, 
, 
• 
MEXICAN "CACALIOID" GENERA ALLIED TO 
SENECIO (COMPOSITAE) 1 
By RICHARD W. PIPPEN' 
Introduction 
This paper represents an investigation of a geographically distinct 
group of North American species of Compositae occurring mainly 
in the highlands of Mexico. In the past these species have been con­
sidered part of the genera Cacalia (DeCandolle, 1838; Gray, 1883; 
Greenman, 1902), Senecio (Schultz 1845; Bentham & Hooker, 1873; 
Hemsley, 1881), and several distinct, smaller genera (Rydberg, 1924; 
Belcher, 1956; Cuatrecasas, 1960). In my study I worked out a taxo­
nomic system for these species, then compared the species individually 
and as a group with superficially similar species of Senecio, and con­
sidered their relationships. It is now apparent that these species are 
generically distinct from Senecio, from the species in eastern North 
America and Asia usually referred to Cacalia, and also from the 
type-species of Cacalia; instead, four separate genera are represented­
Digitacalia, Odontotrichum, Pericalia, and Psacalium. 
The materials used in this study consisted primarily of specimens 
obtained from the following herbaria: Field Museum of Natural His­
tory, Chicago; Gray Herbarium of Harvard University, Cambridge, 
Massachusetts; Botanische Staatssammlung, Miinchen; University of 
Michigan, Ann Arbor; New York Botanical Garden, New York; 
University of Texas, Austin; and United States National Herbarium, 
Smithsonian Institution. I wish to express my gratitude to the cura­
tors of these herbaria who made these specimens available. Abbrevi­
ations of herbarium D8JDes used hereafter are those of Lanjouw and 
Stalleu (1964). The abbreviation "Field Mus. Neg." refers to Field 
Museum of Natura! History negatives of a series of type photographs 
taken in European herbaria by J. F. Macbride during the period 1929 
to 1939. "U. of MICH. Neg." refers to a collection of negatives of 
type specimens maintained at the University of Michigan Herbarium. 
J Part of a dissertation presented to the Horace H. Rackham School of Grad­
uate Studies, the University of Michigan, in partial fulfillment of the requirements 
for the degree of Doctor of Philosophy. 
I Department of Biology, Western Michigan University. 
365 
366 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Observations were made in the field, and preserved cytological 
materials and dried specimens were obtained during two seasons 01 
field work in Mexico (in the summer and autumn 01 1958, and the 
autumn 01 1960), and from a survey trip through the southeastern 
United States in the summer of 1962' Observations were also made 
01 living plants grown from seeds and rootstocks in the University 
01 Michigan Botanical Gardens. 
All measurements were obtained fwm dried specimens. But flower­
ing material 01 all the species was studied both in the dry stllte and 
after soaking, the latter state inducing no significant change in size. 
Dimensions 01 the heads include Irom the base of the phyllaries to 
the tip of the syngenesious anthers. 
The chromosome counts were obt"ined by the standard acet,o­
carmine squash technique on pollen spore mother cells killed and 
fixed in Newcomer's solution. Exeept for Odontotrichum decompositum 
the chromosome numbers herein reported are new reports. Table 1 
summarizes this information. Voucher specimens lor these counts ft.re 
listed in table 1 and IIlso indicated with an asterisk (*) in the list of 
specimens examined under the species. Voucher specimens are pre­
served in the University 01 Michigan Herbarium. 
For advice and assistance my sincere thanks and appreciation 
fire extended to Dr. Rogers McVaugh, who initially c!tlled my atten­
tion to tbis problem and who collect.ed specimens and cytological 
material for me on several occasions; to Dr. B. L. Turner, the Uni­
versity of Texas, for permitting me to cite unpublished chromosome 
counts from his cytological investigations 01 the Mexican composites; 
to the late Mr. N. Y. Sand with, the Royal Botanic Gardens, Kew, 
who sent me special information regarding the type 01 Cacalw. cir­
siifOlw. Hook. & Arn. ; to Dr. A. G. Norman, formerly Director of 
tbe University of Michigan Botanical Gardens, lor the use of the 
flleilities there for growing living plants; to Mr. J. R. Millar, formerly 
Chief Curator of the Herbarium, Field Museum of Natural History, 
for muking available the botanical collections of Sess6 and Mocino 
(then on lonn to Chicago from Madrid); and to Dr. Charles Feddema, 
U.S. F,west Rervice Herbarium, for the line drawings of Hower types, 
Morphological Considerations 
Although these species, previously referred to Cacalw., do apparently 
represent several genera, they have in common It number of morpho­
logical leatures aTld generalizations regarding these may be made. 
Except for D0itacalw. (here desc"ibcd as Tlew) nil the genem ill the 
S Made possible by a grlUlt from the Graduate Student Research fund of the 
Horace 1£, Rackham School of Graduate Studies, the University of Michigan. 
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 367 
TABLE I.-Meiotic ch1'omosolltc COttnts of Bome species of klexican caealioids 
Species Voucher Specimens I Number 
- , 
Digitacalia tridactylitis MeV.ugh 21903 330 
Odontotrichum amplum Pippen 4 ' 30 
Odontotrichum brachycomum MeV.ugh 21934 330 
Odontotrichum cirsii/olium Pippen 52 ' 30 
Odontotrichum decompositum - • 30 
Odontolrichum muUilob'Um Pippen 30 • 30 
Odontotrichum palmeri Pippen 12 Cft, 2 25 
Odonlolrichum pringlei Pippen 37 ' 30 
McVaugh 21765 • 30 
Odontotrichum 3inuatum Pippen 9 • 30 
Pericalia michoacana Pippen 63, 65 ' 30 
Pericalia se8sili/olia. Pippen 61 ' 30 
Psacalium eriocarpum Pippen 62 ' 30 
Psacalium holwayana MeVaugh 21953 ' 30 
Psacalium megapkyllum Pippen 8 • 30 
Psacalium peUatum v. peUatum Pippen 49 I ' 30 
Psacalium species MeV.ugh 21913 , , 30 
-
1 Voucher specimPlls retaim'd ill the Herbarium of t.he Uni"crbit.y of Michigan, 
Ann Arbor. 
2 Determined by Pippen. 
I Determined by B. L. Turner, here published. 
, Detennined by Kruckeberg in Ornduff et aI. 1963. 
Senecioneae, including the cacalioid ones, are contrasted by Belcher 
(1956). 
All of these species are herbaceous perennials, flowering from July 
to December. The perennial caudex and/or roots survive the dry 
season but the aerial portions wither soon after flowering. The first 
season of growth is strictly vegetative. A basal rosette of leaves 
forms in Ockntotrichum and Psacalium; in Digitacalia and Pericalia 
a nonflowering, leafy stem develops. Flowering may occur during the 
second and succeeding years. 
Leaves in all of the genera are petiolate (except certain specimens 
of Pericalia ses8ilifolia). The leaves are usu.lly fleshy or coriaceous 
with prominently convex veins usually more pronounced on the 
abaxial surface. In several species of Odontotrichum (0. pachyphyUum, 
O. platylepis, and O. amplum) the two surfaces of the leaves are 
similar with the venation equally pronounced on both surfaces. 
The leaf shapes and patwrns of lobing are relatively constant, al­
though wide ranges of variation may occur within the limits of some 
species (e.g., Odontotrichum sinuatum) and occasional parallel leaf 
forms between species (0. silphiifolium and O. go/d,'mithii). In general, 
however, each species may be recognized by a characwristic lear shape 
368 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
and pattern of lobing. The venation is mostly pinnate, 8ubpalmate, 
or radiate (the latter in Psacalium only). 
In Odontotrichum the form of the leaves exhibits a series ranging 
from entire or toothed (as in O. amplum, O. pachyphyUum, and O. 
silphiifolium) to lobed (0. sinuatum, O. pringlei, and O. multilohum) 
and pinnatisect (0. decompositum). This series in general parallels a 
reduction in the size of heads and size of inflorescence bracts . 
The inflorescences are monochasial (Rickett, 1955) with at least 5 
heads. They may be corymbose, paniculate, or less frequently race­
mose. The bracts subtending the branches of the infloresconce and the 
calyculate bracts are relatively constan t \\i thin a species and either 
small, linear-subulate or larger and leafy. The leafy inflorescence bracts 
are usually vaginate or naviculate and clasping the stem or in some 
species, petiolate. 
Heads are always discoid with perfect and fertile flowers. The size 
of the heads, as determined by the number of phyllaries and flowers, is 
relatively constant ,vithin each species. 
The flowers are nearly white or cream colored, often "ith a greenish 
or purplish tint but never yellow or orange. The corolla tube is narrow­
ly cylindrical and usually as long as or longer than tho lobes of the 
limb. The limb is divided into 5 equal lobes that are usually free to the 
base of the limb or, in some species, connate for part of their length 
into an ampliate, campanulate, or funnclform throat. The throat is 
usually shorter than the lobes or, less frequently, as long as the lobes 
(longer than the lobes only in Pericalia michoacana). 
This condition of tbe corolla is referred to as the "cacnlioid condi­
tion" to distinguish it from that found in Senecio where the corolla 
throat is mucb longer than the lobes, the latter forming only five small, 
deltoid, ascending teeth above the thro"L--referred to as the "sene­
cioid condition" (fig. 1). 
The achenes vary from ellipsoid, obovoid and dorsally compressed, 
elliptic or ovate in cross section (Odontotrichum and Psacalium) to 
oblong, subcylindrical, and subterete in cross section (Digitacali4 and 
Pericalia). The surface is usually narrowly striated or often promi­
nently 10- to 12-ribbed. In sevcral species the achenes are pubescent 
with simple unicellular hairs but none of the species bas hygroscopic 
trichomes like those found on achenes of several species of Senecio. 
The pappus consists of a single row of many, persistent (deciduous 
in Od<mtotrichum cirsiifolium) , white or cream colored, stifr, barbellate, 
capillary bristles (as in most of the Senecioneae). They are usually as 
long as the corolla or occasionally much shorter and stiffer (0. bra­
chycomum and O. palmeri), or even entirely lacking (Psacalium ca1vum 
and P. nanum). 
A study of pollen grains of representative species (mounted in lactic 
PIPPEN-MEXICAN "CACALIOID" GENERA 
a b 
• 
• 
• 
• 
• 
• 
• 
• 
• • 
• 
• 
• 
• 
• • • 
• • • • • 
• • • 
• 
d e 
• 
• 
! 
• 
• 
• 
• 
• , 
c 
FIGURE I.-Types of discoid corollas (all X 10): a. Ptricalia uJJilijoiia (PipPIn 64, MICH); 
b. P,acali.m pI/tatum var. typo (PiPPN' 49, MICH); c. Odontotrichum sinuatum 
(Pipptn 22, MICH); d. Digitacalia jatropAoidts CL. C. Smith 123, MICH); I. S~tio 
anguli/oli.., (aect. PaImatinnTlit) (McYaugh 9949, MICH); /. &Mcio callolw (sect. 
Mul"di/oli.) (H ... d 36, MICH). 
370 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
acid or Hoyer's medium) revealed a similarity in all four genera. The 
grains are spherical, tricolpate, 42-45 microns in diameter. The surface 
is ornamented with simple spines 7 to 9 microns long. 
Several species exhibit various degrees of pubescence, the presence 
or absence of which on organs such as leaves, achencs, and phyllaries 
is relatively cons(nnt for each species. The trichomes are mostly non­
glandular, conical-attenuate, white or hyaline, multicellular, frequen tly 
with purple cross walls. They vary in length, number of cells, stilTne"" 
and density. Occasionally the distal end of the hair is prolonged into 
" caudate tip as long as or longer than the body. A minute s(,ipitnte 
:.:lllnd, ca. 0.5 mm. long, consisting of a stalk of several cells in a sin­
;;le row, topped with a multicellular, globose, purple gland OCClirs 
mixed among the longer, nonglandul"r hairs in severnl species of 
Psacalium. 
Distribution and Ecology 
The genera Diyitacalia, Odoniotrich'um, Pericalia, and Psacalium 
nre nil endemic to Mexico, except thllt one species, Odontotrichum 
decompositum, extends northward into the mountains of southern 
Arizona. The species of all four genera are restricted mostly to the 
oak-pine zone, which is extensively developed in the higher mountain 
regions of Mexico. Within this zone the species of Odontotrich71.m and 
TJigitacalia are usually associated with the grasslands or oak savannahs 
and Pericalia and Psacalium with the open oak-pine woods or their 
borders. But species of the different genera do not usually grow to­
gether in the same community. 
The annual seasonal pattern in the oak-pine woodland zone is one 
of alternating wet and dry periods. The rainy season often begins 
in May and continues as late as October in certain areas, the peak 
reached during July and August. The dry season lasts from about 
mid-November until mid-May. The highest temperntures are usually 
reached during the months just prior to the peak of the rainy season, 
April through June (Contreras Arias, 1942). The active growing 
period corresponds to the rainy season with little plant growth 
during the dry season when the plants die or remain dormant. 
Most of these tuxa occur at altitudes between 1,000 and 3,000 
meters, although a few specimens have been collected at elevations 
as low as 350 meters. These plants seem to thrive in disturbed sec­
ondary associations such as grazed grasslands, burned-over wood­
lands, or roadsides. Originally they probably grew along the margins 
of the oak-pine woods or in natural openings within the woods. The 
majority of the species grow in well-drained J'()('l.;y, day goil~ but n 
few spceies do well in poorly drained, similar soils. 
The region of greatest contentration of these spceies corresponds 
closely to the Traus-Mexican volcanic belt (Clausen, 1959) or Neo-
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 371 
volcanic Pluteuu (Ruisz, 1959). This zone extends from Lhe region 
of Orizaba, in Veracruz, through the region of Mexico CiLy to west­
ern Michoacan, thence curving northward toward Guadalajara and 
again northwestward toward Tepic. In all directions away from this 
zone the number of species diminishes rapidly. Other centers of 
concentration for Digitacalia, Odontotrichum, and Psacalium fife in 
southern Durango and central Oaxaca, in the latter region are several 
endemic taxa. 
The general distributional patterns within Mexico are illustrated 
in figures 2 and 3. 
Generic Relationships and Evolution 
The genera Digitacalia, Odontotrichum, Pericalia, and Psacalium are 
apparently natural genera, possibly having closer relationships to sep­
arate elements within the genus Senecio than to each other. Still, these 
segregate cacalioid genera may be distinguished as a group from 
Senecio by a nnmber of relatively constant characters if the characters 
are applied collectively (table 2). Odontotrichum is the largest genus 
in the complex. It may be distinguished from the other cacalioid 
genera by the subscapose habit, the erect, nonpeltate basal leaves, and 
TABLE 2.-Comparison of cacalioid genera and Senecio 
CacaUold Genera Senedo 
(a) Heuds always discoid, the flowers (a) Heads radiate or discoid, the ray 
all regular and perfect. flowers pistillate; t.he discoid 
flowers regular and perfect. 
(b) Corollas always creamy or nearly (b) Corollas mostly yellow 01" purple, 
white, never yellow. occasionally creamy or nearly 
white. 
(c) Corollas deeply lobed, the lobes (c) Corollas mostly shallowly lobed, 
extending to the base of the limb the lobes usually lcss than half as 
or connate into a throat rarely as long as the throat. 
long as the free lobes. 
(el) Achenes mostly obovate or ellip- (el) Achenes mostly cylindric, subtercte 
soid, elliptic in cross section, lcss in cross section; glabrous or pu-
frequently cylindric and subteretc bescent, the hairs in mnny species 
in cross section; glabrous or pu- hygroscopic. 
bescent, the hairs never hygro-
• SCOpiC. 
(e) Involucre mostly campanulatc or (c) Involucre cy lindrical, less fre-
turbinate, less frequently qucntly campanulate or turbinate. 
cylindrical. 
(I) Plants all hcrbaceous perennials. (I) Plants annuals, biennials, herba-
ceous perennials, shrubs or trees. 
.. ----~ ~--
372 CONTRlliU1'IONS FROM TH£ NATIONAL HERBARIUM 
, 
_. 
~....,""-':--=-'-o- ~-1'" ____ " 
r .... __ ''' _ 
-~-.... --
-'.'--:--:,,-,--""",~--. , • - lO __
, 
\ 1 
'h 
.. _- , 
I 
===:,' 
• 
2 
FIGUaE 2.-Distribution mapa of the Mexican cacllioid genera: 
1. DigitQcQlia; 2. Odontot,i,hllm . 
• = Collection of one or more species . 
• 
I 
I 
I 
I 
! 
I 
I 
._----
. , 
, 
• , 
• 
• 
I'lPPENr-MEXICAN "CACALIOm" GENERA 
~. - --
~ ...... -- ... 
'""-''''' ­_ .. __ ._-
, 
. -
• 
- -­
... ---, ..• ' ....... ,' 
-----..- ,-
• 
• .~-
..' , 
• 
• 
\. 
• 
'. 
• 
4 
FIGUllB 3.-Diatribution mapa of the Mexican cacalioid genera: 
3. Pnicalia; 4. PJacalium. 
e= Collection of one or more specie •• 
373 
374 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
the corolla lobes free to the base of the limb (fig. 1). Taxa of this de­
scription apparently have no counterpart in Senecio unless it is in S. 
sect. MUlgedijotii. The species in this section have discoid heads usu­
ally of purple flowers (although some white-flowered forms do or('nl') 
but the corolla throat is much longer thfln the lobes, the lobes forming 
only five small, deltoid teeth above the throat (fig. 1). The leaves are 
mostly membranaceous, narrowly ovate or oblong, regularly cllllose­
dentate, and often runcinate or Iyrate-pinnatifid. Several of these 
species have been described as cacalias (sensll latu), e.g., S. caUos118 
Greenm., S. coulteri Greenm., S. runcinatus Less., S. pltrpurascens 
Klatt, and S. d~formans Klatt. 
Psacalil1m is the most distinctive genns of the complex because of 
the centrally peltate leaves. It is further distinguished by the frequent 
occurrence of stipitate, glandular trichomes in the inflorescence and 
the fusion of the corolla lobes into a short throat ca. 1 mm. long (fig. 1). 
No group of species in Senecio is known to be comparable to the species 
of Psacali1J.m, although several species in S. sect. Palmatinenni appear 
closely allied. Senecio 8ubpeltatm Schultz Bip. and S. cordobensis Hems!. 
have peltate leaves but the flowers are typically senecioid and the 
plant., are woody shrubs. Senecio angulifoliu., DC. is similar in several 
respects to P. pelf4tum; the two have Icnfy inflorescence hmcts, beads 
of approximately the slime size, glanduhlr-pubescent stems, and a 
chromosome number reported 8S n=30. However, S. ang1l1ijolim is a 
woody shrub, the leaves are not peltate, the beads are usually radiate 
(occasionally discoid), and the flowers are always yellow. 
Since this group of senecios appenes to be somewhat transitional 
between Senecio and Psacalium, it is possible that the Intter was de­
rived from 1m ancestral stock that also gave rise to species like those 
in S. sect. Palm4tinermi. 
In Digitacalia the plants are leafy stemmed and the upper leaves 
reduced but not bracteirorro. There is neither a basal rosetle of 
leaves nor subterranean tubercles. The corolla is mostly deeply deft 
(fig. 1), but the achenes are subcylindric, suhterete in cross section, 
and thus tending toward the senecioid type. Senecio roldana DC. 
and S. jaliscana S. Wats. resemble the species of Digitacalia in habit, 
although the leaves are less deeply lobed, but in floral characters 
they are distinctly senerioid; S. jaliscana has discoid heads of orange 
flowers and S. roldana radiate heads of yellow or white fiowers. In 
both species the corollas are senecioid. 
Conceivably Digitacalia may have been derived from the same 
anrestral stock that n.lso gave rise to species in Senecio like those just 
diRcmssed. 
Finally, Pericaha is unique ill the cacalioid eomplex ue(,Hllse of its 
leufy-stemmed habit, the distill brartciform leaves, the fleshy suh-
• 
PIPPEN-MEXICAN "CACALIOlD" GENERA 375 
terrtlllMn tuberele" lbe JllImer()U, (S-J5) calyclilate brad" and the 
well developed, long corolla throat with relatively long, recurved 
corolla lobes. This genus contains only two species and these have 
no apparent counterpart in Senecio. But S. hartwegii Benth. and S. 
seemanii Schultz Bip. (sect. Palmatinervii) have certain similarities 
in vegetative structure to the speciCll of Pericalia, although the heads 
of these senecios are much smaller and distinctly senecioid. 
A Central American species, Senecio cooperi Greenm. (sect. Multi­
nervii), has disk florets similar to the discoid flowers of the cacalioid 
genera, with the corolla lobes much longer than the throat. The heads, 
however, are radiate, th~ flowers yellow, the phyllaries narrowly 
oblong and barely overlapping, and the leaves lyrate-pinnatifid. This 
combination of characters definitely associates the species with 
Senecio. 
The hypothetical relationship of the cacalioid genera to each other 
and to the closely allied species of Senecio is summarized in figure 4. 
It is also apparent that the Mexican cacalioid taxa are not the same 
as those in the eastern United States (Cacalia sect. Conophora DC.) 
or those in eastern Asia (C. sect. Eucacalia DC.). Further studies of 
these groups are necessary to determine their proper rela.tionships 
and taxonomic position. 
The 7 or 8 species usually referred to Cacalia sect. Conol'hora DC. 
form a natural group endemic to the eastern United States and 
southern Canada. These are easily distinguished from the taxa in­
cluded in this paper as well as from Senecio by: The presence of a 
prolonged chaffy scale forming a beak up to two millimeters long in 
the center of the receptacle; the consistently small heads with 5 
phyllaries and 5 or 6 flowers; the entire or dentate but not lobed, 
glabrous or glaucous leaves; and the haploid chromosome number of 
usually 25. 
The 8 to 10 species that most earlier workers referred to C. sect. 
Eucacalia comprise a natural group mostly endemic to eastern Asia, 
with tho exception of C. suaveolens L. which is found only in the 
eastern United States. These differ from the Mexican members of 
this complex in: The lack of tufts of hair at the base of the basal 
leaves; the narrowly paniculate inflorescence; the short corolla tube; 
and the throat of the corolla equaling or exceeding the corolla lobes 
in length. 
The genus Cacalia proper, a small group of central European 
species typified by C. alpina L. (see discussion to follow) differs from 
the taxa discussed in this paper primarily as follows: The style 
bmnches are long-attenuate, the abaxial surface uniformly densely 
glandular-papillose; the corolla is usually 4-parted; the chromosome 
number is reported as n=19 (Langlet, 1936); and these taxa, under 
376 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
SENECIO HARTWEGII 
S. SEEMANNII 
I 
I r PERI CALIA 
I / 
I I 
I / 
I I 
I I I 
/ I / 
DIGITACALIA 
S. JALISCANA 
S. ROLDANA 
I I I 
I I I 
I I / 
I I I / 
SESSiLiFOLIA 
MICHOACANA 
HETEROIDEA 
CHIAPENSIS 
JATROPHOIOES 
NAPEIFOLIA 
TRIDACTVlITiS 
.LU / 
--- V / S. SECT. MULGEDIFOLII \--// 
ANCESTRAL ODONTOTRICHUM 
PROTOTYPES ~ 
..... -- -< " \ \ \, PSACALIUM 
\ \ 
\ \ 
\ \ S. SECT. \ 
\ 
\ 
\ 
PALMATNERVII 
\ 
\ 
~ S. ANGULIfOI.lUS I S. COOPERI 
NELSONII 
PELTATUM 
LAXIFLORUM 
HOLWAYANUM 
ERIOCARPUM 
SP. 
TA8ULARE 
MEGAPHYLLUM 
POCULIFERUM 
PELTIGERUM 
NANUM 
CALVUM 
PACHYPHYLLUM 
AMPLUM 
SILPHIlFOlIUM 
GOLDSMITHII 
AMPLIFOLIUM 
GLOBOSUM 
PURPUSII 
BRACHYCOMUM 
PALMERI 
PLA TYLEPIS 
PRINGLEI 
PAUCICAPITATUM 
SINUATUM 
MULTILOBUM 
RADULIFOLIUM 
TUSSILAGINOIOES 
NAPELLIFOLIUM 
CERVINUM 
FlLlCIFOLIUM 
CIRSIIFOLIUM 
DECOMPOSITUM 
FIGURE 4.-Hypothetical relationships of the .pecica of the cacalioid genera in Mexico to 
each other and to certain closely allied species of Stnl(jo. Vertical gaps and positions 
represent relative morphological diversity among species and groups of species. 
the name Adeno8tylu, have been referred to the tribe Eupatorieae 
(Hoffmann, 1890). 
Nomenclature of Cacalia 
One of the major poin Is of difficulty concerning the generic nomen­
clature of the te.xa considered in this paper as well as that of Garolia 
has been the typification of the latter, since no type-species was desig­
nated by Linnaeus. The problem of the typification of Gacalia has 
• 
• 
PIPPEN MEXICAN "CACALIOID" GENERA 377 
been reviewed several times (Rydberg, 1924; Hitchcock & Green, 1947; 
Shinners, 1950; Cuatrecasas, 1960) and therefore does not need to be 
repeated in detail here. It seems clear that O. alpina L. is the most 
logical lectotype of Oacalia. This species, named O. alpina by Linnaeus 
(1753), embodied the Linnaean and pre-Linnaean concept of Oacalia 
in that essentially all of the species of Oacalia described by the pre­
Linnaean botanists (L'Obel, 1581; Clusius, 1601; Bauhin, 1623; 
Morison, 1699; Tournefort, 1700) actually represented the same 
• 
specIes. 
Linnaeus's concept of Oacalia was an inclusive one, and II number 
of the species he considered in this genus were transferred to other 
genera (Haworth, 1812; Cassini, 1816, 1826, 1827). Nomenclatural 
difficulty began when Cassini (1816) created a new genus, Adenostyles, 
for a small group of central European plants including O. alpina L. 
as the type-species. Adenostyles was actually superfluous when pub­
lished since the name Oacalia should have been used for the genus 
containing O. alpina. 
The name Adenostyles has been generally accepted (DeCandolle, 
1838; Bentham and Hooker, 1873; Hoffmann, 1890; Hegi, 1917; Fiori, 
1933) for O. alpina and four or five other species endemic to central 
Europe. 
DeCandolle (1838) summarized the work of Cassini and the other 
post-Linnaean authors. He completely revised the cacalioid complex 
and in effect retypified Oacalia, although he designated no type-species. 
He divided Oacalia in to four sections: Eucacalia and Oonophora and 
two others that he considered doubtfully valid, Aulacophora and Gis­
sampelopsis (the latter two comprised of several shrubby paleo tropical 
yellow-flowered species now mostly referred to Senecio). In the 
emended genus Oacalia, he retained only three Linnaean species, O. 
hastata and O. suaveolens in Eucacalia, and O. atriplicifolia in Oono­
phora. DeCandolle recognized Cassini's genus Adenostyles, therebyex­
cluding from Oacalia the Linnaean type of the genus; he used the 
name Psacalium Casso for the Mexican species with peltate leaves; 
and he referred to Oacalia various species now properly referred to 
the genera Digitacalia, Odontotri<Jhum, Peri<Jalia, and Psacalium. 
In the sense of DeCandolle, therefore, Oacalia was to be typified 
by one of three Linnaean species (but not the logical generic type). 
It included a variety of species, occurring in Asia, eastern North 
America, and Mexico, that differed consistently from Senecio in a 
number of respects and was regarded"by most authors (Gray, 1883; 
Hoffmann, 1890; Greenman, 1902; Fernald, 1950; Blake, 1960; Makino, 
1958; Cronquist and Gleason, 1963) as comprising a single widespread 
genus. On the other hand, several authors (Schultz, 1845; BentlIam& 
Hooker, 1873; Hemsley, 1881) did not accept the Candollean Oacalia, 
378 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
prefen·ing t" relegate all of the species to Senecio. Some authors (e.g., 
Rydberg, 1924; CURt.recasas, 1960) touk the position accepted ill this 
paper, that of recognizing several valid genera in the Candollean 
Oacalia. 
Systematic Treatment 
Key to Mexican "Cacalioid" Genera Allied to SeneciQ 
1. Basal rosette present, the caulinc leaves reduced upward, the uppermost 
usually bracteiform. 
2. Basal leaves centrally peltate . . . . . . . . . . . . . . 4. Psacalium 
2. Basal leaves not centrally peltatc, the petiole attached to the blade a.t the 
edge (subpeltate in O. raduliJolium) . . . . . . . . 2. Odontotriehum 
1. Basal rosette lacking, the plants with only caulinc leaves, the uppermost leaves 
reduced but not bracteiform. 
3. Plants with underground tubercles attached to the base of the stem; heads 
8ubtended by 8-15 linear bracts as long as or longer than the phyllaries. 
3. Pericalia 
3. Plants lacking underground tUbercle.«; heads sUbtcnocd by 3-5 Bubulate 
bracts shorter than the phyllaries •...... .... 1. DI,Uacalia 
1. Digitacalia 
Diuitacalia Pippen, gen. nov. 
Plantae C3ulescentes, tuherculis slIbterraneis nullis, caulibus 
foliisque glabris vel puhescentihus, foliis omnibus cau linihus, nlternis, 
numerosis, laminis profunde lobatis; inflorescentia corymbosa nmpb 
30 cm. longa vellongior; capitula numerosa, breves, corollis profunda 
5-lobatis, lohis saepe hasin versus connatis; achaenia oblonga, suh­
teretia; pappus sordidus. 
Typus : Caealia jatrophoides H.B.K. 
Perennial, lenfy-stemmed herhs to 3 m. tnll ; roots fibrous, fleshy; 
stems annual, erect, stou t, terete or less frequently angulate, gh,brouH 
or pubescent, not tuberclilate at ba~ej leaves cauline , not forming It 
basal rosette, alternMa, numerous, uniformly distributed along the 
stem, not clustered toward the middle, the uppermost not bracteiform; 
blades shaped variously, deeply pinnately subpalmately lohed; in­
florescence eorymhose, large, at least 30 em. long and hrulld, the 
branches sub tended by small, linear-suhulate or large, leRfy bracts; 
heads many, slllall (exccpt D. heteroidta), calyculate, the bracts 3- 7, 
u"lmlly shorter than the "hyllaries ; involucre campanulate or turbi­
nate, the phyllaries 5-8, biseriate appearing uniseri"te, imbricate, the 
overlapped margins sllbsenrio"s, the exposed Sllrface glabrous or 
pubescent, the tips frin ged ",ith minllte, Honglandlll"r trichome. "I' 
to 0.3 mm. long; reccp lncie flat, 01' nearly so, ulveolato , not paleneeolls; 
Oowers all dL,coid , pel'feet, ferlil e; corollus glahrolls, (·.reamy or nearly 
white, 00cnsionally greenish or purplish-tinted bllt ne\"er yello\l' , the 
, 
PIPPEN MEXICAN" CACALIOID" GENERA 379 
tube narrowly cylindrical, ca. 0.5 mm. in diameter, the limb divided 
into 5 equal, 3-nerved, narrowly triangular or linear, ascending or 
spreading lobes, the lobes free to the tube or connate into an ampliate 
throat up to 1.5 mm.long and broad; anthers 1.5-3 mm.long, exserted 
from the corolla throat, the bases rounded or slightly sagittate, the 
terminal appendages trullate or ovate, 0.5-0.7 mm. long; style 
branches 0.7-2 mm. long, slightly flattened, recurved or spreading, 
stigmatic over most of the abaxial surfaces, the adaxial surfaces 
glabrous or hispidulous below, the tips truncate, rounded or obtuse, 
fringed with unicellular, nonglandular "pollen producing" trichomes, 
unappendaged; achenes narrowly oblong, subterete in cross section, 
ribbed; pappus a single row of creamy white, stiff, barbellate, capil­
lary bristles. 
This genus differs from Psacalium and Odontotrichum in that the 
leaves are strictly cauline with no basal rosette formed and the 
&chenes are essentially cylindrical. It differs from Pericalia in that the 
leaves are not concentrated toward the middle of the stem but more 
uniformly distributed, there are no underground tubercules and the 
corolla lobes are not recurved. 
Key to Species of Digitarolia 
1. Heads large, 1.5-2 em. high at maturity; flowers 40 or more; corolla tube at 
least twice 88 long as the lobes. . . . . . . . . . . . 2. D. heteroidea 
1. Heads small, 1 em. high or less; flowers 13 or fewer; corolla tube and lobes 
about equal. 
2. Achenes pubescent; corollas short, 3-3.5 mm. long .. 1. D. chiapensIs 
2. Achenes glabrous; corollas 5-7 mm, long. 
3. Leaves 3-lobed, the lobes forming an angle, with the midrib, of less than 
45 degrees . . . . . . . . . . . . . . . . . . . 5. D. tridaetylitis 
3. Leaves 5- to 7-1obed, the lobes or at least some of the lobes forming an 
angle with the midrib greater than 45 degrees. 
