eF PHYTOLOGIA

An international journal to expedite plant systematic, phytogeographical
and ecological publication

Vol. 69 July 1990 No.1

CONTENTS

4%. G.L. NESOM, An additional species of Gnaphaliothamnus
(Asteraceae: Inuleae) and further evidence for the integrity of
Rt ie oe Sao o ih op oer disiaon'- otheiAduenarwecGacadtidunstwinpnanndn 1

~ B.L. TURNER, Taxonomy of Varilla (Asteraceae, Heliantheae) ..... 4

B.L. TURNER, New species, names, and combinations in Mexican
a a oa daa sumphecebh ats toh 14

-T. MELCHERT & B.L. TURNER, New species, names, and combina-
tions in Mexican Bidens (Asteraceae: Coreopsideae) .......... 20

- G.L. NESOM, Two new species of Mexican Baccharis (Asteraceae:
ee eM Re MCS AE Chines cain We cstct 2 da Leek Giiesotnaas ti omnaszhvinevak oases vasbnse 32

G.L. NESOM, Infrageneric taxonomy of North and Central American
Baccharis (Asteraceae: AStereae) o..........ccececceeeeeseeeeeeeeeeeeeeeeees 40

— §.D. JONES, G.D. JONES, & J.K. WIPFF, Carex fissa, section
Multiflorae (Cyperaceae), NEW tO TEXAS uu... eeeeeeeeeeeeeeeeeeeeeeees 47

- B.L. TURNER & T.M. BARKLEY, Taxonomic overview of the Senecio
flaccidus complex in North America, including S.
CE MTS RD IES ID Ge 0 A NRE te PA beet oe 51

PEDIC ALON Gates TOF. VOIIME GD | osc. cicicccecctockesahconsacnodsedeondsvanlediucesca 56

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Phytologia (July 1990) 68(6):1-3.

AN ADDITIONAL SPECIES OF GNAPHALIOTHAMNUS (ASTERACEAE:
INULEAE) AND FURTHER EVIDENCE FOR THE INTEGRITY OF THE GENUS

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

ABSTRACT

Gnaphaliothamnus durangensis spec. nov. is described from
southeastern Durango, México, bringing to 10 the number of species in
the genus. The new species is most similar to G. concinnus (A. Gray)
Nesom of San Luis Potosi and related to other species with eglandular
leaf surfaces. The occurrence in central Oaxaca of putative hybrids be-
tween the strongly divergent species G. eleagnotdes (Klatt) Nesom and
G. aecidtocephalus (Grierson) Nesom strengthens the hypothesis that
the genus is monophyletic.

KEY WORDS: Gnaphaliothamnus, Asteraceae, Inuleae, México.

Shortly after completion of a systematic study of Gnaphaliothamnus (Ne-
som 1990), a new species has come to light in the course of a study of Mexican
Inuleae at MEXU. The new species is known from only a single collection in
southeastern Durango.

Gnaphaliothamnus durangensis Nesom, spec. nov. TYPE: MEXICO.
Durango: Mpio. Durango, 65-75 km SW of Durango City on road to
La Flor, high ridge with_meadows and forests of Pinus, Quercus, and
Pseudotsuga, 2620 m, 17 Sep 1979, D.E. Breedlove 44285 (HOLOTYPE:
MEXU!; Isotype: CAS).

Gnaphaliothamnus concinno (A. Gray) Nesom similis sed phyl-
lariis ovatis apicibus rotundatis subroseis differt.

Suffrutescent herbs 1.5-2.5 cm tall, stems and leaves densely woolly to-
mentose. Leaves bicolored, glabrescent above, but remaining pubescent to a
degree, eglandular, the upper spreading at right angles to the stem, senescent
and deflexed on the lower half of the stem, elliptic-obovate, 1-2 cm long, 3-5
mm wide, with a terminal mucro, slightly decurrent, the margins very slightly
revolute. Heads campanulate, short pedicellate in compact clusters of 8-12,

1

2 PAY? OL GEA volume 69(1):1-3 July 1990

barely raised above the leaves; phyllaries 24-28, ovate to broadly ovate, the
distal 1/3 pinkish and not narrowed-elongated or strongly opaque, strongly
graduated in 5-6 series, the inner 4.0-4.5 mm long, the outer loosely and per-
sistently woolly on the proximal 2/3. Mature corollas and fruits not seen;
pappus bristles of hermaphroditic flowers with swollen clavellate apices.

The eglandular leaves of Gnaphalhiothamnus durangensis ally it with G. sali-
cifolius (Bertol.) Nesom and its relatives, as opposed to those with glandular
upper surfaces. The new species is similar in leaf shape to G. concinnus (A.
Gray) Nesom and G. eleagnoides (Klatt) Nesom. It is most similar to the for-
mer in its persistently pubescent upper leaf surfaces and outer phyllaries, and
its relatively few heads barely lifted above the leaves. It differs from both of
these species in its ovate phyllaries with pinkish apices that are not indurated
opaque and not narrowed into an appendagelike apical extension. The only
other species of Gnaphaliothamnus in western México is G. salicifolius, which
has longer (2-8 cm long) leaves with strongly glabrate upper surfaces, more
numerous heads in dense corymbs held above the leaves, longer phyllaries with
thicker and narrowed apices, and pappus bristles with only slightly dimorphic
apices. Heads on plants of the type collection of G. durangensis are not mature
enough to enable a count of hermaphroditic and pistillate flowers, but both
types apparently are present.

Evidence for the generic integrity of Gnaphaliothamnus.

The woody, “central-sterile,” gnaphalioid plants of México and Central
America with red corollas and petaloid phyllary apices have been considered
by Anderberg & Freire (1989) to be in two genera, Chionolaena and Gnaphalio-
thamnus. In contrast, a rationale for including all 9 (now 10) species of these
taxa in Gnaphaliothamnus was recently presented (Nesom 1990). The occur-
rence of morphological intermediates between two strongly divergent species
of this group, G. aectdiocephalus (Grierson) Nesom (a member of Chionolaena
fide Anderberg & Freire) and G. eleagnoides, strengthens the hypothesis that
they are congeneric.

Both Gnaphakothamnus aecidiocephalus and G. eleagnoides are endemic to
central Oaxaca. The former is known only from the Cerro del Humo (the type
locality) and the Sierra de Juarez; the latter has only a slightly wider range
and is at least partially sympatric with the former. The only other species of
Gnaphaliothamnus that occurs in the same area is G. salicifolius. I recently
considered the closest relative of G. aecidiocephalus to be G. concinnus on the
basis of its similarity in leaf morphology and incipient dioecism (Nesom 1990).
Gnaphaliothamnus eleagnoides was hypothesized to be most closely related to
G. salicifolius (the type of the genus), and all these species were considered
to be more closely related among themselves than to a group of four species
within the genus that have glandular leaf surfaces.

Nesom: New species and taxonomic notes on Gnaphalothamnus 3

Gnaphaliothamnus aecidiocephalus is strictly dioecious and the plants have
densely crowded, overlapping, strongly deflexed, obovate leaves 2.5-4.5 mm
long and densely close tomentose above, sessile heads in terminal clusters of
2-3, and phyllaries with a distinctly purplish red medial area. Plants of G.
eleagnoides have strictly heterogamous heads, spreading, oblanceolate leaves
15-42 mm long and glabrous above, distinctly pedicellate heads in corymboid
clusters of 15-40, held well above the leaves, and phyllaries with white apices
but without any red coloration.

Two duplicate sheets of a collection made in the Cerro del Humo bear
branches of plants that appear to represent two separate taxa (27 Jan 1963,
3000 m, MacDougal s.n. [MEXU]). Several of these branches have heterog-
amous heads and can be clearly identified as Gnaphaliothamnus eleagnozdes,
although they have relatively shorter leaves (10-15 mm long), a slightly re-
duced number of pistillate flowers (11-13), and close to a 1:1 ratio of pistil-
late:hermaphroditic flowers. The other branches on these two sheets appear to
be intermediate between G. eleagnoides and G. aeczdiocephalus in a number
of features, prominently including leaf size (5-8 mm long), shape (obovate-
oblanceolate), arrangement (spreading-deflexed, not so densely overlapping as
in G. aecidiocephalus), and vestiture (tomentose above but not densely so).
The heads are sessile in compact, terminal clusters of 8-12. Some heads are
all staminate, but some have 1-3 filiform-tubular flowers in the outermost se-
ries. These flowers are without anthers and have pappus bristles with barely
expanded apices, as is characteristic of pistillate flowers in the group, but they
are immature and it is not possible to tell if they are producing fertile ovaries.
The phyllaries have white apices but are faintly pink below that.

The evidence strongly suggests that these flowering branches of intermedi-
ate morphology represent hybrid plants and that the parental taxa should be
considered congeneric.

ACKNOWLEDGMENTS

I thank Drs. Billie Turner and Andrew McDonald for their review and
comments on the manuscript and the staff at MEXU for their help during a
recent visit there.

LITERATURE CITED
Anderberg, A.A. & S.E. Freire. 1989. Transfer of two species of Anaphalis
to Chionolaena. Notes Royal Bot. Gard. Edinburgh 46:37-41.

Nesom, G.L. 1990. Taxonomy of Gnaphaliothamnus (Asteraceae: Inuleae).
Phytologia 68:366-381.

