostly erased with age); these 
have been often identified as S. glabra, a species with unlobed leaves which is 
confined to central and northwestern México. The latter taxon has orange or 
orange yellow corollas (vs yellow) and a mostly leafless capitulescence. 


6. Stnclatria sericolepis (Hemsl.) Rydb., N. Amer. Fl. 34:294. 1927. Li- 
abum sericolepis Hemsl., Biol. Centr. Amer., Bot. 2:232. 1881. TYPE: 
MEXICO. Veracruz: Valley of Cordoba, 10 Mar 1865, Bourgeau 2177 
(HOLOTYPE: K; Isotype: P; fragment of isotype: F!; Photoholotype: 


Turner: Revision of Sinclazrza including Liabellum 185 


x 

= 

jet 

e — 

e ro) 

e c 

no) 

5 

= 

= 

5 2 

° 5 fo) fo) =) 
Ro 2 
w 

io) . 

2 

ic 


186 PHY? OO GPA volume 67(2):168-206 September 1989 


GH!; Photoisotypes: F!,GH!). 


Clambering shrub or small tree to 3 m high. Leaves bicolored, 8-25 cm 
long, 3-9 cm wide; petioles 1-5 cm long; blades ovate to elliptic ovate, white 
tomentose beneath, drying blackish above, the margins minutely serratulate to 
nearly entire. Heads eradiate, numerous in stiffly branching, terminal, corym- 
bose, panicles, the ultimate peduncles mostly 3-12 mm long. Involucres cam- 
panulate, 8-10 mm high, 8-12 mm wide, the bracts 4-5 seriate, gradate, closely 
appressed, the apices decidedly acute. Ray florets absent. Disk florets 18-22 
per head, the corollas ca 8 mm long, glabrous, the tube ca 4 mm long, grading 
into the throat, the lobes linear, ca 1.5 mm long. Achenes ca 2 mm long, 
glabrous below, hirsute-pubescent below the apex, often densely so, the pap- 
pus of numerous dirty white or tawny bristles 8-9 mm long, the outer series of 
short linear scales but not differentiated. 

DISTRIBUTION (Figure 4): Veracruz, Oaxaca and Chiapas, tropical rain 
forests, 100-700 m; Mar-Apr. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Chiapas: Mcpio. 
Ocozocoantla de Espinosa, SW site of Presa de Malposa, 2200 ft, 2 Mar 1968, 
Ton 3803 (F,LL). Oaxaca: ca Valle Nacional, 22 Mar 1964, Quintero 648 
(TEX). Veracruz: Mcpio. Hidalgotitlan, 4 km from Hermanos Cedillo, 150 m, 
10 Apr, 1975, Ortiz 106 (F); 10.3 km E of La Laguna, 140 m, 24 Mar 1981, 
Wendt 3048 (CHAPA,TEX). 

A very distinct species, not easily confused with another, although the 
leaves and habit are similar to S. discolor. 


7. Sinclairia vagans (S.F. Blake) H. Robins. & Bret., Phytologia 28:62. 1974. 


Liabum vagans S.F. Blake, Brittonia 2:354. 1937. TYPE: GUATEMALA. 


Quiche: “Thicket on mountainside along Nebaj-Aguacatan trail,” 2470 
m, 12 Dec 1934, A.F. Skutch 1918(HOLOTYPE: AA!; Isotypes: F!,LL!). 


The holotype is without collection data, but the isotype, collection of which 
Blake also annotated, gives the site as published. Clambering shrubs or small 
trees to 5 m high, probably also hemiepiphytic. Leaves 12-22 cm long, bi- 
colored, 8-15 cm wide; petioles 3-6 cm long; blades broadly ovate, 3 ner- 
vate, densely white tomentose beneath, drying blackish above, the margins 
minutely serrulate. Heads radiate, numerous in terminal corymbose rounded 
panicles, the ultimate peduncles mostly 3-6 mm long. Involucres mostly 5- 
7 mm long, the bracts appressed through maturity, those of the inner series 
mostly pubescent, acute or narrowly obtuse, often with a small apical apicu- 
lation. Ray florets 5-8, the ligules yellow, 6-9 mm long. Disk florets 4-6, the 
corollas yellow, glabrous except for puberulent tufts at the apex of each lobe, 


7-8 mm long, the tube ca 3 mm long, the lobes 1.5-2.0 mm long. Achenes ca | 


2 mm long, pubescent throughout, the pappus of numerous yellowish white or 


| 
| 


Turner: Revision of Sinclairia including Liabellum 187 


tawny bristles 5-7 mm long, the outer series of small linear scales 1.0-1.5 mm 
long. 

DISTRIBUTION (Figure 6): Southeastern Chiapas and adjacent Guatema- 
la in montane rain forests, 2000-2700 m; Nov-Jan. 

ADDITIONAL SPECIMENS EXAMINED: GUATEMALA. Chimaltenan- 
go: above Las Calderas, 2250-2400 m, 16 Dec 1940, Standley 80193(F). Jalapa: 
ca Soledad, 2000-2400 m, 4 Dec 1939, Steyermark 32612 (F,GH). Guatemala: 
Volcan de Pacaya, 1800-2300 m, 20 Dec 1940, Standley 80633 (F). Quezalte- 
nango: Volcan Santa Maria, 8000-11,500 ft, 24 Jan 1896, Nelson 3722 (F,GH). 
San Marcos: 6 km N of San Marcos, 2700 m, 13 Dec 1963, Willams 25855 
MEXICO. Chiapas: Mcpio. Siltepec, ridge above Siltepec, 2000-2400 m, 
18 Jan 1973, Breedlove 31848 (TEX); 26 Nov 1981, Breedlove 55917 (TEX). 
Mcpio. Motozintla de Mendoza, SW side of Cerro Mozotal, 2100 m, 23 Nov 
1988, Breedlove 55800 (TEX). 

A weakly differentiated species, closely related to the widespread S. polyan- 
tha, but the latter occurring at mostly lower elevations and having inner in- 
volucral bracts mostly glabrous, scarious at the apex and becoming loose and 
somewhat recurved at maturity. Sznclairia hypochlora, with glabrous leaves, 
also appears very closely related and is perhaps but a populational form of S. 
vagans. 


8. Sinclairia polyantha (Klatt) Rydb., N. Amer. Fl. 34:299. 1927. Liabum 
polyanthum Klatt, Bull. Bot. Belg. 31:209. 1892. TYPE: COSTA RICA: 
forests near General, Jan 1891, Pittier 4319 (HOLOTYPE: C; Isotype: 
FY). 


Liabum tonduzi B.L. Robins., Proc. Boston Soc. Nat. Hist. 31:270. 1904. 
Sinclatria tonduzii (B.L. Robins.) Rydb., N. Amer. Fl. 34:299. 1927. 
TYPE: COSTA RICA. San José: banks of Rio Virillo, San José, Jan 
1896, Tonduz 9859 (LECTOTYPE, selected here: GH!; Isolectotype: 
F!). 


Sinclatria pittiert Rydb., N. Amer. Fl. 34:300. 1927. TYPE: COSTA RICA. 
Alajuela: ca Alajuelita, Dec 1894, Pittter 9098 (HOLOTYPE US; Iso- 
types F!,GH!) 


Liabum dimidium Blake, J. Washington Acad. Sci. 22:385. 1932. Sinclazria 
dimidia (Blake) H. Robins. & Bret., Phytologia 28:61. 1974. TYPE: 
GUATEMALA. Peten: Tikal, 12-15 Apr 1931, Bartlett 12602 (HOLO- 
TYPE US; isotypes, F!,GH!,LL!,TEX!). 