4. Phyllarics and flowers 5 or 6 per head . . . . . . 4. D. napei(olia 
4. Phyllaries 7; flowers 8-10 per head . . . . . . 3. D. jatrophoides 
1. Digitacalia chiapensts (Hems1.) Pippen, comb. nov. 
Senecio chiapensis Hemsl. Bioi. Centro Amer. Bot. 2:238. 1881. Holotype: 
Chiapas & c., Ghiesbreght 537 (K, not seen; isotype Gil!). 
Cacalia chiapensis (Hemsl.) A. Gray, Proc. Amer. Acad. 19:53. 1883. 
Odontotrichum chiapenoi. (Hemsl.) Rydb. Bull. Torrey Bot. Club 51:418. 1924. 
Lower portion of plant not seen; stems terete, glabrous; leaves 
presumably all cauline, petiolate, oblong-ovate in outline, 10-14 CIll. 
long, 9-14 cm. "ide, pinnately veined; blades deeply pinnately 
5-lobed ca. two-thirds the distance to the midrib, the lobes ovate 
or oblong-acute, 4-6 cm. long, 2-3 cm. "ide, about half as wide as 
long, the lowermost pair of lobes rounded proximally or "ith two 
smaller deflec(,ed lobules, the margins entire, the abaxial smf"ce 
266-836--68 2 
380 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
puberulent mostly along the veins, the adaxial surface glabrous; 
petioles short, to 4 cm. long, glabrous, terete; inflorescence broadly 
coryrnbose-panicula te , to 30 em. long and broad, the branches gla­
brous, sub tended by sessile, ovate bracts; calyculate bracts ca. 3, 
subulate, 1.5-2 mm. long; heads many, 7-8 mm. long; phyllaries 
(6 -) 8, glabrous, narrowly ovate, 3.5-5 mm. long, [-1.5 mm. wide; 
flowers 9; corolla 3-3.5 mm. long, the tube 1.5-2 mm.long, the lobes 
linear or narrowly elliptic, 1.5 mm. long, spreading or reflexed, the 
throat none; anthers 1.5 mm. long; style branches 0.7-[ mm. long, 
the tips truncate or rounded with a terminal fringe of trichomes; 
mature achenes not seen, the immature achenes brown, sparsely 
pubescent; pappus bristles 2-3 mm. long, white. 
DISTRIBUTION: Knolm only from Chiapas. Apparently flowering 
in the fall but the exact time not known. 
This plant has been very poorly collected. It is known from only 
two collections in Chiaras, one is the type. 
CHIAPAS: Nelson 3467 (GIl, US). 
2. Digitacalia heteroidea (Klatt) Pippen, comb. nov. 
Senecio heleroideua Klatt, Leopoldina 24: 125. 1888. Holotype: Me.~ioo; be~ 
tweeD San ADdr~s and San ~ligucl, Liebmann 178 (e, not seen; tracing of 
type in Klat t's herb. Gil!). 
Cacalia laeviuata Schultz Bip. Leopoldina 24 : 125. 1888, in synonymy. 
Cacalia longipetiolo.ta Rob. & Grcenm. Amer. Journ. Sci. III. 50:157. 1895. 
Hoiotype: Oaxaca; Sierra de San Felipe, 7,000 ft., Oct. 10, 1894, C. C. 
Pringle 5828 (Gil!). 
Plants to ca. 2 m. (Pringle 6176) tall; lower portion of the plant 
not seen; stems glabrous, glaucous; leaves pubescent on both surfaces 
mostly along the veins, presumably all cauline, long-petioled, sub­
palmately veined with 5-7 principal veins, transversely broadly 
elliptic in outline, tho larger leaves 6-13 cm. long from the basal sinus 
to the apex, 9.5-22 cro. wide; blades cordate-based, deeply sub­
palmately 5- to 7-lobed, the lobes oblong or triangular, the larger 
ones 5-6 cm. long, 1-3 cm. broad, the margins toothed or lobulate; 
petioles glabrous, terete, 8-12 cm. long; inflorescence paniculate 
corymbose, the major branches sub tended by leafy, narrowly elliptic, 
petiolate bracts, 3-12 em. long, 0.5-3 em. broad, glabrous; pedicels 
5-[3 em. long; calyculate bracts 5- 7, subulate, 5-7 mm. long; heads 
12-40, 16-18 (-20) mm. long; phyllaries 7 or 8, ovate, 11-12 (-14) 
mm. long, 3-4 (-6) mm. wide, glabrous; receptacle ca. 7 mm. across; 
flowers ca. 40; corolla 9- 10.5 mm. long, the tube 5.5-6 mm. long, the 
throat short, campanulate, 1-1.5 mm.long, ca. 1.5 mm. across at the 
apex, the lobes 2-3 mm. long, narrowly triangular, 0.5 mm. wide, 
loosely reflexed; anthers 3 mm. long; style branches ca. 1 mm. long, 
• 
• 
, 
PIPPEN MEXICAN "CACALIOID" GENERA 381 
minutely hispid on the abaxial surface, the tips obtuse with a sparse, 
subterminal fringe of trichomes; mature achenes ellipsoid, 3.5-5.5 
mm. long, 1.5 mm. wide, 1 mm. thick, glabrous, brown, with 10-12 
prominent, paler ribs; pappus 7-8 mm. long, white. 
DISTRIBUTION: Known only from central Oaxaca. Presumably 
flowering in September or October. 
This species is placed in Digitacalia because its general habit is like 
that of the other species in this genus, although the inflorescence 
with few, large heads of 30 or more flowers is more like the species in 
Pericalia. It differs from Pericalia in the short corolla throat and the 
fewer calyculate bracts. D. heteroidea appears to be somewhat inter­
mediate between this genus and Pericalia. 
OAXACA: Conzatti 725 (GH): Pringle 6176 (GH, NY, US): C. L. Smith 287 
(MICH, NY, US). 
3. Digitacalia jatrophoides (H.B.K.) Pippen, comb. nov. 
Cacalia jalrophoide8 H.B.K. Nov. Gen. et Sp. 4:169. 1820. (Folio ed. 4:132. 
1820.) Holotype: Michoacauj nr Lake Cuitzeo, 900 hex'1 Humboldt & 
Bonpland 4298 (P, not seen; Field Mus. Neg. 37869!). 
Bem';o jalrophoide8 (H.B.K.) Schultz Bip. Flora 28:498. 1845. 
Cacalia digitala Sesse & Moe. PI. N. Hisp. ed. 1. 132. 1889. Lectotype 
without locality (presumably Hin Purulandiri montibus") Se88~ & Moe. 
2826 (MAl; Field Mus. Neg. 42288). 
Odontotrichum jatrophoide. (H.B.K.) Rydb. Bull. Torrey Bot. Club 51:419. 
1924. 
• Plants to 3 m. tall; stems to 1 cm. thick at the base, mostly hollow, 
often purplish, glabrous, glaucous; leaves sparsely pubescent on both 
surfaces mostly along the veins and margins, numerous, long-petiolate, 
subcircular in outline, 6-10 cm.long and broad, pinnate or sub palmately 
veined with 5-7 principal veins fused into a common trunk above the 
base of the blade; blades deeply 5- to 7-lobed, the lobes subpinnate or 
subpalmate, the central one larger, linear or narrowly ovate, 6-8 cm. 
long, 1-1.5 em. wide, the margin subentire or distantly toothed (lobes 
occasionally larger and the leaves similar to those of D. napeifolia); 
petioles terete, glabrous, purplish, to 9 cm. long; inflorescence broadly 
corymbose, loose spreading, usually more than 30 cm. long and broad, 
the branches sparsely pubescent or tomentose, the proximal branches 
in the axiJs of the 3-lobed upper leaves, the distal branches subtended 
by petiolate, narrowly ovate bracts to 2 cm. long; calyculate bracts 
3-7, subulate, 2-5 mm. long, heads many, 1-1.5 em. long; phyllaries 
7 or 8, narrowly ovate, 4-5.5 mm. long, 1-1.5 mm. wide, glabrous; 
flowers 8-10; corolla 6-8 mm. long, the tube greenish, 3-5 mm. long, 
the lobes 3 mm. long, 0.5 mm. wide, linear or narrowly elliptic, the 
throat none; anthers ca. 2 mm. long; style branches 1 mm. long, re­
curved, the tips rounded or obtuse with a subterminal fringe of tri­
chomes; mature achenes narrowly oblong appearing subterete in crOBB 
382 CONTRIBUTIONS FROM THE NATIONAl, HEHBARIUM 
section, 4-5 mIll. long, 1.5 llllll. wide, glahrou8, pale t.an, strungly 10· to 
12-ribbed; pappus 5~7 nUll. long, creumy, becOIHing tawny in age. 
DISTRIBUTION: Southern Zaeatecas and Jalisco, eastward through 
Michoacan and GUllnajuato and south to Oaxaca. Rocky places in 
highland pastures and grasslands. }'Io\\'ering in July and August. 
The specimens collected in Zacatecas and one from Oaxaca have 
much broader leaf lobes than the others. These leaf forms are similar 
to those of D. napeijolia DC., but these specimens are considered with 
D. jatrophoides bectLUse the floral characters are more like the latter. 
GUANAJUATO: Pringle 4262 (GH, NY, US). JALISCO: McVaugh 17160 (MICH). 
MICHOACAN: Arsene, s.n., Sept. 9, 1909 (US); Pippen 46 (MICH). OAXACA: L. C. 
Smith 123 (GH); C. L. Smith 388 (MICII, NY, US); Pringle 5741 (GH). ZACATE­
CA8: Rose 3040 (GII, US). 
4. Digitacalia napeifolia (DC.) Pippen, comb. nov. 
Cacalia napeaefolia DC. Prod. 6:328. 1838. Hoiotype: Oaxacaj summit of 
San Felipe, in 183-, Andrieux 280 (0, not seen; Field Mus. Neg. 33809!). 
Senecio napeaejoli-um (DC.) Schultz Bip. Flora 28:498. 1845. 
Odonlotrichum napeaejolium (DC.) Rydb. Bull. Torrey Bot. Club 51 :418. 1924. 
Plants to 2.5 m. tall (Pringle 4778); lower portion of the plant not 
seen; stems terete, purplish, sparsely pubescent; leaves presumably all 
cauline, numerous, petiolate, pinnately veined, oblong-ovate, 15-29 
em. long, 16-23 em. wide, progressively smaller upward; blades deeply 
5- to 7-lobed to within 0.5-1 rm. of the midrib, the lobes oblong­
attenuate, narrowly ovate, to 14 em. long, 4 cm. wide, the margins 
finely or coarsely toothed, the proximal pair of lobes usually deflected 
or at right angles to the midrib, the adaxial surface of the blnde 
glabrous or sparsely pubescent mostly along the veins, the abaxial 
surface pale, sparsely pubescent or tomentose; petioles subterete, 
glabrous below becoming pubescent toward the blnde; inflorescence 
('orymbose, to 45 em. long .llld broad, the branches tomentose, the 
proximal ones subtended by the uppermost le''''es, the distal branches 
HI.btended by linear-subulate bracts, 1.5 cm. long; clllyculate brads 
subulate, 1.5-3 mm.long; heads mllny, 1 cm. long; phyllaries 5 (rarely 
G), oblong or narrowly ovate, 3.5-5.5 nun. long, 1.5-2 mm. wide, 
glabrous or very sparsely puberulent; flowers 5 or 6; corolla 6-7 mm. 
long, the tube 3-3.5 mm. long, the lobes 3-4 mm. long, linear or 
narrowly elliptic, 0.5 rom. wide, the throat none; anthers 2-3 lUl11. 
long; style branches 1-2 mm. long, minutely hispid on the abaxial 
surface, the tips obtuse with a subterminal fringe of trichomes; mature 
nehenes narrowly oblong, appearing subtereie in cross seetion, 4-5 
mm. long, 1-1.5 mm. wide, glabrous, plU'plish brown, strongly 10-
ribbed; pappus 6-7 mm. long, creamy, becoming tawny in age. 
DISTRIBUTION: Known mostly from central OnxlLca with one col-
PIPPEN MEXICAN "CACALIOID" GENERA 383 
lection from Michoacan. Openings in oak-pine woodlands. Flower­
ing in September and October. 
MICHOAc1N: Hinlon el al. 12572 (MICH, NY). OAXACA: MeVaugh 21823 
(MICH); Nelson 1133 (GIl, US); Pringle 4778 (GH, MICH, NY, US). 
5. Digitacalia tridactylitis (Rob. & Greenm.) Pippen, comb. nov. PLATE 1 
Caealia tridaelylili. Rob. & Greenm. Amer. Journ. Sci. III. 50:159. 1895. 
Holotype: Oaxaca; Sierra de San Felipe, 6,000 ft., Nov. 19, 1894, C. G. 
Pringle 5841 (GH!). 
Odonlolriehum tridac!ylitis (Rob. & GreeDm.) Rydb. Bull. Torrey Bot. Club 
51:419. 1924. 
Plants to at least 2.5 m. tall; stems to slightly over 1 cm. thick at 
the base, angulate or terete, sparsely pubescent below becoming denser 
above; leaves cauline, numerous, pinnately veined, obtrullate or 
rhombic in outline, the larger ones 14-20 cm. long, from two-thirds 
as broad to as long as broad, sparsely pubescent adaxially, gray 
tomentose on the abaxial surface; blades deeply 3-lobed at least half 
way to the midrib, the lobes ovate, narrowly ovate or narrowly tri ­
angular, variable in size from 2- to 6-times longer than broad, the 
angle of the lateral lobes to the midrib less than 45 degrees, the margin 
of the lobes usually entire, occasionally toothed, the base of the blade 
decurrent along the petiole for a short distance; petioles 2-3.5 cm. 
long, triangular in cross section, pubescent; inflorescence corymbose, 
ca. 30 cm. long and broad, the branches tomentose, subtended by the 
uppermost cauline leaves; pedicels 3-7 mm. long; calyculate bracts 
~-7, subulate, 1.5-3.5 ffim. long; heads many, 9 mm. long; phyllaries 
7 or 8, narrowly ovate, 3.5-4 (-5) mm.long, 1-1.5 mm. wide, glabrous; 
flowers (7-) 8-10; corolla 5-6 mm.long, the tube 2.5-3 mm. long, the 
throat usually obsolete or very short, 0.5 mm. long and wide, occa­
sionally to 1.5 mm. long, ca. 1 mm. wide (Me Vaugh 21903), the lobes 
narrowly elliptic, 2-3 mm. long, 0.5 mm. wide; anthers 2-3 moo. long; 
style branches 1-1.5 mm. long, the tips rounded with subterminal 
fringe of trichomes; achenes narrowly oblong, appearing subterete or 
slightly compressed in cross section, 3.5-4 mm. long, ca. 1 moo. wide, 
glabrous, pale tan, 10- to 12-ribbed; pappus (3.5-) 4-5.5 moo. long, 
creamy, slightly clavate distally; chromosome number, n=30 (Turner, 
ined.). 
DISTRIBUTION: Morelos, Oaxaca, and Guerrero. Growing in open­
ings in the pine-oak zone. Flowering in October and November. 
The specimens from Morelos tend to have deeper, more narrowly 
lobed leaves and less pubescence on the abaxial surface than the other 
• 
specunens. 
GUERRERO: MeVaugh 21903' (MICH). MORELOS: Pringle 6164 (GH, MICH, 
NY, US); Prinyle 9877 (GH, MICH, NY, US). OAXACA: Nelson 2080 (US); 
C. L. Smith 380 (MICH, NY); D. C. Smith 966 (GH). 
384 CONTRmUTIONS FROM 1'HE NATIONAL HERBARIUM 
2. Odontotrichum 
Odontotrichum Zuee. Abhandl. Baier. Akad. WI ... 1: 311. 1832. 
Perennial, subscapose herbs, 0.5-2 m. tall; roots fleshy, fibrous, 
caudex tough, to 5 cm. long, 3 cm. thick, the nodes and terminal 
buds densely covered with tufts of tawny, long, multicellular hairs; 
stems usually solitary, annual, erect, slender, terete or less frequently 
angulate, glabrous or pubescent; radical leaves forming a basal 
rosette of two to several, the petiole attached to the base of the 
blade, not centrally peltate; blades pinnately or subpalmately veined, 
variously lobed or occasionally subentire, glabrous or pubescent; 
cauline leaves similar to the basal but progressively smaller upward, 
the uppermost bracteiform; inflorescence paniculate or corymbose 
often pubescent but not glandular, the branches sub tended by small, 
linear-subulate or large, leafy, often vaginate bracts; heads few to 
many, calyculate, the bracts usually 1-5, usually as long as or shorter 
than the phyllaries; involucre cylindrical, campanulate or turbinate, 
the phyllaries 5-17, imbricate, biseriat.e appearing uniseriate, the 
overlapped margins subscarious, the exposed surfaces glabrous or 
pubescent, the tips fringed with minute trichomes up to 0.3 mm. 
long; receptacle flat or nearly so, alveolate, occasionally sparsely 
pubescent, not palcaceous; lIowers nil discoid, perfect, fertile, 5-80 
per head; corollas glabrous, creamy or nearly white, occasionally 
greenish or purplish tinted but never yellow, the tube narrowly 
cylindrical, ca. 0.5 mm. in diameter, the limb divided into 5 equal, 
3-nerved, narrowly elliptic, narrowly triangular or linenr, ascending 
or spreading, not recurved lobes, the lobes usually free to the tube, 
occasionally connate into a short, ampliate throat up to 1 mm. long; 
anthers 2-4 mm. long, exserted above the corolla, the bases rounded 
or slightly sagittate, the terminal IIppendnges trullate or ovate, 0.5-
0.7 mm. long; style branches 0.7-2 mm. long, slightly lIattened, 
recurved or spreading, stigmatic on most of the abaxial surface, the 
adaxial surface glabrous or hispidulous below, the tips unappendaged, 
rounded or obtuse, fringed with minute, unicellular, nonglandular, 
"pollen producing" trichomes; achenes oblong, ellipsoid or obovoid, 
somewhat dorsally compressed, elliptic or broadly elliptic in cross 
section, glabrous or pubescent, striate or prominently 10- to 12-
ribbed; pappus a single row of white or creamy, still', barbellate, 
capillary bristles; chromosome numbers 25,30. 
TYPE-SPECIES: Odontofrichum cirsiiJolium ZUCCo 
This is the largest genus treated in this paper, with 21 species. 
It is distinguished from Psacalium by the Jack of centrally peJtate 
leaves, the blades being attached at their bases to the petiole. In 
most DC the species the leaves are vertically oriented, standing erect. 
, 
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 385 
Key to Specie. of Odonwtri.chum 
1. Pappus sbort, 2 mm. long or less. about balf 8B long 88 the corolla tube. 
2. Petioles flattened, winged, the margin up to 1 em. wide; basal leaves lobed 
about one-half the distance to the midrib, lobes usually longer than 
broad. . • • • • • • . . . . • • . . • • • • • 3. O. brachycomum 
2. Petioles 8ubterete, Dot winged; basal leaves coarsely toothed or lobulate, 
but not lobed, lobules broader tban long. • . • • . . • 13. O. palmeri 
1. Pappus at least 3 mm. long, as long 88 or longer than the corolla tube. 
3. Basal leaves toothed or 8ubentire, not lobed (80me species with two auriculate 
basal lobes formed by the deeply cordate basal sinus). 
4. Inflorescence very compact, globose, 2.5-3 em. long and broad; phyllaries 
5 or 6 per head. • • . . . . • . . . • • • • • . • 8. O. rlobosum 
4. Inflorescence tightly or loosely corymboee, not globose, at least 4 em. 
long and broad; phyllaries at least 7. 
5. Heads large with 30 or more flowers; phyllaries 13 or 14. 
6. Bracts 8ubtending the involucre oblong or obovatc, distally laciniate, 
3 mm. wide; corolla tube 9-10 mm. long, twice as long as the lobes 
(5-6 mm. long); phyllaries 12-15 mm.long ..•• I. O. amplum 
6. Bracts subtending tbe involucre linea.r-8ubulate, tapering to a point 
distally; corolla tube 5-6 mm. long, slightly longer than the lobes 
(3.5-4 mm. long); phyllaries 7-9 mm. long. 12. O. pa.hyphylium 
5. Heads small with 15 or fewer flowersj phyllaries 8 or fewer. 
7. Basal leaves deeply cordate-basedj petioles of the cauline leaves 
broadly leafy-margined, the base auriCUlate, clasping the stem; 
corolla tube 5-6 mm. long. . • • . . • • • 19. O. silphiifolium 
7. Basal leaves truDcate- or obtuse-based, rarely slightly cordate; 
petioles of the cauline leaves usually tcrete, the base dilated but 
not auriculate or clasping the stem (occasionally the petiole nar­
rowly leafy-margined or the leaves sessile); corolla tube 3-4 mm. 
long. 
8. Plants 0.6 m. tall or I ... ; basal leaves (including the petIole) 
II-I7 cm. long •.••...••..•••• 17. O. purpu8U 
8. Plants at least I m. tall; basal leaves (including the petiole) (15-) 
21}-39 cm. long. 
9. Phyllaries :> or 6; flowers 6 or 7; leaves sparsely puberulent. 
9b. O. ioldsmlthii var. rowelW 
9. Pbyllaries 7 or 8; flowers 9-13; leaves glabrous. 
9 •• O. goldsmlthll var. ,oldomlthll 
3. Basal leaves pinnately or subpalmately divided into several lobes, the lobee 
as long as broad or longer. 
10. Leaves deeply dissected or pinnatisect into linear segments, the ultimate 
segmenta linear or narrowly triangular, at least 6 times longer than 
broad. 
11. Median and dietal cauline leaves broadly auricul&te-beged, the 
margin laciniate; proximal inflolCscence bracts ovate, navicular, 
1.6-3 em. long, 2-3 em. broad, the margin distally laciniate; pappus 
deciduous before maturity of the achenes • • • • 5. O. eirsiifolium 
11. MedJan and distal cauline leaves often dilated at the base but not 
broadly auriculate~baaed or laciniate-marginedj proximal infiores~ 
ocnee br&ete linear 8ubulate, up to 1.6 cm. long, 1 mm. wide or len j 
pappus pera!otent. 
12. Achenes glabro .... 
386 CONTRIBUTIO:-iS FROM THE NATIONAL HERBARIUM 
13. LCl\\'cs 3- to 4-pillnllt.iecct.; hcad!:t with G-S flOWN6 aUtI [) or 6 
phyllarics ...... .. .. ... .. 6. O. dec:ompositum 
13. Leaves deeply dissected or pinnatifid with the primuy divisions 
again 3- to 5-1obed but not cleft to the midrib; heads with 10 
flowers and 8 phyllarics. . . . . . . . . 10. O. multilobum 
12. Achenes pubescent. 
14. Heads with 12-25 flowers; phyllaries sparsely pubescent. 
II. O. napelfilolium 
14. Heads with 10 or fewer flowers; phyllarics glabrous. 
15. Pappus 7 mm. long, about as long as the corolla (lobes plus 
tube) ; phyllaries 6 or 6. . . . . . . . . 7. O. filicifolium 
15. Pappus 3.&-4 mm. long, about as long as the tube of the corollaj 
phyllaries 7. . . . . . . . . . . . . . . 4. O. ceninum 
10. Leaves shallowly or deeply lobed but not 6ncly dis;cctcd or pinnatisect 
into linear segmeuts, the ultimate segments triangular or deltoid, 
usually less than 3 times longer thnn broad. 
16. lIeads large, 1.5-2.5 em. high with 40 or marc flowers; corolla tube 
7-10 mm. long, twice as long as the lobes. 
17. Brll.Cts subteuding the involucre leafy, oblong or ooovate, distally 
laciniate, 3-5 mm. wide; leaves and stem sparsely puberulellt; 
basal leavcs 18-75 em. long and broad. . . . 15. O. platyiepis 
17. Bracts Bubleudiug the involucre linear·subulatc, simplc, entire­
margined, 1 mm. wide or less, not leafy; h~a\'cs and stem densely 
covered by an interwoven mass of gray white, long trichomes; 
b!l8alleaves 14-16 em. long, cn. 10 em. wide. 
14. O. paucicapitatum 
16. lIeads small, 1- 1.2 cm. high or less; Howers 14 or fe\l"er ; corolla tube 
about equaling the lobes (ca. 5 mm, long). 
18. Achenes pubescent. 
19. Basal lca.ves Bubpeltate, the petiole attached to the blade 2-10 
(-30) mm. above the base, the blades 10-12 (-18) em. long. 
18. O. radulifolium 
19. Basal leaves attached at base to petiole, not peltate, the blades 
20-30 crn. long. 
20. Heads wit.h 9-14 flowcrs find 7 or 8 phyllarics; caulill(~ lca\'('s 
redu ced, bructcifol'lll, narrowly ovate, lllociniatc-Iou"d, 8 cm. 
long or h!S::J, 1-~ l'ln. broad, the basc not auriculat..c. 
16. O. pringlei 
:W. Heads with.5 or (j !lowel's and 5 phylbric'8; cauline leaves broadly 
ovate, lcafy, auriculate-based, 11-:20 cm. long, 11-20 em. 
broud, the ma.rgin d( ~ nt!~te or lobed . . 21. O. tU8sila&inoides 
J 8. AchelleB glabrous. 
21. Leaves shallowly lobed, the lobes 88 long as broad or broader 
(3-7 eln. broad), rounded at the tip, the margin regularly 
toothed; petiole of the cauline lt~nvj ~::I I(·afy.margined, broadly 
ovate, auriculate-based; lower infhm~scellcc bracts leafy, 4-U Olll. 
long, 1-3 cm. wide, ovate-naviculatc. , . . 2. O. amplifolium 
21. Leaves deeply lobed, the 10bf!::J longer tho.n broad (1-3 cm. broad), 
oblong or obovatc, each ()f t.he 101)('8 irrcp:ularly 3- to ;,)·Iobcd; 
petiole of the cauline lea.vC8 dilated or often c1uspillg the stem 
but not broadly leafy-margined or a.uriculate-bascd ; lower 
inflorescence bracts narrowly ovate or linear-8ubulate, 1-1.5 cm. 
long, 3 mm. wide. , . . . • . . . . . . . 20. O. slnuatum 
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 387 
1. Odontotrichum amplum Rydb. Bull. Torrey Bot. Club 51 :417. 1924. Holotype: 
Jalisco; road between Huejuquilla and Mesquitic [Mezquitic], Aug. 25, 
1897, J. N. Ros6 3575 (US!). 
Plants 0.7-1 m. tall, glabrous; stem usually one, 5-8 mm. thick at 
the base, terete, glaucous; basal leaves 2 or 3, coriaceous, long­
petioled, deltoid or transversely triangular, the apex rounded, 9--10 
cm. long, (ll-) 18-19 (-22) cm. wide; blades sagittate-based, coarsely 
dentate or lobulate, the teeth or lobules many, deltoid or 3-angled, 
5-8 (-12) mm. long, 8-10 (-15) mm. wide, ciliate, the abaxial and 
adaxial surfaces very similar; petioles to 28 (-40) cm. long, terete; 
cauline leaves 5-6, the lowermost one-half to two-thirds as large as 
the basal ones and similar to them, the petiole proximally dilated, 
clasping the stem; upper cauline leaves gradually smaller upward, 
the petioles broadly leafy-margined, the uppermost forming leafy 
inflorescence bracts; inflorescence corymbose, 9--23 cm. long, 15-20 
em. broad, glaucous; pedicels 2-6 cm. long; calyculate bracts 3-5, 
leafy, 1-1.5 cm. long, 3-6 mm. wide, distally 5- to 9-toothed; heads 
9-15, 18-25 mm. long; phyllaries 13 (-15), ovate or narrowly ovate, 
12-15 mm. long, 4-5.5 mm. wide, glabrous; receptacle 10--12 mm. 
across; flowers 40--50 or more; corolla 14-16 mm. long, the tube 
9--10 mm. long, Ca. 0.5 mm. wide, the lobes oblong, acute-tipped, 
5-6 mm. long, the throat none; anthers 3.5 mm. long; style branches 
1.5 mm. long, the tips truncate with a terminal fringe of trichomes; 
mature achenes ellipsoid, somewhat dorsally compressed, 6--7 mm. 
long, 2 mm. wide, ca. 1.5 mm. thick, glabrous, tan, 20- to 22-ribbed; 
pappus creamy, 8-11 mm.long, chromosome number, n=30 (Pippen). 
DISTRIBUTION: Western Aguascalientes and southwestern Zacate­
cas. Grazed grassland areas. Flowering in September. 
Known only from four collections. This species is very similar to 
O. pachyphyUum but differs in having larger heads with more flowers 
and leafy calyculate bracts. 
AGUASCALIENTES: McVaugh &; Kook 57 (MICH); Pippen 2, 3, 4 (MICH). 
ZACATECAS: McVaugh 17726 (MICH). 
2. Odontotrichum ampllfolium (DC.) Rydb. Bull. Torrey Bot. Club 51:416. 1924. 
Cacalia ampliJolia DC. Prod. 328. 1838. Holotype: Oaxaca; Sierra de San 
Felipe, on the summit, July 1832, G. Andrieux 293; (0, not seen; Field 
MUB. Neg. 338081). 
Senecio amplifoliu8 (DC.) Schultz Bip. Flofa 28:498. 1845. 
Senecio rumici/olius Klatt, Leopoldina 24:126. 1888. Holotypc: Mexico; 
Cumbre de Estepa, September 1842. Liebmann 237 (C, not seen; tracing of 
type in Klatt's Herb., CH!). 
Cacalia rumici/olia Schultz Bip. Leopoldina 24:126. 1888, in synonymy. 
Odonlotrichum Tumici/olium (Klatt) Rydb. Bull. Torrey Bot. Club 51 :416. 
1924. 
388 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Plants to 1.5 m. tall; stem 0.5--1 cm. thick at the base, terete 
arachnoid pubescent, the hairs multicellular, 0.5 mm. long, white, 
interwoven; leaves sparsely pubescent adaxially, densely pubescent 
abaxially; basal leaves 2-5, occasionally more, long-petioled, pin­
nately veined, ovate-cordate or oblong-cordate, (13-) 17-27 (-30) 
cm. long or longer, (12-) 13-19 (-24) em. wide or ,,;der, usually ca. 
two-thirds longer than wide; blades shallowly lobed, the lobes 12-16, 
rounded, usually broader than long, (1.5-) 3--4 (-5) em. long, (2.5-) 
3-5 (-7) cm. ,,~de, each lobe often shallowly 2- to 3-lobed or coarsely 
toothed; petioles usually longer than the blade, 20-35 cm. long; 
cauline leaves 3 or more, the lowermost similar to the basal ones 
but the petiole leafy-margined and more dilated and clasping at the 
base; upper cauline leaves progressively smaller upward, the upper­
most leaves bracteiform, forming leafy, ovate-naviculate inflorescence 
bracts, (3-) 4-9 em. long, 1-3 em. wide, distally toothed; inflorescence 
corymbose, (6-) 16-30 cm. long, 10-20 cm. broad, the branches 
tomentose; pedicels 0.5-1.5 cm. long; calyculate bracts 1-3, subulate, 
3-6 mm. long; heads 50 to over 100, 6-10 mm. long; phyllaries 7 or 8 
ovate or narrowly ovate, 4-5.5 mm. long, 1-2 mm. wide (specimens 
from Oaxaca with phyllaries 6-7 mm. long, 2-3 mm. wide), glabrous; 
flowers (8-) 10-13, corolla (5-) 6.5-7.5 mm. long, the tube (2.5-) 
3.5--4 (-5.5) mm. long, the lobes linear, (2-) 3-3.5 mm. long, the 
throat none; anthers 2-3 mm. long; style branches 1-1.5 mm. long, 
hispid on the abaxial surface, the tips truneate or obtuse ,,~th a 
terminal or subterminal fringe of trichomes; mature achenes ellipsoid 
or obovoid (often the marginal ones falcate), 5-5.5 mm. long, 2-3 
mm. wide, 1.5-2 mm. thick, glabrous, light brown, striate; pappus 
creamy, becoming fuscous with age, 4-6 mm. long. 
DISTRIBUTION: Oaxaca, Guerrero, and f\.1e:\,r]co. Grassy openings in 
the oak-pine-fir woodlands. Flowering from August to October. 
This species is closely related to O. 8ilphi~folium and o. goldsmithii. 
The specimens from Guerrero and Mexico differ from those in Oaxaca 
in that the size of the heads (phyJ\aries and flowers) tend to be smaller, 
the inflorescence more open and spreading, and the plants less 
pubescent. 