Phytologia (July 1990) 68(6):4-13.

TAXONOMY OF VARILLA (ASTERACEAE: HELIANTHEAE)
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

A taxonomic treatment of Varilla is rendered. Two species are rec-
ognized, V. mezicana and V. tezana, the former having two varieties,
one of these, Varilla mexicana var. gypsophila B. Turner, is newly
described. Both species occur in northeastern México, V. tezana also
occurring in adjacent Texas. Illustrations and distribution maps for
both species are provided.

KEY WORDS: Varilla, Asteraceae, Heliantheae, México.

The genus Varilla was first established by Asa Gray with his description
of V. mezicana from collections obtained by Dr. Wislizenus in northeastern
Durango and Dr. Gregg in southernmost Coahuila. Gray subsequently added a
second, very different species, V. tezana, based on collections made by Charles
Wright in Texas. In the present paper, with the description of V. mezicana
var. gypsophila, I have added a third taxon to the genus, a rather distinct
regional element largely confined to gypseous soils of west central Coahuila.

CHROMOSOME NUMBERS

All of the taxa have been investigated chromosomally, and all have chro-
mosome numbers of n = 18 pairs. References to these are provided below. It is
my assumption that the haploid number, n = 18, is of ancient origin, probably
a polyploid on an ancestral base of z = 9.

Turner: Taxonomy of Varilla 5

GENERIC RELATIONSHIPS

Turner & Powell (1977) have discussed the likely relationships of Varulla,
concluding that it belongs to the tribe Heliantheae, in the subtribe Varillinae
along with several other genera, in particular Clappia (xz = 16), Pseudoclappia
(z= 18) and perhaps the anomalous genera Bebbia (z = 9) and Dyscritotham-
nus (x = ?). In short, they felt that Varilla represented an ancestral “proto-
type” from which these several genera might have arisen, all or most of them
occurring in dry, mostly saline habitats of northern México. Robinson (1981)
accepted the subtribal position of Varilla, but positioned both Clappia and
Pseudoclappia as the only members of a newly erected, but closely related,
subtribe Clappiinae. He correctly noted that Bebbia and Dyscritothamnus are
“unquestionably members of the Galinsoginae and are not closely related to
Varilla ...”.

TAXONOMY

Varilla A. Gray, Mem. Amer. Acad. Arts 4:106. 1849. Type species: Varilla
mezicana A. Gray.

Shrubs or succulent shrublets 0.5-3.0 m high. Stems glabrous or nearly so,
brittle and erect or recumbent and subsucculent or carnose. Leaves alternate or
mostly opposite, filiform and succulent or linear lanceolate and flattened, often
with punctate glands. Heads discoid, 5-50 in terminal subfasciculate cymules,
or solitary and terminal, borne on elongate peduncles. Involucres turbinate to
hemispheric, 4-8 seriate, markedly imbricate to subimbricate, the bracts ovate
lanceolate to lanceolate, glabrous, well bestowed with oil bearing striations
or raised linear ducts. Receptacle ovoid or conical, paleate, the pales like
the involucral bracts. Ray florets absent. Disk florets numerous. yellow, the
throat tubular with orange, oil bearing, ducts or striations. Achenes prismatic
or columnar, black, with 7-9 ribs, the pappus absent or of a few short bristles.
Base chromosome number, z = 18.

KEY TO SPECIES

1. Low succulent shrublets with recumbent stems to 50 cm high; leaves mostly
alternate, narrowly terete, ca. 1 mm wide ................... V. terana

6 PHYTOL Quan volume 69(1):4-13 July 1990

1’ Erect nonsucculent shrubs 0.7-3.0 m high; leaves mostly opposite, linear
lanceolate, flattened, mostly 2-10 mm wide ............... V. mezicana

Varilla mezicana A. Gray, Mem. Amer. Acad. Arts 4:106. 1849. Figure 1.
TYPE: MEXICO. Durango: between Pelayo and Cadena, without date,
Dr. Wislizenus 275 (LECTOTYPE (selected here]: GH!).

The protologue lists two collections, the above and “Valley east of Parras,
Dr. Gregg; April.” Both are mounted on a single sheet at GH and both fall
within the concept of the typical variety as conceived here (Figure 1). The
lectotype is the better preserved of the two collections, containing both flower
and fruiting material, matching closely the description provided by its author.

Shrubs 1-3 m high. Stems glabrous, brittle, with age the bark somewhat
corky; leaves mostly opposite, those of primary shoots linear, mostly 7-8 cm
long, 2-4(-5) mm wide, glabrous, the apices gradually narrowed to very slender
apices. Heads 5-15 in terminal corymbs, the ultimate peduncles mostly 5-15
mm long. Involucre, in fruit, decidedly turbinate, somewhat higher than wide,
5-6 mm high, the bracts 1-2 seriate, subgraduate, linear lanceolate, with 1-
2 orange linear ducts adorning the dorsal surfaces, similar to and grading
into the receptacular pales. Florets numerous, the corollas 3.5-5.0 mm long,
yellow, the tube ca. 1.5 mm long, sparsely glandular pubescent, the throat
abruptly tubular, 1.5-3.0 mm long, the lobes ca. 0.75 mm long. Achene 2-
3 mm long, 8-9 ribbed, black at maturity, sparsely pubescent with crinkly
hairs, the pappus of 5-8 poorly developed awns 0.2-0.7 mm long, sometimes
epappose. Chromosome number, n = 18 pairs (Turner & Flyr 1966).

Two varieties of this species are recognized; these are contrasted in the
following couplet.

1. Achenes 1.5-1.9 mm long, glabrous or nearly so; leaves of primary shoots
mostly 4-8 mm wide; mostly growing on gypseous soils . var. gypsophila

1’ Achenes 2.0-3.9 mm long, to some extent pubescent; leaves of primary
shoots mostly 2-4 mm wide; mostly growing on calcareous
BONG. 5h. Wai: « palrdeilic las ways sieieaahere Stee Vine ee var. mezicana

Varilla mexicana A. Gray var. gypsophila B. Turner, var. nov. TYPE:
MEXICO. Coahuila: ca. 85 air miles W of Cuatro Cienegas, ca. 2 miles
S of Salinas del Rey del Norte, on NW side of Laguna del Rey; on
gypsum sand above lake, 1050 m, 19 Sep 1974, James Henrickson 1414
(HOLOTYPE: LL; Isotype: MEXU).

Varilla mezicana var. mezicana similis sed acheniis brevioribus
plerumque glabrisque, surculis primariis foliis latioribus, et habi-
tationibus in terris gypsorum differt.

Turner: Taxonomy of Varilla

Fig. 1. VARILLA MEXICANA VAR. MEXICANA (POWELL
1867, TEX).

PHYTOLOGIA volume 69(1):4-13 July 1990

aye 4°10'W 99°45wr
30°O5N 30°05'N
TEX
CHI

DUR O NUE

FiG. 2. DISTRIBUTION OF VARILLA MEXICANA: VAR.
MEXICANA (e);
VAR. GYPSOPHILA (0 ).

Turner: Taxonomy of Varilla 9

DISTRIBUTION (Figure 2): west central Coahuila, mostly gypseous allu-
vial or sand blown soils, 700-1400 m; flowering all seasons.

The taxon is largely confined to gypseous soils, being especially common
on the gypseous dunes in the vicinity of Cuatro Cienegas. As indicated in
Figure 2, the two varieties are not known to occur together, but an occasional
intermediate between the two taxa may be found, although most such plants
are probably late flowering specimens of var. gypsophila in which the leaves of
secondary shoots become narrower, or else these are late flowering specimens
of var. mezicana in which the heads become somewhat smaller and possess
somewhat shorter, less pubescent achenes.

REPRESENTATIVE SPECIMENS: MEXICO. Coahuila: Bacon & Lev-
erich 1258 (TEX); Chiang, et al. 9511c (LL); Gieschen s.n. (TEX); Henrick-
son 6027 (LL); Henrickson 12108 (LL); Henrickson 12224 (LL); Henrickson
12254 (LL); Henrickson 12518 (LL); Henrickson 12550 (LL); Henrickson &
Lee 15904 (TEX); I.M. Johnston 7819 (GH); I.M. Johnston 8690 (GH, LL);
M.C. Johnston, et al. 10337 (LL); M.C. Johnston, et al. 10849 (LL); M.C.
Johnston, et al. 10875 (LL); M.C. Johnston, et al. 12163 (LL); M.C. John-
ston, et al. 12178 (LL); Leverich & Turner 1 (TEX); Marroquin 1087 (TEX);
Nesom, et al. 5237 (TEX); Pinkava 5260 (LL); Powell & Turner 2239 (TEX);
Purpus 4463 (GH); Stewart 2657 (GH); Turner 6195 (TEX); Wendt & Lott
(LL).

Varilla mezicana A. Gray var. mezicana Figure 1.

As described in the above account, the variety gypsophila not included;
chromosome number, n = 18 pairs (Powell & Powell 1977).

DISTRIBUTION (Figure 2): northeastern Durango and closely adjacent
southern Coahuila and northwestern Zacatecas, pine-oak-juniper woodlands
in calcareous soils, 1300-2000 m; flowering all seasons.

In his description, Gray noted the lectotype to have been collected in the
state of Chihuahua. The localities given, however, are both from Durango. In-
deed, I have not seen collections of Varilla mezicana from Chihuahua, although
it occurs close to that state in west central Coahuila.