Clambering shrubs or woody vines to 30 m high. Leaves ovate to broadly 
elliptic, bicolored, 10-20 cm long, 3-14 cm wide; petioles 2-5 cm long; blades 


188 


9 ‘Bi 


‘SUBDENTS pur TUUEWIGST S ‘TWESPS jo uonduosep jeseuen 


PHY T.0:,.0. GLA 


volume 67(2):168-206 


September 1989 


Turner: Revision of Sinclairia including Liabellum 189 


densely white tomentose beneath, the margins serrulate to nearly entire. Heads 
radiate, numerous in terminal corymbose panicles, the ultimate peduncles 
mostly 0-10 mm long. Involucres 6-9(-12) mm high, the bracts usually ap- 
pressed through maturity, obtuse or rounded at the apices and mostly glabrous, 
rarely pubescent. Ray florets 5-8, the ligules yellow, 3-6 mm long. Disk florets 
10-15 per head, the corollas yellow, glabrous, 6-8 mm long, the tubes 1-2 mm 
long, the lobes ca 1 mm long. Achenes 1.5-2.5 mm long, densely pubescent 
throughout, the pappus of numerous bristles 5-7 mm long, the outer scales 
few, poorly developed or seemingly absent. 

DISTRIBUTION (Figure 7): Veracruz, Oaxaca and adjacent Chiapas along 
the Gulf slopes, southwards to Panama and Colombia, tropical and montane 
rain forests, 100-1600 m; Jan-Apr. 

REPRESENTATIVE SPECIMENS: MEXICO. Oaxaca: Belleville, 23 Feb 
1910, Orcutt 3064 (F); 49 mi N of Ixtlan de Juarez, 1450 m, 10 Jan 1982, Bacon 
1754 (F). Veracruz: Mcpio. Catemaco, Cerro de Buena Vista, ca Catemaco, 
ca 300 m, 12 Feb 1975, Calzada 1744 (LL,MEXU); Mcpio. Hidalgotitlan, 30 
km E of La Laguna, 100 m, 26 Feb 1981, Wendt 2933 (CHAPA,TEX); Mcpio. 
San Andres Tuxtla, Volcan San Martin Tuxtla, 1630 m, 15 Feb 1972, Beaman 
5725 (F); Mcpio. Zongolica, Nacaxtla, ca 1050 m, 25 Feb 1976, Torres W-137 
(TEX,XAL). 

GUATEMALA. Dept. Izabal: Rio Duke, 1-3 m, 16 Apr 1940, Steyermark 
39559 (F). Dept. Peten: San Diego, 13 Apr 1935, Aguilar 517 (LL); Lacandon, 
16 Mar 1962, Contreras 3534 (LL); Tikal, 24 Mar 1959, Lundell 15784 (LL); 
Aguada Las Cucas, 5 Mar 1961, Lundell 16901 (LL). 

BELIZE. Toledo District: ca river beyond Columbia, 3 Feb 1947, Gentle 
6154 (LL); Temesh River, 28 Feb, 1945, Gentle 5239 (F,LL); Cero Hills, 25 
Mar 1950, Gentle 7007 (FLL). 

NICARAGUA. Boaco: Cerro Membachito, ca 900 m, 8 Feb 1983, Graalva 
2263 (F). Jinotega: NE of Wiwili, 13 Mar 1980, Aragquistan 1814 (F). Man- 
agua: Sierra de Managua, 1932, Garnier A489 (LL). Matagalpa: Los Angeles, 
900 m, 1 Mar- 1983, Moreno 21049 (F). Rivis: Isla Ometepe, 21 Feb 1983, 
Moreno 19875 (F). Zelaya: Cerro Walawas, 100-268 m, 16 Mar 1978, Stevens 
7428 (F). 

COSTA RICA. Alajuela: ca La Laguna, 1200 m, 19 Feb 1966, Molhna 
17514 (GH). Guanacaste: El Silencio, Tilaran, 900 m, 12 Feb 1963, Juménez 
s.n. (F). Punta Arenas: Rio Esquinos, ca 100 ft, 8 Dec 1949, Allen 5437 
(AA,F). San José: Las Nubes, 6000 ft, 24 Jan 1940, Hunnewell 17038 (GH); 
ca El General, 1035 m, Feb 1939, Skutch 4187 (LL). 

PANAMA. Chiriqui: ca New Switzerland, 1800-2000 m, 6 Jan 1939, Allen 
1415 (GH). Cocle: ca El Valle de Anton, ca 1000 m, 2 Dec 1941, Allen 2866 
(GH). Panama: Cerro Campana, 17 Aug 1960, Ebinger 935 (F). 

COLOMBIA: Dept. Caldas: La Selva, 1600-1900 m, 8 Jan 1946, von Snez- 
dern 5246 (F). 


190 PHYTOLOGIA volume 67(2):168-206 September 1989 


‘BUJUBATOS ETTEPOUTS jo uonnquisiq “2 “B13 


Turner: Revision of Sinclatria including Liabellum 191 


A widespread variable species but not readily divisible into regional vari- 
ants. It is, nevertheless, easily recognized by a combination of characters, 
including densely pubescent achenes, rayed heads, involucral bracts which are 
mostly appressed at maturity and bicolored, nearly entire leaves, the upper 
surface of which darkens appreciably upon drying. Vegetatively it superfi- 
cially resembles S. discolor but the latter is readily recognized by its glabrous 
achenes (or nearly so). 


9. Sinclairia deamiu (B.L. Robins. & Bartl.) Rydb., N. Amer. Fl. 34:299. 
1927. Liabum deamii B.L. Robins. & Bartl., Proc. Amer. Acad. Arts 
43:60. 1907. TYPE: GUATEMALA. Tacapa: Gualan, 420 ft, Jan 1905, 
C.C. Deam 194 (HOLOTYPE: GH!; Isotype: F!). 


Liabum subglandulare S.F. Blake, Contr. U.S. Natl. Herb. 24:31. 1922. Sin- 
clairia subglandularis (S.F. Blake) Rydb., N. Amer. Fl. 34:298. 1927. 
TYPE: HONDURAS. Copan: “Hacienda La Zumbadora,” between El 
Paraiso and La Florida, 13 May 1919, S.F. Blake 7386 (HOLOTYPE: 
US). 


Clambering shrubs or slender trees to 15 m high, probably also hemiepi- 
phytic. Leaves bicolored, 10-20 cm long, 6-12 cm wide, deciduous at the time 
of, or soon after, flowering; petioles 3-4 cm long; blades broadly ovate, white to- 
mentose beneath, moderately blackening upon drying, the margins minutely 
serrulate. Heads eradiate in numerous terminal or subterminal, congested, 
corymbose panicles, the ultimate peduncles mostly 1-3 mm long, the branches 
brown tomentose or glandular hirsute, or both. Involucres mostly 4-5 mm 
long, the bracts 2-3 seriate, ovate, pubescent, the apices mostly obtuse. Ray 
florets absent. Disk florets mostly 5, the corollas pale yellow or creamy white, 
ca 7 cm long, glabrous except for the hispid tufted apices of the lobes, the 
lobes ca 2 mm long. Achenes 2-3 mm long, densely pubescent throughout, 
the pappus of numerous dirty white or tawny bristles 6-8 mm long, the outer 
scales narrow,-1.5-2.0 mm long. 