DISTRITO FEDERAL: Afatuda 26624 (Ny). GPERRERO: Sharp 441465 (NY) j 
Nelson 2243 (CH, US, this specimen has unusually large leaves); Rzedowski 1597·1, 
16015 (MICH). MEXICO: Maluda 27730, 29313, 29609 (Ny); H;nton, et al. 8349 
(MICH). OAXACA: Purpu, 3139 (GII, NY, US); Conzalti &: Gonzalez 397 (GH); 
L. C. Smith 124 (GH, this specimen has large cauline leaves and small inflorescence 
bracts); Nelson 1061 (US); C. L. Smith 391 (MICIl, US); MeVaugh 21824 (MICH). 
3. Odont.trichum brachyc.mum (Blake) Rydb. Dull. Torrey Dot. Club 51:415. 
1924. 
Cacalia brachycoma S. F. Blake, Contr. Gray Herb. II. 52:58. HH7. .Lectotype: 
Michoacan; wet ravines near Uruapan, 1,525 m., Nov. 13, 1905, C. G. 
Pringle 10126 (Gm). 
PIPPEN MEXICAN "CACALIOID" GENERA 389 
Plants 1-2 m. tall; stems terete, 7-9 mm. thick at the base, densely 
villous below, becoming subglabrous upward, the hairs 0.5-1 mm. 
long; leaves minutely puberulent mostly along the veins and margins 
with short, white hairs ca. 0.25 mm. long; basal leaves 3 or 4, cori­
aceous, long-petioled, mostly deltoid or ovate, 13-19 em. long or 
longer and broad, pinnately veined; blades sinuately lobed about 
one-half the distance to the midrib, the lobes 3-6 em. long, 2-3.5 cm. 
wide at the base, each distally divided into 3 lobes, the secondary 
obes deltoid, 3-angulate, the base of the blade truncate, decurrent 
along the entire length of the petiole; petiole as long as or longer than 
the blade, 13-19 cm. long, leafy-margined, the margin up to 5 mm. 
wide on each side; cauline leaves 3 or 4, the lowermost usually smaller 
than the basal ones but Rimilar (occasionally larger, more deeply 
dissected, up to 30 cm. long, 20 cm. wide), the petiole broadly leafy­
margined, ovate, dentate, clasping the stem; upper cauline leaves 
progressively smaller upward, the uppermost bracteiform forming 
ovate, naviculate inflorescence bracts, 5-8 cm. long, 3-5 em. wide; 
inflorescence corymbose, 13-28 cm. long and broad; pedicels to 1 
em. long; calyculate bracts 1-3, subulate, 1-2.5 em. long, 1-5 mm. 
wide; heads 50-100, 10-12 mm. long; phyllltries 9, narrowly ovate 
or oblong, 6-7 mm. long, 1.5-2 mm. wide, glabrous; flowers 10-12 
(-14); corolla 6-8 (-10) mm. long, the tube 3-4 mm. long, the lobes 
linear or narrowly elliptic, 3.5-4 (-6) mm. long, the throat none; 
anthers 2-3 mm. long; style branches 1 mm. long, the tips truncate 
with a terminal fringe of trichomes; mature achenes oblong or ellip­
soid, (3.5-) 4-6 mm. long, 1 mm. wide, glabrous, brown, 10- to 12-
ribbed; pappus 1.5-1.6 mm. long, about half as long as the tube of 
the corolla, creamy, the bristles unequal; chromosome number, n=30 
(Turner, ined.). 
DISTRIBUTION: Known only from the mountains of western central 
Michoacan. Grasslands, openings and wet ravines in the oak-pine­
woodland zone. Flowering in October. 
Blake cited both the Pringle 10126 collection at the Gray Herbarium 
and at the U.S. National Herbarium as the type. Pringle 10126 (GH) 
is chosen as the lectotype because it is annotated in what appears to be 
Blake's writing, and he was associated with the Gray Herbarium at the 
time this species was named. 
As the name suggests this species has a very short pappus. In 
Odontotrichum palmeri the pappus is as short as in O. brachycomum but 
O. palmeri differs in having fewer phyllaries and flowers per head and 
toothed, but not lobed, basal leaves. 
MICHOA"CAN: Hinton et al. 12146 (MleH, this specimen has an unusually large 
leaf, NY, US); Hinton et aI. 15288 (GH, NY, US); Hinton et aI. 15624 (GH, NY, 
TEX, US); King & Soderstrom 4904 (MICH); MeVaugh 21934, 21959 (MICH). 
390 CONTRIBUTIONS FROM ~'HE NATIONAL HERBARIUM 
4. Odontotrichum cervinum Hydb. Bull. Torrey Bot. Club 51:420. 1924. 11010-
type: Nayarit; between Pedro Paulo and San Bisscito, Aug. 4, 1897, 
J. N. Rose 1986 (U81). 
Plants ca. 0.5 m. tall; stem slender, 3-4 mm. thick at t he base, sub­
p:],lbrous proximally, becoming sparsely pube.'Cent to tomentose to 
densely tomentose upward , the hairs white, multiceUular; leaves glabrous, 
t he abaxial surface paler; basal leaves :1-5, long-petioled, rhombic or 
ovute or subeircular in outline, the lurger ones about. 6-9 cm. long and 
wide, pinnately veined; blades deeply pinnat.ely dissected into linear 
divisions, 3--4 mm. wide, t,ile mnjor divisions subdivided several times, 
the secondary divisions Illso subdivided; petioles 20-25 em. long, 
slender, glabrous, purplish; clluline leaves 3 or 4, the 100\'el'll1Ost simiIllr 
to the bltsul ones; upper cauline leaves reduced, bract.eirorm, the upper­
mo~t fonning inflorescellce bra.cts ; inflorescence corymbo~e , to 10 em. 
long lLnd broud, the hmnches tOlllentose; pedicels subtended by a lin­
ear-subulnte bmct to I CIlI. long; culyculate bracts linellr-subulate, 4-6 
111m. long; heuds Cll. 25 or fewer, 9-10 mm.long; phyllaries 7, oblong­
ovate, 5-6 mm. long, 1.5-2.5 IIlIll. wiue, glabrous; flowers 8-10; corolla 
7-8 mill. long, t he tube 3.5-4 mill., the lobes 3.5-4 mm. 10ng,Iinear or 
nurrowly elliplic, the throat essentially none; anthers dark brown or 
blnck, 3 mm. long; style branches 0.7-1 mill. long, the tips truncate 
wit.h u terminul fl'inge of trichomes ; mature nchenes not seen, the 
immature achenes greenish, sparsely pubescent with white hllirs; 
pnppus 3.5-4 lIlm. long, creamy. 
DISTUlBUTION: Known only from Nayarit. Grasslnnds and borders 
or fielus. Flowering in July. 
This taxon is known only from one collection besiues the type. It 
is unlL<u,,1 because of the small, almost subternate leaves. It is related 
to O. filiciJolium. 
NAYARIT: MeV.ugh 15313 (MICH). 
5. Odontotrichum cirsiifolium Zucco Abhandl. Baier. Akad. Wiss. 1:311. 1832. 
Holotype: A specimen of a garden plant grown from seeds SC!at by Ka.r~ 
winski from Mexico (M!; r"Hch. Neg. 1392). Not Cacalia cirsiifolia Hook. 
& Arn. 
Cacalia cervariifolia l"ccrvariac(olia" ) DC. Prod. G:328. 1838. Holotype: 
Oaxaca; SR.n Bartolo, Andrieux 8}>(..'Cimcn in the nco herb. CG, not Been; 
Field Mu,. Neg . 2888R'). 
Senecio cervariae/olius (DC.) Schultz nip. Flora 28:498. 1845. 
Sciadm~eri8 vaginala Kunze, Bot. Zei~ . 3411. 1851. Holotypc: specimen grown 
in the gardons at Herb. Bcrol. (not 8cen). 
Mesoneuris bipinnatijida A. Gray, Proe. Amer. Aead. 8:661. 1873. Holotype: 
Mountain region of Chiapas, Ghiesbreghl 805 (GIl!). 
Odontotrichum bipinnalijidum (A. Gray) Rydb. Bull. Torrey Bot . Club 51 :413. 
1924. 
PIPPEN-MEXICAN "CACALIOID" GENERA 391 
Plants to 1.5 m. tall; stem 4-7 mm. thick at the base, arachnoid­
pubescent or tomentose (often flocculent), the hairs white, to 2 mm. 
long or longer; leaves sparsely pubescent on the adaxial surface, 
densely pubescent on the abaxial surface; basal leaves 2 or 3, petiolate, 
elliptic or ovate in outline, 18-25 em. long, 14-22 cm. wide, usually 
2-5 (-10) cm. longer than wide; blades 2 or 3 times pinnatifid, the 
primary segments 17-21, narrow, opposite proximally becoming alter­
nate distally, 8-12 cm.long, ca. 0.5 cm. wide at the base, each divided 
into 5-9 lobes, the secondary lobes often lobed or toothed, the ulti­
mate segments narrowly triangular, ca. 1-2.5 cm. long, 1-3 (-7) mm. 
wide, the rachis narrowly leafy-margined, the margin ca. 1-3 mm. 
wide; petiole and midrib of the blade not clearly distinguished, the 
petiole 8-15 cm. long, often narrowly leafy-margined almost to the 
base, occasionally lobed; cauline leaves 2 or 3, the lowermost similar 
to the basal ones; upper cauline leaves progressively smaller upward, 
the bases broadly leafy-margined, anriculate and sheathing the stem, 
the blades pinnatisect or lobed; inflorescence corymbose, 20-30 cm. 
long, 10--20 cm. wide, the branches subtended by leafy, ovate­
naviculate bracts, 1.5-3 cm. long, 1-2 cm. wide, distally laciniate; 
pedicels 1.5-4 cm. long; calyculate bracts linear, 3-5, up to 1 cm. long; 
heads (10-) 30-40 (-50), 10-12 mm. long; phyllaries 11-13, narrowly 
ovate, (5-) 6--7 (-9) mm.long, 2-3 mm. wide, glabrous; receptacle ca. 
3 mm. across; flowers 20--30 (-40); corolla ca. 7 mm. long, the tube 
3-3.5 mm.long, the lobes narrowly oblong, (3.5-) 4 (-5) mm.long, the 
throat none; anthers 3 mm. long; style branches 1-1.5 mm. long, hispid 
on the abaxial surface, the tips truncate or rounded with a subterminal 
or terminal fringe of trichomes; matnre achenes obovoid, 5.5-6 mm. 
long, 2-3.5 mm. thick, glabrous (in specimens from Oaxaca and 
Chiapas) or pubescent with short, unicellular hairs (in specimens from 
Mexico, Guerrero, and Michoacan), light brown; pappus creamy or 
tawny, 3-3.5 mm. long, the bristles stiff, deciduous; chromosome 
number, n=30 (Pippen). 
D,STRIBUTION: Michoacan, Mexico, Guerrero, and east to Chiapas. 
Habitat variable: roadside ditches, grasslands, and openings in the 
oak-woodlands. Flowering in August and September. 
This species has two forms distinguished by the presence or absence 
of pubescence on the achenes and apparently by their somewhat 
distinct geographical ranges. The specimens from Oaxaca and Chispas 
have glabrous achenes whereas those specimens from the north and 
west of Oaxaca have pubescent achenes. Since no other correlative 
differences are evident I do not feel that these forms are worth nomen­
clatural status. 
392 CONTRIBUTIONS FROM THE NATIONAL H "RBARIUM 
CHIAPAS: Ghi •• breght 805 (NY, the leaves of this specimen are not as deeply 
dissected as the other specimens) j Nelson 3117 (GH, US). GUERRERO: Hinton 
el al. 9435 (MICH, NY). MhICO: Gilly et aI. 81 (MICH); Hinton.1 al. 8211 
(MICH, NY); Matuda 28821 (NY); Karwinski 8.n., September 1827 (M, probable 
parent of type of Odontotrichum cirsiiJolium Zucc.). MICHOAcA.N: Pringle 5272 
(GH); Pippen 52* (MICH). OAXACA: Pringle 4984 (GIl, NY, US); N.lson 1420 
(US); GaleoUi 2491 (US). 
6. Odontolrichum decompositum (A. Gray) Rydb. Bull. Torrey Bot. Club 
51:414. 1924. 
Cacalia decomposita A. Gray, PI. Wright. 2:99. 1853. Hoiotype: Sonors; 
mountains east of Santa Cruz, September 1851, Charles Wright 1286 (GRI). 
Senecio grayanus Hemsl. BioI. Centro Amer. Bot. 2:241. 1881. Name 
illegitimate. 
Me.adenia decomposita (A. Gray) Staodley, Contr. U.S. Nat. Herb. 19:749. 
1915. 
Plants up to 1-1.5 m. tall; stem slender, 3-5 mm. thick at the 
base, terete, densely pubescent below the lowest leaf with long, 
multicellular hairs in flocculent patches, becoming subglabrous or 
sparsely pnbescent "bove the lowest laM; leaves mostly glabrous or 
sparsely pubescent, the hairs multicellular, 0.4-1 mm. long, mostly 
along the veins and margins; basal leaves 2-4, long-petioled, pin­
nately veined, ovate or elliptic in outline, 20-30 cm. long, 12-27 cm. 
wide, about as wide as long or usu"lly 2-5 cm. longer th"n wide; 
blades 3 or 4 times pinnatisect, the primary divisions 10-15 (-17), 
opposite below, becoming alternate dist"lly, each 2 or 3 times pin­
natisect, the ultimate segments narrowly triangular or narrowly 
oblong and acute-tipped or falcate, 0 .. 5-2 cm. long, 2-5 mm. wide, 
the laminar portion of the blade reduced to a narrow margin ca. 1 
mm. wide on each side of the main veins; petioles subterete or tri ­
angular in cross section, 17-25 cm. long, pubescent below becoming 
subglabrous or sparsely pubescent distally; cauline leaves 3 or 4, the 
lowermost one-half as large or as large as the basal ones and similar; 
upper cauline leaves reduced upward, bracteiform, 4-9 cm. long, 
1-4 cm. \\;de, 1-3 times pinnatisect into linear segments, the upper­
most cauline leaf bracts 1-6 cm. long, 5 mm. wide; inflorescence 
paniculate or corymbose, 11-20 cm. long, 12-22 cm. broad, the 
branches densely pubescent, the major branches subtended by nar­
rowly ovate or linear-subulate bracts ca. 1.5 cm. long; pedicels 1-1.5 
mm. long; calyculate bmcts 1-3, 2-4 mm. long, linear-subulate; 
heads ca. 100 or fewer, 10-12 mm. long; phyllaries 5 or 6 (rarely 8), 
narrowly ovate, obovate or oblong-acute, (4-) 5-6 (-7) mm. long, 
(1-) 1.5-2 mm. wide, mostly glabrous or sparsely pubescent; recep­
tacle 1.5 mm. across; flowers (5-) 6-7 (-8); coroll" 7-8 (-9) mm. 
long, the tube 3.5-4 (-5) mm. long, the lobes linear-acute, 3.5-4 mm. 
long, 0.5 mm. wide, the throat none; anthers 2-3 mm. long; style 
branches 1-1.5 mm. long, the tips truncate or obtuse with a terminal 
PIPPEN-MEXICAN "CACALIOlD" GENERA 393 
fringe of trichomes (or rarely a cluster of trichomes covering the 
entire tip); mature achenes ellipsoid, often falcate, 4-5 mm. long, 
1.5-2 mm. wide, 1 mm. thick, glabrous, dark brown, contrasted 
with paler ribs, the ribs 15-18; pappus white or creamy, (4-) 5.5-6 
(-7) mm. long; chromosome number, n=30 (Kruckeberg in Ornduff 
et aI., 1963). 
DISTRIBUTION: Northwestern Durango, northward into Sonora, 
Chihuahua, and Arizona. Its southern limit appears to be northern 
Durango as it has not been found farther south. Grasslands, pastures, 
and borders of oak-pine woodlands. Flowering in August and 
September. 
This species is unusual because of its several times pinnatisect 
leaves which have the appearance of being compounded. Of the 
species of Odontotrichum growing in Mexico this one is the most 
northern in range, extending into southern Arizona. 
Odontotrichum decompositum, or "lvlatarique," as it is called by 
the natives, is the only species of Odontotrichum of known economic 
importance. It is used medicinally in the treatment of diabetes and 
other ailments (Martinez, 1959). 
ARIZONA: Barkley 14A624 (TEX); DaTTow et 01. 1235 (NY); Eggleston 10765 
(US); Blumer 1378 (GH, NY, US); Kruckeberg 4644 (MICH); Darrow, s.n., 
July 4, 1937 (NY); Harrison & Kearn'y 6194 (US); 6045 (GH); Kearney & 
Peebles 10092 (US); Goodding, s.n., Aug. 28, 1912 (NY); Goodding 784 (GH, NY); 
Jane. 25053 (GH, NY); Palmer s.n., July 1890 (GH, US); Wilcox s.n., in 1891 
(US). CHIHUAHUA: Gentry 1959, 2815 (GH, US); Hewitt 68 (GH); Knobloch 
1293 (MICH); Pringle 767 (GH, MICH, NY, US); Mearns 2219 (NY, US), 
527, 531 (US, appear to be stunted plants, collected near the summit); MuUer 
3420 (Gil); Nelson 6095 (GH, US); Townsend & Barber 158 (GH, NY, US); 
flarde LeSueur 145 (GH, TEX); Tucker 2456 (US); Townsend s.n., in 1909 (US); 
Gentry 536M (MICH). DURANGO: Nel.on 4758 (NY, US). SONORA: Phillips 
651 (GH, MICH); White 2741, 3289, 4353 (GH MICH). 
7. Odontotrichum filicifolium Rydb. Bull. Torrey Bot. Club 51 :414. 1924. 
Holotype: Nayarit; between Santa Gertrudis and Santa Teresa, Aug. 8, 
1897, J. N. Rose 2101 (USI). 
Plants up to at least 0.5 m. tall; stem glabrous proximally, sparsely 
pubescent to tomentose distally; leaves glabrous; basal leaves 2-4, 
long-petioled, pinnately veined, deltoid or ovate-cordate in outline, 
4-5 cm. long and broad; blades deeply and finely dissected, the major 
divisions several, each divided into smaller segments or lobes, the 
ultimate segments narrow, linear-attenuate, laminar portion of the 
blade only a narrow margin, ca. 3-5 mm. wide on each side of the 
major veins; petioles slender, 11-12 em. long; cauline leaves 3 or 4, 
the lowermost smaller than the basal ones but similar; upper cauline 
leaves bracteiform, dilated at the base, distally laciniate; inflorescence 
eorymbose, the branches tomentose, sub tended by narrowly ovate 
394 CONTRIBUTIONS FIlOM TH~; NATIONAL HERBARIUM 
brads; ealY(,lIln.t..e bracts 2 or ;J, lineur-suhulll.t.e, 2-4 111111. lung; 
heads 15~25, "hout 10 III Ill. long; pbyl]';rie, 5 or 6, IllllTowly u"ute, 
6-7 mill. long, 2 Ullll. wide, glabrous; flowers 6~8; cornlla cn.. 7.5 mm. 
long, the tube 4 mill. long, the lobes ~l.5 mm. long, the throltt none; 
anthers ell. 3 mill. long; style bmnches O.7~1 Illlll. long, hispid (Ill 
the llbaxilll surface, the tips obtuse with . u\)termillul Cringe of tri­
cbomes; mllture IlcilenM nut seen, tile illlmature achenes densely villous; 
pappus fuscous, 7 mm. long. 
DISTRIBUTION: Known only from tbe type specimen. 
Tbis taxon is quite similar to Odontotrichum cermnum and WliS 
collected in tbe s,une region. It differs from O. cermnum in baving 
smltller bends and the larger leavC8. Field observation. Illld study of 
future collections Illl'y indicate that tbese two tam should be cun­
. idered as one species. 
8. Odontotriehum ilobosum (Rob. & Fern.) Rydb. Bull. Torrey Bot. Club 51:418. 
1921. 
Caralia globoSG Rob. & Fern. Proc. Amer. Acad. 30:119. 180'i. Hoiotype: 
Chihuahua; in & moist meadow, Gus.chuchic, JUlie 25, 1892, Hartman 52:2 
(GII!). 
Odontotrichum $cabrum Rydb. Uull. Torrey Bot. CLub 51:416. 1024 . 
CacaJia pTaknsis Staodl. Pub!. Field !\:Ius. Bot. 22:61. 1940. Holotype: 
Chihuahua; wet meadows ncar San Juanito, 2,430 m., July 26, 1937, F. 
Shreve 8031 (Fl, isotype MICHl) . 
Odontotrichwn pralense (StandI.) J. CU!l.tr . llrittonia 8:157. J955. 
Plants 0.5~0.7 (~1.2) m. tall; stem "lender, 5 min. thick or less at 
tbe base, mostly subglabrous below becoming sparsely pubescent 10 
araehnoid-tomentose above; leaves glabrous or very sparsely puber­
ulent mostly along tbe veins; b'1SI11Ieaves 2-4, long-petioled, pinnately 
veined, cordate or cordate-ovate, 4~7 (~8) CIlI. lung and broad; blades 
shallowly dentate or entire, the teet.h rounded or triangular, the 
IlIllrgin of the blade ciliate; petioles glabrous, slender, 12~21 (~:JO) 
CIIl. long; cauline leaves 3 or 4, tile lowermost similar to and as large 
as the basal ones; upper cauline leaves progressively smaller upward, 
the upperlJlost braeteiform, leafy, ovate, clasping the stem; inflores­
cence globose, very compact, 2.5~3 em. long and broad, the branches 
arachnoid-tomentose; pedicels 5 111111. long or less; calyculate bracts 
1- 5, Ilarrowly ovate or linen,r-suhulatc, 4-5 mm. long, I mm . wide or 
less; heads about 20, 7~10 mm. long ; phyllaries 5 or 6, narrowly ovate 
or o\rate, 5-7 mm. long, 1.5-2 mm. wide, glabrous; flower:i (4-) fj~6 
(~8); eorolla 6~7 (~ IO) mm. long, tbe tube 3~3.5 (~5) mill. long, the 
lobes 3~3.5 (~5) 1ll1II. long, 0.5 111m. wide; anthers 2~3 (~3.5) mIll. 
long; style branches ca. 1 mm. long, the tips truncn.te or obtuse with 
" Ruhterminal fringe of tricbomes; mature achenes pale brown, gla­
brous; papllUS creamy, 3~3.5 (~5) mm. long. 
, 
, 
PIPPEN-MEXICAN "CACALIOID" GENERA 395 
DISTRIBUTION: Chihuahua and Durango. Wet marshy .. reas in 
pastured grasslands. Flowering in July and August. 
This species is unique because of its habitat-most of the other 
species in this genus grow in dryer situations-and the very tightly 
globose inflorescence. 
Comparison of the type of Oacalia pratensis StandI. with tbe type 
of Odontotrichum globosum indicates that they are definitely con­
specipc because they are similar in all respects. 
Evidence of natural hybridization between O. globosum and O. 
llinuatum was observed by Dr. McVaugh in southern Durango in 1962 
and a sampling of these populations was made (voucher specimens, 
Mcraugh 21732-21739, MICH). The hybrid population sbowed 
various degrees of ecological and morphological intergradation be­
tween the two very dissimilar parents. 
Odontotrichum 8cabrum, described by Rydberg from a specimen 
collected in 1897 (Rose 3471, US), is one of these intermediate bybrid 
forms, resembling O. globo8um in leaf shape but with a more open 
inflorescence like O. llinuatum. It is not distinct enough to deserve 
nomenclatural status. 
CUIHUAHUA: Knobloch 1273 (MICH); Shrl'llc 8031 (US); Gentry 2785 (MICII, 
OH, isoparatype of C. prat,"';. Stand!.). DURANOO: Graham 99 (MICH); 
MeVaugh 21732, 21734-21739 (MICH, intermediate forms between O. globo.um 
nnd O .• inuatum); Penn,U 18532 (NY); Ne!.on 4769 (NY); Rose 3471 (US, ho10-
typo of O. Bcabrum Rydb.). 
9. Odonlolrichum go!dsmlthll (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51:416. 
1924. 
Cacalia goldsmithii B. L. Rob. Proc. Amer. Acad. 43:45. 1907. 
Slender plants to I m. tall; stems up to 5 mm. thick at the base, 
mostly glabrous or sparsely pubescent above; leaves mostly glabrous 
Or sparsely puberulent a.long the veins and margins; basa.l leaves 
2 or 3, erect, long-petioled, pinnately veined, variable in shape from 
ovate to broadly ovate, 7-14 (-19) cm. long, 6-12 (-16) cm. wide, 
as long as wide to twice as long as wide; blades mostly truncate or 
obtusely based, occasionally slightly cordate, the margins sinuately 
double-<lentate or lobulate, the teeth deltoid-rounded, in two a.lter­
nating series of large and small respectively; petioles &--14 (-20) cm. 
long, slender; cauline leaves ca. 3, the lowermost usually elliptic, 
as large as or one-half as large as the basal ones and similar; upper 
cauline leaves greatly reduced, the uppermost bracteiform, leafy, 
up to 6 em. long, 0.1>-1 cm. wide, the base of the petiole dilated; 
inflorescence loosely corymbose, 18-23 em. long, 9-11 cm. broad, 
the branches densely pubescent, subtended by subulate or narrowly 
ovate bracts 1-3 cm. long, 1-5 mm. wide, smaller distally; pedicels 
1>-15 mm. long; calyculate bracts 3 or more, subulate, 1.5 4 mm. 
288-836 68 3 
396 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
long; heads 50-75, 10-11 mm. long; pbyllaries (5-) 7 or 8, usually 
ovate-oblong or narrowly ovate, 5-6 (-7) mm. long, (1-) 2-3 mm. 
wide, glabrous; receptacie np to 1.5 mm. across; flowers 7-13; corolla 
mostly 6-8 mm. long, tbe tnbe 3-4 mm., tbe lobes 3-4 (-7, Hinton 
9202) mm. long, narrowly elliptic, strongly 3- (-5-) nerved; anthers 
2-3 mm. long; style brancbes 1 mm. long, usually bispid on 
the abaxial surface, the tips truncate witb a terminal fringe of tri ­
cbomes; mature acbenes ellipsoid, 4-5 mm. long, 2 mm. wide, ca. 1 
mm. or less thick, brown, glabrous, 10- to 12-ribbed; pappus creamy 
wbite, becoming fuscous in age, 3.5-5 (-7) mm. long. 
DISTRIBUTION: Eastern Jalisco to Michol1Cftn and western Guerrero 
and Mexico. Grasslands and pastures. Flowering in July (and 
August?). 
Tbis species is very closely related to O. silphiijolium, and further 
field and garden study may indicate tbat tbey should be considered 
as tbe same species. In tbe present treatment, bowever, they are 
recognized as two species with O. goldsmithii differing from O. siI.­
phiijolium in its slender, mostly glabrous habit, open, loose inflores­
cence with shorter heads, truncate or obtuse leaf bases, and smaller 
cauline leaves that are not auriculate-based. 
Key to Varieties of O. golcl<milhii 
a. Heads with 5 or 6 phyllaries, 6 or 7 flowers; leaves sparsely puberulcnt. 
9b. O. loldsmithii var. rowe1lii 
A. Heads with 7 or 8 pbyllaries, 9-13 flowersj leaves glabrous. 
9a. O. ,oldsmithii var. Itoldsmithii 
9 •. Odontotrichum loldsmithii var. eoldsmithii 
Cacalia gold3mithii B. L. Rob. as to Holotype: Jalisco; Hacienda San Marcos, 
350 m., July 12, 1905, P. Goldsmith 8 (GRI). 
PhylJ",ies mostly 8; flowers 9-13 per head; basal leaves ovate or 
elliptic, glabrous, the blndes truncate or obtuse-b,~<ed, rarely cordate. 
DISTRIBUTION: J alisco, MicboRclin, Guerrero (Mina Distr.), and 
Mexico. 
GUERRERO: Hinton d al. 9202 (MICH, NY, this specimen has larger heads 
than the other specimens). JALISCO: McVaugh 15460J 16132 (MICIl). Mi:xICO: 
Hinton 1266 (MICH); Hinton et al. 8066, 7984 (MICH, NY). MICROACAN: 
Hinton et al. 5078 (GH, NY, US). 
9b. Odontotrichum goldsmithii var. rowellii Pippen var. nov. ])LATE 2 
A varietas goldsmithii floribus phyllariisque paucis, foliis radicalibus, 
puberulibus differt. 
Heads with 5 or 6 phyllaries, 6 or 7 (Jowers; basal leaves ovate or 
narrowly ovate, the blades obtuse or cordate-based, sparsely 
puberulent. 
PIPPEN MEXICAN" CACALIOID" GENERA 397 
HOLOTYPE: Guerrero; 1 mi. N. of Agua del Obispo, 3,300 ft., July 
1, 1952, O. M. Rowell, Jr. 29~1 (MICH!). 
DISTRIBUTION: Knowu only from central Guerrero. 
Thls variety is distinguished from the other in the smaller heads 
with fewer phyllaries and flowers and the somewhat puberulent leaves. 
GUlORRIlRO: Wilkin80n &: Rowell 8381 (MICHl; Rhy .... &: Rowell8BB6 (MICHl; 
Hicks &: Rowell 8604 (MICHl; Rzedow.ki 16078 (MICHl; 8horp 441481 (NY). 
10. Od.ntolrichum muItil.bum Pippen, sp. nov. PLATE 8 
Plantae tenues ca. 1 m. altae, caule foliisque glabris vel sparse 
pubescentibus, foliis radicalibus erectis, latis ellipticis 18-30 cm. longis 
10-17 cm. latis, profunde pinnatifidis, segmentis principalibus 12-15 
linearibus; petiola tenus glabra 10-30 cm. longa; folia culaine bractei, 
forma 1-2; inflorescentia corymbosa, 16-30 cm. long, 11-16 cm. lata­
bracteis 10-20 mm. longis, linearis vel anguste triangularis; capitula 
50-100, ca. 12 mm.longa; phyllaria 8, 5-6 (-7) mm.longa, 1.5-2 mm. 
lata, anguste ovata, glabra; flores 10, corollis 7 mm.longis, tubo lobis­
que 3.5 mm. longis, achaenis ellipsoidea vel obovata, (3-) 3.5-4 mm. 
longa, 1.5-2 mm. lata, glabra, pallide fulva; pappus sordidus (5-) 6 
(-7) mm. longus. 
Plants ca. 1 m. tall; stem and leaves glabrous or sparsely pubescent, 
becoming sparsely or densely pubescent in the inflorescence; flowering 
stem 4-6 mm. thick at the base; basal leaves 3 or 4, long-petioled, 
erect, pinnately veined, broadly elliptic in outline, 18-30 cm. long or 
longer, 10-17 cm. broad; blade deeply pinnatifid, the laminar portion 
narrow, ca. 3-6 mm. wide on each side of the vein, the primary lobes 
12-15, linear, 7-11 cm. long, each divided into 3-5 major lobes, the 
secondary lobes either 1- to 2-lobed or coarsely toothed, the ultimate 
segments linear-acute; angle of the lobes with the midrib less than 90 
degrees; petioles ca. 10-30 cm. long, mostly glabrous, triangular in 
cross section; cauline leaves 1 or 2, similar to the basal leaves but 
much smaller; inflorescence loosely corymbose, 16-30 cm.long, 11-16 
em. broad, the branches subtended by linear or narrowly triangular bracts 
10-20 mm. long, 1-4 wide; calyculate bracts 3-5, subulate, 3-5 mm. 
long; heads 50-100, ca. 12 mm. long; phyllaries 8, narrowly ovate, 
5-6 (-7) mm. long, 1.5-2 mm. wide, glabrous; receptacle 2 mm. 
across; flowers 10; corolla 7 mm. long, the lobes as long as the tube, 
3.5 mm. long, linear, the throat none; anthers 3 mm. long; style 
branches 1 mm. long, the tips truncate or rounded with a subterminal 
fringe of trichomes; mature achenes (3-) 3.5-4 mm. long, ellipsoid or 
obovoid, 1.5-2 mm. wide, 1 mm. thick, glabrous, light brown, ca. 
20-striate; pappus (5-) 6 (-7) mm. long, creamy or tawny; chromo­
some nnmber, n=30 (Pippen). 
398 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
HOLOTYPE: Jalisco; ca. 15 mi. SSE. of Autlan by way of Chante, 
oak-dominated slopes E. of Mamantian, ca. 4,500 ft., Aug. 1, 1949, 
R. L. &: C. R. Wilbur 2094 (MICH!). 
The type specimen is somewhat smaller in st,ature than the other 
specimens of the same taxon, but it is representative. 