REPRESENTATIVE SPECIMENS: MEXICO. Coahuila: Gonzdlez 310
(TEX); Gregg s.n. (GH); Palmer 680 (GH); Pringle 42 (GH, LL); Pringle
310 (TEX); Rodriguez & Carranza 2368 (TEX); Rodriguez & Carranza 2404
(TEX); Shreve & Tinkham 9852 (GH, LL); Thurber 855 (GH). Durango: Flyr
146 (TEX); M.C. Johnston, et al. 10991 (LL); Powell & Turner 1867 (TEX);
Turner 15049 (TEX). Zacatecas: M.C. Johnston, et al. 11521 (LL).

Varilla tezana A. Gray, Pl. Wright. 1:133. 1852. Figure 3. TYPE: UNITED
STATES. Texas: “Saline plains, from the Nueces to the Rio Grande,
Texas; Sept.” Wright s.n. (LECTOTYPE (selected here): GH!).

10 PHYTOLOGIA volume 69(1):4-13 July 1990

Fic. 3. VARILLA TEXANA (LEVERICH 22, TEX).

Turner: Taxonomy of Varilla 11

In the protologue a collection by M. Trecut from the same district is also
cited; I have designated the Wright collection as lectotype.

Low succulent shrublet to 50 cm high, the stem sarcose and recumbent;
leaves narrowly terete, succulent, 15-30 mm long, 0.5-1.5 mm wide; heads soli-
tary, terminal, on peduncles 10-20 cm long; receptacles ovoid, 1.8-2.0 times as
long as wide (ca. 9 mm high, 4-5 mm wide); disk florets 200+; achenes colum-
nar, black, epappose; chromosome number, n = 18 pairs (Powell & Turner
1963).

A very distinct species often forming relatively “pure” stands in saline river
bottoms. The species has been amply described by a number of workers.

DISTRIBUTION (Figure 4): Southernmost Texas and adjacent northern
Mexico in saline flats, 10-200 m; flowering all seasons.

REPRESENTATIVE SPECIMENS: UNITED STATES. Texas: Dimmit
Co., Correll & Johnston (LL). Hidalgo Co., Clover (LL). Maverick Co., Seigler,
et al. 1401 (TEX). Starr Co., Wood 745 (TEX). Webb Co., Correll & Johnston
19744 (LL). Zapata Co., Correll 35441 (LL). Zavala Co., Shinners 7386 (LL).

MEXICO. Nuevo Leon: Barkley 14350 (LL); Garcia 75 (TEX); Johnston
4361 (TEX), Johnston 5012 (TEX); Johnston & McMillan 6070 (TEX); John-
‘ston & McMillan 6072 (TEX); Levertch & Turner 22 (TEX); Nesom 6815
(TEX); Smith M644 (TEX). Tamaulipas: Heard 4 (LL, TEX); Johnston 4340
(TEX).

ACKNOWLEDGMENTS

I am grateful to Dr. Guy Nesom for the Latin diagnosis and to both him
and Dr. Timothy Ramamoorthy for reviewing the manuscript. Nancy Webber
provided the illustrations.

LITERATURE CITED

Powell, A.M. & S.A. Powell. 1977. Chromosome numbers of gypsophilic
plant species of the Chihuahuan Desert. Sida 7:80-90.

& B. Turner. 1963. Chromosome numbers in the Compositae VII.
Additional species from the southwestern United States and Mexico.
Madrono 17:128-140.

Robinson, H. 1981. A revision of the tribal and subtribal limits of the
Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51:1-102.

July 1990

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Turner: Taxonomy of Varilla 13

Turner, B.L. & D. Flyr. 1966. Chromosome numbers in the Compositae X.
North American species. Amer. J. Bot. 53:24-33.

Turner, B.L. & A.M. Powell. 1977. Helenieae - systematic review. in Brology
and Chemistry of Compositae 2:699-737 (V.H. Heywood, et al., editors,
Academic Press, London).

Phytologia (July 1990) 68(6):14-19.

NEW SPECIES, NAMES, AND COMBINATIONS IN MEXICAN ASTERACEAE
B.L. Turner
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

The following new species, names and combinations of Mexican
Asteraceae are proposed: Brickellia oligadenia (B.L. Robins.) B.
Turner, stat. nov.; Helianthus petiolaris Nutt. var. fallax (Heiser)
B. Turner, comb. nov.; Helianthus praecox Engelm. & A. Gray var.
runyonii (Heiser) B. Turner, stat. nov.; Hemizonia greeneana Rose
var. peninsularis (Moran) B. Turner, stat. nov.; Hymenoxys ursina
(Standl.) B. Turner, comb. nov.; Iva ambrosiifolia (A. Gray) A.
Gray var. lobata (Rydb.) B. Turner, comb. nov.; Microseris dou-
glasii (DC) Schultz-Bip. var. platycarpha (A. Gray) B. Turner, stat.
nov.; Otopappus pittieri (Greenm.) B. Turner, comb. nov.; Peryme-
nium fayi B. Turner, spec. nov.; Pseudogynoxys chenopodioides
(H.B.K.) Cabrera var. cummingii (Benth.) B. Turner, comb. nov.;
Viguiera superaxillaris (S.F. Blake) B. Turner, comb. nov.; Zin-
nia angustifolia H.B.K. var. littoralis (B.L. Robins. & Greenm.) B.
Turner, comb. nov.; Zinnia maritima H.B.K. var. palmeri (A. Gray)
B. Turner, comb. nov.

KEY WORDS: Asteraceae, México, Brickellia, Helianthus, Hemi-
zonia, Hymenozys, Iva, Microseris, Otopappus, Perymenium, Pseu-
dogynorys, Viguiera, Zinnia.

Preparation of a treatment of the Asteraceae for México (Turner & Nesom,
in prep.) has necessitated the following new species, names, and combinations.

Brickellia oligadenia (B.L. Robins.) B. Turner, comb. nov. BASIONYM:
Brickellia squarrosa (Cav.) B.L. Robins. var. oligadenia B.L. Robins.,
Mem. Gray Herb. 1:92. 1917.

McVaugh (1984) followed Robinson (1917) in treating this as a variety
of Brickellia squarrosa (Cav.) B.L. Robins. (nom. illegit.; = B. cavanillesi
iCass.| A. Gray), to which it is certainly related. He noted, however, the
geographical isolation of the taxon and the several characters that mark it, all

14

Turner: New names and combinations in Mexican Asteraceae 15

of which suggest that it is worthy of specific rank. It is readily distinguished
from B. cavanillesi by its larger, eglandular heads, appressed, more scarious
involucral bracts and larger, longer petioled, leaves. It also approaches B.
argyrolepis B.L. Robins. and occasional specimens with eglandular peduncles
might be mistaken for that taxon.

Helianthus petiolaris Nutt. var. fallax (Heiser) B. Turner, stat. nov. BA-
SIONYM: Helianthus petiolaris Nutt. subsp. fallar Heiser, Rhodora 60:279.
1958.

Helianthus praecox Engelm. & A. Gray var. runyonii (Heiser) B. Turner,
stat. nov. BASIONYM: Helianthus debilis Nutt. subsp. runyonw Heiser,
Madrono 13:161. 1956.

Heiser (Heiser, et al. 1969) subsequently placed this taxon as a subspecies
under Helianthus praecor Engelm. & A. Gray.

Hemizonia greeneana Rose var. peninsularis (Moran) B. Turner, stat.
nov. BASIONYM: Hemizonia greeneana Rose subsp. peninsularis Moran,

Trans. San Diego Soc. Nat. Hist. 15:286. 1969.

Hymenoxys ursina (Standl.) B. Turner, comb. nov. BASIONYM: Actznea
ursina Standl., Field Mus. Publ. Bot. 22:126. 1940.

Iva ambrosiifolia (A. Gray) A. Gray var. lobata (Rydb.) B. Turner, stat.
nov. BASIONYM: Cyclachaena lobata Rydb., N. Amer. Fl. 1:3310.
1972.

This variety is largely confined to the montane regions of eastern México
(type from near Monterrey, Nuevo Leon); westward it grades into the var.
ambrositfolta (type from trans-Pecos, Texas, U.S.A.). Jackson (1960) treated
these regional taxa as subspecies under Jva ambrostifolia; the supravarietal
classification seems unwarranted considering the degree of morphological in-
tergradation observed in regions of peripheral allopatry.

Microseris douglasii (DC.) Schultz-Bip. var. platycarpha (A. Gray) B.
Turner, stat. nov. BASIONYM: Calais platycarpha A. Gray, Pacific RR.
Rep. 4:113. 1857.

Chambers (1955) treated the present taxon as a subspecies of Microseris
douglaszz, noting its localized occurrence in southernmost California and ad-
jacent Baja California, México.

Otopappus pittieri (Greenm.) B. Turner, comb. nov. BASIONYM: Zez-
menia pittier: Greenm. in W.W. Jones, Proc. Amer. Acad. Arts 41:156.
1905.

16 PHY T OcL O1G LA volume 69(1):14-19 July 1990

Villasenor & Strother (1989) erected the monotypic genus Tuztla to ac-
commodate this species. In spite of their reasoned treatment which included
comparisons with species of Otopappus, Zermenza, and Verbesina, I find their
phenogram showing the relative isolation of this species unconvincing. Until its
position can be established with more certainty, it would appear more prudent
to position this within Otopappus where I perceive its immediate relationships.