DISTRIBUTION (Figure 6): Lowland tropical forests of northern Guatema- 
la and adjacent México, Belize, Honduras and El Salvador 100-700 m; Feb-Mar. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Campeche: Tux- 
pena, 23 Mar 1932, Lundell 1437 (F). Chiapas: Mcpio. Copainala, 60 km 
NW of Soyala, 550 m, 13 Mar 1973, Breedlove 34048 (F,LL). Tabasco: Mc- 
pio. Tenosique, ca puentes Boca del Cerro, 270 m, 11 May 1984, Cowan 4692 
(TEX). 

BELIZE. El Cayo: El Cayo, 13 Apr 1931, Bartlett 12934 (F,GH,LL); hill- 
side, Camp 6, 23 Mar 1938, Gentle 2395 (GH,LL). 

GUATEMALA. Alta Verapaz: Pantin, ca 600 m, 5 Apr 1939, Standley 
70872 (F). Peten: ca La Libertad, 9 Mar 1935, Aguilar 484 (LL); Remate, 
14 Mar 1960, Contreras 662 (LL); Lake Peter Itza, 28 Mar 1961, Contreras 


192 PHY TODBOG LA volume 67(2):168-206 September 1989 


2068 (LL); Quexil, 7 Feb 1970, Contreras 9615 (LL); Tikal, Mar 1961, Ibarra 
1 (LL); Tikal, on ruins, 3 Mar 1961, Lundell 16843 (LL). 

EL SALVADOR. Santa Ana: ca Santa Ana, 655-800 m, 8 Jan 1922, Stand- 
ley 19695 (GH). La Libertad: W of Santa Teda, 750 m, 19 Jan 1946, Carlson 
174 (F). 

HONDURAS: Comayagua: Comayagua, ca 500 m, 20 Feb 1933, Edwards 
P568 (AA). Cortes: San Pedro Sula, 21 Feb 1987, Thieme 5332 (F). Santa 
Barbara: Quebrado de Limones, 150 m, 17 Apr 1947, Standley 7541 (F). 

A very distinct taxon, largely confined to lowland tropical rainforests, easily 
recognized by its small involucres (3-5 mm high), sessile heads, each with 
5 pale yellow florets, the lobes of which have tufts of hispid hairs. Blake 
described forms with mostly glandular hairs as L. subglandulare. Williams 
(1976) correctly submerged this under the present species, as did Robinson 
(1983). 


10. Sinclairia andromachioides (Less.) Sch.-Bip. ex Rydb., N. Amer. FI. 
34:298. 1927. Vernonia andromachioides Less., Linnaea 6:397. 1831. 
Liabum andromachioides (Less.) Hemsl., Biol. Centr. Amer., Bot. 2:231. 
1881. TYPE: MEXICO. Veracruz: forests ca Misantla, w/o date, Schiede 
1284 (HOLOTYPE: B; Photoholotypes: F!,GH!). 


Clambering shrubs or hemiepiphytes. Leaves mostly 10-25 cm long, 6- 
18 cm wide; petioles 1-4 cm long; blades broadly ovate to ovate elliptic, not 
markedly bicolored, if at all, the lower surface decidedly brownish or dingy 
puberulent or tomentose, 3 nervate from above the base, the margins entire. 
Heads discoid, numerous in terminal corymbose panicles. Involucres 5-7 mm 
high, 3 seriate, the bracts gradate, the inner series glabrous with obtuse or 
rounded apices. Ray florets absent. Disk florets 15-25, the corollas yellow, 5-6 
mm long, the tube ca 2.5 mm long, the lobes ca 1.5 mm long. Achenes ca 1.5 
mm long, pubescent throughout, or nearly so; the pappus of numerous tawny 
bristles, the inner series 5-6 mm long, the outer scales similar to the inner but 
ca 0.7 mm long. 

DISTRIBUTION (Figure 2): Known only from Veracruz, montane cloud 
forests in barrancas bordering streams along the eastern slopes of Pico de 
Orizaba and Cofre de Perote, ca 1200 m; Jan-May. 

ADDITIONAL SPECIMENS EXAMINED: Veracruz: Orizaba, w/o date, 
Botteri s.n. (F,GH); valle de Cordoba, Feb 1866, Bourgeau 1848 (GH); Fortin, 
Feb 1883, Kerber 277 (GH); Mirador, Feb 1843, Liebmann 399 (F,GH); ca 
Orizaba, 23 Jan 1895, Pringle 5905 (GH); Zacuapan, Jan 1908, Purpus 2942 
(GH); Zacuapan, May 1919, Purpus 8162(GH); Zacuapan, Mar 1932, Purpus 
14233 (AA,F); w/o locality, May 1935, Purpus 16452 (LL); Orizaba, Oct 1855, 
Schaffner 368 (GH). 

The species is closely related to the widespread S. polyantha but can be 
distinguished by its faintly, if at all, bicolored leaves and smaller involucres 


Turner: Revision of Sinclairia including Liabellum 193 


(5-7 mm high vs 7-9 mm). The two taxa apparently do not coexist. It seems 
noteworthy that S. andromachiozdes has not been collected for over 50 years, 
suggesting that many of the original populations have been destroyed, probably 
for the cultivation of coffee. 


11. Sinclairia deppeana (Less.) Rydb., N. Amer. Fl. 34:300. 1927. Andro- 
machia deppeana Less., Linnaea 6:401. 1831. Liabum deppeana (Less.) 
Hemsl., Biol. Centr. Amer., Bot. 2:232. 1881. TYPE: MEXICO. Ver- 
acruz. Cuesta Grande del Jalacingo, Dec 1819, Schiede 1239 (HOLO- 
TYPE: B; Photoholotypes: F!,GH!). 


Clambering shrubs, small trees, or hemiepiphytes to 5 m high. Leaves 
bicolored, 12-26 cm long, 6-14 cm wide; petioles 4-10 cm long; blades broadly 
ovate to broadly elliptic, densely white tomentose beneath, drying blackish 
above, the margins minutely serrulate to nearly entire. Heads radiate, few to 
humerous in terminal, rounded, corymbose panicles, the ultimate peduncles 
stiffly divaricate, mostly 1-2 cm long. Involucres mostly 12-15 mm high, 3- 
4 seriate, unequally gradate, the inner bracts mostly glabrous with broadly 
rounded apices, these becoming loose and somewhat squarrose with age. Ray 
florets mostly 8, the ligules yellow, 8-12 mm long. Disk florets 15-20, the 
corollas yellow, 8-10 mm long, glabrous except for puberulent tufts on the 
lobes, the tubes ca 3 mm long, the lobes ca 1.5 mm long. Achenes 1.5-2.0 mm 
long, pubescent throughout, the pappus of numerous yellowish white bristles 
9-11 mm long, the outer scales ca 0.5 mm long. 

DISTRIBUTION (Figure 2): Veracruz and Oaxaca along the Gulf slopes 
in pine-oak montane cloud forests, 1300-2500 m; Oct-Feb. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Oaxaca: 70-75 km 
S of Tuxtepec on road to Oaxaca City, 1370 m, 5 Jan 1982, Breedlove 56743 
(TEX); 35 mi E of Teotitlan, 1650 m, 22 Feb 1979, Croat 48249 (F). Ver- 
acruz: Mcpio. Orizaba, Orizaba, w/o date, Botteri 1197 (F); Mcpio. Tatatila, 
between Tatalita and Escalona, 14 Feb 1986, Chazaro 3972 (TEX,WIS,XAL); 
Mcpio. Xico, between Tonalco and Oxtlapa, 2500 m, 8 Oct 1986, Chazaro 
4028B (TEX,WIS,XAL); between Xico and Buena Vista, 2000 m, 25 Dec 
1987, Chazaro B. 1988 (WIS); Mcpio. Zongolica, Nacaxtla, ca 1050 m, Torres 
W141 (TEX,XAL); Mcpio. Acajete, Plan de Cedeno, 1740 m, 10 Dec 1971, 
Ventura 4677 (MEXU,TEX); Mcpio. Yecuatla, El Cajon, 1380 m, 14 Dec 1971, 
Ventura 4701 (MEXU,TEX). 