This species is related to O. pringlei by the shape of the leaf, but 
differs in that the achenes are glabrous; and to O. sinuatum, differing 
in the more cylindrical heads, ratber than turbinate, and in the lower 
pair of lobes on the leaves not deflected. 
This species is sympatric with O. prinUlei and O. palmeri, possibly 
intergrading with the latter. Two specimens from western Jalisco 
(Pippen 29, sheets 1 and 2, MICH) are intermediate between O. 
multilobum and O. palmeri, suggesting that hybridization may be 
occurring. At this locality both O. multilobum and O. palmeri as well 
as the internlediate forms were growing together, the latter plants 
mistaken for a third species at the time of collection. Future field 
studies and examination of additional material are necessary to 
determine if integration has occurred. 
JALIBCO: Pippen 25, 30*, 29 (possible hybrid between O. multilobum and O. 
palm",;), 40 (MICH). 
11. Odontotrithum napellifolium (S. Schauer) Rydb. Bull. Torrey Bot. Club. 
51:419. 1924. 
Cacalia napellifolia S. Schauer, Linnaea 19:732. 1847. Holotype: Mexico; 
about 1840, Aschenborn 312 (ll, not seenj tracing or type in Klatt's Herb., 
OHl). 
Plants 0.5-0.6 m. tall; stem slender, 2-4 mm. thick at the base, 
densely pubescent below, becoming sparsely pubescent upward; 
leaves mostly glabrous, the margin ciliate; basal leaves 4-7, long­
pet.ioled, subpalmately veined with 5-7 principal veins, subcircular 
in outline, the base cord"te, 2.5-5 em. long from tbe basal sinus to 
the tip, :\-8 cm. wide; blnde, deeply lobed, the primary lobes 3-5 
(-7), each deeply divided into 2 or 3 (-4) lobes, the secondary lobes 
again divided into 3-5 lobules, the ultimate segments narrowly tri­
angular or deJt.oid, 3-10 mm. long, the proximal pair of lohes de­
flected, parallel t.o t.he petiole, t.he base of the blade decurrent on the 
petiole for a short, distance; pet.ioles 7-15 em. long, slender; c"uline 
leaves 1 or 2, t.he lowermost similar to the basal ones; upper cauline 
leaves bracteiform, reduced, 2-3 cm. long, 0.5-1 em. wide, 5- to 7-
lobed; inflorescence corymbose, the branches tomentose, the pedicel. 
0.5-3 cm. long; calyculate bracts about 3, subHla!e, ca. 4 mm. long; 
heads 5-8, 10-14 mm. long; phyllaries 10, narrowly ovate, 5-6 mm. 
long, 1.5-2 mm. wide, sparsely puhescent" the t.ips purplish; flowers 
12-25; corolla 6-7 mm. long, the tube 3-3.5 mill. long; anthers 2.4-3 
mm. long; .t,yle branches 0.7-1 mm. long, hispid on t.he abaxial 
PIPPEN-MEXICAN "CACALIOID" GENERA 399 
surface, the tips truncate or obtuse with a subterminal fringe of 
trichomes; mature achenes ellipsoid, 5 mm. long, 1.5 mm. wide, 
1 mm. thick, pale tan or creamy, 10- to 12-ribbed, the ribs pubescent 
with ascending hairs; pappus creamy, 3-5 mm. long. 
DISTRIBUTION: Guerrero and Hidalgo. Grasslands and openings 
in oak-pine woodlands. Flowering in June and July. 
This taxon is distinct because of the small, deeply lobed basal 
leaves. It is closely related to O.fili.cijolium and O. cervinum but differs 
from both in the larger number of flowers per head. 
GUERRERO: Kubicek & Rowell 3779 (MICH); Rhymes & Rowell 3896 (MICH). 
HIDALGO: Pringle 10024 (MICH, NY, TEX, US); Salazar s.n., Aug. 1, 1913 (US). 
12. Odontotriehum paehyphyllum (Schultz Bip.) Rydb. Bull. Torrey Bot. Club 
51:417. 1924. 
Cacalia pachyphylla Schultz Bip. ex Seem. Bot. Yay. Herald. 310. 1856. 
Hoiotype: Northwestern Mexico; Seemann 1999 (K, not seen; MICH Neg. 
6221). 
Senecio 8clerophyllu8 Hemsl. BioI. Centro Amer. Bot. 2:247. 1881. Based on 
Cacalia pachyphylla Schultz Bip. not S. pachyphyllo8 Remy. 
Plants 0.5-0.7 m. tall; stem 3-4 mm. thick at the base, pubescent, 
the hairs white, multicellular, often flocculent; leaves mostly glabrous 
or sparsely pubescent along the veins; basal leaves 2 or 3, coriaceous, 
long-petioled, reniform or rounded-deltoid, rarely subcircular in out­
line (6-) 8-10 (-ll) em. long from the basal sinus to the apex, (10-) 
13-18 (-20) em. wide, the tip broadly rounded, the base truncate or 
usually shallowly cordate; blades variable, shallowly repandly dentate 
or subentire, the teeth rounded or obtuse, the adaxial and abaxial 
surfaces very similar; petioles (4-) 8-18 (-35) em. long; cauline leaves 
2 or more, the lowermost smaller than the basal ones but similar; 
upper cauline leaves becoming bracteiform upward with the petiole 
broadly leafy-margined, clasping the stem; inflorescence corymbose, 
7-20 em. long, 7-15 em. broad, the branches tomentose or \~llous 
(glabrous in the type and isotype); pedicels 2-6 em. long; calyculate 
bracts 3-5, subulate, 9-13 mm. long, 1 mm. wide, sparsely pubescent; 
heads 9-15, 1-1.5 em. long; phyllaries 13 or 14, narrowly ovate, 7-9 
mm. long, 1-3 mm. wide, glabrous; flowers 30-35; corolla (7.5-) 8-10 
mm. long, the tube (4-) 5-6 mm. long, the throat short, ampliate, up 
to 0.5 mm. long, 1 mm. wide at the apex, occasionally lacking, the 
lobes 3.5 4 mm. long; anthers 2-3 mm. long; style branches 1.5 mm. 
long, the tips truncate or obtuse with a terminal or subterminal fringe 
of trichomes; mature achenes ellipsoid or oblong, 4-5 mm.long, 2 mm. 
wide, 1.5 mm. thick, glabrous, brownish, 15- to 16-ribbed; pappus 
white or tawny, 6-8 mm. long. 
DISTRIBUTION: Known only from southern Durango. Openings in 
pine-oak woodlands. Flowering in August and September. 
400 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
The type is mostly glabrous with linear inflorescence bracts and 
large basal leaves with acute basal sinuses. These characteristics are 
somewhat different from the other specimens studied but the speci­
mens are all similar enough in size of heads, nllm bel' and size of 
flowers, texture and general shape of the leaves to be considered 
part of the same taxon. 
This species is closely related to O. amplum, differing from it in 
having smaller heads with narrower inflorescence bracts and basal 
leaves rounded at the base. 
DURANGO: Gentry 6969 (MICH, US); MeVaugh 21659 (MICH); P,nnelll8332 
(NY); lVaterfall12668 (MICH); MeVaugh 21731 (shows possible intergradation 
with O. globo.um), 21729 (MICH); Dumbey &; Ownbey 1874 (MICH, U8); May,iU" 
7784 (MICH). LOCAUTY UNKNOWN: Seemann 8.D. (probably isotype of Seemann. 
1999, G11). 
13. Odontotriehum palmeri (Grcene) Hydb. BUll. Torrey Bot. Club 51:417.1924. 
Cacalia palmeri E. L. Greene, Pittonia 1:219. 1888. Bolotype: Jaliscoj 
Hio Blanco, in 1886, E. Palmer 168 (NDI, Isotypes GHI, USI, NYI). 
Plants 0.7-1 (-1.5) m. tall; stems 3-5 (-7) mm. thick at the base, 
sparsely or densely pubescent, rarely subglabrous, hairs 1.5-3 mm. 
long, stili, white; leaves sparsely or densely pubescent, the pubescence 
denser on tbe abaxial surface; basal leaves 2-3 (-5), pinnately veined, 
corillceous, broadly ovate or occllSionally subcircular in outline, 9- 17 
(-23) cm. long and broad (broadest below the middle); blades coarsely 
dentate or shalh)wly lobulate, the lobules up to 1 cm. long, 2 cm. 
wide, each 3-allgulllte, mucronate; petioles (5-) 6-15 (-30) cm. long; 
cauline leaves several, the lowermost as large as or one-half as large 
as the basal ones and similar, the petiole dilated and clasping the stem 
at the bllSe, often leafy-margined; upper cauline leaves reduced and 
bractei(orm, the petiolar portion hroadly leafy-margined and c1llSping 
the stem, the blade portion ovate, dentate; inflorescence corymbose, 
9-12 (-18) cm. long, 10- 15 cm. broad, the branches sparsely pubes­
cent, the distal branches subtended by narrowly ovate or subulate 
brncts up to 1.5 mm. long, the proximlll brnnches in the axils of the 
uppermost leaves; clllyculate bracts usually 3, subuillte, 5 mm. long 
or less; heads lip to 100, en. 1 cm.long; phyllllries 5 or 6 (-7), narrowly 
ovllLe, 5-7 (- 8) mm. long, 1.5 (-3) mm. wide, glabrous or sparsely 
pubescent, tightly imbricated fonning II narrow cylindriclll involucre, 
tbe tips attenuate; reccptacle often sparsely pubescent with long, 
mult cellular hairs; flowers 5 or 6 (-8); corolla 6.5-9 mm. long, the 
tube (3-) 3.5-4 mOl. long, the lobes 3.5-5 (-5.5) mm. long, the throat 
Ilone; anthers 2-3 10m. long; style branches 1-1.5 mm. long, the tips 
truncate or rounded with a tenninal fringe of trichomes; mature 
achones 3.5--4 mm. long, 1.5-2 mm. wide, ellipsoid, glabrous, promi­
nently 10- to 12-ribbed, the ribs dark red purple, sharply contrasted 
• 
Pll'PEN MEXICAN "CACALIOID" GENERA 401 
by pale golden-tan bands separating them; pappus creamy or wbite, 
(1-) 1.5 (-2) mm. long, about one-halI as long as the corolla tube; 
chromosome number, n=about 25 (Pippen). 
DISTRIBUTION: Southern Durango, Nayarit, Jalisco, and Micho­
acan. Common in grasslands, pastures, and borders of oak-pine 
woodlands. Flowering in August and September. 
This species is similar to O. brachycomum in having an extremely 
short pappus but differs in having smaller heads with fewer flowers 
and phyllaries, and the leaves not lobed. 
Rydberg described Odontotrichum nephrophyllum from a specimen 
collected in southern Durango (Rose 2248, US). It differs from O. 
palmeri in that the pappus is as long as the corolla tube and the 
petioles of the basal leaves are densely pubescent. Examination of 
future collections may indicate that this is distinct from O. palmeri, 
but at the present I do not feel that it is significantly different. 
DURANGO: Rou 2248 (US). JALISCO: Diguel 8.n ., July 1912 (MICH); Graham 
124 (MICH); MeVaugh 20481,12929 (MICH); MeVaugh &: Ko'!. 441 (MICH); 
King 3659 (MICH, NY, TEX, US); Powell &: Edmond.,on 869 (TEX); Pringle 
2304 (GH, NY, US); Pringle 11500 (GH, MICH, US); Pippen 10, 20, 27 A, 32, 
28 (MICHl. MICROAcAN: Arsine 5132 (GH, NY, US); Feddema 183 (MICH); 
Leavenworth 539 (NY, MICH) ; Pippen 43 (MICH); MeVaugh 21960 (MICHl. 
NAYARIT: Feddema 308 (MICH); Pippen 12* (MICHl. 
14. Odontotrlchum paaclcapitatum (Rob. '" Greenm.) Rydb. Bull. Torrey Bot. 
Club 51:414.1924. 
Cacalia pau.cicapitata Rob. &: Greenm. Amer. Journ. Sci. III. 50:158. 1895. 
Holotype: Oaxaca.; Sierra de Clavellinas, 7)000 ft., Oct. 25, 1894, C. G. 
Pringle 6018 (GHI, Isotypea GH!, US!). 
Plants to 1 m. tall; eaudex apparently fleshy, globose, at least 2 
em. long, 2.5 cm. wide; stem ca. 5 mm. thiek at the base, densely 
covered with a flocculent interwoven mass of long, gmyish-wbite 
hairs persisting through tbe inflorescence; leaves glabrous on tbe 
adaxial surface, the abaxial surface arachnoid or tomen tose; basal 
leaves 2 or 3, petioled, pinnately veined, oblong in outline, 14-16 cm. 
long, ca. 10 em. wide; blades deeply pinnately lobed, the primary 
lobes 9 or 10, at right angles to the axis but ascending distally, extend­
ing ca. four-fifths of the distance to the midrib, each irregularly 3- to 
5-lobed; petioles 5-12 cm. long, slender; cauline leaves ca. 3, tbe 
lowermost about one-half as large as the basal ones and similar; 
upper cauline leaves reduced and bracteiform, 4-6 cm. long, ca. 1 em. 
wide; inflorescence subraeemose; pedicel. (2-) 4-6 cm. long; calycu­
late bracts 3-5, linear-subulate, 1-2 cm. long; head. 5-7, 2-2.5 cm. 
long; phyllaries 12 or 13, oblong or narrowly ovate, 10-12 mm. long, 
3-4.5 mm. wide, densely pubescent; flowers 40 or more; corolla 11-14 
mm. long, the tube 7-10 mm. long, the throat narrowly eampanulate, 
1-1.5 mm. long, equally broad at the apex, the lobes ca. 3 mm. long, 
402 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
lineal'; anthers 3 mm. long; style bran(,hes 1.5 mm. long, hispid dis­
taJly on the abaxial surface, the tips truncate with a terminal fringe 
of triehomes; mature achenes ellipsoid, ohovoid or 3-angulate, 5--6 
mm. 101lg, 2-3.5 mm. wide, 1.5 mm. thick, densely villous, the hairs 
ascending, grayish-white or creamy, up to I mm. long; l"'pP"S 
crcl1my, 7-8 mm. long, davat .. distally. 
DISTRIBUTION: Known ollly from the type specimens. 
This species is most unusual because of the dense, gl'llyislHdlite, 
flocclIlent pubescence covering the entire plant except the adaxiul 
sUl'fll(,e of the leaves. The leaves are very much like those of O. 
sinuatum but the heuds are much larger and the flowers hu \"e a short, 
but evident throat. The apparently fleshy cauuex is "Iso uuique. 
15. Odontotrichum pl.tyl.pis (Rob. & Scat.) Rydb. Bull. Torrey 1I0t. Club 51: 
417. 1924. 
Cacalia plafylepis Rob. & Rent. Proc. Amer. Acad. 28:110. 1893. Lectotype: 
Jalisco; Rio Blanco, October 1886, E. Palmer 689 (CR!). 
Ca('.alia coriacea IV£. Eo Jones, Extracts from Contr. West. Bot. 18:RO. 1933. 
Hoiotype: Jalisco; Orendain, No\,. 2R, 1930, M. E. Jones s.n. (POM, not 
BCCll; isotype GH!). 
l'lants to 1.5 m. t,aU; stem 6-12 Illlll. t,hick at t,he !>n.,e, densely 
.h"ggy-villous below, becoming Ies., dense upward; leaves "p"rsely 
pubescent mostly along the veins, the abnxilll surface densely pubes­
eent; b"sal leaves 3 or 4, long-petioled, erect" corinceons, broadly o"ute 
in outline, IS-35 (-75) cm. long and broad or slig-ht.Jy broader; blndes 
deeply pinnately lohed two-thirds to three-fonrths the distllnce to the 
midrih, the primnry lobes ca. 11, the proximlll p"ir (6-) 10-16 (- 22) 
cm. long, 2-6 cm. wide at the base, the uistal lohes progrc.'Sively 
smaller upw"rd, each lobe irregul"rly divided into 2-4 lo!>es, the major 
~ccondary lobes oft.en ~ttill lohed , t.he ultimate segments triu,nguln.r or 
deIt.oid or ohlong-ohtuse, the base of the blade decurrent on tbe petiole 
for a short distance Iormin~ a leafy margin, the nbaxillJ n.nd adaxial 
surfaces very similar; p~t,ioles 16~32 em. long, trillllgulnr' in eros~ sec­
tion, densely pubesrent. below becoming sparsely pubescent distnlly; 
cauline leaves several, the lowermost smaller than the hasal ones but 
similar; upper cauline leaves hracteifol'm, the petiol"r portion leafy­
margined, clasping the stem, the hlade portion sinul\tely lobed, tho 
uppermost forming inflorescence brncts, the I"rger ones 3-6 em. long, 
1.5-4 ern. wide, distally laciniate; inflorescence corymbose, 16-27 cm. 
long, 16-18 em. broad, the branches sparsely or densely arachnoid 
jlubescent, the hairs frequently "'it,h purple crosswalIs; pedicels 1.5--3 
(-9) em. long; calyculute bl'l1cts 5-8, narrowly obovate, 1.5- 2 cm. 
long', 5-10 HUH. wide, luciniate-Iobed ; heads 15 or fewer, 1.5- 2 em. 
long; phylluries 13 or 14 (-17), nllrrowly ovute or elliptic, 13- 15 (- 16) 
mm. long, 4-5 mill . wille, mostly g],,],rous, often sparsely pllberlllent; 
PIPPEN-MEXICAN "CACALIOID" GENERA 403 
receptacle to 1 em. across; flowers 40-80 or more per head; corolla 13-
15 (-16) mm.long, the tube 10-11 mm.long, the lobes (3.5-) 4-5 (-6) 
mm. long, the throat none; anthers 3.5-4 mm. long; style branches 
1-1.5 mm. long, the tips truncate with a subterminal fringe of tri­
chomes; mature achenes ellipsoid, 6-7 mm. long, 3 mm. wide, 1.5-2 
mm. thick, pale with ca. 10 contrasting dark ribs, the intervals between 
the ribs pubescent with hyaline, ascending hairs; pappus white, (8-) 
10-11 mm. long, clavate distally. 
DISTRIBUTION: Mostly in the central grassland regions of Jalisco 
around Guadalajara and extending northward to Aguascalientes. 
Open, rocky grasslands, scrub and pastured areas. Flowering from 
mid-October to December, the flowers emitting an unpleasant odor. 
In the original description two specimens were cited as syntypes: 
Jalisco; plains of Guadalajara, Nov. 26, 1888, C. G. Pringle 1816 
(GH!) and JaHsco; Rio Blanco, October 1886, E. Palmer 689 (GH!). 
The latter was designated the lectotype because it is a more complete 
specimen than Pringle 1816. The lectotype specimen consists of a 
basal leaf ca. 18 cm. long, a stem in three pieces, with 6 cauline leaves 
below the 8-headed inflorescence (Mich. Neg. 1253). There is also 
a second sheet at the Gray Herbarium (an isolectotype) that has a 
basal leaf ca. 28 cm. long, the base of the plant with the roots, and 
an inflorescence with ca. 15 heads, but no portion of the stem. 
This species is spectacular because of its extremely large basal 
leaves, more than one-half meter long, that stand erect. 
AGUABCALIENTEB: McVaugh & Koel, 48 (MICHl. JALlSCO: Jone3 27692 (NY); 
Palmer 689 (US, 3 sheets, isotypes); McVaugh 22123 (MICH); Pringle 1816 
(US, 2 sheets, NY, MICH, isoparatypes); Pringle 9873 (GIl, US, NY); Safford 
1432 (US); Jones s.n., Nov. 27, 1930 (GH, isotype of C. coriacea Jones); Pippen 
23, 58 (MICH); Jany s.n., in 1892 (US). 
16. Odontotrichum pringlei (S. Wats.) Rydb. Bull. Torrey Bot. Club 51: 415. 
1924. PLATE 4 
Cacalia cirsHJoUa Hook. & ArD. Bot. Beech. Vcy. 436. 1841. Holotype: 
Nayarit; between Tepic and San BlaB, Sinclair s.n., in Hooker Herb. 
(K, not seen; Mich. Neg. 6231). Not Odoniotrichum cirsii/oZium Zucco 
Senecio cirsiifolia (Hook. &: Arn.) Schultz Bip. Flora 28:499. 1845. 
Cacalia pringlei S. \Vats. Proc. Amer. Acad. 25:156. 1890. Lectotype: Jalisco; 
gr88By slopes of the barranca near Guadalajara, November 1888, C. G. 
Pringle 1749 (GIll). 
Plants to 1.5 m. tall; stem 4-10 mm. thick at the base, densely 
villous, flocculent below the lowest leaf and sparsely puberulent or 
becoming glabrous above; leaves sparsely pubescent with 8- to 10-
celled hairs 0.3-0.5 mm. long, slightly longer and denser along the 
veins and on the abaxial surface; basal leaves 2-4, erect, coriaceous, 
long-petioled, pinnately veined, oblong or elliptic in outlioe, 20-35 
(-40) cm. long, 13-30 cm. wide, as long as broad to twice as long as 
404 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
broad; blades deeply pinnately lobed, the primary lobes 1O~16, (4~) 
5~10 (-15) cm. long, 3-4 cm. wide at the base, each usually divided 
into 3 major lobes and 2 or 3 lesser lobules or teeth, the latter usually 
narrowly triangular, the secondary lobes triangular or 3-angulate or 
again divided into 2 or 3 triangular lobules, the lobes, including the 
proximal pair, forming an angle with the midrib of less than 90 
degrees; blades obtuse or attenuate, not cordate at the base, decurrent 
along the petiole for 1-2 cm. below the proximal lobes; petioles 20-30 
(-40) cm. long, triangular in cross section, villous proximally, be­
coming sparsely pubescent distally; cauline leaves 3-7, much reduced 
in size, bracteiform, the lowermost leaflike, sinuately lobed, the 
petiole dilated at the base and clasping the stem, up to 15 cm. long, 
3 cm. broad; inflorescence loosely paniculate or corymbose, 18-25 
cm. long, 16-20 cm. broad, the branches subglabrous to sparsely 
pub",cent, subtended by narrowly oVllte bracts (2-) 3~5 (-7) em. 
long, 0.5-1.5 cm. wide; calyculate bracts 5-8, linear-subu\',te, 3.5-5.5 
mm. long; heads ca. 100, 10-12 mm. long; phyllaries 7 or 8 (-9), 
ovate or elliptic, 5-7 mm. long, 2-3 (-4) mm. wide, glabrous; recep­
tacle 1.5-2 mm. across; flowers 11-14; corolla 6-8 (-10) mm. long, 
the tube 3-4 (-5) mm. long, the lobes :1-4 (-5) mm. long, linear, the 
median nerve strong, the throat none; anthers 2-3 mm. long; style 
branches I (-1.5) mm. long, usually hispid on the abaxial surface, the 
tips truncate or obtuse with a subterminal or terminal fringe of 
trichomes; mature achenes ellipsoid (3-) 4-5 10m. long, 2 mm. wide, 
1-1.5 mm. thick, pubescent with minute, creamy or ta\vuy hairs 
between the ribs, the ribs about 20, darker than the area between 
the ribs; pappus (4-) 6-7 mm. long, white, the tips slightly clavate; 
chromosome number, n=30 (Pippen; Turner, ined.). 
Two specimens were cited in the oribrinal description as syntypes: 
Jalisco; grassy slopes of the barranca near Guadalajara, November 
1888, O. G. Pringle 1749 (GH) andlS11 (GH). Pringle 1749 is chosen 
as the lectotype because it is " more complete specimen. 
DISTRIBUTION: Nayarit, Jalisco, and Colima. Grasslands, pastured 
areas, roadsides, and less frequently rocky slopes in oak-pine wood­
lands. Flowering from October to early December. 
This species is closely related to O. l'latylel'is and O. sinuatum, 
differing from the former in having smaller heads; f!'Om the latter in 
having pubescent achenes and differently shaped leaves. Oacalia 
cirsiifolia Hook. & Arn., known only from the very incomplete type 
specimen consisting of just a portion of the inflorescence and one 
upper eauline leaf, is undoubtedly conspecific with O. pringlei because 
of the similarity of the heads and the pubescent aehenes. 
COLIMA: Palmer 1234 (Gil, NY, US). hLiseo: Me Vaugh &: Koelz 1253, 1104 
(MICHl; MeVaugh 13616, 21765* (MICH); Pippen 37*, 41, 59 (MICH); Pringle 
PIPPEN-MEXICAN "CACALIOID" GENERA 405 
1811 (US, IBoparatype of C. pringki S. Wats.); Pringle 11499 (GH, MICH, US). 
NATAR[T:FeTTi85881 (US); Afexia845 (GIf, US); McVaugh & Koe/z601 (MICH); 
Palmer B.n., Jan. 5-Feb. 6, 1892 (US); Barclay •. n. (GH, probably collected in late 
December 1837 at the same time and place as the type of Cacalia cirsii/olia 
Hook. &: Arn., since Barclay and Sinclair were together at that time, attached to 
the H.M.S. Sulphur (McVaugh, manuscript ined.I). 
17. Od.nt.triehum purpusU (GreeDm.) Rydb. Bun. Torrey Bot. Club 51:418. 
1924. 
Cacalia purpmii GreeDm. UDiv. Calif. Pub!. Bot. 4:95. 1910. Holotype: 
Puebla; Cacalote, AuguBt 1909, C. A. Purpu. 3845 (FI, iBotype NYI). 
Plants to 0.6 m. tall, pubescent with white, stiff, multicellular hairs; 
stem slender, ca. 3 mm. thick at the base; basal leaves 2-4, pinnately 
veined, petioled, subcoriaceous, ovate or broadly elliptic, to 8 cm. 
long, 6 cm. wide, the margin of tbe blade doubly dentate with two alter­
nating series of large and small teeth, respectively, the base of tbe 
blade truncate or obtuse; petioles terete, 3-9 em. long, about as long 
as the blade; cauline leaves ca. 4, the lowermost smaller than the basal 
ones but similar to them; upper cauline leaves reduced, bracteiform; 
inflorescence compact or spreading, corymbose, 3-14 cm. long, 3-19 
cm. broad, the branches arachnoid-pubescent or tomentose, subtended 
by linear-subulate bracts ca. 5 mm. long; calyculate bracts 3-7, 
linear-subulate, 3-4 mm. long; heads 15- 30, 10-12 mm. long; phyl­
laries 7 or 8, oblong or ovate, 5.5-7 mm. long, 1.5-2 mm. wide, gla­
brous; flowers 10-13, corolla ca. 7 mm. long, the tube (3.5-) 4 mm. 
long, the lobes (3-) 3.5 mm.long; antbers 2.5 mm.long; style branches 
1.5 mm. long, the tips truncate witb a terminal fringe of trichomes; 
mature achenes oblong, ca. 3.5 mm. long, 1 mm. wide, glabrous, 
brown; pappus 4-6 mm. long, creamy or tawny. 
DISTRIBUTION: Puebla and Oaxaca. Known only from three collec­
tions. Flowering in JW1e to August. 
This taxon is quite closely related to the O. sUphiijolium-goldsmithii 
group but is much smltJler and does not occur within the present known 
geographical range of either of the others. When more material of 
this taxon becomes available and can be studied it may indicate that 
O. purpusii should not he considered as a distinct species. It is con­
sidered distinct here because of the general small habit, distinct geo­
graphical distribution, and lack of correlative evidence to combine it 
with any other taxon. 
OAXACA: Conuuti 4015 (US); Con .. tti & Gonzal .. 1237 (GH, this specimen has 
no leaves). 
18. Odontotrlchum radullf.llum ("raduloefollum"l (H.B.K.) Rydb. Bull. Torrey 
Bot. Club 51 :415. 1924. 
Cacalia radulifolia ("rodulacfolia"] H.B.K. Nov. Gen. et Sp. 4: 169. 1820. 
(Folio ed. 4:132. 1820) . lIolotype: GuaDojuato; between the village of 
Temeeeatio and Gua.najuato, DO date, 1050 hex., Humboldt c.t Bonpland 
B.n. (P, not .... n; Field Mus. Neg. 378751) . 
406 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Senecio radulaefolius (II.B.K.) Schultz Bip. Flora 28:498. 1845. 
Cacalia schaffneri A. Gray, Proc. Amer. Acad. 19:53. 1883. Lectotype: ncar 
San Luis Potosi in 1876, Schaffner 294 (Gm) 
Odon/otrichum schaffneri (A. Gray) Rydb. Bull. Torrey Bot. Club 51:415. 
1924. 
Plants to 1.5 m. tall; entire plant pubescent except the phyllaries, 
the hairs white, 5- to 7-celled; stem 3-5 (-8) mm. thick at the base, 
green or purplish; basal leaves 2 or 3, long-petioled, pinnately veined, 
ovate-cordate or subcircular in outline, (6-) 10-12 (-18) cm. long, 
(8-) 12-14 (-22) cm. wide, usually 3-5 cm. broader than long or as 
long as broad, subpeltate, the petiole attached to the blade 2-10 (-30) 
mm. above the base; blades pinnately lobed three-fifths to four-fifths 
the distance to the midrib, the primary lobes 7-9 (-11), each divided 
into usually 3 (2-5) lobes, the secondary lohes often lobed, 3-angulate 
or coarsely dentate, the lowermost pair of primary lobes deflected, 
parallel to the petiole; petioles (7-) 12-21 (-30) cm. long; cauline 
leaves 2-4, the lowermost about as large as the basal ones and similar, 
but the base of the petiole more dilated and often leafy-margined; 
upper cauline leaves progressively smaller upward, bracteiform, auric­
ulate-based, elliptic or ovate, distally sinuately 7- to 9-lobed; in­
florescence loosely corymbose, (3-) 12-30 cm. long and broad, the 
branches subtended by linear-subulate bracts (5-) 7-10 mm. long; 
calyculate bracts 3-5, subulate, 2-5 mm. long; heads (25-) 50-75, 
10-11 mm. long; phyllaries 5, oblong or narrowly ovate, 5-7 mm. 
long, 1-2 (-3) mm. wide, glabrous; flowers 5 or 6; corolla 6.5-7.5 mm. 
long, the tube 3-3.5 (-4) mm. long, the throat very short, ampliate, 
0.5-0.7 mm. long, 0.7-1 mm. wide at the apex, the lobes 3-3.5 mm. 
long; anthers 2-3 mm. long; style branches 1-1.5 mm. long, hispid on 
the abaxial surface, the tips truncate or obtuse wit.h a subtenninal 
fringe of trichomes; mature acbenes ellipsoid, 4 mm. long, 1.5 mm. 
wide, brown, ca. 14-ribbcd, pubescent, the bairs 0.5 mm. long, greenish 
or tawny; pappus creamy or fUSCOllS, 4-5.5 mm. long. 
DISTRIBUTION: Sinaloa, Durango, San Luis Potosi and Guanajuato 
to Tamaulipas. Grassy openings in oak-pine woodlands and pastures. 
Flowering in July and August. 
Comparison of the t.ype specimen of Cacalia 8chaffneri A. Gray 
with the photograph of the type of C. radulifolia H.B.K. and the 
similarity of the original descriptions of these two taxa indicate that 
they are conspecific, even though the subpeltate condition of the 
leaves was not mentioned in the original description of C. radulifolia. 
Gray mentioned in his description of C. 8chaffneri that possibly this 
was the little known C. radulifolia of H.B.K. This is the only species 
of OdlYntotrichum with subpeltate leaves. 
DURANOO: Graham 100 (MICH); Feddema 1821 (MICH); May,illes 7587 
(MICH). GUANAJUATO: Kenoyer 2110, 2300 (GH); Schumann 148 (US). SAN 
PIPPEN-MEXICAN "CACALIOID" GENERA 407 
LUIS POTosi: Manning &: Manning 53560 (GIl): Mickel 501, 561 (MICH): 
PQrry &: Palmer 543 (NY): Pringle 3566 (GH, US): Pringle 4095 (NY, US): 
Schaffner 294 (NY, US, i80type of C. schaffner; A. Gray: the US specimen also 
bears the number 724 assigned to the ebeet by A. Vigener who purchased many 
of the Scbaffner collections {J. Rzedowski, pers. comm.n. SINALOA: PoweU ct 
Edmondson 929 (MICH, TEX). NUEVO LEON: M eyer &: Rage" 2608 (MICH) 
TAlIAULlPA8: Stanford, R' tMrford &: NortMr_f! 818 (GH): Blanford, 7'aylor, &: 
Lauber 2421 (MICH, NY, TEX, US). 