Perymenium fayi B. Turner, spec. nov. TYPE: MEXICO. Sinaloa: along
highway 40, 53 mi NE of Mazatlan, on side of mountain, 10 Sep 1965,
Raymond C. Jackson 7230 (HOLOTYPE: TEX).

Perymenium pringlei B.L. Robins. & Greenm. similis sed foliis
minute strigillosis in paginis inferis, capitulis majoribus, et bracteis
involucro luteo-scariosis differt.

Shrubs to 1-2 m high. Stems (upper) tetragonal, deeply grooved on each
side, strigose. Leaves ovate lanceolate to linear lanceolate, 6-14 cm long, 0.5-
3.0 cm wide; petioles 2-10 mm long; blades trinervate from or somewhat above
the base, green and sparsely to moderately appressed strigose below, the mar-
gins remotely serrate. Heads campanulate, ca. 10 mm high, 8-10 mm wide
(excluding rays) arranged terminally in 4-8 flowered, subfasciculate corymbs,
the ultimate (mature) peduncles mostly 3-5 cm long. Involucres 3-4 seriate,
graduate, 6-8 mm high, the innermost bracts broad, yellowish, only sparsely
ciliate. Receptacular pales 5-7 mm long. Ray florets mostly 8, the ligules yel-
low, 8-12 mm long, 2-3 mm wide. -Disk florets 20-30; corollas 5.5-6.5 mm long,
the lobes hispidulous. Anthers brown, the appendages white. Achenes 3.5-4.0
mm long, ca. 2 mm wide, wingless, ciliate along the margins, the pappus of
ca. 30 deciduous bristles, mostly 2-3 mm long.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Sinaloa: 3.5 mi SW
of El Palmito, highway 40, ca. 6000 ft, 8 Nov 1964, D. Flyr 307 (TEX); 53 mi
NE of Mazatlan, 10 Sep 1965, Jackson 7233 (TEX).

The holotype has narrowly lanceolate leaves and is superficially markedly
different from the other two collections cited, the latter having ovate leaves.
Nevertheless all are very similar as to vestiture and details of head and floret
structure. While compared with Perymenium pringlei, the present species
might ultimately find its closest relationship with P. hintonaa McVaugh of
Michoacan and adjacent state of México; both of the latter species possess
similar large heads with large florets, but P. fayz has leaf blades acute to obtuse
at the base and the vestiture is strictly appressed strigillose throughout with
very short hairs.

It is a pleasure to name this species for Dr. John J. Fay, in recognition of
his scholarly treatment of this difficult genus.

Turner: New names and combinations in Mexican Asteraceae NG

Pseudogynoxys chenopodioides (H.B.K.) Cabrera var. cummingii (Benth.
ez Oerst.) B. Turner, comb. nov. BASIONYM: Gynorys cummingi
Benth. ez Oerst., Kjoeb. Vidensk. Med. Dask. Nat. Foren. 1852:106.
1852.

Robinson & Cuatrecasas (1977) treated this widely distributed taxon at
the specific level within Pseudogynorys. It appears to be exceedingly close
to P. chenopodioides, distinguished primarily by its pubescent foliage. Only a
single collection is known from Mexico (Oaxaca, 32 km N of Puerto Escondido,
Martinez, et al. 2720; MEXU, TEX), although the variety is common from
Guatemala southwards to Colombia. The var. chenopodiozdes is largely con-
fined to the Gulf slopes of México. One might make a case for the inclusion
of var. cummingii as an infraspecific category of P. haenke: (DC.) Cabrera,
the only other species of Pseudogynorys native to México, but | think its rela-
tionship is closer to P. chenopodioides. Detailed monography may ultimately
show that all of these are but allopatric regional units of a very variable P.

cordifolia (Cass.) Cabrera.

Viguiera superaxillaris (S.F. Blake) B. Turner, comb. nov. BASIONYM:
Hymenostephium superazillare S.F. Blake, Proc. Biol. Soc. Washington
37:57. 1924.

The type of this species (US!) is given as “La Bojada, Tamazula, Durango,
México, altitude 300-600 m, Nov. 1921, by J.G. Ortega (no. 4437).” Blake
positioned the species in Hymenostephium (which I consider to be part of
Viguiera, as do Robinson {1981] and McVaugh [{1984]), noting that it was
nearest to the widespread, highly variable V. cordataS.F. Blake of Blake (1918)
where it does not appear to have any close relatives. This is discussed in more
detail in Turner (1987) where I needlessly described Viguzera vorobikae B.
Turner, which is clearly a synonym of the present taxon.

Zinnia angustifolia H.B.K. var. littoralis (B.L. Robins. & Greenm.) B.
Turner, comb. nov. BASIONYM: Zinnia littoralis B.L. Robins. & Greenm.,
Proc. Amer. Acad. Arts 32:16. 1896.

This taxon, which is largely confined to the coastal areas about Mazatlan,
Sinaloa, was maintained at the species level by both Torres (1963) and Strother
(1979). In my opinion it is a localized coastal ecotype of the widespread
Zinnia angustifolia, with which it appears to intergrade. Strother distinguished
the latter (in key form) from Z. littoralis by the color of receptacular bracts
(“stramineous to tip” in Z. httoralis vs. “metallic yellow to bright, coppery
orange distally”), but I do not find these compelling distinctions; in nearly all
other characters it is like Z. angustifolia, the most obvious distinction being
the somewhat larger heads and shorter, broader leaves in Z. littoral.

18 PERT Os Orel A volume 69(1):14-19 July 1990

Zinnia maritima H.B.K. var. palmeri (A. Gray) B. Turner, comb. nov.
BASIONYM: Zinnia palmeri A. Gray, Proc. Amer. Acad. Arts 22:423.
1886.

As indicated by McVaugh (1984), Zinnia maritima is distinguished from Z.
palmerz by its “somewhat woody or almost shrubby” habit and petiolate leaves
which are “rounded to acute at base and the blades sometimes elliptic.” Such
plants grade into what Strother (1979) accepts as Z. palmert, although both
Torres (1963) and McVaugh (1984) treat the two taxa as synonymous. I agree
with Strother (1979) that “in spite of considerable morphological variation,
two distinct modes are discernible and distinguishable.” In short, typical var.
marituma appears to be a coastal ecotype of the widespread allopatric var.
palmeri, the former presumably confined to coastal regions from Acapulco,
Guerrero northwards to Jalisco; the latter is largely a taxon of the more interior
montane habitats. This relationship is similar to that noted for Z. angustifolia
var. littoralis and Z. a. var. angustifolia, the former being a coastal ecotype,
the latter being a more widespread montane element.

ACKNOWLEDGMENTS

Iam grateful to Guy Nesom and Carol Todzia for reviewing the manuscript;
the former also provided the Latin diagnosis.

LITERATURE CITED

Blake, S.F. 1918. A revision of the genus Viguiera. Contr. Gray Herb.
54:1-199.

Chambers, K.L. 1955. A biosystematic study of the annual species of Mi-
croseris. Contr. Dudley Herb. 4:207-312.

Heiser, C., D.M. Smith, $.B. Clevenger, & W.C. Martin. 1969. The North
American sunflowers (Helianthus). Mem. Torrey Bot. Club 22:1-218.

Jackson, R.C. 1960. A revision of the genus Jva L. Univ. Kansas Sci. Bull.
41:793-876.

McVaugh, R. 1984. Asteraceae, in Flora Novo-Galiciana 12:1-1157. Univer-
sity of Michigan Press, Ann Arbor, MI.

Turner: New names and combinations in Mexican Asteraceae 19
Robinson, B.L. 1917. A monograph of the genus Brickellta. Mem. Gray
Herb. 1:3-151.

Robinson, H. 1981. A revision of the tribal and subtribal limits of the
Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51:1-102.

& J. Cuatrecasas. 1977. Notes on the genus and species limits of
Pseudogynozys (Greenm.) Cabrera (Senecioneae, Asteraceae). Phytolo-
gia 36:177-192.

Strother, J.L. 1979. Extradition of Sanvitalia tenuts to Zinnia (Compositae-
Heliantheae) Madrono 26:173-179.

Torres, A.A. 1963. Taxonomy of Zinnia. Brittonia 15:1-25.

Turner, B.L. 1987. New taxa and combinations in Viguzera (Asteraceae,
Heliantheae). Phytologia 63:434-436.

Villasenor, J.L. & J.L. Strother. 1989. Tuztla, a new genus for Zermenia
pwttiert (Compositae: Heliantheae). Syst. Bot. 14:529-540.

Phytologia (July 1990) 68(6):20-31.