The species appears to be a very large headed form of S. discolor, but it 
is readily distinguished from the latter by a number of characters including 
much larger, broader involucral bracts and pubescent achenes. 


12. Sinclairia tajumulcensis (Standl. & Steyerm.) H. Robins. & Bret. Phy- 
tologia 28:62. 1974. Lzabum tajyumulcense Standl. & Steyerm., Publ. 


194 Pon Yolo E.G A volume 67(2):168-206 September 1989 


Field Mus. Nat. Hist., Chicago, Bot. Ser. 23:27. 1943. TYPE: GUATE- 
MALA. San Marcos: moist thicket bordering stream, barrancas SW of 
Tajulmulcom, 2300-2500 m, Feb 1940, J.A. Steyermark 36543 (HOLO- 
TYPE: F!). 


Clambering shrub or small tree 4-5 m high. Leaves unknown, presumably 
deciduous shortly after anthesis. Heads numerous in congested, somewhat 
pyramidal, corymbose panicles, the ultimate peduncles 3-i0 mm long. In- 
volucres campanulate, 4-5 seriate, 14-17 mm high, 18-22 mm wide, the bracts 
gradate, linear lanceolate, acuminate, densely tomentulose or lanuginose. Ray 
florets absent. Disk florets 20-40, the corollas yellow, 10-12 mm long. Achenes 
ca 4-5 mm long, densely pubescent throughout, the inner pappus of 30-40 bris- 
tles 10-12 mm long, the outer pappus of numerous narrow scales 1.0-1.5 mm 
long, similar in color to the inner bristles. A very distinctive species, known 
only by the type collection (Figure 4). 


13. Stnclatria hebmanni (Klatt) Sch.-Bip. ex Rydb., N. Amer. Fl. 34:300. 
1917. Liabum hebmannw Klatt, Leopoldina 23:146. 1887. TYPE: 
MEXICO. Oaxaca: “Bartolo,” 1842, F.M. Liebmann 357(HOLOTYPE: 
C; possible fragment of holotype: GH!). Liebmann apparently collected 
type material at San Bartolo, Oaxaca, in 1842, along the main route 
between México City and Tehuantepec. Sousa (1979) showed the main 
route between these cities, which was traversed some 8 years earlier by 
the collector Andrieux. McVaugh (1972) noted that Liebmann collected 
in Oaxaca along the Pacific Coast during the year concerned. There 
is but little question that the packet material I examined at GH, cited 
above, belongs to what previously has been called S. caducifolia and is 
probable fragmentary type material of it S. liebmannii. 


Vernonia hypoleuca DC., Prod. 5:27. 1836. Liabum hypoleucum (DC.) S.F. 
Blake, Proc. Biol. Soc. Washington 39:144. 1926. 


Sinclairia blakei H. Robins. & Bret., Phytologia 28:60. 1974. [not Sinclairia 
hypoleuca (Greenm.) Rydb.] TYPE: MEXICO. Guerrero(?): between 
Acapulco and México City, Nov 1790-Dec 1791, Haenke s.n. (LECTO- 
TYPE: G-DC, microfiche!). 


Liabum caducifolium B.L. Robins. & Bartl., Proc. Amer. Acad. Arts 43:59. 
1907. Stnclairia caductfolia (B.L. Robins. & Bartl.) Rydb., N. Amer. Fl. 
34:299. 1927. TYPE: MEXICO. Guerrero: Acapulco and vicinity, Oct 
1894-Mar 1895, E. Palmer 245 (HOLOTYPE: GH!; Isotype: F!). 


Slender, often arching, sometimes clambering shrubs or small trees to 10 m 
high which become deciduous at the time of, or soon after, flowering. Stems 


Turner: Revision of Sinclairza including Liabellum 195 


slender and whip like above, but 5-7 cm thick below. Leaves 6-15 cm long, 
3-12 cm wide, bicolored; petioles 1-4 cm long; blades mostly to some extent 
triangular with basal lobes or flanges, 3 nervate, white tomentose beneath, the 
margins weakly serrulate to nearly entire. Heads eradiate, numerous in rather 
stifly branched corymbose panicles. The ultimate peduncles 3-8 mm long, at 
first loosely arachnoid but soon glabrate. Involucres 5-8 mm high, 2-3 seriate, 
gradate, the bracts 15-20, closely appressed with acute apices. Ray florets 
absent. Disk florets 6-12, the corollas glabrous, creamy white or pale yellow, 
8-12 mm long, the tubes 3-4 mm long, grading into an ill defined tubular 
throat, the lobes ca 2 mm long. Achenes 3-4 mm long, moderately pubescent, 
more so above, the pappus of yellowish white or tawny bristles 7-9 mm long, 
the outer scales 1-2 mm long, scarcely differentiated. 

DISTRIBUTION (Figure 6): Jalisco to Oaxaca along the Pacific slopes in 
tropical deciduous forests, 50-1200 m; Oct-Dec. 

REPRESENTATIVE SPECIMENS: MEXICO. Colima: 13 mi N of Santi- 
ago, 200-300 m, 10 Dec 1959, McVaugh 1642 (LL,MICH). Jalisco: 6 mi SW 
of Pihuamo, 2200 ft, 29 Oct 1962, Cronquist 9764 (GH,NY,TEX). Guerrero: 
Taxco, 16 Jan 1936, Abbott 43 (GH). Michoacan: 10 km SE of Aquila, 50 m, 
24 Nov 1963, Feddema 2725 (MICH,TEX). Morelos: Cuernavaca, 3 Jan 1899, 
Deam 7(F,GH). Oaxaca: Mcpio. Jalapa del Marques, Cerro de la Huerta, 620 
m, 17 Nov 1978, Koch 783381 (CHAPA,TEX). 

This fairly common species is noteworthy for its caducous leaves which 
drop soon after flowering commences. Unfortunately, the specific name long 
applied to this species by most workers, including McVaugh (1984), must be 
replaced by the earlier specific name, Liabum hebmannu, as noted above. 

Sinclairia hebmannit appears most closely related to the largely Central 
American species, S. sublobata, both possessing similar, usually lobed, rather 
thick leaves which tend to drop shortly after flowering. The latter can be 
distinguished by its larger heads with obtuse involucral bracts and generally 
larger, less deltoid, leaf blades. 

Liabum adenotrichum is positioned in synonymy here with some trepida- 
tion. The only plant known (the type) appears to be a hybrid derivative, 
perhaps a backcross from S. klatti into S. hebmanni. Both of the latter occur 
in the region of the type of L. adenotrichum. Most of the characters are those 
of S. hebmanni, but the larger heads with a few glands and the somewhat 
drooping capitulescence strongly suggest that an influx of genes from S. klattu 
has occurred. 


14. Sinclairia similis (McVaugh) H. Robins. & Bret., Phytologia 28:62. 1974. 
Liabum similis McVaugh, Contr. Univ. Michigan Herb. 9:468. 1972. 
TYPE: MEXICO. Nayarit: “Shaded places in pockets of trees (notably 
Bursera spp.) on rough lava flow at the base of Volcan Ceboruco,” 
ca 6 mi NW of Ahuacatlan, 3200 ft, 4 Oct 1962, A. Cronguzst 9602 


196 PHY TOLOGIA volume 67(2):168-206 September 1989 


(MICH,TEX!). 