19. Odontotri.hum silphiifolium (Rob. &: Greenm.) Rydb. Bull. Torrey Bot. 
Club 51:416. 1924. 
Cacalia silphiiJolia Rob. &: Greenm. Amer. Journ. Sci. III. 50:158. 1895. 
Holotype: Mbico; Sierra de las Cruces, Aug. 21, 1892, C. G. Pringle 
5251 (GH!). 
Plants to ca. 1.5 m. tall; roots exuding an amber-colored ftuid 
when cut; stem 1-1.5 cm. thick at the base, subglabrous or puberu­
lent upward; leaves minutely puberulent especially along the veins; 
basal leaves 2 or 3, erect, long-petioled, pinnately veined, ovate­
cordate, (13-) 15-20 (-45) cm. long from the tip to the basal sinus, 
(11 - ) 14-21 cm. wide, usually 2-4 cm. longer than wide, occasionally 
as wide as long; blades sinuately double-dentate, the teeth in two 
alternating series of large and small, respectively, the base of the 
blade cordate or sagittate, rarely truncate, the basal lobes 4-7 cm. 
long, 5-9 cm. wide, rounded; petioles (15-) 2(}-30 cm. long; cauline 
leaves 2-4, the lowermost similar to the basal ones but smaller, 
occasionally deltoid and truncate-based with the petiolar portion 
broadly leafy-margined, auriculate-based and clasping the stem; 
upper cauline leaves smaller, bracteiform; inftorescence tightly 
corymbose, usually 6-11 cm. long and broad, occasionally up to 28 
cm. long, cu. 20 em. broad, the branches tomentase, subtended by 
linear or narrowly ovate bracts 1.5 cm. long, 1 mm. wide; pedicels 
0.3- 3 em. long; calyculate bracts 1-5, linear-subulate, (3.5-) 5-7 
mm. long; heads 50-75; 10-15 mm. long; phyllaries 8, narrowly 
ovate or ovate, (6-) 7-8 mm. long, 1.5-2 (-3) mm. wide, glabrous; 
flowers 9-13; corolla 8.5-9 (-11) mm. long, the tube 5 (-6) mm. 
long, the lobes 3.5-4 (-5) mm. long, the throat none; anthers 2-3 
mm. long; style branches 1- 1.5 mm. long, the tips truncate with a 
sparse terminal fringe of trichomes; mature achenes ellipsoid, 4-5.5 
mm. long, 1.5-2 mm. wide, 1-1.5 mm. thick, glabrous, light brown, 
inconspicuously 16- to 18-ribbed; pappus creamy, (5-) 7-8 mm.long. 
DISTRIBUTION: Michoacan, east to M~xico and Hidalgo, south to 
Morelos. Grasslands, pastures, borders, and openings in oak wood­
lands. Flowering in August and September. 
This species is very closely related to O. goldsmithii and may be 
distinguished from it by its coarser habit, larger, puberulent, cordate 
leaves and more compact, tight inftorescence with larger heads. 
408 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
DISTRITO FEDERAL: Matuda 19184 (US). IlWALOO: R()$ •• t aI. 9187 (GIl, NY, 
US). Mhleo: Matwla 19417 (NY); PriDgle 6453 (GH, MICH, NY, US). 
MlenoAc.( N: King 3615 (MICH, NY, TEX, U8); Pippen 54 (MICH). MOKELO.: 
Moore 3421 (Gil); Orcult 3746 (GIl, U8); Rose &: Hay 53lt (GIf, U8); Pringle 
9080 (Gn, US). 
20. Odontotrichum oinuatum (Cerv.) Rydb. Bull. Torrey Dot. Club 51 :415. 1924. 
Cacalia tl'nuala CeTV. 1'" Llave & Lex. Nov. Veg. Vesco rac. 1:29. 1824. 
Holotype: DiBtrito Federalj mountains of the desert of San Angel, near 
Mexico City, Cen.'antes 8.n. (not scen). 
Senecio albo-Iutesan& Schultz Bip. Flora 28:498. 1845. Based on Cacalia 
B1'nuata Cerv. not S. sinuatu8 H.B.K. 
Senecio calophyllus Hemol. BioI. CeDtr. Amer. Bot. 2:237. 1881. Name 
U1egitlmate. 
Slender plants to 1.5 (-2) m. tall; stem 4-6 mm. thick at the base, 
light green, pubescent below becoming subglabrous above; leaves 
sparsely puberulent, mosily along the veins and margins; basal leaves 
3-5 (-7). erect, 10ng-peLioled, oblong-cordate in outline, (8-) 12-19 
cm.long, (8-) 12-17 cm. wide, usually 2-3 cm.longer than broad or as 
long as broad; blades deeply lobed to within 1-3 cm. of the midrib, the 
primary lobes 8-11 (-13), 1-3 cm. broad at the base, mostly at right 
angles to the midrib, each lobe distally divided into 3 major secondary 
lobes and 2 or 3 lesser lobules or coarso teeth, the proximal pair of 
primary lobes usually deflected, more or less parallel to the petiole; 
petioles 10-40 em. long, usually longer than the blade, slender, gla­
brous, green, often mottled with purple; cauline leaves 3-5 Or more, 
the lowermost similar to the basal ones and about one-third as large, 
the petiole more clasping at the base; upper Gauline leaves greatly re­
duced, bracteiform, the petiolar portion leafy-margined and clasping 
the stem, distally lobed; inflorescence corymbose (10-) 15-27 em. long, 
(6-) 12-21 cm. broad, the branches sparsely to arachnoid-pubescent, 
sub tended by narrowly ovate bracts 1-1.5 cm. long, 3 mm. wide, 
occasionally larger; pedicels 3-5 mm. long; calyculate bracts 3-7, lin­
ear-subulllte, 3-7 mm. long; heads 75-100, 10-12 mm.long; phyllarics 
mostly 8 (7-9), elliptic or narrowly ovate, 5-6 (-7) mm.long, 2-3 mm. 
wide, glabrous, receptacle often slightly pubescent, up to 2 mm. across; 
flowers (9-) 10-11 (-13); corolla 6-7 mm. long, the tube 3-3.5 (-4) 
mm. long, the lobes 3-3.5 mm. long with a strong median nerve; 
anthers 2-3 mm.long; style branches 1-1.5 mm. long, the tips truncnte 
with a terminal fringe of trichomcs; mature achenes ellipsoid, 3.5-4 
mm.long, 1.5-2 mm. wide, ca. 1-1.5 mm. thick, glabrons, light brown, 
ca. 20-striated; pappus white, 4-5.5 (-7) mm. long; chromosome 
number, n=30 (Turner, ined.). 
DISTRIBUTION: From southern Chihuabua south and enst to 
Mexico. Common in grasslands, pastures, and borders of oak-pine 
PIPPEN-MEXICAN "CACALIOID" GENEllA 409 
woodlands, occasionally in poorly drained areas. Flowering from 
August to October. 
This is one of the most common species in western and central 
Mexico and exhibits considerable morphological variation. The basal 
leaves on the same plant as well as within the same population may 
vary in size, shape, and depth of lobing. 
The holotype of this species has not been located. McVaugh and 
Rzedowski searched in Mexico City and McVaugh in Madrid but no 
Cervantes specimen was located. It is possible that no type specimen 
exists, as few other La L1ave and Lexanza specimens are known. The 
original description, however, is sufficiently diagnostic to accept this 
name. 
AGUASCALIENTE.: MeVaugh & Koelz 49 (MICH); Pippen I, 7 (MICH). Dro­
TruTO FEDERAL: Bourgeau 716 (GH, US); Smyth 172 (US). DURANGO: Gould 8989 
(MICH); King 3734 (NY, TEX); Me Vaugh 21733 (MICH, part of hybird populo.. 
tion between O. 8inuatum nnd O. globosum); Mallsilles 7572, 7793, 7880, 8448, 
(MICH), 8485 (MICH, NY); Palmer 651 (GIl, NY, US); Po .... 11 & Edmond.on 
948 (TEX); G,nlry 8562 (MICH). GUANAJUATO: Dug .. 451 (GH). JALroco: 
Hartweg 125 (GH, NY); MeV.ugh 12810 (MICR); Me Vaugh & Koelz 276 (MICH); 
Pippen 9", 22, 57 (MICH). MtxICo: Matuda 19706, 21808,26882 (NY); Pippen 
56 (MICH); Pringle 4272, 9875 (GIl, NY, US); Rose & Painter 7808 (GH, US). 
MrcHoAcAN: Arsine 5740 (US); Feddem. 10 (MICH); Pippen 45,50 (MICH). 
PUEBU: Ar.ln, 93A, 334, 93, lI81 (US). WrTROUT LOC.'LITY: Viseher 58 (Ny). 
21. Odonlolrlchum lussil.einold.o (H.B.K.) Rydb. Bull. Torrey Bot. ClUb 
51:416.1924. 
Cacalia t"""aginoid .. H.B.K. Nov. Gen. et Sp. 4:168. 1820. (Folio cd. 
4:132. 1820.) Holotypc: Hidalgo; Real del Monte, 1420 hex., no date, 
Humboldt & Bonpland 8.n. (P, not seen; Field Mus. Neg. 378781). 
Senecio tU8.ilaginoide. (H.B.K.) Schultz Blp. Flora 28 :498. 1845. Not S. 
tU3silaginoides Wa1 t. 1788. 
Senecio Ja1'Jarus Hemsl. BioI. Centro Amer. Bot. 2:239. 1881. Based on 
Cacalia tUlsilaginoides H.B.K., not S. tU88ilaginoides Walt. 1788. 
Cacalill ampulla~a Greenm. Proe. Amer. Acad. 34:577. 1899. Holotype: 
Hidalgoi Sierra de Pachuca, 9,000 ft., July 17, 1898, C. G. Pringle 1617 
(GHI). 
Odonwtr1'chum ampullaceum (Greenm.) Rydb. Dull. Torrey Bot. Club 51 :416. 
1924. 
Roots and lower portion of the stem not seen; erect herbs up to 
1.3 m. tall (acc. Balls 5428); stem pubescent; leaves puberulent on 
the adaxial surface, the abaxial surface gray-tomentulose, the tomen­
tum often flocculent; basal leaves presumably 2 or more, pinnately 
veined, ovate~ordate in outline, 20-30 (-35) cm. long from the apex 
to the basal sinus, 17-35 cm. wide, usually 2-4 cm. longer than broad 
or occasionally as broad as long; blades lobed one-half the distance 
to the midrib (at least on the distal portion of the blade), the pri­
mary lobes 10-12, 5-6 (-14) cm. long, 4-5 (-ll) cm. broad at the 
410 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
base, each sinuately and irregularly divided into at least 3 lobes, the 
secondary lobes usually coarsely dentate; petioles at least as long as 
the blades, usually longer (to 0.5 m.) ; lower e,wline leaves not seen; 
ullper cnuline leaves leafy, bracteiform, o\'ate, naviculat,e, clnsping 
the stern, 11-20 ern. long, 10-12 ern. wide, the margin toothed or 
sinnately lohed; inflorescence tightly eorymbose, 17 (-21) cm. long, 
12-17 em. broad, the brancbes tomentose, the lowermost bmnches 
in the "xits of the uppermost leaves, the distal branches subtended 
by Iinear-snbulate bracts 1.5-4 cm.long, 1- 2 (-4) mm. wide; cnlycu­
late bracts 4 or 5, linear-subulate, 3- 5 mm. long; heads 100 or more, 
7-10 mm. long; phyllaries 5, narrowly ov!\te, 5-7 mm. long, \.5-2 
(- 3) wide, glabrous, often purplish; flowers 5 or 6, corolt.. 6- 7 1ll1lI. 
long, the tube 3-3.5 mm. long, the lolles 3-3.5 mm. long, the throat 
none; anthers 2 (-2.5) mm. long; style bmnches 1 mm. long, hispid 
on the abaxial surface, the tips truncate or obtuse with a subterminal 
fringe of trichomes; mature fichenes not seen, immature achenes 
3.5-4 mm. long, pale tan, densely pubescent; ptippUS creamy or 
tawny, 4-5 (-7) mrn. long. 
DISTRIBUTION: Known only from Hidalgo and Veracruz. In wet 
mountain meadows. Flowering in August (?) (Kunth stated in the 
description that the plants flowered in May. The specimens I have 
examined flowered in August). This is one of the rew species of Odon­
totriehum found in eastern Mexico. 
This species is somewhat similar to O. amplijalium, differing mostly 
in t.he smaller heads with fewer flowers and the pubescent achenes. 
Com parison of the description and type specimen of Cacalia ampul­
laeea Greenm. with the description and type photograph of O. t118-
silaginoides indicates that they are conspecific because of their 
similarity in appearance and their many common clunacters. Green­
man apparently reached this conclusion nfter the public.\tion of C. 
amp1tllacea because the type specimen is annotated as C. t1t88ilagi­
noides H.B.K., in what appears to be Greenman's writ.ing. 
HIDALGO: Rose & Hay 5.574 (US); Rose &: Pal:tller 6738 (CH, US) j Pringle 
9868 (GH, NY, US). V'"ACRUZ: Ball, 5428 (US). 
$. Periealia 
Pericalia (Cass. Diet. Sci. Nnt . 48: 459. 1827, not validly published) Rydb. Bul1. 
Torrey Bot. Club 51: 376. 1924. 
Perennial, leafy-stemmed herbs, 1.5- 2 m. tall; roots fibrous, fleshy; 
stems usually solitary, erect, annual, terete to often 6- to 8-angulate 
distally, pubescent or glabrous, tuberculate at base, the clustered 
suhterrane,m tubercles el>ch up to 2.5 cm. thick, 5 cm. long, pubescent 
with minute, multieellul:lr, tan hn,irs; leaves call line, not forming n. 
basal rosette, succulent when fresh, coriaceous when dry, petiolate or 
PIPPEN-MEXICAN "CACALIOlD" GENERA 411 
sessile, alternate, clustered toward the middle of the stem but con­
tinuing upward to the inflorescence, becoming smaller and widely 
separated distally, not usually becoming bracteiform; inflorescence 
paniculate or corymbose; heads usually few, nodding, 15-20 mm. 
long, sub tended by 10-15, linear or narrowly triangular, calyculate 
bracts as long as or longer than the phyllaries and loosely surrounding 
them, especially in bud; involucre campanulate to turbinate, the 
phyllaries suhequal, biseriate appearing uniseriate, imbricate, the 
overlapped margins subscarious, the exposed surfaces glabrous, the 
tips fringed with minute trichomes up to 0.3 mm. long; receptacle 
flat or nearly so, alveolate, not paleaceous; flowers all discoid, perfect, 
fertile, 25-50 or more per head; corollas glabrous, creamy or nearly 
white but never yellow, the tube narrowly cylindrical, ca. 0.5 mm. in 
diameter, the limb divided into 5, equal, narrowly triangular, 3-
nerved, recurved lobes connate into a campanulate throat, 2-5 mm. 
long, as long as or twice as long as the lobes; anthers 3-4 mm. long, 
exserted beyond the corolla throat, the bases rounded or slightly 
sagittate, the terminal appendages trullate, rounded or ovate, 0.5-0.7 
mm. long; style branches 1-2 mm. long, slightly flattened, recurved, 
the stigmatic area covering most of the adaxial surface, the abaxial 
surface glabrous or hispidulous below, the unappendaged tips truncate, 
to obtuse, fringed with nonglandular "pollen-producing" trichomes; 
achenes narrowly cylindrical, subterete in cross section, glabrous, 
purplish brown, 10-ribbed; pappus creamy or white, distally clavate. 
TYPE SPECIES: Pericalia sessilijolia (Hook & Arn.) Rydb. 
Cossini suggested that Cacalia cordifolia H.B.K. should be con­
sidered as either a separate genus or subgenus under the name Peri­
calia but he did not validly publish the name as either. DeCandolle 
cited Pericalia as a synonym of C. cordijolia H.B.K. and Rydberg 
used Pericalia as a genus in 1924. 
Microscopic examination of freehand sections of the unique tubercles 
revealed their stemlike nature by the presence of discrete collateral 
vascular bundles. The parenchyma cells contain needle-shaped crys­
tals. The tubercles function not only in storage but also in vegetative 
reproduction, since in some circumstances they give rise to new 
flowering stems. 
Key to Species of Pericalia 
1. Flowers 40-50 per head; lobes of the corolla as long a.s the throat. 2-3 mm.long; 
leaves broadly ovate or elliptic, the margin eonr::;ely toothed or lobulate, the 
lobules broader than long . . . . . . . . . . . . . . 2. P. sessilifolia 
1. Flowers 25-35 per head; lobes of the corolla less than half as long as the throat, 
1-1.5 mm. long, the throat 4-5 mm. long; leaves deltoid or 8uhcircular, 3- to 
7-1obed, the lobes longer than broad. . . . . . . . . . 1. P. michoacana 
266-836--68 i 
412 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
1. Peric_n_ mlch._can_ (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51:377. 
1924. PLATE 5 
Cacalia michoacana B. L. Rob. Proe. Amer. Acad. 43:46. 1907. Holotype: 
Michoac:1n; Uruapan, on pine covered crater cone, 1,080 m., Oct. 31, 
1905, C. O. Pringle 10117 (GII!). 
Cacalia trigonaphyUa S. F. Blake, Journ. Wasb. Aead. 19:260. 1929. IIolotype: 
Jalisco; San Sebastian, trail to mine La Sabala., Feb. 10, 1921, Ynex Alexia 
1656 (US!). 
Slender herbs 0.6-1.5 (-2) m. tall; underground tubercles 1.5-2.5 
cm. thick, to 5 cm.long; stem 2-5 (-7) mm. thick at the base, greenish, 
often mottled with purple, glabrous to sparsely or densely pubescent 
proximally, becoming glabrous and oft.en glaucous abo\'e; leaves 
10-15 or more, long-petioled, snbpalmately veined with 3- 5 principal 
veins with the central vein stronger, densely to sparsely pubescent 
to glabrous; blades variable in shape from subcircnlar-eordate-bllSed 
with 5-7 deltoid or triangular lobes, to broadly deltoid-hastate with 
3 major, narrowly triangular lobes, 4- 7 (- 10) cm. long from the 
basal sinus to the tip, 5.5-10 (-16) cm. wide, usually 2-3 cm. wider 
than long, progressively smaller upword, the margins minutely or 
coarsely dentate; petioles 5-10 (-15) mm. long, purplish; inflorescence 
racemose or paniculate, 10-20 em. long, 8-12 cm. broad or larger, the 
distal brancbes sublended by linear-subulate bracts 1-1.5 cm. long, 
tbe proximal branches often in the axils of the uppermost leaves; 
pedicels 2-5 (-7) cm.long; calyculate bracts 10-15, linear or narrowly 
triangular,4- 5 (-7) mm. long; heads mostly 6-12 (-50), nodding, 15-
20 mm. long; phyllaries (10-) 12 or 13, narrowly ovate or narrowly 
oblong, (7-) 10-13 mm. long, 1-2 mm. wide; receptacle 4-5 mm. 
across; flowers 25-35; corolla 9-12.5 mm. long, the tube (3.5-) 4-4 .5 
(-5.5) mm. long, the throat funnelform or eampanulate, 4-5.5 mm. 
long, 1.5 mm. wide at the apex, tbe lobeg 1-1.5 mm. long, triangular, 
strongly recurved; anthers 3.5-4 mm. long; style branches ca. 1.5 
mm. long, the tips truncate or rounded with a sparse subterminal 
fringe of trichomes; achenes narrowly oblong or narrowly obovoid, 
terete in cross section, 2.5 mm. long, 0.5 mm. tbick, glabrous, dark 
brown with 10 pale ribs; pappus (8-) !}-IO (-11) mm. long, white, 
slightly clavate distally; chromosome number, n=30 (Pippen). 
DISTRIDUTION: From northwest Jalisco ellStward to Mexico. 
Growing mostly on rocky slopes in oak-pine woods. Flowering in 
November and December. 
The specimens examined show a wide range of variation in leaf 
shape and pubescence between the two extreme forms described 
above. Some correlation between leaf shape and pubescence is apparent 
since the plants with 5- to 7-10bod leaves tend to be more pubescent 
aDd tbose plants witb 3-10bed leaves tend to be glabrous and glaucous. 
A specimen from eastern Jalisco (Me Vaugh &\ Koelz (35) bas more 
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 413 
rounded lobes than the other specimens. Despite these variations nll 
of these specimens form only one taxon. 
Comparison of the types of Pericalia michoacana and Oacalia tri­
gonophyUa and examination of a large sample of these taxa indicate 
that they are conspecific. They have the same Dumber of phyllaries 
and flowers in a head and the morphological structure of the flowers 
is the "OIDe. The only observable difference is in the pubescence and 
the number of lobes. Blake based O. trigonophylla on a collection in 
which the leaves were 3-lobed and subglabrous. No geographical or 
ecological differences are noticed between these two extremes in 
vegetative form and therefore no nomenclatural status is given to 
these variations. 
JALISCO: Mexia 1656 (US, MICH, isotypes of C. Irigonophylla Blake); Me­
Vaugh 14290, 21355, 14136, 14093, 22071 (MICH); Pippen 26, 38, 63*, 65* 
(MICH); McVaugh &: Ko,/z 435 (MICH). MEXICO: Hinton ,I 01. 2755 (US), 
4993 (NY, US); Moore &: C,tto 5469 (MICH). MICHOAcAN: Hinton ,I al 12718, 
13469 (MICH, Ny); Pringl, 10117 (US, isotype). 
2. Peric.li. sesoilifoli. (Hook. '" Am.) Rydb. Bull. Torrey Bot. Club 51 :376. 
1924. 
Cacalia cordi/alia H.B.K. Nov. Gen. et Sp. 4:168. t. 360. 1820. Holotype: 
M6xicoj Santa Rosa, Humboldt & Bonvland s.n. (P, not scen; Field Mus. 
Neg. 378651), not Cacalia cordi/alia L.f. 
Cacalia .... ili/olia Hook. '" Am. Bot. Beech. Yoy. 436. 1841. Holotype: 
Nayarit; between San BIas and Tepic, in 1837, Sinclair B.n. in Hooker 
Herb. (K, not seen; MICH Neg. 6211). 
Senecio otJatijolitU Schultz Bip. Flora 28:498. 1845. Based on Cacalia cordi­
folia H.B.X. not 8. cordifoliu8 L.r. 
Senecio beecheyanua Schultz Dip. Flam. 28:499. 1845. Based on Cacalia 
aC88ilifolia Hook. &; Arn. Name illegitimate. 
Senecio eardiophyllm Hemsl. BioI. Centro Amer. Bot. 2:237. 1881. Based 
on Cacalia cordijolia H.B.K. Name illegitimate. 
Senecio 3es3ilijolim (Hook. &: Arn.) Hemsl. BioI. Centro Amer. Bot. 2:247. 
1881. Based on Cacalia sessilijolia Hook. &: Arn. 
Cacalia nutans Sesse & l\.loc. PI. N. lIisp. ed. 1. 132. (FI. Mex. ic. ined. 
145j Field Mus. Neg. 30705!) 1889. Lectotype: Without locality (pre­
sumably "in humidis unbrosisque Tepalpae circuitibus propeMexicanum"), 
SessA & Moe. 1488 or 28221 (both numbers appear on the specimen), 
Field Mus. Neg. 42285. 
Pericalia ovali/olia (Schultz Bip.) Rydb. Bull. Torrey Bot. Club 51 :377. 1924. 
Plants 0.5-1.5 (-1.8 m., Balls 5575) m. tall; tubercles 2-2.5 em. 
thick, to 5 em. long; entire plant glabrous and glaucous, often sparsely 
pubescent when young; stems 3-9 mm. thick at the base; leaves 10-15, 
petiolate or sessile, subpalmately veined with often 3 principal veins, 
the central one usually dominant; blades cordate-based, broadly ovate 
or broadly elliptic, the larger ones 4-13 em. long from the basal sinus 
to the apex, 5-12 (-14) em. wide, usunlly 1-3 em. wider than long 
(often as long as wide to 1)\ times longer than wide), the margin 
414 CONTRIBUTIONS FROM THE l'ATIONAL HERBARIUM 
variable from subentire to dentate to 9- to 14-lobulate, the lobules, 
when present, usually broader than long; petioles (when present) sub­
terete, purplisb green, 3-9 em. long; inflorescence corym bose or pa­
nieulate, 1(}-30 em. long, 7-18 cm. broad, the proximal branches often 
in tbe axils of the distal leaves, the other branches subtended by sub­
ulate or narrowly ovate bracts 7-10 mm. long, I mm. wide; pedicels 
(2-) 3-6 (-9) em. long; calyculate bmcts 1(}-16, linear or narrowly 
triangular, 6- 11 mm. long; heads 6-15 (-20), 15-20 mm. long; phyl­
laries 12-14 (- IS), narrowly ovate or narrowly triangular, (9- ) 11-14 
mm. long, 2-3 mm. wide, greenish or purplish; receptacle ca. 7 mm. 
across; flowers 4(}-SO 01' Illore; corolla (7-) 8- 12 (-13) mm. long, the 
tube (3.5) 4-5 (-7) mm. long, the throat 2- 3 mm. long, 1-1.5 mm. 
across at the top, the lobes about as long as the throat, 1.5- 3 mill. 
long, strongly recurved; anthers 3-3.5 (-4) mm. long; style branches 
1-1.5 (-2) mm. long, recurved; mature achenes oblong or ellipsoid, 
subterete in cross section, 3.5 mm. long, 1-1.5 mm. wide, ca. 0.5-1 
mm. thick, glabrous, purplish brown, I O-ribbed; pappus white (7-) 
8-10 mm. long, distally clavate; chromosome !lumber, n=30 (Pippen). 
DISTRIBUTION: Durango, NIlyarit, and Jalisco, east to Michoacan 
and Mexico. Common on rocky slopes in the oak-pine woodlands. 
Flowering in September "nd October. 
Comparison of Oacalia. cordijolia. H.B.K. and P. sessilijolia. (Hook. 
& Arn.) Rydb. indicate that they fire conspecific. They differ only in 
the presence or absence of the petiole. Several specimens from Nayarit 
have both sessile and petiolate leaves on the same plant. The heads, 
flowers, and other morphological characters are similar. The sessile­
leaved plants are apparently vegetative variations that occur several 
times within tbe range of the petiolate forms. This difference is not 
significant enough for nomenclatural status. 
This species is a victim of nomenclatural niles. Since Oacalia cordijolia 
H.B.K. was superfluous when published because of the earlier O. 
cordijolia L. f., il. different taxon, the next oldest valid nallle for this 
taxon is Oacalia sessilijolia [Pericalia MSi.ilijolia (Hook. & Al'Il.) 
Rydb.]. This is unfortunate since the petiolate-Ie,,,,ed plants are by 
far more common than the sessile-leaved lorms. 
AUGUASCALIENTES : McVaugh & Kocu 147 (MICII); Pippen 5 (MICH). 
DISTRITO FEDER.U: Balla 5575 (US) i Bourgeau 715 (GH); Pringle 7983 (OR) j 
Lyonnet 194 (GH, NY, US); Purplls 5625 (OTI, NY, US). DURANGO : Gentry 6968 
(Gil, l\IICH, US); MeVaugh 21658 (MICH); Maysilles 7579 (MICH); Pennell 
18345 (NY, all of tho specimens from Durango are of slightly smaller sta.ture with 
more, smaller leaves). GUANAJUATO: Duges 481 (OlI, US). JALISCO : McVaugh 
13559, 13625, l7ll8, 17183, 18121 (MICll) ; MeV.ugh & Kae/z 920 (MICH); 
A{exia 1450 (US); Palmer 576 (GIf, NY, US)j Pippen 17, 18,31,33,34, 61.,64 
(MICH); Pringle 1736 (GH, MICH, NY, US), 9869 (GH, NY, US), 11497 (GH, 
MICH, US); &fford 1451 (US); /las, 2828 (GH, US). MEXICO: Matuda 21825 
• 
PIPPEN-MEXICAN "CACALIOID" GENERA 415 
(NY). MICHOAC.(N: ArB"', 8 .n ., Sept. 11, 1909, Sept. 14, 1911,5729,7347 (US); 
Feddema 161 (MICH); Pippe-n 51 (MICH). NAYARIT: Feddema 381, 1345 (MICH): 
McVaugh &: Ko.lz 479, 792, 656 (MICH); Mexia 635 (US); Ortega 5727 (US) ; 
Palmer 1832 (GIl, NY, US) ; Pippen 13, 15 (MICH) . Q UERETARO: Matuda 29787 
(NY) . SAN LUIS POTosi: Palmer 47 (GR, NY, US); Parry&: Palmer 541 (GH, 
US); Schaffner 255, 33 or 371 (G II). 
Three specimens in the herbarium at the University of Michigan 
do not fit precisely into either of the above taxa, P. 8essilifolia Or P. 
michoacana. Two of these, McVaugh 13469 and 13825 (from northeast 
slopes of the N evado de Colima, below Canoa de Leoncito, 2,25()-
2,550 m.), have leaves similar to P. michoacana in that they are sub­
circular, 7-lobed, and pubescent, but they are about 0.5 times larger 
with much longer petioles "nd the inflorescence has larger heads with 
more flowers. The phyllaries are finely pubescent with fuscous hairs 
and the heads are subtended by about 15 calyculate bracts as wide 
as but longer than the phyllaries. 
These specimens may represent a different species but, because of 
the paucity of material available and its rather close similarity to P. 
michoacana, I will only mention their existence for the present. 
The third nonconforming specimen, McVaugh 13862 (Sierra de 
Mananthtn, 15--20 miles southeast of Autlan, near Asseradero EI 
Cuart6n, 2,500 m .), differs from the above in that the leaves are less 
deeply lobed, less pubescent, and the trichomes are mostly restricted 
to the leaf margins; the phyllaries are glabrous, the subinvolucral 
bracts no longer than the phyllaries and somewhat narrower. This 
specimen approacbes P. sessilifolia but differs in the longer petioles, 
the more subeircular than ovate, 7-lobed leaves, and the presence of 
pubescence. Future collections and field observations in this region 
will be necessary to properly interpret this specimen. 
4. Psacalium 
PSQcalium Casso Dist. Sci. Nat. 43:461. 1826. 
Perennial, subscapose herbs, 0.1-3 m. tall; roots fleshy, fibrous; 
caudex tough, t.o 5 em. long, 3 em. tbick, the nodes and terminal 
buds densely covered with masses of tawny, multicellular hairs, 1-2 
em. long; stems usually solitary, annual, erect, slender or stout, 
usually terete, glabrous or pubescent; radical leaves forming a basal 
rosette of two to seveml, centrally peltate with several principal 
radiating veins, each pinnat.ely subdivided, the blades variously lobed, 
rarely subentire, m ostly pubescent; cauline leaves alternate, pro­
gressively smaller upward, the uppermost bracteiform, usually non­
peltate witb a broad lenfy-margined petiole and palmately 3- to 
7 -lobed blade; inflorescence paniculate or corym bose, tbe branches 
often stipitate glandular, with minute, multicellular glands, 0.5 mm. 
416 CONTRIBUTIONS FROM THE KATION AL HERBARIUM 
long, subtended by small, lincll.r.subulate or large, leafy, orten vagi. 
nate bracts; heads few to many, calyculate, the bracts 3-5, as long 
as the phyllaries or shorter (longer in P. pellatum) .. involucre cam· 
panulate, turbinate or less frequently cylindric"l, the phyllaries 
5-17, subequal, biseriate appearing uniseriat.e, the overlapped mar· 
gins subscarioUll, the exposed surfaces pubescent or glabrous, the 
tips fringed with minute trichomes up to 0.3 mm. long; receptacle 
flat or nearly so, alveolate, not paleaceous, occasionally sparsely 
pubescent; flowers all discoid, fertile, 5-80 per head; corollas glabrous, 
creamy or nearly white, occasionally greenish or purplish tinted but 
never yellow, the tube narrowly cylindrical, 0.5 mm. in diameter, 
the limb divided into 5, equal, narrowly.triangular, 3·nerved, ascend. 
ing or spreading, not recurved lobes, connate into a short, ampliate 
or campanulate throat, about 1 mm. long, 1-1.5 mm. wide (or occa· 
sionally lacking), the lobes longer than the throat and as long as to 
one·half as long as the tube; anthers 2-4 mm. long, exserted beyond 
the corolla throat, rounded or slightly sagittate at tbe base, the 
terminal appendages trullate or ovate 0.5-0.7 mm.long; style branches 
0.7-2 mm. long, slightly flattened, recurved or spreading, the stig. 
matic surface covering most of the adaxial surface, the abaxial surface 
glabrous or bispidulous below, the unappendaged tips truncate to 
obtuse with a fringe of nonglandular "pollen producing" trichomes; 
achenes obovoid or ellipsoid, somewhat compressed and elliptic or 
broadly elliptic in cross section, glabrous or pubescent, striate or 
prominantly 10· to 12.ribbed; pappus a single row of white or creamy, 
stiff, barbellate, capillary bristles (lacking in P. nanum and P. calvum); 
cbromosome number, n=30. 