NEW SPECIES, NAMES, AND COMBINATIONS IN MEXICAN BIDENS
(ASTERACEAE: COREOPSIDEAE)

Tom E. Melchert & B.L. Turner

Department of Botany, University of Iowa, Iowa City, Iowa 52240 U.S.A.
&
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

In the preparation of a taxonomic treatment of Bidens for México
we have found it necessary to describe three new species: B. balsana
T. Melchert (from Guerrero), B. oaxacana T. Melchert (from Oaxaca),
and B. saltillensis T. Melchert (from Coahuila); two new varieties: B.
aequisquama (Fern.) Sherff var. guanajuatensis T. Melchert (from
Guanajuato) and B. sharpii (Sherff) T. Melchert var. tamazulapana
T. Melchert (from Puebla and Oaxaca); four new specific combina-
tions: B. cronquistii (Sherff) T. Melchert, B. hintonii (Sherff) T.
Melchert, B. pueblensis (Sherff) T. Melchert, and B. sharpii (Sherff)
T. Melchert; and four new varietal combinations: B. acrifolia Sherff
var. langlassei (Sherff) T. Melchert, Bidens chiapensis Brandegee
var. feddemana (Sherff) T. Melchert, comb. nov. B. odorata Cav.
var. rosea (Schultz-Bip.) T. Melchert, and B. schaffneri (A. Gray)
Sherff var. wrightii (A. Gray) T. Melchert. When appropriate, the
relationships of these various taxa are discussed. Especially notewor-
thy is the treatment of B. crongutstt1 and B. hintonii, both of which
were originally positioned in the genus Coreocarpus, and B. acrtfolta
var. langlasset which is transferred from Cosmos.

KEY WORDS: Asteraceae. Coreopsideae, Bidens, Coreocarpus,

Mexico.

The senior author and several of his former students have conducted exten-
sive cytotaxonomic and comparative flavonoid studies within and among the
more difficult genera of the Mexican and Central American Coreopsideae (Cos-
mos, Melchert 1967, 1968; Dahlia, Sorensen 1969; Giannasi 1972; Coreopsis,
Crawford 1969; Bidens, Hart 1973, Ballard 1975, Roseman 1986; and Theles-
perma, Melchert 1963). The broad taxonomic overview gained through this

20

Melchert & Turner: New names and combinations in Mexican Bidens 21

combination of studies, which were largely based upon populationally oriented
research, augmented by experimental studies on greenhouse grown plants, has
resulted in systematic treatments of the genera Bidens, Cosmos, and The-
lesperma. In order that the revised concepts might be incorporated into the
junior author’s concept of these genera for the Asteraceae of México (Turner &
Nesom, in prep.), the necessary new species descriptions, new names and new
nomenclatural combinations are presented here. More detailed morphological,
chromosomal, flavonoid and experimental crossing data relating to these will

be published later.

Bidens acrifolia Sherff var. langlassei (Sherff) T. Melchert, comb. nov.
BASIONYM: Bidens langlasse: Sherff, Bot. Gaz. (Crawfordsville) 64:24.
1917. = Cosmos langlassez (Sherff) Sherff, Field. Mus. Publ. Bot. 8:425.
1932.

ADDITIONAL SPECIMENS EXAMINED (OTHER THAN THE TYPE):
MEXICO. Guerrero: Distr. Mina, Guayameo-Filo, oakwoods, 1240 m, 12 Sep
1936, Hinton, et al. 9388 (G, MICH, NY, US).

Sherff, in his original description, correctly positioned this taxon in Bidens.
He later transferred it to Cosmos because its general habit was “more that
of a Cosmos than Bidens,” an arrangement which Sherff (1955) retained in
his treatment for the North American Flora. Since a meaningful analysis of
the characteristics distinguishing Cosmos and Bidens would require detailed
comparisons of their flavonoid chemistries, such a discussion cannot be fully
undertaken here. At a morphological level suffice it to note that: (1) Many
Bidens species (e.g., B. pilosa L., B. odorata Cav., B. mollifolia Sherff, B.
pringle: E. Greene, etc.) characteristically bear square or tetragonal stems,
whereas only terete stems are known in Cosmos; 2) while linear-tetragonal
achenes are basic in both genera. only those of Bidens display 3 or more,
nearly equally developed, elongated ribs on each face; and 3) while the fila-
ments of Bidens are glabrous and somewhat elastic (1.€., readily stretched by
the emerging style branches), those of Cosmos are invariably pilose and rela-
tively inelastic. Since Cosmos langlassei has all of the traits characteristic of
Bidens, there is little doubt that it is properly positioned in Bzdens. More-
over, “Cosmos” langlasset is morphologically all but indistinguishable from B.
acrifolia Sherff (including B. polyglossa Sherff) both of which Sherff himself
correctly positioned within Bidens (1955; 1964). All have sharply squared
stems arising from a short woody crown; well spaced, deeply 2-3 pinnatisect
leaves that are broadly triangular in outline; distinctive heads with 8-12 nar-
towly oblanceolate, two toned yellow rays subtended by (8-)11-16 narrowly
linear-subulate outer involucral bracts; exaristate, linear-tetragonal achenes
(some with 2 tiny smooth or weakly barbed awns), the innermost of which be-
comes purplish brown and slightly recurved apically (when mature). Indeed,

22 PAY TOL O'G LA volume 69(1):20-31 July 1990

the only characters which distinguish between Bidens acrifola (including B.
polyglossa) and “Cosmos” langlassez are leaflet size and shape, and stem/leaf
pubescence patterns as follows:

1. Ultimate segments linear oblanceolate to 2.5 mm wide; stems with scat-
tered appressed hairs, especially above; the leaves obscurely
hispidulose”. 2). lvls ese nes se ass eee ss eae ee var. acrifolia

1’ Ultimate segments narrowly linear to linear filiform, 0.5-0.8(-1.0 mm) wide
throughout; stems glabrous and leaves much more remotely
PUDESCEDE oe oisie seals teie les wale slotelopeue'snrne sible siete ae er var. langlasse1

Given the fact that leaf dissection patterns are generally highly variable
in Bidens (3-5 partite and 2-3 pinnatisect leaves commonly segregating within
single populations), var. langlasse: may ultimately prove to be nothing more
than a narrow leafleted, nearly glabrous form of B. acrifolia. However, since
var. langlassei is known definitely only from southwestern Guerrero (and per-
haps adjacent Michoacan), with var. acrifolza occurring in Jalisco, Sinaloa, and
Durango, I have chosen to treat them provisionally as geographical variants of
a single species.

The chromosome number of var. acrifolia is n = 10, a relatively uncom-
mon number in Bidens (reported as B. polyglossa); that of var. langlassez is,
unfortunately, unknown.

Bidens aequisquama (Fern.) Sherff var. guanajuatensis T. Melchert, var.
nov. TYPE: MEXICO. Guanajuato: Route 110, 8.6 mi E of Cd. Guana-
juato, near km 87, locally abundant in undisturbed areas of oak covered
hillside, 11 Oct 1971, Melchert, Ballard & Hart 71-94 (HOLOTYPE:
TEX; Isotype: MEXU).

Bidens aequisquamae (Fern.) Sherff var. aeguzsquamae similis
sed caulibus singulis (vs. fasciculatis), foliis 3(-5) pinnatisectis, seg-
mentis lanceolati-ovatis pubescentibus, in superficiebus ambabus,
et involucrorum bracteis exterioribus plerumque 7-10 (vs. 11-13).

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Guanajuato: Mpio.
de Dolores Hidalgo, 8-10 km above (northeast of) Santa Rosa, steep slopes
in oak forest, red clay soil with abundant humus cover, 2500-2600 m, 17 Sep
1967, McVaugh 28954 (MICH).

In addition to the characters referred to in the above diagnosis, var. gua-
najuatensis possesses weakly trimorphous achenes, 1.¢., though mostly black,
linear-subtetragonal and biaristate as in var. aequisquama, several achenes near
the periphery of each head are 4.0-5.5 mm long and bear prominent yellow
ribs (2 lateral, 1 ventral and sometimes 1 dorsal), plus scattered small yellow

Melchert & Turner: New names and combinations in Mexican Bidens 23

papillae on the blackish, inter-rib areas, such achenes occur just interior to
shorter (3.5-4.0 mm long), totally yellow, linear-clavate, peripheral achenes of
the fruiting heads (similar yellow peripheral achenes have been observed in a
few individuals from var. aequisquama populations).

It is entirely possible that var. guanajuatensis is a distinct species. In
addition to its obvious relationship to var. aequisquama, its overall habit also
suggests a close relationship to the large rayed forms of Bidens mollifolia,
which occur in the states of Michoacan, Jalisco, and México. Although plants
of the B. mollifolia complex develop diagnostic small, black, epappose, clavate
or linear-clavate achenes, studies by the senior author have shown that it is
common for a few achenes in totally mature fruiting heads of B. molhifoha to
develop yellow papillae or small irregular, yellow, calluslike marginal ridges.
Other than being epappose, such B. mollifolia achenes are rather similar to
the short yellow outer, but biaristate achenes seen in var. guanajuatensis.

Additional study is needed.

Bidens balsana T. Melchert, spec. nov. TYPE: MEXICO. Guerrero: Low
mountains 7.4 mi east of Chilpancingo on the road to Tixtla, plants
along roadcuts in dry hilly area with small oak, “annual-like” plants with
awned achenes and yellow rays with or without a red basal anthocyanin
spot (as in 206 A); or awnless perennial plants (as in 206 B); or single
awned, seemingly perennial plants (as in 206 C), 21 Oct 1971, Melchert,
Ballard & Hart 206 (HOLOTYPE: TEX; Isotype: MEXU).

Bidens serrulata (Poir.) Desf. var. sharp Sherff similis sed
differt plantis perennibus (vs. annuis) et involucrorum bracteis in-
terioribus moderate vel dense puberulis (vs. glabris).