Slender arching shrubs 3-4 m high. Leaves 10-20 cm long, 6-10 cm wide; 
petioles 3-9 cm long; blades broadly ovate to deltoid, bicolored, white tomentu- 
lose beneath, 3 nervate from the base, the margins irregularly serrate. Heads 
eradiate, numerous in terminal corymbose panicles, the ultimate peduncles 
mostly 5-20 mm long. Involucres turbocampanulate, 8-10 mm high, 3-4 se- 
riate, the bracts appressed at first, but squarrose with age, narrowly ovate, 
appressed pubescent, the apices acute. Ray florets absent. Disk florets 15-20, 
the corollas white, glabrous, 9-11 mm long, the tubes 2-3 mm long, gradually 
passing into the throat, the lobes 2-3 mm long. Achenes 3.5-5.0 mm long, 
sparsely short pilose, the pappus of tawny white bristles 7-9 mm long, the 
outer scales not much differentiated, 0.5-1.0 mm long. 


DISTRIBUTION (Figure 4): Nayarit and adjacent Jalisco, mostly tropical 


deciduous forests along the Pacific slopes, 0-1000 m; Oct-Nov. 
_ ADDITIONAL SPECIMENS EXAMINED: MEXICO. Jalisco: 3 km SE 
of Puerto Vallarta, ca 50 m, 12 Nov 1963, Feddema 2507 (MICH,TEX). 

This taxon is known by only 2 collections, cited above. It is clearly related 
to S. lhebmannii, but is amply distinct, as noted by McVaugh (1984), who 
provides an excellent illustration of the species. 


15. Sinclazria klatti (B.L. Robins. & Greenm.) H. Robins. & Bret., Phy- 
tologia 28:61. 1974. Liabum klatti B.L. Robins. & Greenm., Amer. J. 
Sci., ser. 3, 1:156. 1895. Sinclairiopsis klatta (B.L. Robins. & Greenm.) 
Rydb., N. Amer. Fl. 34:293. 1927. TYPE. MEXICO. Oaxaca: Monte 
Alban, 6000 ft, 24 Nov 1894, C.G. Pringle 6059 (HOLOTYPE: GH!; 
Isotypes: GH!). 


Shrubs or small trees 2-5 m high, leafless when in flower. Leaves 10-18 cm 
long, 6-11 cm wide; petioles 3-7 cm long; blades broadly ovate, bicolored, dingy 
white tomentose beneath, the margins denticulate. Heads eradiate, mostly 
10-30 in both terminal and axillary pendant clusters, the ultimate peduncles 
glandular pubescent and arachnoid tomentose, mostly 3-25 mm long. Involu- 
cres 15-17 mm high, narrowly campanulate, 2-3 seriate, the bracts lanceolate, 
2-4 nervate, glandular pubescent, the apices narrowly acuminate. Ray florets 
absent. Disk florets 18-30, the corollas yellow, pubescent, tubular throughout 
or nearly so, 13-16 mm long, the lobes ca 3 mm long, decidedly hispid on the 
apices. Achenes 2-3 mm long, densely pubescent throughout, the pappus of 
numerous tawny bristles mostly 8-17 mm long, the outer series of similar, but 
very short scales. 

DISTRIBUTION (Figure 5): Veracruz and adjacent Oaxaca, mostly bar- 
rancas in tropical deciduous forests, 200-1500 m; Nov-Mar. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Oaxaca: 8-10 mi 
N of Oaxaca, along highway 175, 31 Jan 1960, King 2499 (TEX); Estacion 


Turner: Revision of Sinclairia including Liabellum 197 


Almoloyas, 800 m, 3 Mar 1907, Conzatti: 1751 (GH). Veracruz: Mcpio. Ac- 
ultzingo, 3 km W of Acultzingo, ca 5000 ft, 26 Mar 1970, Long 3279 (F); 
Mcpio. E. Zapata, Plan del Rio, 210 m, 14 Feb 1973, Herndndez 1838 (F); 
Mcpio. Naolinco, 6 km ESE of San Antonio Paso del Toro, 650 m, 14 Jan 
1984, Nee 28816 (F,TEX,XAL); Zacuapan, barranca de Tenampa, Feb 1910, 
Purpus 4886 (GH); Zacuapan, Feb 1914, Purpus 7048 (GH). 


II. SUBGENUS MEGALIABUM (species 16-23). 
Sect. Megaliabum (species 16-18). 


Sinclairia subgenus Megaliabum (Rydb.) B. Turner, stat. nov. Based 
upon: Megaliabum Rydb., N. Amer. Fl. 34:293. 1927. TYPE SPECIES: 
Vernonia andrieuzi DC., Prod. 5:16. 1836. 


16. Sinclairia andrieuri (DC.) H. Robins. & Bret., Phytologia 28:60. 1974. 
Vernonia andrieuzit DC., Prod. 5:16. 1836. Lzabum andrieuzu (DC.) 
Hemsl., Biol. Centr. Amer., Bot. 2:231. 1881. Megaliabum andrieuri 
(DC.) Rydb., N. Amer. Fl. 34:293. 1927. TYPE: MEXICO. Oaxaca: 
between Tehuantepec and the river “Guaracoalcos” {Coatzacoalcos], Sep 
1834, G. Andrieur 269 (HOLOTYPE: G-DC, microfiche!; Photoholo- 
type: F!). 


Stiffly erect suffruticose herbs or soft wooded shrubs 1-3 m high. Leaves 
bicolored, mostly 10-25 cm long, 6-14 cm wide; petioles 1.5-7.0 cm long; blades 
deltoid to subcordate, 3 nervate, the lower surfaces densely white tomentose, 
the margins irregularly serrate to rarely lobed. Heads 1-4, very large, radiate, 
arranged in strict terminal cymes, the ultimate peduncles mostly 2-6 cm long. 
Involucres broadly campanulate, 2-3 cm high, 3-4 cm wide, 4-6 seriate, the 
bracts linear lanceolate, tomentose throughout. Ray florets 21-34, the ligules 
orange or yellow orange, 15-25 mm long. Disk florets numerous, the corollas 
15-18 mm long, the tubes 6-7 mm long, the lobes 3-5 mm long. Achenes 3.5- 
5.0 mm long, densely white pubescent throughout, the pappus of numerous, 
tawny to purplish, bristles mostly 1-2 cm long, the outermost series shorter 
but scarcely differentiated from the inner. 

DISTRIBUTION (Figure 8): Oaxaca, Chiapas and adjacent Guatemala, 
mostly Pacific slopes along barrancas in tropical deciduous forests, 300-1500 
m; Jul-Nov. 

REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: Mcpio. Arriaga, 
13 km N of Arriaga, 830 m, 4 Oct 1972, Breedlove 28286 (CAS,LL); Siltepec, 
3 Nov 1940, Matuda 4108 (F,GH,LL). Oaxaca: Mcpio. Pochutla, 8 km NE of 
Chacalapa, 24 Nov 1977, Delgado 692 (TEX); 37 mi SE of Oaxaca, 13 Oct 
1962, Cronguist (GH,NY,TEX). 

GUATEMALA. Huehuetenango: on rocky cliff along River Selegua, 47 km 
from Huehuetenango, 1400 m, 20 Nov 1967, Molina 21411 (F,GH); NW of 
Cuilco, Cerro Chiquihui, 17 Aug 1942, Steyermark 50776 (F). 


September 1989 


volume 67(2):168-206 


PHYTOLOGIA 


198 


Fig. 8. Distribution of Sinclairia (Sect. Megaliabum). 