TYPE SPECIES: Psacalium peltalum (H.n.K.) Rydb. 
This genus is easily recognized by tbe peltate basal leaves and is 
the only genlls discussed here with .tipitate glandular pubescence in 
addition to tbe usual nonglandular trichomes. 
Key to Species of Psacalium 
1. Inflorescence and upper portion or stem minutely stipitate gla.ndular; plants 
also pubescent with nonglandular hairs. 
2. Achenes pubescent. 
3. Corolla tube about twice as long as the lobes, 6-7 mm.long; hends nodding, 
about 40 or fewer; pappus 9-10 mm. long .... 3. P. holwayanum 
3. Corolla tube about the same length as the lobes, 3.5--4 (-5) mm. long; 
heads erect, 50 or morej pappus 6-7 mm. long ... 2. P. erlocarpum 
2. Achenes glabrous. 
4. Flowers 5-6 per hendj phyllarics 5, most1y glabrolls or very sparsely 
pubescent. . . . . . . . . . . . . . . . . . . . 11. P. tabulare 
4. Flowers 8 or more per head; phyllurics at least as numerous 1\8 the flowers, 
densely pubeacen t. 
PIPPEN-MEXICAN "CACALIOID" GENERA 417 
b. Ba.salleaves lobed one-half or less the distance to the center; phyllaries 
(7-) 8; flowers 8-10; bracts Bubtending the phyllaries either 3-5 and 
one-half as long as the phyllarics, or 1 or 2 and as long as the phyllaries. 
6. Lobes of the leaves 4 or 5; leaves 5-6 em. in diameter, pubescent on the 
adaxial surface; phyUaries subt.ended by 1 or 2 bracts as long as or 
longer than thep hyllaries. • . . . • . • . • • 4. P.laxiflorum. 
6. Lobes of the leaves 6 or 7; leaves 20-30 em. in diameter, glabrous on 
the adaxial surface; phyllaries subtended by 3-5 bracts about one­
half as long as the phyllaries. • • • • • • • 12. P. species 
5. Basal leaves lobed about two-thirds the distance to the center; phyllaries 
(8-) 12-15; flowers 20 or more; bracts subtending the phyllaries 3 or 4, 
longer than the phyllaries • • • . • . . . . . • • 8. P. peltatum 
1. Inflorescence and upper portion of the stem laCking stipitate glands; plants 
pubescent with nonglandular trichomes or glabrous. 
7. Plants with fleshy underground tubers attached to the caudex. 
B. Pappus present, 4-6 mm.long; flowers 5-B; phyllaries 5; plants 0.5 m. tall 
or taller . . . • • • • • • . • • • . • . . • • . 9. P. peltigerum 
8. Pappus lacking; flowers 10 or 11; phyllaries 7; plants up to about 15 em. 
tall. . . . . . . . • . . . . . • • • • • • . • . • 6. P. nanum 
7. Plants lacking underground tubers. 
9. Pappus lacking; achenes obovoid, 1(}-12 mm. long, 5-6 mm. wide; 
leaves deeply and finely dissected almost to the center, the major 
segments again divided, the ultimate segments linear. 1. P. ealvum 
9. Pappus present; achenes ellipsoid, up to 7 mm. long, 2 mm. wide; leaves 
lobed as deep as three-fourths the distance to the center, not deeply 
and finely dissected. 
10. Phyllaries densely pubescent. 
11. Bracts 8ubtending the phyllaries 3 or 4, longer than the phyllaries; 
phyllaries 8-10; flowers 12 or 13; receptacle naked, not pubescent. 
8. P. pelt.tum 
11. Bracts subtending the phyllaries 3-5, less than half as long as the 
phyllaries; phyllaries 5; flowers 5 or 6; receptacle sparsely pubescent. 
5. P. megaphyllum 
10. Phyllarics glabrous or very sparsely pubescent. 
12. Phyllaries 9-10 mm. long; achenes glabrous; leaves pubescent, 
densely so on the abaxial surface. 
13. Phyllaries 5; flowers 5 or 6; ranging from Veracruz west to 
Michoacan. . . . . . . . . . . . . . . . 11. P. tabulare 
13. Phyllaries B or 9, flowers 10-12; known only from central Oaxaca. 
7. P. nelsonll 
12. Phyllaries 5-7 mm. long; achenes pubescent; leaves glabrous or 
sparsely pubescent along the veins on the abaxial surface. 
10. P. poeuliferum 
1. Psaealium ealvum (Brand.) Pippen, comb. nov. 
Cacalia calva Brand. Univ. Calif. Pub!. Bot. 4:193. 1911. Holotype: Puehla; 
Cerro de Gavilan, August 1900, C. A. Purpus 4113 (UC, not seen; isotypes 
GHI, USI). 
Odontotrichum cal.um (Brand.) Rydb. Bull. Torrey !lot. Club 51 :420. 1924. 
Lower portion of the plant not seen; plants CR. 0.5 m. tall; stem 
and leaves densely pubescent to tomentose; flowering stem terete; 
418 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
basal (?) leaves 2 or 3, long-petioled, pelLute, ~nbcircular in outline, 
12-14 cm. in diumeter; bludes deeply dissected into 6-8 divisions 
extending almost to the center, the major divisions several times 
subdivided, the ultimate divisions narrow, linear; petioles slender, 
20-25 cm. long; cauline leaves reduced and bracteiform; inflorescence 
corymbo~e, the branches subtended by Jinear-subulate bracts to 1 
cm. long; heads about 20; phyllaries 12, narrowly ovate, 8 mm. long, 
1 mm. wide; flowers 12-14; corolla about 7-8 mm. long (flower meas­
urements from the original description), the tube 3-4 mm. long, the 
throat ca. 0.5 mm. long, 0.5-1 mm. fiCro~s at the apex, the lobes 4 
mm.long; anthers about 3 Illm. long; style branches recurved, hirtel­
lous; mature achenes obvoid, 10-12 mm.long, 5-6 mm. wide, glabrous, 
pale brown; pappus lacking entirely. 
DISTRIBUTION: Known only frOIll the type specimens. 
The specimens examined lacked adequate flowering material, and 
the poorly preserved leaves Illade it difficult to determine whether or 
not they were peltate. 
The most unusual features of this species are the extremely large, 
obovoid achenes and lack of a pappus. :Field examinations and study 
of material collected in the future may indicate a change in the 
taxonomic position of this taxon. 
2. Psaealium eriocarpum (8. F. Blake) Blake Journ. Wash. Acad. Sci. 28:492. 1938. 
Cacalia eriocarpa S. F. Blake Journ. Wash. Acad. Sci. 129. 1929. Holotype: 
Jalisco; Arroyo de Santa Gertrudis, San Sebastian, steep slopes in open 
pine and oak woods, 1,500 m., Jan. 21, 1927, Ynez Mexia 1539 (US!). 
Slender plants up to 1.3 m. tall, stems 5-10 mm. thick at the base, 
purplish green, sparsely or densely pubescent with long, multicellular, 
stiff, attenuate-tipped hairs, 1-1.5 mm. long, Itnd with multicellular, 
stipitate glands, 0.5 mm. long; leaves hirsnte with nonglandular hairs 
like those on the stem, the pubescence denser on the adaxial surface; 
basal leaves 2 or 3, long-petioled, centrally pcltate, subcircular in 
outline, 15-25 (-30) cm. in diameter, the venation radiate with 10 or 
11 principal veins; blades shallowly lobed one-third the distance to the 
center of the blade or less, thc lobes 8-10, 1.5-2 times broader than 
long, each 3-angulate or subdivided into 2 or 3 smaller lobules, the 
margins spinulosc-toothed or entire, the center of the blade flat, not 
depressed; petioles 15-30 em. long, slender; cauline leaves 1 or 2, 
greatly reduced, bractciform, inflorescence paniculate, usually :10 CIll. 
long or longer, 10-15 Clll. broad, the branches sub tended by narrowly 
triangular bracts 3-7 mm. long, 1 mill. wide; pedicel< 3.5-5 (-9) mill. 
long, calyculate bracts 3-5, subulate, 2-5 mill. long; heads 50-70 
(-90), 1.2-1.5 cm. long; phyllaries 5 or 8, narrowly ovate or elliptic, 
(6-) 7-8 (-9) Illm. long, 1-3 mm. wide, purplish tipped, pubescent, 
occasionally sparsely stipitate glandular; flowers 6 (when 5 phyllaries) 
PIPPEN-MEXICAN "CACALIOID" GENERA 419 
or 9-11 (when 8 phyllaries); corolla about 8-9 mm. long, the tube 
(3.5-) 4 (-5) mm. long, the throat short, campanulate, about 1 mm. 
long and broad, the lobes about as long as the tube, 3-4 (-5) mm., 
narrowly triangular; anthers 3-4 mm. long; style branches 1.5-2 mm. 
long, the abaxial surface hispidulous, especially distally, the tips 
rounded or obtuse with a sparse subterminal fringe of hairs; mature 
achenes not seen, the immature achenes pale green, densely villous; 
pappus (5.5-) 6-7 (-8) mm. long, white; chromosome number, n=30 
(Pippen). 
DISTRIBUTION: Western Jalisco. Mostly in pine and, less frequently, 
oak woodlands. Flowering mid-November to December. 
This species has two forms: the more common form has small heads 
with 5 phyllaries and 6 flowers, the other has larger heads with 8 
phyllaries and 9-11 flowers. These two forms are similar in all other 
aspects, including range. The larger headed form is known only from 
an area in between the known populations of the smaller headed form. 
These forms are not given nomenclatural status. This species is 
closely related to P. holwayanum but differs from it in the smaller 
heads, erect inflorescence, and less deeply lobed leaves. 
JALISCO: McVaugh 13954, 14287, 13684, 14165 (MICII); McVaugh & KoeZz 
919 (MICH); Pippen 36, 62*, 66, 67 (MICH). 
3. Psacalium holwayanum (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51: 372. 
1924. 
Cacalia holwayana B. L. Rob. Proc. Amer. Acad. 43: 45. 1907. Holotype: 
Michoacan; Uruapan, Oct. 11, 1899, E. W. D. Holway 2617 (GHI). 
Psacalium langlassei Rydb. Bull. Torrey Bot. Club: 51: 376. 1924. Holotype: 
Guerrero; Sierra Madre, in 1898, E. LanglG8s6 576 (UBI). 
Plants up to 1 m. tall; stems 3-5 mm. thick at the base, densely 
pubescent with stiff, white hairs up to 2 mm. long, 12- to 20-celled, 
and also with 6-celled stipitate glands, 0.1-0.2 mm. long, abundant 
distally; leaves sparsely or densely pubescent with stiff, nonglandular 
hairs only; blts.,1 leltves 2 or 3, long-petioled, peltate, subcircular in 
outline, (9-) 12-14 (-26) em. in diameter, the venation radiate with 
8 principal veins; blades lobed about one-third the distance to the 
center, the lobes 7 or 8, 2.5-4 cm. long, 3-6 cm. wide, each one 3-
angulate or divided into 2 or 3 smaller secondary lobes, the margins 
irregularly and sparsely toothed; petioles (9-) 10-18 (-32) em. long, 
terete; cauline leaves 2 or 3, similar to the basal ones but smaller, 1-5 
cm. in diameter, the uppermost subpeltate with petioles 1-5 (-10) 
cm. long; inflorescence thyrsiform, 16-30 cm. long, 14-16 cm. broad, 
the branches pubescent and stipitate glandular like the slem, sub­
tended by either linear bracts 1-2 em. long, 1 mm. wide or occasionally 
spathulate bracts, 3-toothed Itt the tip; pedicels 5-15 mm. long; 
420 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
calyculate bracts 3-5, subulate, 4-7 mm. long; heads about 40 or 
fewer, nodding, 1.5-3 cm. long; phyllaries 8, narrowly ovate, 9-13 
mm.long, 1.5-3 mm. wide, purplish, pubescent with glands and sparse­
ly hirsute with nonglandular, long, stiff hairs; receptacle 3-3.5 mm. 
long; flowers 10 or 11; corolla 10-11 mm.long, the tube 6-7 mm.long, 
the throat short, campanulate, ca. 1 mm. long, 1.5-2 mm. across, 
the lobes 3-3.5 (-5, Pringle 13294) mm. long, linear, 0.5 mm. wide, 
anthers 3.5-4 mm. long; style branches 1.5-2 (-3) mm. long, hispid 
on the abaxial surface, the tips truncate with a terminal fringe of 
trichomes; mature achenes 5-6 mm. long, 1.5 mm. wide, 0.5-0.7 mm. 
thick, oblong, or ellipsoid, pale, densely pubescent with unicellular, 
adpressed, ascending hairs; pappus 9-10 mm.long, white; chromosome 
number, n=30 (Turner, ined.). 
DISTRIBUTION: Known from west central Michoacan eastward to 
Mexico and from one collection in Guerrero. Mostly in pinewoodlands. 
Flowering mid-November to December. 
An isoparatype, Langlasse 576 (US), was described by Rydberg as 
Psacalium langla8sei, because the leaves were less deeply lobed. Com­
parison of this specimen with the type and other specimens of P. 
holwayanum indicates that they are conspecific because they are 
similar in all characters except loaf lobes. The specimen Rydberg 
examined was merely less deeply lobed. 
GUERRERO: Langlass~ 576 (GII, paratype of C. holwayana, isotype of P. 
langlass.". MEXICO: Gilly 5 (MICH); Maluda 30015 (NY). MICHOACAN: 
King &; Soderslrom 4890 (MICH); Pringle 13297 (GH, US), 13672 (GIl, para­
type; US), 13979 (GH, MICH, US); McVaugh 21953* (MICII). 
4. Past:alium laxiflorum Bcnth. PI. Hnrtw. 41. 1841. Holotype: l\Hchoac:in; 
Marclia, pine woods, in 1836, Hartweg 318 (K, not seen; Field Mus. Neg, 
379491) . 
Senecio moreliae Hemsl. BioI. Centro Amer. Bot. 2:243. 1881. Based on 
Psacalium laxiflorum Benth. not S.laxiflorum Vivo 
Slender plants 6-7 em. tall; stems 3-4 mm. thick at tbe hase, 
pubescent with multicellular stiff hairs, 1-1.5 mm. long, and minute, 
2- to 3-celled stipitate glands ca. 0.1 mm. long, most abundant dis­
tally; leaves pubescent on the adaxial surface with nonglandular 
hairs, subglabrous abaxially; basal leaves 2 or 3, petiolate, peltate, 
subcircular in outline, 5-6 cm. in diameter, the venation radiate with 
5 principal veins; blades lobed about one-half the distance to the 
center, the lobes 4 or 5, 1-2 cm. long and broad or occasionally 1.5-2 
times broader than long, each slightly 3-angulate, the tips rounded 
or obtuse, the margins mucronulate-toothed, the sinuses broadly 
rounded; petioles 6-8 em. long, terete; cauline leaves usually 2, the 
lowermost similar to the basal ones, the upper one nonpeltate, 2.5-4 
em. long, 1-2 cm. wide with the petiolar portion leafy-margined, 
I 
PIPPEN-MEXICAN "CACALIOID" GENERA 421 
ovate, the hlade portion 3-lobed; inflorescence paniculate, 16-26 cm. 
long, 11-12 cm. broad, the branches pubescent and stipitate glandular 
like the stem, subtended by narrowly ovate or ovate bracts 2-2.5 cm. 
long, 0.5-1 cm. wide progressively smaller upward, the bracts glan­
dular on the adaxial surface, glabrous abaxially; pediceis 5-10 mm. 
long; calyculate bracts 1-2, subulate, 8-10 mm. long; heads 30 or 
fewer, 14-15 mm. long; phyllaries 8, narrowly ovate, (6-) 7-8 mm. 
long, 1.5-3 mm. wide, purplish, densely stipitate glandular; flowers 8; 
corolla 8-9 mm. long, the tube 5-5.5 mm. long, the throat ampliate, 
0.5 mm. long, 0.7-1 mm. wide, the lobes 3-3.5 mm. long, linear; 
anthers 3 mm. long; style branches 1.5 mm. long, hispid on the 
abaxial surface, the tips truncate or rounded with a terminal or 
subterminal fringe of trichomes; mature achenes not seen, immature 
achenes glabrous, striate; pappus 6-7 mm. long, white. 
DISTRIBUTION: Known only from the type collection and one col­
lection from southern Durango (Rose 2337, GH, US). In pine-wood­
lands. Flowering in August. 
This species has smaller leaves than the other members in the 
section, and they tend to be 4-lobed and somewhat cross shaped. It 
is related to P. holwayanum and P. eriocarpum. 
5. Psaealium megaphyllum (Rob. & Greenm.) Rydb. Bull. Torrey Bot. Club 51: 
374. 1924. PLATE 6 
Cacalia megaphylla Rob. & Greenm. Amer. Journ. Sci. III. 50:157. 1895. 
Hoiotype: Jalisco; on hillside near Guadalajara, Oct. la, 1889, C. G. 
Pringle 2490 (GHI). 
Cacalia obtusiloba Rob. &: Greenm. Amer. Journ. Sci. III. 50:158. 1895. 
Holotype: Oaxaca; Sierra de San Felipe, 6,000 ft., Nov. 17, 1894, C. G. 
Pringle 5840 (GHI). 
Psacolium obtusilobum (Rob. &: Greenm.) Rydb. Bull. Torrey Bot. Club 
51:374. 1924. 
Stout plants to 3 m. tall; entire plant pubescent, the pubescence 
denser in the inflorescence and on the abaxial surface of the leaves, 
the hairs 0.5-0.7 mm. long, 5- to 15-celled, white; stem to 2 cm. 
thick at the base; basal leaves 3 or 4, long-petioled, peltate, subcircular 
in outline, 30-60 cm. in diameter, the venation radiate with 5-7 
principal veins; blades lobed one-half to two-thirds the distance to 
the center, the primary lobes 5-7, 7-15 cm. long from the base of 
the sinus to the distal edge, 5-7 cm. wide at the base, each usually 
divided into 3 lobes, the secondary lobes (5-) 7-10 Cill. long, 3-5 cm. 
wide at the base, each divided into several smaller, triangular lobules 
1-3 cm. long, the margins coarsely toothed, the center of the blade 
depressed; petioles 25-80 cm. long, terete, purplish green; cauline 
leaves 5-7 or more, the lowermost usually smaller than the basal 
ones but similar to them, 3D-35 cm. in diameter, the upper cauline 
leaves progressively smaller upward, subpeltate or nonpeltate, the 
422 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
uppermost 8-15 cm. long, 10-20 cm. wide, palmately 3- to 5-lobed, 
the petioles 3-7 cm. long; inflorescence broadly paniculate-corymbose, 
to 1.5 m. high, ca. 1 m. broad, the proximal branches in the axils of 
the uppermost leaves, the distal branches subtended by ovate bracts 
2-5 cm. long, 3-7 mm. wide; pedicels about as long as tbe phyllaries; 
calyculate bracts 3-5, subulate, 1.5-2 mm. long; heads at least 150, 
10-12 mm. long; phyllaries 5, narrowly elliptic or narrowly ovate, 
(5.5-) 7-8 (-10) mm. long, (1.5-) 2-3 mm. wide, densely white 
puberulent; receptacle sparsely pubescent with filiform, multicellular 
hairs; flowers 5 (-6); corolla 6-8 mm. long, the tube 3-4 mm. long, 
the throat 0.5 mm. long and broad (or occasionally lacking), the 
lobes narrowly triangular, 2.5-4 (-5.5) mm. long, reflexed; anthers 
2-3.5 mm. long; style branches 1-1.5 mm. long, the abaxial surface 
hispid, denser distally, the tips obtuse with a sparse terminal fringe 
of trichomes; mature achenes oblong or ellipsoid, 5-5.5 mm. long, 
1.5-2 mm. wide, 1 mm. thick, pale green, glabrous, 15- to 20-ribbed; 
pappus 4-5 (-7) mm. long, while; chromosome number, n=30 
(Turner, ined.). 
DISTRIBUTION: From Zacatecas southeast to Oaxaca. Common, at 
least in western Mexico, in rocky grasslands, roadsides, and grazed 
areas. Flowering in September and October. 
This species is striking because of its very large size and especially 
its extremely large basal leaves, depressed in the center. Most existing 
herbarium specimens do not adequately represent the species because 
even one leaf is larger than the average herbarium sheet. 
Robinson and Greenman in describing Cacalia megaphy/la and C. 
obtusiloba stated that the btter differed from the former in having 
smaller heads and more rounded basal leaf lobes. Comparison of the 
types of these two species, however, indicates that they are conspecific 
because of several common characters: Number of phylluries and 
flowers, pubescence of phylluries, leaves and stem, general habit, and 
lenf shape (the latter somewhat variable). 
AGUASCALlENTES: McVaugh & Koelz 39 (MIeII); Pippen 8* (MICH). GUER­
RERO: MeVaugh 21897, 21910 (MICH); Moore 5119 (MICH). JALlSCO: MeVaugh 
13321 (MICH); Pippen 16, 24, 39 (MICH). MEXICO: IIinton 2200 (US), 2355 
(NY); Matuda 26572 (NY). MleIIOAC.(N: Arsene 2386 (US); llinton et al. 13480, 
15629 (MICH, NY); Pippen 47 (MICH); MeVaugh 21928 (MICH). MORELOS: 
Pringle 6165 (GU, NY, US). NAYARIT: lIfexia 877 (GU, MICH, UH). PUEBLA: 
Ar3ene &:: Nicholas 334 (US)j Arsene, B.n., November 1908 (US). ZACATECAS: 
MeVaugh 17650 (MICII). 
6. Psacalium nanum Pippen, sp. nov. PLATES 7, 8 
Plantae nnnae usque ad 12 em. altae, tuherihus pluribus, carnosis, 
fusiformibus, subterraneis, cliulibllS hil'~;utis scaposis, foliis l'adiealibus 
peItatis, suborbieularihus, usque ad 5 em. diametro, profunde 7-
I 
PIPPEN-MEXICAN "CACAIJOID" GENERA 423 
lobatis, supra hirsutis, subtus glabris, petiolis tenuibus 5-10 em. 
longis hirsutis; inflorescentia corymbosa, 2-3 em. longa latasque, 
bracteis foliosis, trullatus, usque ad 2 cm.longis; capitula usque ad 25, 
ca. 7 mm. longa; phyllaria 8, elliptica 3.5 4.5 mm. longa, glabra ; 
flores 10-11, corollis 3.5-4 mm. longis, tubo 1.5-2 mm. longo, lobis 
2-3 mm. longis; achaenia matura non visi; pappus nullus. 
Plants scapose, up to 12 cm. tall with several fleshy, elongate, 
fusiform tubers attached to the caudex beneath the soil; stems, 
petioles, and adaxial surface of the leaves hirsute with tawny multi­
cellular hairs to 1.5 rom. long, the abaxial surface of the leaves gla­
brous; stems slender, 2 mm. thick at the base; basal leaves 7-15,long­
petioled, centrally or subcentrally peltate, subcircular in outline, 
to 5 cm. across, the venation radiate with 7 principal veins; blades 
deeply lobed one-half to two-thirds the distance to the center of 
the blade, the primary lobes 7, each divided into 3 major lobes and 
2 or 3 lesser lobules, the secondary lobes often again lobed, the ulti­
mate segments triangular, mucronate; petioles slender, 5- 10 cm. 
long; inflorescence corymbose, 2-3 cm. long and broad, tightly com­
pact, the primary branches subtended by petiolate, leafy bracts up 
to 2 em. long, the blade portion obtrullate; pedicels mostly glabrous ; 
calyculate bracts usually 3, subulate, up to 1.5 mm. long; heads 25 
or fewer, about 7 mm. long; pbyllaries 7, elliptic, 3.5-4.5 mm. long, 
0.7-1 mm. wide, glabrous ; flowers 10 or 11; corolla, 0.7-1 moo. wide, 
elliptic, strongly 3-nerved, the throat none; anthers 2 mm. long; style 
branches 0.7 mm.long, hispid on the abaxial surface, the tips truncate 
or rounded with a terminal or subterminal fringe of trichomes ; 
mature achenes not seen, the immature achenes brown, glabrous, 
lO-ribbed; pappus en tirely lacking. 
HOLOTYPE: Guerrero; Teitepec (Mina Distr.). Open stunted pine 
forest, on mossy rocks, 3,600 m. (ace. to tho label) , July 17, 1939, 
G. B. Hinton et al. 14464 (US!, isotypes MICH!, NY!). 
DISTRIBUTION: Known only from the type collection. Flowering in 
July. 
This species has affinities with both P. peltigerum (through the 
underground tubers) and P. calvum (through the lack of a pappus). 
The smill size makes this species unique in this genus. 
7. P .. eallum nol800I1 Rydb. Bull. Torrey Bot. Club 51:374. 1924. Holotype: 
Oaxaca; vicinity of Cerro de San Felipe, 9,50~11,OOO ft., in 1894, E. W. 
NellOn 1111 (US!). 
Roots, base of plant and stem below the inflorescence not seen; 
leaves, presumably basal, petioled, peltate, subcircular in outline, 30-
35 cm. in diameter, the venation radiate with 7 or 8 principal veins; 
blades lobed about one-half the distance to the center of the hlade, 
slightly longer than wide, each divided into 2 or 3 secondary lobes 
424 CON1'RIBUTIONS FROM THE NATIONAL H e:RBARIOM 
which are obtusely deltoid, mucronate-tipped, the adaxial surface 
pubescent with multicellular hairs up to 1.5 mm. long, the abaxial 
surface gray-tomentose; cauline leaf (only one seen) similar to the basal 
leaves but subpeltate, the petiole attached 2-3 mm. above the base of 
the blude, the blade 8-lobed; inflorescence paniculate, at least 35 cm. 
long, 15 cm. broad, the branches purple-tomentose, the tomentum 
denser toward the heads, the branches subtended by leafy bracts, 4-10 
cm. long, obovate, palmately 3- to 7-lobed or subentire; calyculate 
bracts about 5, linear-subulate, 4-5 mm. long; heads many, about 12 
mm. long; phyllaries 8 or 9, elliptic, narrowly ovate or oblong, 10 mm. 
long, 2-3 mm. wide, glabrous, purplish tinted; flowers mostly immature, 
10-12; corolla about 7 mm. long, purplish tinted when dry, the tube 
3.5 4 mm.long, tho throat 0.5-1 mm.long, 1-1.5 mm. broad, the lobes 
3 mm. long; anthers 2-3 mm. long; style branches 1.5 mm. long; 
mature achenes not seen, the immature achenes glabrous; pappus 
white, about 5 mm. long. 
D,STRIBUTION: Known only from the type specimen. Flowering 
time unknown. 
The type collection consists only oC a basal leal and a portion oC the 
inflorescence. It is apparently related to P. pellatum because of the 
lealy inflorescence bracts, the number ol phyllaries and Bowers, and 
the glabrous achenes. It differs Crom P. peltalum because oC the gla­
brous phyllaries, short subinvolucral bracts, tomentose condition of 
the leaves, and the outline of the basal leaves. This specimen appears 
to be significantly different from any oC the known species of Psacalium 
and therelore I have followed Rydberg in recognizing it as a distinct 
species, despite the incomplete condition oC the type specimen. 
8. Psacalium pelt.tum (H.B.K.) Casso Diet. Sci. Nat. 43:461. 1826. 
Cacalia pcltaJa H.B.K. Nov. Gen. et Sp. 4:170. t. 361. 1820. (Folio cd. 4:133. 
1820.) 
Senecio peUijeruI Hemsl. BioI. Centro Amer. Bot. 2:245. 1881. Based on 
Cacalia peleato H.B.K., not S. peUatu. DC. 
Plants up to 2.5 m. tall; stems and leaves sparsely to densely pilose 
or hirsute with multicellular, conical-attenuate nonglllndular hairs, 
0.5-2 mm.long, the crosswalls oCten purple, the stems also occasionally 
densely covered with minute, multicellular stipitate glands, 0.3~.5 
mm. long, (lacking in var. pella/um); s tems stout, terete; basal leaves 
3 or 4, long-petioled, peltate, subcircular in outline, 10-30 (-65) em. 
in diameter, the venation radiate with 6-8 principal veins ; blades 
deeply lobed about two-thirds the distance to the center, the primary 
lobes 7-9, each sinuately 5- to 7-lobed, the distal secondary lobes orten 
larger than the others, again lobed or 3-angulate; lowermost cauline 
leaves similar to the basal ones and about as large; upper cauline 
PIPPEN-MEXICAN "CACALIOID" GENERA 425 
leaves progressively sma.lJer upward, subpeltate, the uppermost non­
peltate with leafy-margined petioles, the blade palmately 3- to 5-
lobed; inflorescence racemose, paniculate or corymbose, the branches 
densely stipitate-glandular, sub tended by sessile, leafy bracts, ovate 
or narrowly ovate, less often linear; heads 10-100, 1.5-2.5 cm. long, 
phyllaries 8-10 or 12-14, pubescent; the involucre subtended by 3 or 
4 narrowly ovate or linear bracts longer than the phyllaries; flowers 
12-20 (-40); corolla about 9-14 mm.long, the tube 6-10 mm. long, the 
throat ampliate, 0.5 mm. long, 1-1.5 mm. broad, the lobes narrowly 
triangular, 3-4 mm. long; anthers 3-3.5 (-4) mm. long; style branches 
1.5-2 mm. long, recurved, hispid on the abaxial surface, tbe tips 
truncate or obtuse with a sparse subterminal fringe of trichomes; 
mature achenes oblong or ellipsoid, 5.5-7 mm. long, 1-1.5 mm. wide, 
0.7-1 mm. thick, glabrous, creamy, multistriate; pappus 6-8 (-9) mm. 
long, white. 
This is a variable and complex species usually distinguisbed by the 
sessile, elliptic or lobed inflorescence bracts, the 3 or 4 long, calyculate 
bracts subtending the phyllaries, the creamy, glabrous achenes, and 
the usua.lJy deeply lobed basal leaves. 
Within this complex are three sma.lJer groups, recognized as varieties, 
which seem to maintain themselves both morphologica.lJy and geo­
graphically. 
Of several other specimens that are apparently a part of this com­
plex, two are quite similar to var. conzattii and are discussed with 
that taxon. Another specimen is unique (Guerrero, between Petlacala 
and Buenavista, G. B. Hinton 14877, MICH, NY) and is from a 
poorly explored region where no specimens of P. pellalum have been 
collected. This specimen has much less deeply lobed leaves with 
much broader lobes than P. pel/atum but with an inflorescence like 
var. pellalum, except it is glandular-pubescent. More material from 
this region as well as field observations will be necessary to determine 
the best taxonomic position of this plant. 
Key to Varieties of Psacalium peluuum 
1. Stem and inflorescence glandular pubescent, the glands stipitate, 0.5 mm. 
longj plante also pubescent. with long, stifl', Donglandular hairs. 
2. Heads few, ca. 10, 2- 2.5 em. long; calyculate bracts 15-17 (-25) mm. long i 
pappus bristles 7- 9 mm. long; flowering in September and October. 
8e. P. peltatum var. conzattii 
2. Heads many, at least 50, ea. 1.5 em. long; calyculate braets 20-35 mm. 
tong, pappus bristles 6-7 mm. long) flowering in July and August. 
8b. P. peltatum var. adenophorum 
1. Stem and inflorescence lacking atipitate glands but sparsely to densely b.irsute 
or pilose with Donglandular hairs ... Sa. P. peltatum var. peUatam 
426 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
8a. Psacalium peltatum var. peltatum 
Caca.lia peltata 1I .B.K. as to hoiotypc: .MicboacRn i in wood!:! around Pazcuaro 
[Plitzcuaro1, 1130 hex" Humboldt & Bonpland 4346 (P, not seeD; Field 
l\! us. Neg. 378731). 
Caralia peUata var. couUeri GrL'eom. Proe. Amer. Acad . 40:51. 1904, Lectot.ype: 
Hidalgo (Veracruz on the label) ; Rcal del Monte, Th. Couller 420 (Gill). 
Psacalium eouUeri (Greenm.) Rydb. Bull. Torrey Ilot. Club 51 :373. 1924. 
P8acalium argu/um Rydb. Bull. Torrey Bot. Club 51 :373. 1924. Jlolotypc: 
San Lui. Poto." in 1879, J. G. Schaffner 725 (NY I) . 