Erect perennials 80-100(-160) cm tall, arising from a short rhizome, of-
ten multibranched from the base. Stems terete, mostly glabrous, but some
with scattered pilose hairs near the base (glaucous and totally glabrous in the
greenhouse). Leaves either tripartite with deeply toothed, ovate segments, or
deeply and evenly 2-3 pinnatisect with numerous lance-linear segments (invo-
lute and appearing subfiliform when growing under very dry conditions), the
primary segments at right angles to the midrib. Heads radiate, numerous, 4-5
cm across the expanded rays, each with a tuft of hairs at its base. Ray florets
mostly 5 per head. Ligules two toned yellow with a distinctive irregularly
shaped, maroon-red anthocyanin spot at their base, mostly 20-22 mm long,
8-11 mm wide, the red spot varying considerably in size, essentially absent in
some individuals. Outer involucral bracts (9-)11-15, narrowly linear, 4-6 mm
long, 0.3-0.4 mm wide, ciliate. Inner bracts moderately pubescent. Disc florets
numerous, yellow with a red spot just above the tube. Pales narrowly linear,
rounded apically, reddish brown above, becoming white with stripes below.
Achenes dimorphous, several of the peripheral ones in each head subclavate,

24 REY F/OLO:GihA volume 69(1):20-31 July 1990

3.5-6.8 mm long, at first olivaceous, but becoming yellowish or rubrocasta-
neous; inner achenes blackish, narrowly linear-tetragonal, 6-9(-11) mm long,
tapering slightly toward both ends, short setose apically, exaristate, or the
innermost with 1 or 2 short awns. Chromosome number, n = 11 pairs.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Guerrero: 4.2 mi
E of Chilpancingo on road to Chilapa, 21 Sep 1967, Melchert, Crawford, &
Averett 67-154 (IA, TEX); north slope of Cerro Alquitran, 10-14 km by road
W of Mex. highway 95 and Mazatlan, granitic rocks, 2250-2450 m, 6 Dec
1966, Anderson & Laskowski 4405 (F, G, MICH); Cerro Alquitran, Mpio. de
Chilpancingo, 2250 m, 6 Dec 1966, Rzedowski: 23687 (MICH).

This species closely resembles Bzdens sharpw (Sherff) T. Melchert var.
sharpit (also n = 11), but is distinguished by its perennial habit and densely
puberulent inner involucral bracts. Known only from two sites in Guerrero
(Chilpancingo area); the two collections from Cerro Alquitran all with tripar-
tite leaves and distinct red basal spots on their rays; those from the type local-
ity with 2-3 pinnatisect leaves and ligules with or without the red basal spot.
At the latter site, an extremely steep roadcut, some of the plants appeared
annual-like (perhaps slow growing), while some were definitely perennial. In-
terestingly, the type of the annual, B. sharpzz, the only collection of this annual
species from Guerrero, is also from this area (near Omitelmi).

Bidens chiapensis Brandegee var. feddemana (Sherff) T. Melchert, comb.
nov. BASIONYM: Bidens feddemana Sherff, Brittonia 16:60. 1964.

We would also place Bidens macvaughii Sherff (Brittonia 16:63. 1964)
as synonymous with this taxon; as does McVaugh (1984). Bidens chiapensis
has three geographically based morphological phases. Most collections of this
species have been obtained from Chiapas and Guatemala, where all plants
examined appear to be sprawling lignescent herbs, the radiate heads of which
bear mostly eight, short, pale lemon yellow rays per capitulum. North of the
Isthmus of Tehuantepec, the few available collections are all discoid. One col-
lection is known from Oaxaca. Except for having discoid heads, the latter is
essentially identical to the Chiapas-Guatemala plants (1.e., appears prostrate
and has three awned achenes). In contrast, the discoid plants from Guanaju-
ato, Michoacan (B. feddemana), and Jalisco (B. macvaughit), are stout, erect
shrubs with biaristate achenes. The erect shrubs are distinguished as var. fed-
demana, whereas the prostrate collection from Oaxaca is included here as a
discoid form of the usually short rayed var. chiapensts. Chromosome numbers
from the short rayed, prostrate, Chiapas-Guatemala populations are diploid
with n = 12 pairs; those of the discoid populations are unknown.

Bidens cronquistii (Sherff) T. Melchert, comb. nov. BASIONYM: Coreo-
carpus cronqutsti Sherff, Brittonia 16:433. 1964.

Melchert & Turner: New names and combinations in Mexican Bidens 25

Sherff presumably positioned this square stemmed, white rayed, “Bidens
odorata like” “annual” (Smith, 1981 notes the plant to appear perennial in
greenhouse grown material) in Coreocarpus because its small, obcompressed,
marginally incurved, epappose achenes superficially resemble those of Coreo-
carpus, i.e., have thick, corky wings along their somewhat tnrolled lateral mar-
gins. In total characters, however, this Guerrero endemic undeniably belongs
to Bidens, being particularly close to (and perhaps conspecific with) B. gracil-
lima Sherff, B. mznensis Sherff, and B. oligantha Brandegee (including B.
anthriscoides DC. var. decomposita Sherff). Like B. cronquzstz, these are all
small headed “B. odorata like” plants of the Sierra Madre del Sur with special-
ized achenes, neutral rays, and strongly dimorphic involucres (not possessing
pistillate rays and essentially monomorphous involucres as in more “typical”
Coreocarpus species).

In a recent biosystematic revision of Coreocarpus, Smith (1989) retained
C. cronguistu (and its close relative C. hintoni) in Coreocarpus; but quite
convincingly showed them to be morphologically and genetically distinct el-
ements within the genus. Speculating on their phylogeny he noted that the
plants of this distinct species pair “bear a striking resemblance to some species
of Bidens,” in particular, suggesting a possible relationship with the recently
described B. clavatus Ballard which, like B. oligantha, etc., is yet another
specialized member of the B. odorata species complex (with clavate achenes).
Despite the many features which Smith listed that ally B. cronqutsti (and
B. hintont) to Bidens (quadrangular stems, leaf dissection, ligule color, ach-
ene shape and chromosome number) he concluded that “their winged achenes,
however, keep them in Coreocarpus.” In our view, the inrolled marginal wings
on B. cronquisti achenes were most likely developed through the enlargement
and coalescence of yellow papillae and/or callus like ridges, such as occur along
the introrse margins of many mature achenes of B. oligantha, B. mznensis, and
B. gracillima. Achenes intermediate between these extreme forms are known:
MEXICO. Guerrero: Temisco, 350 m, 5 Nov 1937; Mezia 8748 (F, G, MO,
NY) PEX UE: US).

The transfer of Coreocarpus cronquisti and C. hintoni to Bidens, leaves
Coreocarpus congregatus (S.F. Blake) E.B. Smith (formerly Coreopsis congre-
gatus) as the only somewhat discordant (sterile rayed, dimorphic involucred)
element in Coreocarpus.

Bidens hintonii (Sherff) T. Melchert, comb. nov. BASIONYM: Coreocarpus
hintonit Sherff. Brittonia 16:58. 1964.

Sherff originally described this species as a Coreocarpus, no doubt because
all, or at least the outer, achenes in each fruiting capitulum have yellow to
brown, corky pectinate wings along their somewhat inrolled lateral margins
(the innermost achenes in some heads being much narrower, slightly attenu-
ated and yellowish at the tip, their margins and ventral midribs smooth or with

26 PHY EOL. Giz volume 69(1):20-31 July 1990

scattered small tuberculae). In its total characteristics, however, this species
is very similar to members of the Bidens mollifolia polyploid complex, par-
ticularly the populations occurring from the state of México, westward. Like
B. hintont, these are square stemmed, white to rose rayed perennials with
tiny black epappose achenes hidden among the chaff of the fruiting capitula.
While most B. mollifolia achenes are clavate to linear clavate and wingless,
our studies have shown that in totally mature fruiting heads it is common for
a few achenes to develop yellow papillae and/or yellow callus like marginal
ridges. It is very likely that the corky wings of B. hintonzi are essentially an
elaboration of the less ornate marginal outgrowths that occur in B. mollifolia.

Smith (1989) retained Bidens hintoni and B. cronquisti) in Coreocarpus,
but noted their morphological and genetic isolation within Coreocarpus and
their close relationship to certain Bidens species (see above discussion under
B. cronqutstit).

Bidens hintoniz was known to Sherff only by the type, a Hinton collection
from Guerrero (Distr. Mina). We have examined an additional collection,
as follows: MEXICO. Guerrero: San Antonio, Montes de Oca, 20 Oct 1937,
G.B. Hinton, et al. 11510 (GH, MICH, NY, US). The MICH specimen was
inexplicably annotated by Sherff as B. aequisquama (Fernald) Sherff. Smith
(1989) did not examine the above collection, but cited a recent additional
collection from Guerrero, 6 km SE of Guayameo, 820 m, Villasenor & Soto
s.n. (UARK, MEXU).

Bidens odorata Cav. var. rosea (Schultz-Bip.) T. Melchert, comb. nov.
BASIONYM: Bidens rosea Schultz-Bip., in Seem. Bot. Voy. Herald 308.
1856.