Turner: Revision of Sinclairia including Liabellum 199 


This is a common species in Chiapas. Most all field collectors have de- 
scribed the species as having orange or orange yellow corollas. 


17. Sinclairia moore: (H. Robins. & Bret.) H. Robins. & Bret., Phytologia 
28:61. 1974. Lzabum moore: H. Robins. & Bret., Phytologia 27:252. 
1973. TYPE: MEXICO. Guerrero: 36-38 km from Iguala on road to 
Teloloapan, streamsides and slopes by Rio de los Sabinos near Los Sabi- 


nos, 5 Nov 1949, H.E. Moore, Jr. 5518( HOLOTYPE: US; Isotype: GH!). 


Erect suffruticose herbs or shrubs to 3 m high. Leaves bicolored, 10-12 cm 
long, 6-8 cm wide; petioles 2-3 cm long; blades broadly ovate, white tomentose 
beneath, the margins minutely serratulate to nearly entire. Heads radiate, 
2-5 in strict terminal cymes, the ultimate peduncles 1-3 cm long. Involucres 
campanulate, 20-22 mm high, 24-30 mm wide, the bracts 4-5 seriate, gradate, 
densely tomentose throughout. Ray florets 11-13, the ligules yellow, ca 20 mm 
long. Disk florets 25-32, the corollas yellow, 18-20 mm long. Achenes 3.5-4.5 
mm long, densely pubescent throughout, the pappus of numerous bristles in 
2-3 series. 

ADDITIONAL SPECIMEN EXAMINED (Figure 8): MEXICO. Guerrero: 
few miles W of Chilpancingo on dirt road towards Omiltemi, 16 Oct 1976, 
Stuessy & Gardner 4218 (WIS). 

The species appears to stand somewhere between S. andrieuzi and S. 
pringler, but closer to the former, as noted by its authors. 


18. Sinclarria pringlei (B.L. Robins. & Greenm.) H. Robins. & Bret., Phy- 
tologia 28:61. 1974. Liabum pringle: B.L. Robins. & Greenm., Proc. 
Amer. Acad. Arts 32:49. 1896. Megaliabum pringle: (B.L. Robins. & 
Greenm.) Rydb., N. Amer. Fl. 34:294. 1927. TYPE: MEXICO. Jalisco: 
“rocky slopes, mountains near Lake Chapala,” 7000 ft, 18 Oct 1895, C.G. 
Pringle 6214 (published, in error, as 6215) (HOLOTYPE: GH!; Isotype: 
FY). 


Suffruticose perennial herbs or soft wooded shrubs 1-2 m high. Leaves 
bicolored, mostly 8-15 cm long, 5-12 cm wide; petioles 0.3-0.8 cm long; blades 
ovate to somewhat deltoid, 3 nervate, dirty white tomentose beneath, the 
margins irregularly serrate to denticulate. Heads 3-12, eradiate, arranged in 
strict terminal cymes, the ultimate peduncles mostly 1-4 cm long. Involucres 
15-20 mm high, 25-35 mm wide, 3-5 seriate, the bracts ovate lanceolate, 3-5 
mm wide, often squarrose with age. Ray florets absent. Disk florets 70-80, 
the corollas 10-12 mm long, glabrous, creamy white, the tubes 2-3 mm long, 
grading into the throat, the lobes 3-4 mm long. Achenes 3.5-5.0 mm long, 
densely pubescent throughout (rarely sparsely so), the pappus bristles tawny, 
10-12 mm long, the outer scales well defined, 0.5-1.5 mm long. 


September 1989 


volume 67(2):168-206 


PHYTOLO GIA 


200 


Fig. 9. 


Distribution of 


(Sect. Liabellum). 


Turner: Revision of Sinclairia including Liabellum 201 


DISTRIBUTION (Figure 8): Jalisco, Pacific slopes in tropical deciduous 
or oak forests, 1500-2000 m; Aug-Nov. 

REPRESENTATIVE SPECIMENS: MEXICO. Jalisco: ca 25 mi SW Gua- 
dalajara, 1650 m, 1 Oct 1952, McVaugh 13300 (MICH,TEX); mountains N 
of Autlan, 1500-1650 m, 5 Oct 1960, McVaugh 19913 (MICH,TEX); hills of 
Etzatlan, 6000 ft, Pringle 10824 (F,LL,TEX,WIS); 8.7 mi W of Atenquillo, 30 
Sep 1984, Sundberg 2951 (TEX). 

This species is closely related to S. andrzeurzz and was positioned with that 
taxon in the segregate genus Megaliabum by Rydberg (1927), which would 
presumably also include the recently described Sinclairia moore. All of these 
stand somewhere “between” the subgenera Liabellum and Sinclairia and con- 
stitute the principal reason for the submergence of the former within the latter. 


Sect. LIABELLUM (species 19-23). 


Sinclairia Sect. Liabellum (Rydb.) B. Turner, status nov. Based on: Li- 
abellum Rydb., N. Amer. Fl. 34:294. 1927. (TYPE SPECIES: Liabum 
palmer: A. Gray) 


19. Sinclairia hintoniorum B. Turner, sp. nov. (Figure 10.). TYPE: 
MEXICO. México: Distr. Temascaltepec, Chorrera, hill, 23 Sep 1935, 
G.B. Hinton et al. 8482 (HOLOTYPE: LL!; Isotype: GH!). 


Liabum palmeri (A. Gray) Rydb. similis sed differt plantis ro- 
bustis 1-2 m altis herbaceis sed aliquantum suffruticosis, capitulis 
majoribus, acheniis majoribus (5-6 mm longis vs 3-4 mm), et pappo 
setis porphyreis serie exteriori et interiori similari. 


Stiffly erect suffruticose herb to 2 m high (according to label data). Stems 
purplish, pubescent throughout with glandular trichomes. Leaves (only the 
uppermost examined) deeply 3 lobed, the lobes in turn 3 lobed, densely to- 
mentose below, 5-9 cm long, 3-7 cm wide. Heads 5-30 in a strict terminal cyme, 
the ultimate peduncles thick, mostly 2-9 cm long. Involucres broadly campan- 
ulate to hemispheric, 15-20 cm high, the bracts 3-4 seriate, linear lanceolate, 
both appressed sericeous and glandular pubescent. Receptacle convex, decid- 
edly pubescent, 6-9 mm across. Florets 50-60 per head, the corollas yellow, 
15-16 mm long, the lobes ca 5 mm long, a few hairs at the apices. Achenes 
4-5 sided, with 10-12 ribs, 5-6 mm long, appressed pilose, the pappus bristles 
tusty brown in 3-4 series, the inner series about 15 mm long, the outer series 
similar to those of the inner, but shorter (3-8 mm long). 

The type locality is at ca 18° 52’ N, 100° 11’ W (Hinton & Rzedowski 
1975). 


202 PHYTOL O GPA volume 67(2):168-206 September 1989 


Fig.10. Sinclairia hintoniorum, 


from holotype. 


Turner: Revision of Sinclairia including Liabellum 203 


DISTRIBUTION (Figure 9): Known only from the type locality, as de- 
scribed above, having been collected on two different dates at that site. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. México. Distr. Te- 
mascaltepec, Charreua, 1230 m, 10 Sep 1932, G.B. Hinton 2038 (F,GH). 

The species is superficially similar to S. palmerz, but differs markedly from 
that species in its robust habit (ca 2 m high vs 30-75 cm) very large heads 
with more numerous florets, larger achenes (5-6 mm vs 3-4 mm long) with 
pappus rusty brown, the outermost series similar to those of the inner, not 
differentiated into short, flattened, scales. While not noted on the collectors’ 
label, it is likely that the stems arise from subterranean tuber-like roots as in 
the other species of sect. Liabellum. 