Stems and leaves sparsely or densely pilose or hirsute ,,~th non­
glandular, multicellular trichomes, 0.5-2 mm. long, the stem "nd 
inflorescence not stipitate glandular; basal leaf lobes (7-) 10-12 
(- 20) cm.long, 3-5 cm. wide at the base, the sinuses obtuse or brolldly 
rounded; petioles (10-) 20--40 em. long, t,he petiole of the lowermost 
cauline leaf more dilllted than those of t,he I",sttl leayes, and lenfy ­
marginedj inflorescence panieulatc or corymbose, u.bollt 1 Ill. long 
and broad, the leafy brac!..q oYate, 3-4 (-7) cm. long, 2- 3 cm. broad, 
occasionally 3-lobed llear the distal end; calyculate uraets nurrowly 
ovate or ovate, (1 1-) 12-17 (- 25) nun. long, one of these usually 
broader thun the others surrounding the sume head; heads many, 
CR. 15 mm. long; phyllaries 8-10, narrowly oyute, 10- 12 (-15) mm. 
long, 2-3 nun. wide, pubescent but not stipitllto glandular; 80wers 
12-17 (-20); corolla 8.5-10.5 mm. long, the tube (5.5-) 6-7 mm. 
long, the lobes 3-3.5 mm. long, I Illm. wide; anthers 3.3- 5 mm. long, 
pappus 7- 8 mm. long; chromosome number, n=30 (Pippen). 
DISTlllBUTION: l<rom southern Chihuahua south through Durango 
to Jalisco, eastward across the Tl'I1lls-Mexiciln volr.anic belt in 
Miehoacan and Mexico to Puehla. Grasshmd openings and borders 
of the pine woodlands. Flowering September and October. 
The pubescence vllries from sparsely short puberulen t to densely 
pilose or villous. Greenmun di:.; tingu lshed the villo us ~ peeimell :-; a:; 
var. coulteri. Apparently this WIlS the only character used and it 
docs not appeal' to be consistent or correlated with any other chul'ue­
ters, therefore, I have not recognized the var. coulteri. 
Rydberg dist,inguished J>sacalium argui.·um, it group of r-:;pccimcn:'i 
from San Lui.s Potosi, Il.S differing (rom P. peltatum hy the presence 
of longer calycnh,te bracts Ilnd 1II0re sharply pointed basal leaf lobe.s. 
Although such n. tendency is noted ill these spec·imens, it is neither 
l'.onsllitent nor restricted to this rChrioll . There ure no other significallt 
differences in the'e specimens and 1 ba" e not supported this .cpam­
tion. 
CHIHUAHUA: Gentry, Correll, & Arguelle..'f 180 16 (US). DURANGO: NelsQn 4954 
(N Y, US); Pennell 18343 (NY) ; [ berra Garcia 414 (US). DISTR1TO FJomERAL: 
Matuda 26175, 26632 (NY). HIDALGO: Pring" 9870 (GH). JALI SCO: MeVaugh 
13821 (l.nCII). Mtxlco : H inl.n 1908 (US); Matuda 28343,29483,29591 (N Y); 
Ros, &: Painter 7939 (GH, US, paratypes of v. coulteri Greeum.) j Lllonnd 3122 
• 
• 
• 
, 
PIPPEN-MEXICAN "CACALIOID" GENERA 427 
(US). MWHOAcAN: Nel80n 6582 (GH, NY, US); Pippen 44, 49', 53 (MICH); 
Pringle 3340 (GH, NY, US); Seier 1259 (GH, NY, US). MORELOS: Lyonnet 794 
(US); Pringle 9871 (GIl, MICH, NY, US). PUEBLA: Beaman 3619 (TEX); 
Purpu8 3038 (NY, US). SAN LUIS POTosI: Schaffner 294 (GH); Parry &: Palmer 
543 (GIl, NY, US). TLAXCALA: Sharp &: Hernandez X. 44508 (NY). VERACRUZ: 
Seaton B.n., August 1891 (GH, paratype of v. cQulteri Greenm.). 
8b. PaaeaHum peitatum var. adenophorum S. F. Blake, Journ. Wash. Acad. 
Sci. 32:150. 1942. Holotype: Nuevo Leon: Municipio de Rayones, upper 
west slopes of Sierra de Cebolla, above 2,750 m., Aug. 21, 1939, C. H. 
Muller 2911 (U81). 
Plants ca. 1.5 m. tall; stems and leaves sparsely or densely pilose 
or hirsute with multicellular, nonglandular hairs, the stem and in flores­
cense also densely covered with minute stipitate glands; stem 5-9 
mm. thick at the base; leaves similar to those of the typical variety; 
inflorescence paniculate or corymbose, ca. 30 cm. long, 11-17 cm. 
broad, the bracts up to 6 em. long, 1.5 cm. broad; calyculate bracts 
2.0-3.5 cm. long, 3-10 mm. wide; heads ca. 40, about 2 em. long; 
phyllaries (8-) 12 or 13, narrowly ovate, (10-) 15-16 mm. long, 2-3 
mm. wide; flowers ca. 20; corolla 13-14 mm. long, the tube (7-) 
9-10 mm. long, the lobes ca. 4 mm. long; anthers 3-3.5 mm. long; 
mature achenes not seen; pappns 6-7 mm. long. 
DISTRIBUTION: Known only from the southern region of Nuevo 
Leon and Tamaulipas. Flowering in July and August. 
This taxon is known from only one collection in addition to the 
type (Tamaulipas: Stanford, Taylor, &: Lauber 2535, NY, TEX, US). 
The longer subinvolucral bracts and presence of stipitate glands 
separate it most easily from the typical variety. Time of flowering 
and geographical range separate it from var. conzaUii. 
Be. Psacalium peltatum var. conzaUii (Rob. & Greenm.) Pippen, comb. nov. 
Cacalia peltata var. conzattii Rob. &: Greenm. Proc. Amer. Acad. 32:49. 
1896. Lectotype: Oaxaca; Sierra de Sa.n Felipe, 10,000 ft., Dec. 13, 1895, 
C. G. Pringle 6238 (GH!). 
Psacalium conzattii Rydb. Bull. Torrey Bot. Club 51:372. 1924. 
?Psacalium mollifolium S. F. Blake, Journ. Wash. Acad. Sci. 28:491. 1938. 
Ho\otype: Guerrero; Taxco, Aug. 12, 1937, Ruth Q. Abbott 353 (GHI). 
Plants usually 1 m. tall or less; stems and leaves sparsely or densely 
pilose or hirsute with nonglandular trichomes, these especially dense 
on the abaxial surface of the leaves, the stem and inflorescence also 
densely stipitate-glandular; stem 4-10 mm. thick at the base; basal 
leaves subcircular, 14-25 cm. in diameter; blades deeply lobed usually 
more than three-fourths the distance to the center, the primary lobes 
4-10 em. long, 2-4 cm. wide at the base; inflorescence subpaniculate 
10-20 (-30) cm. long, 11-16 cm. broad, the branches sub tended by 
bracts about 3 em. long, 0.7-1.5 em. wide; calyculate bracts narrowly 
elliptic, 15-17 (-25) mm. long, 1-2.5 mm. wide; heads usually 10 
266-836-68-8 -
428 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
(-30),2-2.5 em. long; phyllaries 12-15, narrowly ovate, 12-15 (-IS) 
mm.long,2-3 (-4) mm. wide, pilose with nonglandnlar hairs, stipitate­
glandular; fiowers (20-) 25-40; corolla 10-12 mm. long, the tube 
(6-) 7- S (-9) mm. lung, the lobes 3.5-4 mm. long, 1 mm. wide; an­
thers 3- 3.5 (-4) 111111 . long; lIIature achene. 6-7 mm. long, 1-1.5 mm. 
wide, ca. lO-ribbed; pappus 7-S (-9) mm. long. 
DISTRIBUTION: Known mostly from central Oaxaca and question­
ably from Guerrero and Durango. Openings in pine wood lands. 
Flowering in September and October. 
Two specimens were cited in the original description as syntypes: 
Oaxaca; Cerro de San Felipe, 9,000 ft., Nov. 29, 1895, O. Oonzatti 
27 (GH!), Dec. 13, 1895, 0. G. I'ringle 6238 (GH! ). Despite the fact 
the taxon was named for Conzatti, possibly indicating his specimen 
as the lectotype, Pringle 6238 is a more com plete and more repre­
sentative specimen and is thus a better choice for the lectotype. 
Two specimens have been studied that are obviously closely 
related to this taxon but difTer in certain respects. One specimen 
(15 mi. W. of El Saito, U. T. Waterfall 12688, MICH) has heads 
and stipitate glands like val'. conzauii but the leaves are smaller 
with obovate lobes; most significant is its uccnnence in Dllrango 
which is quite far from Oaxaca. SlIch a separation in range is difficult 
to explain if this specimen is a p"rt of this taxon. 
The other specimen (Guerrero: Taxeo, Ruth Q. Abbott 353, GH) 
was the basis of Psacalium mol/ijalium S. F. Blake. It differs from 
var. conzattii in that the basal leaves are lobed less th"n half the 
distance to the center and the ab"xial surface is densely tomelltose. 
The inOorescenC6 is similar to val'. conzaUii, including the long calycu­
late bracts. These differences do not appear significant enough to 
give this specimen nomenclatural status; at least not until field 
observations and study uf future material call be made. 
OAXACA: Pringle 6238 (US, isotype); Camp 2401 (NY); MeVaugh 21826 
(MICH). 
9. PBacalium peltigerum (Rob. & Scat.) Rydb. Bull. T orrey Bot. Club 51: 314. 
1924. 
Cacalia peltigera Rob. & Seat. Proc. Amer. Acad. 28:111. Ma.y 13J 1893. 
Plants up to 1.5 Ill. tall with several fleshy underground tubers up 
to 5 CIII. long, 3 em. thick attnched to the eaudex; stems 5-12 mill. 
thick at the base, mostly glabrolL' or sparsely hirsute; leaves spursely 
or densely pubescent with stiff, white hairs up to 1 mIll. long; busul 
leaves 2 or 3, !ong-petioled, peltate, subcircuiar in outline, (10-) 20-30 
cm. in diameter, the venation radiate with 7-9 (-12) principal veins; 
bludes anguiate or lobed one-third to four-filths the distance to the 
center, the primary lobes 9-11, each divided into 2 or 3 maj or lobes 
and several lesser lobules, the secondary lobes usually lobed or coarsely 
, 
PIPPEN-MEXICAN "CACALIOID" GENERA 429 
toothed, or if the blade angulate the margins mucronate-tipped; 
petioles 29-30 em. long, pubescent proximally, becoming glabrous to­
ward the blade; cauline leaves 3 or 4, the lowermost similar to the 
basal ones but smaller; upper cauline leaves progressively smaller up­
ward, either nonpeltate with the blades 3- to 7-palmately lobed or 
remaining peltate and angulate, not lobed; inflorescence mostly corym­
bose, 9-30 em. long and broad, the branches subtended by linear­
subulate bracts, 1.5-2 em. long, 1-2 mm. wide; pedicels 1-2 em. long; 
calyculate bracts 3-5, subulute, 2 (-5) mm. long; heads many, 12-15 
mm. long; phyllaries 5, narrowly oblong, narrowly elliptic, or occa­
sionally spathulate, (6-) 7-8 (-10) mm.long, 1-2 mm. wide, glabrous; 
receptacle 1-1.5 mm. across; /lowers 5 or 6; corolla 9-10 mm.long, the 
tube (3.5-) 4-5 mm. long, the lobes as long as the tube or slightly 
longer, narrowly elliptic, the throat none; anthers 3-3.5 mm. long, 
style branches 1-1.5 mm. long, distally hispid on the abaxial surface, 
the tips truncate or rounded with a subterminal or terminal fringe of 
trichomes; mature achenes ellipsoid, 4-5.5 mm.long, 1.5-2 mm. thick, 
glabrous, tan, prominently 10-ribbed; pappus (4-) 5-5.5 (-6) mill. 
long, creamy or tawny. 
This species is easily distinguished by the presence of fleshy sub­
terranean tubers. Microscopic examination of freehand sections reveals 
that these tubers are modified stems with discrete vascular bundles. 
Druse-shaped crystals were found in the parenchyma cells. In green­
house-grown plants these tubers formed during the first season of 
growth and persbted through the dormant period. There is no evidence 
that they function in vegetative propagation as do the tubercles in the 
genus Pericalia. 
Three varieties are recognized and may be separated primarily by 
the outline of the basal leaves and geographical distribution. These 
three varieties exhibit all interesting range in leaf shapes from almost 
entire (var. hintonii) , to broadly lobed (var. laxi,wbum), to deeply 
linear-lobed, the lobes five times longer than broad (var. peltigerum). 
Key to Varietie. of Psacalium pehigerum 
1. Basal leaves angulatc, not lobed, the margin with mucronate tips j stem hie· 
aute . . . . . . . . . . . . . . . . . 9b. P. peUia:erum va.r. hintonii 
1. Basal leaves lobed at least half the distance to the center, the margins toothed, 
the teeth triangular, not mucronate; stems mostly glabrous or very sparsely 
hirsute. 
2. Lobes of the basal leaves extending two-thirds to four-fifths the distance 
to the center of the blade, 1-2 em. wide at the base, much longer than 
broad, the ultimate segments of the lobes narrowly triangular or linear­
attenuate, about 5 times longer than wide; stem glabrous. 
9 •. P. pcltigerum var. peltigerum 
430 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
2. Lobes of the basal leaves extending less than one-half the distance to the 
center of the blade, 5-7 em. wide at the base, about 8'3 long as wide, the 
ultimate segments of the lobes obtusely deltoid or triangular, about as 
long as broad; stem hirsute at least on the proximal half. 
9t. P. peltigerum var. latilobum 
98. Psacalium peltigerum var. peltigerum 
Cacalia peitigera Rob. & Seat. as to IIoiotype: Jalisco i bluff of the barranca 
near Guadalajara, Sept. 14, 1801, C. G. Pringle 5154 (GIl!). 
Blades of the leaves deeply lobed two-thirds to four-fifths the 
distance to the center, the primaq lobes 9-1J, 1-2 em. wide at the 
base, much longer than wide, each subdivided into 2 or 3 major lobes 
and several lesser lobules, the secondary lobes usually lobed or coarsely 
toothed, the ultimate segments narrowly triangular or linear-atten­
uate, ca. 5 times longer than wide. 
DIS'fRIBUTION: Centml and western Jalisco. In the dry grasslands 
around Guadalajara and the pine woodlands to the west. Flowering 
in July and August, 
In the original description two specinlens were listed as syntypes: 
Jalisco; Rio Blanco, July 1886, E. Palmer 171 (GH!); bluff of the 
barranca near Guadalajara, Sept. 14, 1891, C, G, Pringle 5154 (GH!). 
Pringle 5154 is chosen as the lectotype because it is more complete, 
The name was published in a paper concerning plants discovered by 
Pringle, and this specimen is annotated in what appears to be Robin­
son's writing. 
The leaves of this variety exhibit considerable latitude in shape and 
lobing, but are always deeply lobed with the ultimate segments very 
narrow, The inflorescence is more elongate and broader than in the 
otber varieties, 
hLlseo: Pringle 4627, 9872 (GH, NY, US); King 3656 (MICII, NY, TEX, 
US); MeVaugh 18603, 15094,20280 (MICH); Wilbu, & Wilbu, 1996 (US). 
9b. Pscalium peltigerum var. hintonii Pippen, var. nov. PLATE 9 
Varietas peltigera foliis profunde lobatis, caule glabro, infloreseentia 
majore differt, 
Plants to ca, 0.5 m, tall; stems hirsute with multicellular white 
hairs to 2 mm. long; blades of the basal leaves sballowly 9- to 11-
angled but not lobed, the margin mucronate-toothed; upper cauline 
leaves similar to but smaller than the lower ones; inflorescence tightly 
eorymbose, 10 crn. long and hroad (those examined not mature). 
HOLOTYPE: Mexico; Carbon eras, Toma, July 21, 1935, G, B. Hinton 
et al. 7966 (MICHl, isotype NY!), 
DISTHIBUTION: Known only from the type collection. 
The floral characters are similar to var. peltigerum and the plants 
have similar flesby subterranean tubers, They differ in that the leaves 
are angulate, not deeply lobed, and the stem is hirsute. Field obser-
, 
PIPPEN-MEXICAN "CACALIOID" GENERA 431 
vations and study of future material may indicate that var. hintonii 
deserves specific rank, hut for the present it is considered only as a 
variety. 
9co Psacalium peltlgerum var.latilobum Pippen var. nov. PLATE 10 
Varietas peltigera foliorum lobis longioribus quam latioribus, caule 
glabro differt. 
Plants to ca. 0.5 m. tall; stems sparsely pubescent; blades of the basal 
leaves lobed less than one-half the distance to the center, the primary 
lobes 10 or 11, 5-7 cm. wide at the base and equally long, the lobes 
subdivided into 2 or 3 secondary lobes, obtusely triangular or deltoid, 
about as broad as long; margins of the lobes coarsely, triangular­
toothed; inflorescence corymbose, tightly compact, ca. 9 em. long 
and broad (inflorescence not fully matured). 
HOLOTYPE: Guerrero; 1)\ miles W. of Omiltemi, 7,800 ft., June 11, 
1953, T. W. McCorcle & C. M. Rowell, Jr. 3453 (MICH, sheet 11; 
isotype, sheet 2!). 
D'STRIBUT,ON: Known only from the type collection. 
10. P •• <alium po<uliferum (S. Wats.) Rydb. Bull. Torrey Bot. Club 51:375. 
1924. 
Slender herbs ca. 1 m. tall; roots exuding a reddish-orange sap when 
cut; stems 4-7 mm. thick at the base, densely puberulent proximally 
becoming subglabrous upward and again puberulent in the inflo­
rescence; leaves glabrous or sparsely pu berulen t, mostly along the veins 
with 6- to IO-celled hairs ca. 0.5 mm. long; basal leaves 2 or 3, long­
petioled, centrally or subeentrally peltate, subeireular in outline, 15-
25 (-60) cm. in diameter; blades lobed one-third to two-thirds the 
distance to the depressed center, the prinIary lobes 7 or 8 (-10), 4-8 
cm. wide at the base, usually 2-3 em. longer than broad, each divided 
into 2 or 3 secondary lobes or merely triangulate, the margins irregu­
larly toothed; petioles \5-30 (-40) em. long, terete, slender, usually' 
sparsely pubescent; cauline leaves 3 or 4, the lowermost similar in 
size and shape to the basal ones except the petiole leafy-margined 
and dilated at the base; cauline leaves progressively smaller upward; 
the uppermost bracteiform, nonpeltate, the petiolar portion leafy­
margined, auriculate-based, the blade portion 5- to 7-lobed; inflo­
rescence paniculate or corymbose, 12-30 em. long and broad, the lower 
branches in the axils of the upper leaves; pedicels tomentose; calycu­
late bracts 3, subulate, about 1 mm. long (or lacking); heads about 
100, 9-10 mm. long; phyllaries 3-5 (usually 4), varying on the same 
plant, elliptic or obovate, (4-) 5-6 (-7) mm. long, 1-3 mm. wide, 
glabrous, the tips purplish; flowers 3-5, opposite the phyllaries; 
corolla 6-8 mm. long, the tube as long as the lobes, 3--4 mm., the 
lobes linear, 0.5 mm. wide, the throat none; anthers 2-3 mm. long; 
432 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
"tyle branches ca. 1 mm. long, lhe tips truncate or rounded with a 
terminal or subterminal fringe of trichomes; mature Rchenes oblong 
or narrowly ellipsoid, 5 mm. long, 1-1.5 mm. wide, pubescent with 
fuscous, unicellulnr hairs; pappus (3.5-) 5- 6 (-8) mm. long, creamy. 
DI5TRIDUTIO >l : Western J.lisco to Nayarit, north to Sinaloa. 
Rocky, grllss-oltk domint\tcd slopes and pastured areas. Flowering in 
mid-July and August. 
This species is unusllal because of the very small heads wilh mostly 
4 phyllaries and flowers. It is closely related to P. tabulare but. differs 
in its sparser pubescence and smaller heads. Both, howe,'er, have 
similnr leafy, auriclliate-based, cauline leaves. 
JALlSCO: Pippen 19,21,27, 60 (MICJl); 1','ngle 9874, 44\4 (G il, !'iY, US); 
lVilbu, & lVilbur 2017, 2166 (MICII, US) . NAYARIT: /(Q88 !!I35 (GIl, US) ; 
Kino 3677 (MICII, NY, TF.X, US); Pippen II C'vIICH); McYa1luh 1f>406, 18838 
(MICH). SUUI.OA: Ortega 7146 (US, the lenJ of this specimen is somewhat 
atypical bllt this inflorescence is very similar to P. poculiferum) , 
11. PSBcalium tabulare (IIemsl.) Rydb. Bull . Torrey Bot. Club 51 :375. 1921. 
Senecio tabularis Hernsl. BioI. Centr. Amer. Bot. 2 :24S. lRS1. Holotypc: 
Veracruz ; valley of Orizaba, Escarnl1l1a., Aug. 16, IS66, HouTgeau 2926 
(I{ , not , <.en ; isotypcs GIll and C, Field Mus. Neg. 2263\1). 
Cacalia labulaTi .. (HemAL) A. Gray, Proc . Amer. Acad. 19:52. ISR:l. 
Plnnts lip t.o 2 m. tnll (Rolls 5350); st.ems nt least 7 mm. thick at 
the base, purplish, dengely pubescent; leaves sparsely pubescent on 
the adaxial surfuce, densely pubescen t on the abaxial surface, the 
hairs multicellul"r, 1- 1.5 mm. long, 15- t.o 20-celled; basal leaves 2-4, 
long-pet,ioled, peltate, suhcircular in outline, 16-27 (- 60, Balls 5:l50) 
cm. in diamet.er, the venation rnoiate with 8 principal veins; blades 
lobed abuut half way to tile center, the primary lobes 7 or 8, 5- 8 ern. 
long und broad, each usually divided into 3 triangular secondary lobes, 
1-2.5 em. long, 2-3 cm. broad , the mar~in. coarsely del toid-toothed; 
petioles up to 35 cm . long, terete, pubescent; cauline lenve., 2- 4; the 
lowermost subcentrally peltate, the hlades broadly ovate or subcir­
cular, 14-16 ern. long, 15-17 cm. wide, palmately 6- to 7-lohed, tho 
petioles attached t,o the blade about 3 cm. ahove the base, leafy-mar­
gined , auriculate-based; uppermost calliine leaves sessile, auriculate­
based, 3-lobed distltlly; inflorescence paniculate, 30-40 cm. long, 8-30 
cm. broad, tbe branches often sparsely stipitate glandular, subtended 
by narrowly triungular bracts 1- 5 cm. long, 0.3-1 cm. wide; pedicels 
5-10 mm. long, with 2-5 subulate brncts, 2-6 mm. long; heads 75 to 
over 100, about 12 mm. long; phyllaries 5, narrowly oYate, 9- 10 mm. 
long, 2-3 mm. broad, mostly glabrous or occasionally sparsely pu­
bescent; receptacle 1.5 mm. across; flowers 5, opposite the phyllaries; 
corolla 7-9 mm. long, the tube (3.5-) 4-5 mm. long, the throat cam­
panulate 0.5-1 mm. long and wide, the lohes 3 mm . long:, linear; an­
thers 3 mm.long; style branches 1- 1.5 mm. long, hi.' pid eHl the Ilbaxial 
• 
, 
, 
PIl'PEN-MEXICAN "CACALIOID" GENERA 433 
surface, the tips truncate or obtuse with a tenninal or subterminal 
fringe of trichomes; mature achenes narrowly ellipsoid, 5-5.5 mm. 
long, 1.5 mm. wide, 1 mm. or less thick, glabrous, pale, about 18-
ribbed; pappus (5.5-) 6-7 mm.long, white. 
DISTRIBUTION: Known mostly from western Veracruz and eastern 
Puebla and one specimen collected in Michoacan. Openings in oak­
pine woodlands. Flowering from August to October. 
This species is very closely related to P. pocu1ijerum, but differs 
in that the heads always have 5 phyllaries and flowers, and glabrous 
achenes. 
Some of the specimens examined are sparsely stipitate-glandular 
on the inflorescence branches but this is not constant in this species. 
PuEBLA: Purpu. 5630 (GH, NY, US). M,CROACAN: Pippen 55 (MICH). 
VERACRUZ: Ball. 5350 (US); Muller •. n., August 1843 (NY); Sharp 44823 (NY). 
12. PsaeaHum species 
Plants to about 1 m. tall; stems slender, to 5 mm. tbick at the 
base, subscapose, densely hirsute below with white, nonglandular 
multicellular hairs up to 1 mm. long, distally sparsely hirsute witb 
nonglandular bairs as well as minutely stipitate glandular; leaves 
glabrous on tbe adaxial surface, sparsely pubescent along tbe veins 
on the abaxial surface; basal leaves 2 or 3, long-petioled, peltate, 
subcircular in outline, 20-30 cm. across, venation radiate with about 
8 principal veins; blades lobed one-third to one-fourtb (rarely one­
balf) the distance to the center of the blade, the primary lobes 6 or 
7,5-8 cm. wide at the base about as long as wide, eacb usually divided 
into 3, triangular attenuate-tipped secondary lobes; petioles slender, 
to 30 cm. long, pubescent; cauline leaves 1 (or 2), similar to the 
basal leaves but much smaller; inflorescence paniculate, ca. 35 em. 
long, 25 cm. broad, the branches densely stipitate-glandular, sub­
tended by petiolate, leafy bracts, the blade to 6 cm. long, elliptic, 
entire or palmately 3- to 5-lobed; pedicels purple, to 3 cm. long; 
calyculate bracts 3-7, subulate, 4-5 mm. long; heads few, immature; 
phyllaries (7 or) 8, oblong or elliptic, ca. ) 0 mm. long, 2-3 mm. wide, 
hirsute, occasionally sparsely glandular; flowers ca. 9, achenes gla­
brous; pappus white, chromosome number, n=30 (Turner, ined.). 
DISTRIBUTION: Known only from Guerrero. Flowering probably in 
November. 
These specimens appear to be somewhat intermediate between 
Psacalium eriocarpum and P. holwayanum. Tbey differ from both of 
these species by the glabrous achenes, the leafy inflorescence bract..q, 
the mostly glabrous basal leaves, as well as the distinct geographical 
range. 
These specimens probably represent either a distinct species or a 
variety of one of the above. The material examined was immature 
434 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
and could not be studied thoroughly. Perhaps after future collections 
have been studied and field observations made it will be possible to 
• 
assIgn a name. 
GUERRERO: MeVaugh 21913* (MICH); Sharp 441496 (Ny). 
Doubtful Names and Excluded Taxa 
Caealia albicans Sesse & l\Ioc. Fl. ~Icx. ed. 2. 180. 1894. Known only from the 
type description. This cannot be associated with any species of the genera 
discussed in this paper. According to the description the leaves are Innceolatc 
and sinuately pinnntifid and therefore this probably should be referred to 
Senecio. 
('acalia amplexicaulis Schultz Bip. ex Klatt, Leopoldina 24:125, in synonymy, 
1888. Listed as a synonym of Senecio bractealus Klatt. 
Cacalia berlandieri DC. Prod. 6:328. 1838. =Seneclo desortum Hemsl. (BioI. 
Centro Amer. Bot. 2:239.1881). 
Ca<a1i. <.Iotri<h. S. F. Blake, Contr. Gray Herb. II. 52:58. 1917. This should 
be referred to Senecio. The structure of the corolla is similar to that of S. 
angultjoIiu8 in that the throat is longer than the recurved lobes. The plant 
is apparently a shrub. It is known only from the type. 
Cacalia eriopoda Schultz Rip. ex Klatt, Leopoldina 24:125, in synonymy, 1888. 
Listed as synonym of Senecio eriopodu8 Klatt. 
Cae.Iia guatemalensis StandI. & Steyermark, Publ. Field Mus. Bot. 23:99. 1944. 
This should probably be referred to Senecio sect. Afulgedifolii since the 
description states that the flowers are purple and the inflorescence is glan­
dular pubescent. This species is not a Psacalium, as suggested by Cuatrccasas 
(Britton in 8:157. 1955) since, according to the description, the leaves arc 
not peHate. 
Cacalia incana Scss6 & Moe. PI. N. IIisp. ed. 1. 132. 1889. A later homonym of 
C. incana L., 1753. 
Cacalia liebmannii Schultz Bip. ex Klatt, Leopoldina 24:125, in synonymy, 188ft 
Listed as a synonym of Senecio liebmannii Buchinger. 
CacaIia Iyrata Scss6 & Moe. PI. N. Hisp. cd. 1. 131. 1889. Known only from the 
type description. This is probably refern.ble to Senecio scct. Mulgedifolii 
because, according to the description, the flowers arc purple and the leaves 
are lyrate-pinnatifid. 
Cmalia macrophylla Sess6 &: Moe. PI. N. Hisp. cd. 1. 132. 1889. Later homonym 
of Cacalia macrophylla Bicb. lS08. 
Cacalia pallescens Schultz Bip. ex Klatt, Leopoldina 24:126, in synonymy, 1888. 
Listed as a synonym of Senecio pallescens Klatt. Rydberg transferred this 
species to Odontotrichum based on a tracing in Klatt's herbarium. 
Cacalia parasitica Schultz Bip. Bot. Zeit. 15:759. 1Sti7. According to Greenman 
(Contr. U.S. Nat. Herb. 23:1635. 1924) t.his is Senecio parasiticu8 Greenm. 
It is a shrub. No specimens have been seen by me. 
Cacalia peitata Sesse & Moc. PI. N. Hisp. ed. 1. 132. 1889. A later homonym of 
Cacalia peltata H.B.K. [Psacalium peitatutn (H.n.K.) Cass.] 
PIPPEN-MEXICAN "CACALIOlD" GENERA 435 
Cacalia pinnatifida Sess6 & Moe. PI. N. Hiap. cd. 1. 132. 1889. A later homonym 
of C. pinnatifida L., 1767. The description and photograph of the type (an 
illustration) indicate that thIs is probably Odontotrichu.m amplifolium (DC.) 
Rydb. 
Cacalia prenanthoick, II .B.K. Nov. Gen. el Sp. 4:167. 1820. =ErechUt.,. .. lerl. 
IDI.folia f. preDlntboldes, Beleher (Ann. Mo. Bot. Gard. 43:30. 1956). 
Cacalia purpura8cen3 Schultz Bip. ex Klatt, Leopoldina. 24:126, in synonymy, 
1888. Listed as a synonym of Senecio purpurascen8 Klatt. 
C.calla pudica Slandi. & Slcycrmark, Publ. Field Mus. Bot. 23:255. 1947. This 
is not in any of the genera herein considered, beea.use the flowers are purple 
and the throat of the corolla is much longer than the lobes. It is more closely 
related to Senecio Sect. Mulgedi/olii. 
Cacalia runcinata H.B.K. Nov. Gen. ct Sp. 4:168. 1820. -Senecio roseus Schultr. 
Bip. (Flora 28:498. 1845) . 
Carolin runcinata Less. Linnaea 5:162. 1830. = Senedo c:oulterl Greenm. (Ann. 
Mo. Bot. Gard. 1:272. 1914). 
Cacalia sarracenica Sesse & Moe. Fl. Mcx. cd. 2. 180. 1894. A later homonym 
of Cacalia sarracenica I.., 1753. 
Cacalia llinuata Ses8~ « Mac. PI. N. Hisp. ed. 1. 1889. A later homonym of 
Cacalia 8inuata Cerv. 1824 (Odontotrichum sinualum (Cerv.) Rydb.]. 
Cacalia spathulata Besse &: Moc. ex D. Don, Trans. Linn. Soc. 16:250. 1833. 
=Chaptalla nutaDs (according to Blake). 
Cacalia 8ubdecurrens Schulz Bip. ex Klatt, Leopoldina 24:125, in synonymy, 
1888. Listed as a synonym of Senecio dejormis Klatt. 
Cacalia 8utruita Grcenm. Proc. Amer. Acad. 32:310. 1897. This probably should 
he referred to Senecio. It is not a. member or the genera. herein discussed, 
because the flowers are bright orange, the throat of the corolla is longer than 
t.he lobes, and the achcnes have hygroscopic t.richomes. 
Caenlia tepicana M. E. JoUC5. Contr. West. Bot. 15:156. 1929. =Senecio hartwerli 
Benth. according to Blake (Conlr. U.S. Nat. Herb. 29:136. 1945). 
Cacalia toluecana DC. Prod. 6:328. 1838. =Seneclo c:allosus Schultz Bip. according 
to Greenman (Ann. Mo. Bol. Gard. 1:271. 1914). 