This is the correct varietal name for what heretofore has been called Bidens
odorata var. calcicola (Greenm.) Ballard er T. Melchert (1975). Sherff (1955),
however, referred the latter name to varietal status under B. pilosa. Under the
current Code of Botanical Nomenclature (1988), the varietal name rosea was
automatically created with the publication of B. rosea var. calcicola Greenm.
(1905); under article 26.2 of the present Code, the correct name should be var.
rosea.

Bidens oaxacana T. Melchert, spec. nov. TYPE: MEXICO. Oaxaca: Route
190, 39.8 mi SE of Totolapan (15.9 mi NW of Rio Hondo bridge and 8.2
mi SE of E] Camaron, adjacent to small roadside chapel; plants scattered
on rather open dry, oak-pine covered hillsides, 18 Oct 1971, Melchert,
Ballard & Hart 71-166 (HOLOTYPE: TEX; Isotype: MEXU).

Bidens steyermarkii Sherff similis sed differt foliis 2-3 pinnati-
sectis segmentis ultimis lineari-filiformibus et acheniis minutis ni-
gris sin pappo.

Melchert & Turner: New names and combinations in Mexican Bidens 27

A rather small, often somewhat tufted, rhizomatous perennial with small,
white rayed heads, young plants easily mistaken for annuals. Stems decid-
edly square. Leaves deeply 1-2 pinnatifid with linear filiform segments, these
only 0.5-1.0 mm wide. Flowering heads with ca. 5 tiny white rays, these 4-5
mm long, obovate, apically truncated. Disc florets yellow. Fruiting capitula
with numerous tiny, clavate achenes, these only 2.5-4.0(-5.0) mm long, black,
glabrous, epappose, hidden by the pales. Chromosome number, n = 12 pairs.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Oaxaca: ca. 105 km
WNW of Tehuantepec, ca. 65 km SE of Totolapan, ca. 1300 m, 6 Nov 1970,
Cronquist & Fay 1-880 (IA, NY).

While other square stemmed, white rayed species of Bidens have small,
clavate, epappose achenes (B. mollifolia, B. clavata Ballard, and B. steyer-
markit), B. oazacana is the only Mexican Bidens with such fruits, and with
deeply bipinnatisect leaves with narrowly linear to linear filiform leaf segments,
these mostly 0.5-1.0 mm wide. It’s closest relative appears to be B. steyer-
marku, a poorly known, once-collected species from western Guatemala.

Bidens pueblensis (Sherff) T. Melchert, comb. nov. BASIONYM: Bidens
bigelovr A. Gray var. pueblensis Sherff, Bot. Gaz. (Crawfordsville)
88:287. 1929.

SPECIMENS EXAMINED: MEXICO. Jalisco: ca. 28 mi W of Ayutla
and ca. 70 mi NW of Autlan, 3 Nov 1963, A. Cronquist 9743 (MICH, MO,
NY, US). Michoacan: 4.5 mi E of Cojumatlan, cliffs overlooking SE shore
of Lake Chapala, 7 Oct 1965, A. Cronquist 10292 (NY). México: route 55,
0.5 mi N of Ixtapan de Sal, 23 Oct 1971, Melchert, Ballard & Hart 71-288
(IA, TEX). México D.F.: Pedegral de San Angel, Sep 1927, E. Lyonnet s.n.
(US). Guerrero: Mpio. de Tlacatepec, Cerro Tlacatepec, near village of Agua
Fria, ca. 40 km N Coyuca de Benitez, 4 Dec 1963, R. Feddema 2904 (MICH);
Cerro Tlacatepec, near village of Agua Fria, ca. 40 km N Coyuca de Benitez,
4 Dec 1963, Rzedowski 18184 (MICH). Oaxaca: route 190 ca. 35 mi NW Cd.
Oaxaca, 19 Oct 1971, Melchert, Ballard, Hart 71-185 (IA, TEX); 10 km S
of Suchixtepec and 95 km N of Puerto Angel, 8 Nov 1970, A. Cronguist, J.
Fay 10897 (NY, US); Cerro San Felipe, 18 Oct 1908, C. Conzatti s.n. (FM).
Puebla: Hacienda Baton, vicinity of Puebla, Bro. G. Arsene s.n. (US).

While the overall morphology of this taxon is very much like Bidens bigelovi,
it is distinguished absolutely by the diploid chromosome complement (n = 12)
and its five, tiny (4-5 mm), two toned yellow rays, each with a minuscule red
anthocyanin dot at their base. Whether discoid or short radiate, B. bigelovi
is always tetraploid (n = 24), and the ligules, when present, are either white,
white with a red dot, or pale lemon yellow, the rays frequently varying from
2-5 per head within a given population.

Bidens saltillensis T. Melchert, spec. nov. TYPE: MEXICO. Coahuila:

San Lorenzo Canyon, “5.1 km (3.2 mi) en terraceria de la carreteria

28 Pra ret OG tA volume 69(1):20-31 July 1990

Saltillo a Zacatecas (54). 1 hora a pie arriba en el canon,” 2700-2800
m, forest of Quercus, Cupressus, and Buddleja, 18 Aug 1982, Clark P.
Cowan 3559 (HOLOTYPE: TEX; Isotype: MEXU).

Bidens odoratae Cav. var. rosea (Schultz-Bip.) T. Melchert
similis sed differt duratione perenni (vs. annui) et caulibus teretibus
(vs. quadratis).

Perennial herbs 20-40 cm high. Stems glabrous or nearly so, seemingly
terete (not clearly 4 sided), arising from a twisted, cordlike rhizomatous root
(root present on only one specimen. Leaves simple or 3-4 parted, 3-8 cm
long, the divisions linear lanceolate, mostly 2-4 mm wide, glabrous and entire
(rarely a few marginal hispidulous hairs). Heads radiate, mostly 1-3 per stem,
the ultimate peduncles 5-8 cm long. Involucres 5-6 mm high, the inner series
purple, glabrous, the margins prominently white scarious, the outer series
with mostly eight lanceolate, ciliate bracts 3-4 mm long. Receptacular bracts
about as long as the florets, similar to the inner involucral bracts but linear
lanceolate and narrowly acute. Ray florets 3-5, neuter, sterile (rarely a few
reduced stamens or style branches), the ligules 9-11 mm long, 5-7 mm wide,
white with 6-8 rosaceous veins. Disk florets 12-20, the corollas yellow, ca. 5mm
long, the lobes markedly short papillose pubescent. Anthers purplish brown
with yellow appendages. Achenes (somewhat immature) linear, 8-12 mm long,
brown, hispidulous, tetragonal below, somewhat tapering above, the pappus
of mostly 2 awns 0.5-1.0 mm long, sometimes deciduous or absent.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Coahuila: from above -
location at the same date, Cowan 3565 (MEXU, TEX); mountains near Saltillo,
3 Sep 1948, J. Greg 488 (G, MO).

The species superficially resembles the white rayed annual, Bidens odorata
var. rosea, but differs markedly because of its perennial habit and nearly terete
stems and 3-5 rayed heads. Indeed, it is the only white rayed perennial Bidens
known to have terete stems.

Bidens schaffneri (A. Gray) Sherff var. wrightii (A. Gray) T. Melchert,
comb. nov. BASIONYM: Bidens heterophylla Ortegies var. wraghtw A.
Gray. Proc. Amer. Acad. Arts 19:16. 1883.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Chihuahua: 2 mi
N of Madera, plants in moist ditch and adjacent cultivated fields, 2 Sep 1966,
Melchert, Sorensen, & Crawford 6270 B (leaves deeply pinnatisect); 2 mi N
of Madera, plants in moist ditch and adjacent cultivated fields, 2 Sep 1966,
Melchert, Sorensen, & Crawford 6270 A (leaves linear lanceolate) (IA, TEX);
ca. 10 mi N of Madera, above village of Bravo on west edge of Lago de Babicora,

1 Sep 1966, Melchert, Sorensen, & Crawford 6261 (1A, TEX).

Melchert & Turner: New names and combinations in Mexican Bidens 29

This taxon (which also includes Bidens insolita Sherff) has long resided in
synonymy under B. aurea (Ait.) Sherff and was so treated by Sherff (1955)
in his treatment for the North American Flora. While Bidens aurea is mostly
tetraploid with n = 23 pairs of chromosomes, B. schaffneri var. schaffnerz and
var. wrightw are both diploid with n = 11 pairs. The latter variety, which
occurs mainly in central Chihuahua. differs from the more southern, primarily
streamside, var. schaffnerz in possessing simple and/or tripartite leaves with
narrowly lanceolate blades, as well as deeply pinnatifid leaves and achenes with
barbed pappus awns. In addition, var. schaffneri has proven fully compatible
with var. wraghti in synthetic crosses (Melchert, unpubl.).

Bidens sharpii (Sherff) T. Melchert, comb. nov. BASIONYM: Bidens ser-
rulata (Poir.) Desf. var. sharpu Sherff, Bot. Leafl. 2:6. 1950.

SPECIMENS EXAMINED: MEXICO. Oaxaca: route 190 ca. 14 mi S of
Nochixtlan, King 2517 (TEX); along road from Ixtlan de Juarez to Villa Alta,
14 July 1968, Carman 68-39 (IA, TEX); route 190 ca. 15 mi NW of Yanhuitlan,
14 Sep 1967, Melchert, Averett, & Crawford 67-64 (IA, TEX); route 175, 21.5
mi N of jct with route 190 just N of El Cerezal, 18 Oct 1971, Melchert, Bal-
lard, & Hart 71-179 (IA, TEX); route 190 31.5 NW of Cd. Oaxaca, 19 Oct
1971, Melchert, Ballard, & Hart 71-186 (IA, TEX); route 190 128 mi SE of
Tamazulapan, 20 Oct 1971, Melchert, Ballard, & Hart 71-201 (1A, TEX).