It is a pleasure to name this taxon for the remarkable Hinton family, consti- 
tuting three generations of plant collectors in México: George (father), Jaime 
(son) and George (grandson), the latter two individuals still actively collecting 
out of their hacienda near Cerro Potosi, Nuevo Leon. 


20. Sinclairia palmeri (A. Gray) B. Turner, comb. nov. BASIONYM: Li- 
abum palmeri A. Gray, Proc. Amer. Acad. Arts 22:432. 1887. Liabellum 
palmeri (A. Gray) Rydb., N. Amer. Fl. 34:294. 1927. TYPE: MEXICO. 
Jalisco: Rio Blanco, 1886, E. Palmer 586 (HOLOTYPE: GH!). 


Herbaceous perennials 30-90 cm high, the stems leafy and crowded below, 
arising from well defined tuber-like roots. Leaves mostly 8-16 cm long, largely 
confined to the lower portion of the stems, deeply 3 lobed to pinnately dis- 
sected, the divisions mostly 5-20 mm wide, the undersurfaces densely white 
tomentose, the margins serrulate to nearly entire; petioles 2-8 cm long, winged 
throughout. Heads 1-10, arranged in rather strict terminal cymes, the ulti- 
mate peduncles 2-10 cm long, both arachnoid and densely stipitate glandular. 
Involucres turbocampanulate to campanulate, 15-20 mm high, 3-4 seriate, the 
bracts subimbricate, linear lanceolate, both tomentose and stipitate glandular. 
Ray florets absent. Disk florets 30-40, the corollas yellow, sparsely pubescent, 
ca 15 mm long, the lobes ca 4 mm long. Achenes obpyramidal, 2.5-3.0 mm 
long, densely appressed sericeous, the inner pappus bristles 10-15 mm long, 
tawny white, the outermost pappus composed of white scales 1.0-1.5 mm long. 

DISTRIBUTION (Figure 9): Chihuahua, Sinaloa, Durango, Zacatecas, Na- 
yarit and Jalisco, pine-oak woodlands, grassy hillsides and open rocky places, 
1500-2700 m; Sep-Nov. 

REPRESENTATIVE SPECIMENS: MEXICO. Chihuahua: 4 mi SW of 
Matamoros, 6000-6300 ft, 4 Oct 1959, Correll 22738 (LL); Sierra Chinatu, San 
Juan, 8900 ft, 8 Oct 1959, Correll 22921-A (LL). Durango: Canyon Cantero, 
Sierra Gamon , ca 2000 m, 21 Sep 1948, Gentry 8365 (GH); ca 50 km W 
of Cd. Durango, 2550 m, 27 Sep 1962, McVaugh 21655 (LL); 41 mi WSW 
of Cd. Durango, 27 Sep 1984, Sundberg 2891 (TEX). Jalisco: 8-10 km SE 


204 PHYTO T:0.G 1A volume 67(2):168-206 September 1989 


of El Mortero, 2450 m, 5 Nov 1963, Feddema 2479 (TEX); ca Guadalajara, 
7 Dec 1888, Pringle 2127 (GH); W of Bolanos, 15-17 Sep 1897, Rose 2983 
(GH); Michoacan: 9 mi W of Cd. Hidalgo, 24 Sep 1984, Spooner 2561 (TEX). 
Zacatecas: ca 1 km S of Valparaiso, 2000-2400 m, 11 Sep 1966, Anderson 3603 
(GH). 


21. Sinclairia gentryi (H. Robins.) B. Turner, comb. nov. BASIONYM: 
Liabellum gentryi H. Robins., Phytologia 41:46. 1978. TYPE: MEXICO. 
Nayarit: Arroyo del Obispo, 31 mi SE of Tepic, canyon with running 
stream in oak woodlands, on rocks, 2 Aug 1951, H.S. Gentry 11030 
(HOLOTYPE: DUKE; Isotype: ARIZ!; Photoholotype: TEX!). 


Small, scapose, perennials 15-30 cm high, the 2-3 pairs of leaves all basal 
and seemingly opposite. Leaves bicolored, broadly ovate to ovate elliptic, 8- 
12 cm long, 3-6 cm wide, weakly 3 nervate to pinnately veined, the margins 
serrulate to nearly entire, the undersurfaces white tomentose. Heads 2-3 on a 
naked scape, the ultimate peduncles 2-8 cm long, both tomentose and stipi- 
tate glandular. Involucres campanulate, 10-14 mm high, 12-16 mm wide, the 
involucral bracts subimbricate, 3-4 seriate, linear lanceolate, both tomentose 
and stipitate glandular. Ray florets absent. Disk florets 20-30, the corollas 
yellow, ca 12 mm long, sparsely pubescent below, the lobes ca 3.5 mm long. 
Achenes obpyramidal, ca 2 mm long, densely appressed sericeous, the pappus 
of numerous tawny white bristles in 2-3 series, the inner series ca 8 mm long, 
the outer series similar to those of the inner, but 1-2 mm long. 

DISTRIBUTION (Figure 9): Known only from the type locality, as de- 
scribed above. 

The species is closely related to S. palmeri and was treated as synonymous 
with that species by McVaugh (1984). It differs, however, in a number of 
features, as noted by H. Robinson in his original description, including, scapose 
habit, unlobed leaves, larger heads, longer pappus bristles, the outer series of 
which are essentially like those of the inner, except as to size (S. palmeri has 
an outer series of white, scales, quite unlike the inner bristles). The species 
is known only by type material. Robinson did not have access to the isotype 
(ARIZ); while clearly a different plant, it retains all of the features found in 
the holotype. 


22. Sinclairia cervina (B.L. Robins.) B. Turner, comb. nov. BASIONYM: 
Liabum cervinum B.L. Robins., Proc. Amer. Acad. Arts 29:317. 1894. 
Liabellum cervinum (B.L. Robins.) Rydb., N. Amer. Fl. 34:294. 1927. 
TYPE: MEXICO. Jalisco: “Bluffs of barrancas, San Marcos,” 9 Jun 
1893, C.G. Pringle 4398 (HOLOTYPE: GH!; Isotypes: F!,MU!). 


Perennial herbs 50-80 cm high, the stems leafy throughout and arising from 
tuber-like roots. Leaves mostly 10-18 cm long, 3-10 cm wide, deeply tripar- 
tite, the divisions often again lobed, the ultimate divisions mostly 3-10 mm 


Turner: Revision of Sinclairia including Liabellum 205 


wide with entire or irregularly serrate margins, arachnoid tomentose beneath. 
Heads 3-7 in strict terminal cymes, the ultimate peduncles 2-5 cm long, white 
tomentose or with scattered eglandular trichomes. Involucres turbinate, 16-22 
mm high, 3-4 seriate, the bracts mostly 2.5-3.0 mm wide at midsection, densely 
appressed pubescent. Ray florets absent. Disk florets 50-60, the corollas yel- 
low, or reportedly white, 15-20 mm long, the lobes 4-6 mm long. Achenes 
2.5-3.0 mm long, densely appressed sericeous, the inner pappus bristles 12-14 
mm long, tawny, the outer scales 1-3 mm long, not strongly differentiated. 

DISTRIBUTION (Figure 9): Southern Sinaloa, Nayarit and Jalisco, oak 
woodlands and grassy areas, 1000-2000 m; May-Aug. 