Cacalia tuberosa Sess6 &; Moo. PI. N. Hisp. ed. 1. 132, 1889. A later homonym of 
Caealia tuberosa Nutt. 1818. 
Caealia ,iscosa Sess6 & Moe. PI. N. Hisp. ed. 1. 132. 1889. Known only from the 
type description which states that it is a shrubj therefore, it is most probably 
not a species in the genera discussed in this paper. 
Odontotrl.hum delphinifolium Rydb. Bull. Torrey Bol. Club 51:419. 1924. 
Rydberg saw only the type specimen which waa in fruit. This species has 
since been collected in flower. It is not an Odontotrichum, because the inflo­
rescence has only one large head, yellow radiate flowers, and no pappus. 
This taxon should proba.bly be referred to the Heleneae. 
436 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Odontotrichum liebmannii (Buchinger) Rydb. Bull. Torrey Bot. Club. 51:419. 
1924. Rydberg trnnsferred this species from Senecio on the strength of a 
tracing of a leaf In the Klatt Herbarium at the Gray Herbarium. I have 
Bcen no specimens and cannot associate tho description or the tra.cing of 
Klatt's with any species of the genera of thie paper (see C. li~bmannii Schult z 
Bip. above). 
Psacalium thlJT&oideum DC. Prod. 6:335. 1838. Known only from tbe type de­
scription. It was questionably published by DeCandolle as n Psacalium. 
Thus far the description has not been matched with any other cacalioid 
species. The type specimen is probably in Prague and must he examined 
before the position of this taxon can be determined. 
Collections Cited Excluding Types 
(Abbreviations: D.= Digitacalia: O.=Odontotrichumj Pc.=Pericalia; Ps.= 
PSll.calium.) 
ARS~NE, G. 
8.n. Nov. 1908, Fa. megaphyllum 
s.n. Sept. 9, 1909, D. jatrophoides 
s.n. Nov. 4, 1909, Pe. scssiIifolia 
s.n. Sept. 14, 1911, Pc. sessilifolia 
93. O. sinuatum 
938.. O. sinuatum 
334. O. einuatum 
1181. O. sinuatum 
2386. Ps. megaphyllum 
5132. O. palmeri 
5729. Pe. sessilifolia 
5740. O. sinuatum 
7347. Pe. sessilifolia 
ARB~NE, G., &: NICOLAS, BRO. 
334. Ps. megaphyllum 
BALLS, EDWARD K. 
5299. Pa. peltatum v. pcltll.tum 
5333. O. amplifolium 
5350. Ps. tabulare 
5428. O. tussilaginoidcs 
5575. Pe. scssilifolia 
BARCLAY 
S.n. O. pringlei 
BARKLEY, FRED A. 
14A624. O. decompoeitum 
BEAMAN,JOHN 
3610. Ps. peItatum v. peitatum 
BLUMER, J. C. 
1378. O. decompositum 
BOURGEAU 
715. Pe. sessilifolia 
716. O. sinuatum 
2926. Ps. tabulare 
CAMP, W. H. 
2401. Pa. peltatum v. conzattii 
CONZATTI, C. 
715. D. heteroidea 
4015. O. purpusii 
CONZATTI, C., &; GONZALEZ, V. 
397. O. amplifolium 
1237. O. purpusii 
DARROW, R. 
s.n. July 4, 1937, O. decornpositum 
DARROW, R., PHILLIPS, &: PULTZ 
1235. O. decompositum 
DODDS, DONALD, &: SIMPSON, ROBT. F. 
32. Pe. michoacana 
DIGUET, L. 
S.n. July 1912, O. palmeri 
DUGE8, A. 
451. O. einuatum 
481. Pe. 8eB8ilifolia 
EGGLESTON, W. W. 
10765. O. decompositum 
FEDDEMA, CHARLES 
10. O. sinuatum 
161. Pe. sessilifolia 
183. O. palmeri 
308. O. palmeri 
381. Pe. sessilifolia 
1345. Pe. se5silifolia 
FERRI8, ROXANA S. 
5881. O. pringle! 
GALEOTTI, H. 
2491 bis. O. cirsiifolium 
GENTRY, HOWARD S. 
536M. O. decompositum 
1959. O. decomp08itum 
437 
438 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
2815. O. decompositum 
6968. Pe. sessilifolia 
6969. O. pachyphyllum 
8562. O. sinuatum 
GENTRY, HOWARD S., CORREM" & 
AHGUELI,ES 
18016. Pa. peltatum v. peltatum 
GHlEBBnEGHT 
805. O. cirsiifolium 
GILLY, C. L., SR. 
5. Pa. holwayanllm 
GOLDMAN, E. A. 
132. O. sinuatum 
GODDING, L. N. 
s.n. Aug. 28, 1912, O. decompositum 
784. O. decompositum 
GOULD, FRANK W. 
8989. O. sinuatum 
GRAHAM, SHIRLEY 
94. O. sinllatum 
99. O. globosuID 
100. O. radulifolia 
124. O. palmeri 
HARTWEG 
125. O. sinuatum 
HEWITT, W. P. 
68. O. decompositllID 
HINTON, C. B. 
1266. O. goldsmithii v. goldsmithii 
1908. Pa. peltatum v. peltntllm 
2200. Pa. megaphyllum 
2355. Pa. megapbyllum 
2487. Ps. ho!wayanum 
2755. Pe. michoacana 
4993. Pe. michoacana 
7984. O. goldsmithii v. goldsmithii 
8066. O. goldsmithii v. goldamithii 
8211. O. cirsiifolium 
8349. O. amplifolium 
9202. O. goldsmithii v. goldsmithii 
9436. O. cirsiifolium 
12146. O. brachycomum 
12572. D. napeifolia 
12718. Pc. michoacana 
13463. O. amplifolium 
13469. Pe. michoacana 
13480. P,. megaphyllum 
14464. Pa. nanum 
14877. Pa. pcltatum v. pf'ltatum 
15078. O. goldsmithii v. goldsmithii 
15288. O. brachycomllm 
15624. O. brachycomum 
15629. Ps. mcgaphyllum 
IBARRA GR.o\CJ..\, P. 
407. O. globosum 
414. Pa. peltatum v. pcltatum 
JONES, MARers K 
2.5053. O. decompositum 
27692. O. pringlei 
JONY, P. 
s.n. in 1892. O. platylepis 
KEARNY, T. 11., & HARRISON, G .. J. 
6045. O. dccompositum 
6194. O. decompositum 
KEARNY, T. II., &: PEEBLES 
10092. O. dccompositum 
KENOYER, L. A. 
2110. O. rndlllifolillln 
2330. O. radulifolium 
KINO, ROBERT M. 
3615. O. silphiifolium 
3656. Ps. pcUigcrum v. pdtigerum 
3659. O. palmeri 
3677. Ps. poclllifcrum 
6737. O. sinuatum 
KING, ROBERT M., & SODERSTROl\f, T. R. 
4904. O. brachycomum 
KNOBLOCH, I. W. 
1273. O. globosum 
1293. O. decompositum 
KRUCKERERG, A. R. 
4644. O. decompositum 
LEAVENWORTH, "'1\1. C. 
539. O. palmeri 
PIPPEN-MEXICAN "CACALIOID" GENERA 439 
LESUEUR, HARDE 
145. O. dccompositum 
LYONNET, E. 
194. Pc. aessilifolia 
794. Pa. peltatum v. peltstum 
3122. Pa. peltatum v. pelt-ntum. 
MANNING, W. E. & M. S. 
53560. O. radulifoliUlD 
!l.IATuDA, E. 
19148. O. ailphiifolium 
19417. O. ailphiifolium 
]9706. O. sinuatum 
21808. O. sinuat.um 
21825. Pe. scssiliColia 
26175. Ps. peltatum v. peltatum 
26572. Pa. megaphyllum 
26632. Ps. peltatum v. pcltatum 
26882. O. sinuatum 
27730. O. amplifolium 
28343. Pa. peltatum v. pcltatum 
28821. O. earsHfolium 
29313. O. amplifolium 
29483. Po. peltatum v. pcltatum 
29591. Pa. peltatum v. peltntum 
29609. O. amplifolium 
29787. Pe. 8cssilifolium 
30015. Po. holwayaaum 
MAT8ILLES, JAMES 
7579. Pc. seesilifolia 
7587. O. radulifolium 
7752. O. sinuatum 
7755. Pe. sessilifolia 
7784. O. pachyphyllum 
7793. O. sinuatum 
7880. O. sinuatum 
8448. O. sinuatum 
8485. O. sinuatum 
MeV AUGB, ROQERS 
12810. O. sinuatum 
12929. O. palmeri 
13321. Po. megaphyllum 
13559. Pe. sessilifolia 
13616. O. pringlei 
13625. Pe. Bessilifolia 
13684. Pa. eriocarpum 
13821. Po. pe\tatum v. peltatum 
14093. Pe. michoacana 
14136. Pe. michoacana 
14165. Pa. eriocarpum 
14287. Po. criocarpum 
14290. Pe. michoacana 
14390. Pe. michoacana 
15094. Pa. peltigerum v. peltigerum 
15313. O. cervinum 
15460. O. goldsmithii v. goldsmithii 
16132. O. goldsmithii v. goldsmithii 
16406. Pa. poculiferum 
17118. Pc. scssilifolium 
17160. D. jatrophoidcs 
17183. Pe. eessilifolia 
17650. Pa. megaphyllum 
17726. O. amplum 
18121. Pe. oCB8ilifolia 
18603. Ps. pcltigerum v. peltigerum 
18838. Ps. poculiferum 
20280. Pa. peltigerum v. peltigerum 
20481. O. palmeri 
21355. Pc. michoacana 
21658. Pe. sessUifolia 
21659. O. pnchyphyllum 
21729. O. pachyphyllum 
21731. O. pachyphyllum 
21732. O. globooum 
21733. O. sinuatum 
21734-39. O. globosum x O. slnuatum 
21765. O. pringlei 
21823. D. napeifolia 
21824. O. amplifolium 
21826. Pa. peltatum v. conzsttii 
21897. P8. megaphyllum 
21903. D. tridactylitia 
21910. Pa. megaphyllum 
21928. Po. megaphyllum 
21934. O. brachycomum 
21953. Pa. holwayanum 
21960. O. pRImeri 
21964. O. brachycomum 
MCVAUOH. n .. & KOELZ. W. 
39. Ps. megaphyllum 
48. O. platylepia 
49. O. sinuatum 
57. O. amplum 
147. Pe. acssiliColia 
276. O. sinua.tum 
435. Pe. michoacana 
441. O. palmeri 
479. Pe. sessilifolia 
601. O. pringlei 
656. Pe. 8C88i1ifolia 
440 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
792. Pe. sessilifolia 
919. rs. eriocarpum 
920. Pe. scssHifolia 
1104. O. pringlci 
1253. O. pringlci 
MEXIA, YNEZ 
635. Pc. scssilirolia. 
845. O. pringlci 
877. 1'8. megaphyllum 
1450. Pc. scssilifolia. 
1656. P c. michofl.cnna 
MEAUN8, EVGAH A. 
531. O. dccomposituUl 
2219. O. dccompoeitum 
MEYEH, F. G., I\ud ROGEI18, D. J. 
2608. O. radulilolium 
MICKEL, JOHN T. 
501. O. rtuJulirolium 
561. O. raduliColium 
Moont:, H. E., JR. 
3421. O. s ilphiifolium 
5119. Ps. mcgaphyUllm 
MOOltE, H. E., JB .• & CETTO. M. 
5469. Pc. michoac£l.na 
MULLER, C. H. 
3420. O. dceompositum 
MULLER, F. 
s.n. in 1853. Pa. tabulare 
1619. O. silphiifolium 
NARVAEZ MONETa, M ., &; SALAZAR, A. E. 
5:')7. O. plntylcpis 
N E "'. ELSON, • H 
lOG) . O. amplifolium 
1133. D. napeifolia 
1420. O. cirsiifolium 
2080. D. tridactylitis 
2243. O. l.unplifoliulll 
3117. O. cirsiifolium 
3467. D. chiapensis 
4758. O. decompositum 
4759. O. globosum 
4954. Pa. pcltatum v. pcltatum 
6582. Ps. pcltatum v. pcltatum 
OWNBEY, GERALD B., & OWNBEY, 
FINDLEY 
1874. O. pncbyphyllum 
ORCU-rr, C. R. 
3746. O. silphiifoliuIn 
ORTEOA, J ESuS C, 
fJ7Z1. pc. sC"s!:Iil ifolia 
7146. Ps. poculitcrum 
PALMER, EDWARD 
s.n. July 1890, 0, dccompositulll 
s.n. 1892. O. priugh~i 
47. Pc. s(~ssiliro1iil. 
1 GS. O. palawri 
576. Pc. sessilifolia 
651. O. sinuatum 
689. O. platylepi:l 
1234. O. pringh· j 
1832. Pc. scStiil irolit~ 
PARRY & P ,\I,I\f };H 
M 1. Pc. scssilirolia 
543. O. mdulirolium 
M3. Ps. peltatum v. pf'ltatulIl 
PENNELL, ] i'RANClS 'tV , 
18332. O. pacbyphyUum 
1834:1. Ps, pcltatum v. pclt.atuill 
1834ii. Pc. scss i1ifoliu 
18582. 0. globosum 
PHlLLlP8, EOWIN A. 
651. O. dccompositum 
PIPPEN, H leu .' RD W. 
1. O. sinuatum 
2. O. amplum 
3. O. amplum 
4. O. amplum 
5. Pe. BC'SSilifoHum 
G. Pa. pcltatum v. pcltlltum 
7. O. sinuatum 
8. Ps. mcgaphyllum 
9. O. sinuatum 
10. O. palmcri 
11. Pi>. poculiCcrum 
12. 0, palmeri 
13. Pc. scsailirolia 
15, p(~. sessilifolia 
16. P~. mcguphyUum 
17. Pc. scssilifolia 
18. Pe. sessilifolia 
19. Ps. poculifcrum 
20. O. palmeri 
21. Ps. poculiferum 
22. O. sinuatum 
23. O. pl.tylcpie 
, 
, 
PIPPEN-MEXICAN "CACALIOID " GENERA 
24. P •. mcgaphylIlIm 
25. O. multilobum 
26. Pe . micboacana 
270.. O. palmeri 
27. Ps. poeuliferum 
28. O. palmeri 
29. O. cf. palmeri 
30. O. mllitilobum 
31. Pe. !WssilifoJia 
32. O. palmeri 
33. Pe. sessilifolia 
34. Pe. 8cssiliColia 
36. Pe. criocarpum 
37. O. prillglei 
38. Pe. michoacana 
39. Pa. mcgaphyllum 
40. O. multilobum 
43. O. palmeri 
44. Ps. pcltatum v. pcltatum 
45. O. sinllatum 
46. D. ja trophoides 
47. Pe. megllphyUum 
48. Pe. sessilifolia 
49. Pa. pcltatum v. pcltatum 
[,0. O. sinuatum 
51. Pe. scssiliColia 
52. O. cin;iifolium 
53. P •. pelt.tum v. pelt.tum 
54. O. silphiifolium 
56. O. sinuatum 
57. O. sinuatum 
58. O. platylcpi. 
59. O. pringlei 
60. Pa. poculiferum 
61. Pe. scssilifolill. 
62. Pa. erioearpum 
63. Pe. micboncana 
64. Pe. scssilifolia 
6.1), Pc. michoacana 
66. Pli. eriocarpum 
67. Ps. eriocarpum 
POWELL, A. M., & EDMONDSON, J. 
869, O. palmeri 
929. O. raduHfolium 
948. O. siOlmtum 
PRINGLE, C. O. 
767. O. deeomposituIO 
1736. Pe. scssilifolia 
1811. O. pringlci 
1816. O. platylepis 
2304. O. po.lmcri 
3340. P •. pelt.tum v. pelt.tum 
3566. O. r.dulifolium 
4095. O. radulifolium 
4262. D. j.trophoides 
4274. O. sinuatum 
4414. Ps. poculiferum 
4672. Ps. peltigcrum v. peltigerum 
4778. D. n.peifolio 
4785. O. amplifolium 
4984. O. cirsiifolium 
5272. O. cirsiifolium 
5741. D. jatrophoides 
6018. O. pRucicapitatum 
6164. D. trid.ctylitis 
6165. Ps. meg.phyllum 
6176. O. heteroidea 
6238. Pe. peltatum v. conzattii 
6453. O. silphiifolium 
7983. Pe. scssilifolia 
9080. O .• ilphiifolium 
9868. Pa. tussilaginoides 
9869. Pe. seasilifoli. 
9870. Pa. pcltatum v. pcltatum 
9871. P •. pelt.tum v. pelt.tum 
9872. P8. peltigerum v. peltigerum 
9873. O. platylcpis 
9874. Pa. poculiferum 
9875. O. ainuatum 
9877. D. trid.ctylitis 
10024. O. n.pellifolium 
10 117. Pe. michoacana 
10126. O. braehycomum 
11497. Pe ...... IlIfoli. 
11499. O. pringlei 
11500. O. palmeri 
13297. Pa. holwo.yo.num 
13672. Pa. holwayo.num 
13979. Ps. holwayanum 
PURPU8, CARL A. 
3038. Pa. peltatum v. peltatum 
3139. O. amplifolium 
3845. O. purpusii 
4113. Pa. calvum 
5625. Pe. 8CS8iHiolia. 
5630. Pa. tabula.re 
R08~, J. N. 
1935. Pe. poculiferum 
1986. O. cervinum 
2101. O. filicilolium 
2248. cf. O. palmeri 
2337. Pa. laxifiorum 
441 
442 CONTRIBUTIONS f'ROM THE NATIONAL HERBARIUM 
2828, Pc, scssilifolia 
3040, (), jatrophoidcs 
3471. O. gJobosum x O. sinuntum 
3575. 0. umplum 
7808. O. sinuntum 
ROSE, J. N., & HAY, HOBERT 
5311. O. silphiifolium 
5547. O. tussilaginoidc8 
RosE, J. N., & PAINTER, J. H. 
6738. O. tussilag,inoldcs 
7039. Ps. pcltutum v, pcltatum 
HOSE, J. N" ET At. 
9187. 0. silphiifolium 
ROWELL. CHESnm M., In. 
3387. O. goldsrnithii v. rowdlii 
3604. O. goldsmithii v. rowellii 
3719. 0, nupellifolium 
3886. O. goldsmitbii v. rowcllii 
3896. 0, nnpcllifolium 
ItzEDOWSKI, J. 
15974. O. amplifoliulll 
16015. O. nmplifolium 
16073. O. goldsmithii v. rowcllii 
SAFrORO, \VILLIAM E. 
1432, 0. platylepi. 
1451. Pc.8cssilifolia 
SALAZAR, F. 
s.n. Aug. 1, 1913. O. nnpcUHolium 
ScHAFFNER, J. O. 
B.n. &:pt. 1955. Pc . scssilifolia 
255. Pc. 5Cssi1ifolia 
294. O. TH.dulifolium 
204. Pa. pcltatum v. pcltatum 
ScHUMANN, \V. 
148. 0, rndulifolium 
SE,\TON, HENRY 
B,n.in 1891. Ps. pelLatum v. pel tatum 
SEElfANN 
• ,n. 0. p.chyphyllum 
SELER, Eo. 
1259, 1'., pelt.tum v. pcltatum 
SHARP, A. J. 
44508, P •. pelt..,turn v, pcltat.um 
441431. 0, goldsmithii v. rowcllii 
441465. 0 .• D1plifolium 
SHREVE, FORREST 
8031. 0, globo,urn 
SMITH, CUARLES L. 
287. D, heteroidca 
380, D. tridactylitl.; 
387, D. heteroidca 
388. D. j.trophoid"s 
S~uTn, LUCIUS C. 
123, D. jatrophoides 
124. 0, umplifolium 
g66, D, t.ridnctylitis 
SMY'fIl, E. G. 
172, 0, sinna-tum 
STANFORD, RETIIEUl<'ORO, & NORTHCRAFT 
818. 0. rndulifolium 
STANFORD, TAYLOR, & LAUBER 
2421, O. radulifolium 
2535, Pa. pcltatum v, adcnophorum 
TOWNSEN'I>, C. II. T. 
s.n, in 1909, O. decompol:litum 
TOWNSEND, C. H. T., & BARBER, C. M. 
158. O. dccompositum 
TUCKER, J. 1\1. 
2456. 0, decoDlp<lSitum 
VlscnER 
14. Pc. scssilifoll~ 
58. O. sinuatulU 
WATEnFALL, U. T. 
12668. 0. pachyphyllurn 
WHITJo:: , STEPHEN S. 
2741. 0. de<:ol1l post tum 
3289. 0. decomposi tulll 
4353. 0. dCCOOlpositum 
WILBUR. 11 , L. « C, R, 
2017. PI'!. poculifcrum 
2166. Pa. poculifcrum 
WILCOX, TIMOTIIY E . 
s.n. ill 1891. O. dccompositum 
WRIGHT, CHARLES 
1286. O. dccomp05itum 
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Literature Cited 
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1956. A revision of the genus Erechlites (Compositae) with inquiries into 
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1959. Sedum of the Trans-Mexican volcanic belt. Ithaca. 
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CRONQUIST, A'I and GLEASON, H. H. 
1963. Manual of vascular plants. 
CUATRECASAS, J. 
1955. A new genus and other novelties in Compositae. Brittonia 8:151-163. 
1960. Studies on Andean Compositae IV. Brittonia 12:182-195. 
DECANDOLLE, A. P. 
1838. Prodromus systcmatis naturalis 6:327-336. 
FERNALD, M. L. 
1950. Gray's manual of botany, ed. 8. 
FIORI, A. 
1933. Flora Italiana. Mariano Ricci. Firenze. 
GRAY, A. 
1883. Contributions to North American botany. I. Characters of new 
Compositae with revisions of certain genera and critical notes. 
Proc. Amer. Acad. 19:1-96. 
GREENMAN, J. M. 
1902. Monogmphie der nord· und central·amerikanischen Arten der 
Gattung Senecio I. Bot. Jahrb. Engler 32:1-33. 
HAWORTH, A. H. 
1812. Synopsis plantarum succulentarum. London. 
HEGI, G. 
1917. Illustrierte Flora von Mittel-Europa 6 (1) :403-407. Munehen. 
HEMSLEY, W. G. 
1881. Biologia. Centrali·Americana Bot. 2:235-247. London. 
266-836-07'--8 443 
444 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
HITCHCOCK, A. S., nnd GREEN, ~'L L. 
1947. Supplement species lcctotypicac gcncrum Linnaci. Brittonin 6:114-
118. 
HOFFMANN, O. 
1890. Compositne in Engler and Prantl., Die IlaiUrlichcn rllanzen­
(amllien 4 (5) :225-272. Leipzig. 
LANGLET, O. F. J. 
IH3G. Nllgra Diclrag till Kannedomcn om Kromosomlulen inon1. ~ym­
phaeaceae, Ranunculacc::w, Polcmoniaccacoch Compositae. Svensk. 
Bot. Tid8k. aO:288-2U4. 
LANJOUW, J., and STAFI.EG, F. A. 
1964. Index Hcrbariorum. Part 1. The Herbaria of tho world, cd. 5. Inter­
national Bureau for Plant Taxonomy and Nomenclature. Utrecht. 
LrNNAEu8, C. 
1753. Species Plantnrum. Stockholm. 
L'OBEL, MATTHIAS DE 
1581. Plnntarum seu stirpium ieanes. Antwerp. 
MAKINO, T. 
1958. An illustrated flora of Japan. Tokyo. 
MAHTJNEZ, 1\1. 
IU5H. Plantcs IV[cd icinales de h-Jexico, cd. 4. Edicioncs Botas. l\·l cx ico D. F. 
l\{oruSON, R. 
1699. Pluntarum historiae. Jacob Bobart. Oxford. 
ORNDUr'F, n., ET AL. 
1963. Chromosome numbers in CotnpositA.c. III. Scnccioncnc. Amer. 
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1955. Materinls for a. dictionary of botanical terms. III. Inflorescences. 
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19"14. Some senccioid genera. Bull. Torrey Bot. Club 51:369-378, 409-420. 
SCHULTZ, C. II. 
1845. Ueber einigo mit Senecio zu verbindcndc Gattungen. FlorA 28:497-
498. 
SHINNERS, L. II. 
1950. The Texas species of Cacalia. Field nnd Lab. 18:79-83. 
TOURNEFoR'r, J. P. 
1700. Institutioncs rei harbarine. (Cacalia on 1:451.). Paris. 
INDEX 
(Synonyms in italics. Page numbers of principal entries in boldface.) 
Adenostyles, 376, 377 
Aulacophora (sect. of Cacalin) J 377 
Caoslia, 365, 366, 375, 376, 377, 378 
albicans, 434 
alpina, 375, 377 
ampiexicaulis, 434 
amplijolia, 387 
ampullacea, 409, 410 
atriplicifolia, 377 
sect. Aulacophora, 377 
berlandieri, 434 
brachycoma, 388 
calotricha, 434 
calva, 417 
cervariifolia (lfccrvariaefolia"), 
390 
chiapensis, 379 
cir3iifolia, 403, 404 
sect. Cissampelopsis, 377 
sect. Conophors, 377 
cordi/olia, 411, 413, 414 
coriacea, 402 
decvmposila, 392 
digilata, 381 
eriocarpa, 418 
eriopoda, 434 
sect. Eucacalia, 377 
globosa, 394 
gold3muhii, 395, 396 
guatemalensis, 43" 
hastata, 377 
ho[wayana, 419 
'incana, 434 
jatrQphoides, 378, 381 
laeviqata, 380 
liemnannii, 434 
longipetiolata, 380 
lyrata, 434 
macrophylla, 434 
megaphylla, 421, 422 
michoacana, 412 
napeaejolia, 382 
napellifolia, 398 
Cacalia.-Continued 
nomenclature of, 376 
nutans, 413 
oblusiloba, 421, 422 
pachyphylla, 399 
pallescens, 434 
palmeri, 400 
parasitica, 434 
paucicapitata, 401 
peUata, 424, 426, 434 
var. conzattii, 427 
vsr. couUeri, 426 
peUigera, 428, 430 
pinnatijida, 435 
platylepis, 402 
praiensis, 394,395 
prenanthoides, 435 
pringlet', 403 
pudica, 435 
purpurascens, 435 
purpusii, 405 
raduliJolia C'radulaefolia") I 405,406 
rumicifolia, 387 
funcinata, 435 
sarracenica, 435 
8chaifneri, 406 
se88ilijolia, 413,414 
silphiijolia, 407 
sinuata, 408, 435 
spathulata, 435 
8uaveolens, 375, 377 
3ubdecurrens, 435 
Buffulta, 435 
tabularis, 432 
teplcana, 435 
toluccana, 435 
tridactylitia, 383 
trigonaphy/la, 412, 413 
tuberCMa, 435 
tU88ilaginoides, 409, 410 
viscosa, 435 
cacalioid condition, 368 
Chaptalia nutans, 435 
445 
446 CONTRIBUTIOXS FROM THE NATIONAL HERBARIUM 
CiRsampciopsia (scct. of Cacnlia), 377 
Conophora (sect. of Cacnlin), 375, 377 
Digitucnlia, 365, 366, 367, 368, 370, 371, 
372 (fig.), 374, 377, 378, 379 
chiapensis, 379 
he\eroidcn, 379, 380, 381 
j.trophoidcs, 369 (fig.) , 379, 381, 382 
llapcifolia, 379,382 
tridactyHtia, 379,383 
Erechtit.es valcriunncfolia r. prcnan-
thoidcB, 435 
Eucncalia (sect. of Cscn.lia), 375, 377 
Matariqllc, 393 
Mesadenia decomposita, 392 
Mesoneuris bipinnali,fola, 3nO 
Mulgedifolii (sect. of Senecio), 374,434, 
435 
Multinervii (sect. of Senecio) , 375 
Odontotrichum, 365, 367, 368, 370, 371, 
372 (fig.), 377, 378, 379, 384, 406, 
410, 434 
amplifolium, 386, 387, 410, 435 
amplum, 367, 368, 3SS, 387, 4000 
ampullauum, 409 
bipinnalijidum, 390 
brnchycomum, 368, 38:>, 388, 389, 
401 
calvum, 417 
cervinum, 386, 390, 394, 399 
chiapensis, 379 
cirsiifoHum, 368, 384, 3.~ ,), 390 
decompositllm, 366, 368, 370, 386, 
392, 393 
dclphinifolium, 435 
filicifolinm, 386, 390, 393, 300 
globosum, 38!), 394, 39;; 
goldsmithii, 367, 396, 3D6, 405, 407 
var, goldsmithii, 385, 396 
var, rowcllii, 385, 396 
jatropho1'des, 381 
Liebmannii, 436 
multilobllm, 368, 3S6, 397, 398 
napeae/olium, 382 
napcllifolium, 386, 398 
ncphrophyUum, 401 
pachyphyllum, 367, 36R, 38S, 387, 
399 
palmeri, 368, 385, 389, 398, 400, 401 
paucicapitatum, 386, 401 
pl.tylepiB, 367, 386, 402, 404 
prateruJe, 394 
pringlei, 368, 386, 396, 403 
IlUrpusii, 385, 405 
OdoDtotrichum-Continllcd 
raduliColium, 378, 386, 405 
rumiclfolium, 387 
8cabrum, 394, 395 
8chaffneri, 406 
silphiifolium, 367, 368, 385, 396, 
405, 407 
slnu.tum, 367, 368, 360 (fig.), 386, 
395, 398, 402, 404, 408 
tridaclylitis, 383 
tuss ilnginoidcs, 386, 409, 410 
Pnlmatincrvii (sect, of Senecio), 374, 375 
PcriMlin, 365, 367, 368, 370, 371, 373 
(fig.), 374, 377, :ns, 379, 410, 429 
michoacann, 368, 41 t, 412, 413, 415 
ovaliJolia, 413 
scssilifoli" 367, 369 (fig.), 411, 413, 
414, 415 
Psac.litlm, 361), 367, 368, 370, 371, 373 
(fig.), 374, 377, 378, 379, 384, 
415, 424, 434 
Grgututn, 426 
caivum, 368, 411, 42;1 
conzalli£, 427 
couttcri, 426 
erioC!lTpUm, 416, 418, 421, 433 
holwnynnum, 416, 419, 420, 421, 
433 
langlassei, 419, 420 
laxiflorum, 417, 420 
megnphyUum, 417, 421 
molltfolium, 427, 428 
nanum, 368, 417, 412 
nehmnii. 417, .. 23 
oblusilobutn, 421 
pcltntum, 416, 4l7, 424, 425, 434 
var. ndcnophorum, 42!i, 427 
var. conzattii, 425, 427, 428 
var. peitnturn, 369 (flg.L 42.5, 
426 
peltigerum, 423, 428 
va.r. hintonii, 429, 430 
var. latilobum, 4~O, 431 
var. peI t igerum, 429, 430 
poculiferllm, 4l7, 431, 433 
species, 417, 433 
tabulare, 416, 432 
thyraoideum, 436 
Sciadoseris vaginata, 390 
Senecio, 365, 368, 371, 377, 378, 434, 435 
albo-ltdescen3, 408 
ampli/oiiuB, 387 
angulifoHII!J 36ft (fig .) , 374, 434 
Senecio-Continued 
beecheyanm, 413 
bracteatus, 434 
oallosu8, 369 (fig.), 374, 435 
colophVU "", 408 
corawphyllm, 413 
cervariaejoli'Wl, 390 
chiapen3is, 379 
ciraiiJolia, 403 
cooperi, 375 
cordobensis, 374 
deformans, 374, 435 
dtlJortu m, 434 
eriopo<iu9, 434 
far/arm, 409 
grayanm, 392 
hartwegii, 375, 435 
heteroideUB, 380 
jaiiscana, 374 
jatrophoidM, 381 
liebmannii, 434 
moreliae, 420 
INDEX 447 
Senecio-Contin ued 
sect. Mulgedifolii, 374, 434, 435 
sect. M ultinervii, 375 
napeaejolium, 382 
ovatijoliw,413 
palIesoon8, 434 
sect .. Palmatinervii, 374, 375 
parnsiUcUfl, 434 
peltifcM..ts, 424 
purpurOSCCDs, 374, 435 
radulaefoliw, 406 
roldann., 374 
roseUB, 435 
rumiclfolim, 387 
runcinatus, 374 
!ClerophyUm, 399 
seemanii, 375 
St88ilijolius, 413 
8ubpeitatu8, 374 
tabulare, 432 
tm'ilaginoides, 409 
senecioid condition, 368 
u.s. GOYUI'I.UIT PRIIITIII; 0"1(:' , It ..