23. Sinclairia angustissima (A. Gray) B. Turner, comb. nov. BASIONYM: 
Liabum angustissimum A. Gray in S. Wats., Proc. Amer. Acad. Arts 
22:432. 1887. Liabellum angustissrmum (A. Gray) Rydb., N. Amer. Fl. 
34:295. 1927. TYPE: MEXICO. Jalisco: “Guadalajara,” Jul 1886, E. 
Palmer 215 (HOLOTYPE: GH!). 


Perennial herbs 15-45 cm high, the stems leafy throughout and arising from 
tuber-like roots. Leaves tripartite to the base (rarely not), appearing ternate, 
the divisions entire, mostly 1-3 mm wide, 3-10 cm long, bicolored, the lower 
surfaces densely white tomentose. Heads 1 or 2 at the apices of solitary stems, 
the ultimate peduncles 2.5 cm long, white tomentose or glandular pubescent, 
or both. Ray florets absent. Disk florets 30-40, the corollas yellow, 12-14 mm 
long, the lobes 3-4 mm long. Achenes ca 2.5 mm long, appressed sericeous, 
the inner pappus bristles tawny white, 8-10 mm long, the outer bristles not 
well differentiated, 0.5-1.0 mm long. 

DISTRIBUTION (Figure 9): Endemic to Nayarit and Jalisco, hillsides and 
along bluffs of barrancas, 1500-2000 m; Jul-Aug. 

ADDITIONAL SPECIMENS EXAMINED: MEXICO. JALISCO: hillsides 
near Guadalajara, 2 Jul 1889, Pringle 2501 (F,GH); bluffs of the barranca of 
Guadalajara, 4500 ft, 18 Jul 1902, Pringle 9934 (F,GH,LL). NAYARIT: Santa 
Gertrudis, 8 Aug 1897, Rose 3406 (GH). 

The species appears to be a small version of Sinclatria cervina but the 
leaves are nearly always parted to the base and the divisions are narrowly 
linear, giving the appearance of whorled leaves. The heads are also smaller 
with fewer involucral bracts. McVaugh (1984) notes that S. angustisszma, S. 
cervina and S. palmeri are closely related, all occurring in the basin of the 
Rio Santiago. Nevertheless, from the relatively few collections known, each 
appears to retain their distinct characters and intergrades between them have 
not been discerned. Sinclatria cervina apparently has not been collected since 


1902. 


206 PRY TOLO GEA volume 67(2):168-206 September 1989 


ACKNOWLEDGMENTS 


I am grateful to Dr. Guy Nesom for the Latin diagnoses and for a review 
of the manuscript. Specimens of Sinclairza were borrowed from the following 
institutions: F, GH, LL, MU, TEX, WIS; I am grateful to the Directors for 
this courtesy. 


LITERATURE CITED 


Hinton, J. & J. Rzedowski. 1975. George B. Hinton, Explorador Botanico en 
el Sudoeste de México. An. Escuela Nac. Cien. Biol. 21:1-114. 


McVaugh, R. 1972. Botanical exploration in Nueva Galiciana from 1790 to 
the present time. Contr. Univ. Michigan Herb. 9:205-358. 


. 1984. Liabum, in Flora Novo-Galiciana 12:576-584. 


Robinson, H. 1983. A generic review of the tribe Liabeae (Asteraceae). 
Smithsonian Contr. Bot. 54:1-69. 


, A.M. Powell, R.M. King, & J.F. Weedin. 1985. Chromosome num- 
bers in Compositae, XI:Liabeae. Ann. Missouri Bot. Gard. 72:469-479. 


Rydberg, P.A. 1927. Liabeae, in N. Amer. Fl. 34:289-360. 


Sousa, S.M. 1979. Itinerario botanico de Andrieux in Mexico. Taxon 28:97- 
102. 


Phytologia (September 1989) 67(2):207-208. 


BOOK REVIEWS 


Michael J. Warnock 
Department of Life Sciences, Sam Houston State University, Huntsville, 


Texas 77341 USA 


The Rhododendron Species Vol. II Elepidotes, Part 1 Arboreum - Lacteum. 
H.H. Davidian. Timber Press, 9999 SW Wilshire, Portland, OR, 97225 
USA. 1989, 390 pp., illus, $54.95 + shipping, cloth. ISBN 0-88192-109-2. 


At first glance, this book appears to be of the typical coffee table variety 
with lots of pretty pictures and with its primary use to be as a conversation 
piece. While the photographs and drawings are excellent, and a joy to exam- 
ine, the text itself contains an abundance of information useful to practicing 
scientists and horticulturalists as well. The book begins with a series of maps 
of southeast Asia, where most of the Rhododendron species originate. The 
maps are followed by a brief glossary, with the next several pages occupied by 
an illustrated glossary which is used to clarify many of the terms used in the 
keys and descriptions to refer to leaf and corolla shapes, vestiture and seeds. 
Without the drawings in this illustrated glossary, many of the descriptions 
and keys would be extremely difficult to decipher, since many of the terms 
are not commonly used to describe features of other plants. A series of maps 
of horticultural hardiness zones for the northern hemisphere is included as a 
guide for successful outdoor cultivation of Rhododendron species. Keys to se- 
Ties, subseries and species seem adequate to separate the taxa, although I am 
not familiar enough with the plants to know if they really work. One difficulty 
that I had with the keys was the unnecessary splitting of couplets. For exam- 
ple, in the key to series, the primary lead to the first couplet is on the first 
page of the key and the secondary lead to the first couplet is at the end of the 
key, four pages later. The same is true of several other couplets in the keys. 
Completing short leads first will solve most of these separations and make the 
keys easier to use. Each taxon is described at length, and most of the species 
discussions include some historical background of the plant (when, where and 
by whom it was brought into cultivation-although many are not cultivated 
at the present time) as well as notes to aid prospective growers of the plants 
(particularly in Britain). Each description also includes the describing author 
and publication citation—not at all typical of the usual conversation piece. In 
fact, several new taxa are described and published in this work, making it 


207 


208 PHYTOL OCIA volume 67(2):207-208 September 1989 


an essential for workers interested in the systematics of Rhododendron. This 
book would be a fine addition to the library of any Rhododendron enthusiast, 
whether they were interested in taxonomy, horticulture or simply want to look 
at the pictures. 


McGraw-Hill Dictionary of Scientific and Technical Terms, 4th Ed. 
Sybil P. Parker, ed. McGraw-Hill Book Co., New York, NY. 1989, xvi, 
2088 pp. + 49 pp. appendix, price unknown, cloth. ISBN 0-07-045270-9. 


A monumental and possibly impossible task is attempted in this book. A 
laudable effort has been made to assimilate the terms and definitions from a 
tremendous diversity of fields into one volume (and quite a large one at that). 
In fact, the listing of fields runs the gambit from “Acoustics” to “Zoology.” 
Each term has a brief definition, as in a typical dictionary. Many of the terms 
are quite specialized in use and scope. My impression is that practicing scien- 
tists will find this work of limited usefulness because most of the terms in this 
dictionary will not apply to their field and they will have no opportunity to use 
them, and they will already know the definitions for most of the terms that do 
apply to their particular field. The greatest usefulness for this book might be 
for politicians and administrators who frequently need to deal with scientific 
and technical terms with which they may be unfamiliar. For biologists (and 
botanists in particular), the book is even more limited in its utility since the 
bulk of the terms seem to apply to engineering and the physical sciences. The 
appendices include the familiar metric/ American standard conversion factors, 
physical constants, computational formulae and abbreviations. A limited bi- 
ographical listing is included in the final pages and some of the names which 
are or are not on the list are interesting. For instance, Copernicus and David 
Baltimore are included, while Arthur Cronquist, Asa Gray and Leonardo da 
Vinci are not. You might buy a copy for your favorite bureaucrat. 


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