This taxon, which Sherff misplaced in Bidens serrulata, is very common
across central Oaxaca (one collection, the type, from central Guerrero). In
sharp contrast with B. serrulata, which is centered in the state of Mexico, var.
sharpit mostly possesses a distinctive maroon red anthocyanin blotch at the
base of their two toned yellow ligules. The size of this spot varies consider-
ably. Though usually conspicuous, it may be quite small and difficult to see
when pressed, or rarely even absent. Regardless of ray color, however, var.
sharpii can be distinguished by its unique combination of (9-)11-15(-17) nar-
rowly linear outer bracts, glabrous inner bracts (indeed the plant is glabrous
throughout, except for a tuft of hairs at the base of each capitulum), and
a chromosome number of n = 11. Bidens serrulata has totally yellow ligules
(these also two toned but never with a basal red spot), 8 wide, outer involucral
bracts, pubescent inner involucres, and a chromosome number of n = 12.

Bidens sharpii (Sherff) T. Melchert var. tamazulapana T. Melchert, var.
nov. HOLOTYPE: MEXICO. Oaxaca: Mpio. Tamazulapan, Cerro Peri-
con, al NW de San Pedro Nopala, ca. 2460-2660 m, “Veq. Ecetonia
matorral espinoso-encinar. Suelo cafe rojizo sobre roca ignea,” 21 Oct

1984, P. Tenorio L. 7882 (HOLOTYPE: TEX; Isotype: MEXU).

Bidens sharpu (Sherff) T. Melchert var. sharpiz arcte similis
sed differt bracteis interioribus receptaculi dorsaliter pubescentibus

30 PHYTOLOGIA volume 69(1):20-31 July 1990

(vs. glabris) et bracteis involucralibus ad apices purpuratis, area
purpurata 1-2 plo longiore quam latiore.

Much resembling var. sharpz but the inner involucral bracts pubescent
dorsally (vs. glabrous), the inner receptacular bracts purple apically, the purple
area 1-2 times as long as wide.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Puebla: Mpio. Cal-
tepec, Maguey Manzo, E] Gavilan al NW de San Simon, 3 Oct 1984, Tenorio
L. 7567 (MEXU, TEX); Cerro El Gavilan, SE de Caltepec, 1880-2320 m, 11
Oct 1984, Tenorio L. 7644 (MEXU, TEX).

The var. tamazulapana resembles var. sharpii, but has involucral bracts
moderately to densely pubescent with multicellular hairs; interior phyllaries
with distinctive jet black tips, these rather sharply delineated basally (not
elongated and gradually tapered as in var. sharpzt); rays with or without the
red basal anthocyanin spot (the holotype entirely yellow rayed); some of the
disc florets with tiny dark spherical glands on their teeth. It is known only
from three recent collections (cited above). While treated here as an isolated
variant of Bidens sharpii, the combination of “B. serrulata like” multicellular
involucral hairs, jet black chaff tips, yellowish anther appendages, tiny dark
glands on some of its disc corolla teeth, and its somewhat isolated distribution,
suggest that var. tamazulapana may very well prove to be a distinct local ©
species. Additional study is needed to affirm its biological status.

ACKNOWLEDGMENTS

The senior author is grateful to B.L. Turner and Guy Nesom for their
encouragement in these endeavors, to the latter for the Latin diagnoses, and
to both for their review of the manuscript. Much of the field work connected
with this contribution has been supported by N.S.F. grants, GB-3851 and
GB-6684X.

LITERATURE CITED

Ballard, R.E. 1975. A biosystematic and chemosystematic study of the
Bidens pilosa complex in North and Central America. Doctoral Thesis,
Univ. of Iowa, Iowa City.

Crawford, D.J. 1969. A cytotaxonomic and chemotaxonomic study of Core-
opsis in Mexico. Doctoral Thesis, Univ. of Iowa, lowa City.

Melchert & Turner: New names and combinations in Mexican Bidens 31

Giannasi, D.E. 1972. The flavonoid systematics of the genus Dahha (Com-
positae). Doctoral Thesis, Univ. of Iowa, Iowa City.

Hart, C.R. 1973. Systematics of the Bidens ferulaefolia complex: a cytotax-
onomic and comparative flavonoid investigation. Doctoral Thesis, Univ.
of Iowa, Iowa City.

McVaugh, R. 1984. Bidens, in Flora Novo-Galiciana 12:122-150. University
of Michigan Press, Ann Arbor, MI.

Melchert, T.E. 1963. Systematics of the genus Thelesperma (A cytotaxo-
nomic and chemotaxonomic study). Doctoral Thesis, Univ. of Texas,
Austin.

. 1967. Systematic studies in the Coreopsidineae (Compositae): New
Cosmos from Mexico. Sida 3(3):170-176.

. 1968. Systematic studies in the Coreopsidineae: cytotaxonomy of
Mexican and Guatemalan Cosmos. Amer. J. Bot. 55:345-353.

Roseman, R.R. 1986. A systematic study of Bidens section Greenmaniza in
Mexico, Central America and Jamaica: chemotaxonomy, cytotaxonomy
and phenetics. Doctoral Thesis, Univ. of lowa, lowa City.

Sherff, E.E. 1955. Bzdens, in N. Amer. FI., Ser. II 12:70-130.

Smith, E.B. 1989. A biosystematic study and revision of the genus Coreo-
carpus (Compositae). Syst. Bot. 14:448-472.

Sorensen, P.D. 1967. Systematics of the genus Dahlia (Compositae, He-
liantheae - Coreopsideae). Doctoral Thesis, Univ. of Iowa, lowa City.

Phytologia (July 1990) 68(6):32-39.

TWO NEW SPECIES OF MEXICAN BACCHARIS (ASTERACEAE: ASTEREAE)
Guy L. Nesom
Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

Southeastern Mexican populations previously considered to repre-
sent Baccharis heterophylla Kunth are here recognized as the new species,
B. glandulifera spec. nov. The latter is endemic to Oaxaca and Chi-
apas, and is allopatric with B. heterophylla, now considered to be re-
stricted to central, western, and northwestern México. The type of B.
vaccinioides Kunth, a name traditionally applied to plants of Chiapas
and adjacent Central America, was collected in Hidalgo and is con-
specific with B. conferta Kunth. Since the southern “vaccinium like”
plants have never received a name, they are formally described here as
B. confertoides spec. nov., an epithet chosen to emphasize their close
relationship to B. conferta.

KEY WORDS: Baccharis, Asteraceae, Astereae, México, Central

America.

In connection with taxonomic studies of the Asteraceae- Astereae of México,
two aspects of the systematics of Baccharis are discussed here in some detail.
Hypotheses of the relationships of these taxa in a broader context will be
presented in a separate paper (Nesom 1990).

I. A new species related to Baccharis heterophylla.

Baccharis glandulifera Nesom, spec. nov. TYPE: MEXICO. Oaxaca:
Mpio. Juquila, Puertecillo de Lachoa, km. 180 carretera Oaxaca-Puerto
Escondido, pine-oak woods, 2000 m, 14 Apr 1965, Rzedowski 19619 {pis-
tillate) (HOLOTYPE: TEX!; Isotype: ENCB). (Rzedowski 19618 is a
staminate specimen from the type locality. Rzedowski 19617, also col-
lected at the same locality, is Baccharis trinervis Pers.).

Baccharis heterophylla Kunth similis sed phyllariis ad apices

glandibus fuscatis tumidisque, acheniis minoribus, et pappo setis
brevioribus differt.

32

Nesom: Two new Mexican Baccharis 33

Glabrous, glutinous shrubs or small trees 1-3(-4.5) m tall. Leaves bright
green or yellow green, thick, with sunken resin glands, oblanceolate to oblong
oblanceolate, distinctly or obscurely trinerved, with 1-4 pairs of teeth, usually
in distal half, infrequently entire, 20-50 mm long, 4-15(-20) mm wide, 3-5
times longer than wide, apex blunt to obtuse, base cuneate or attenuate to a
short petiole or petiolar region 1-3 mm long. Heads sessile or subsessile in tight,
terminal clusters, campanulate, 4-5 mm wide, 3.5-5.0 mm long; phyllaries of at
least the outer series with a dark brown or blackish, sharply delimited, swollen
gland on the abaxial apex, stramineous below, margins thin and translucent.
Hermaphroditic corollas 4 mm long, pubescent on the limb and upper tube.
Pistillate corollas tubular, 2-3 mm long, with fimbriate apical extensions; style
branches 0.5 mm long. Achenes 0.8-1.1 mm long, 0.4-0.6 mm wide, 10-11
nerved; mature pappus 4.0-6.5 mm long.

Oaxaca to central Chiapas (Figure 1); pine, pine-oak, or pine woods, dis-
turbed forest of Alnus, evergreen forest of Tarodzum and Ficus; 1000-2950 m;
flowering December through May (-July).

Representative collections examined: MEXICO. Chiapas: Mpio. Venus-
tiano Carranza, at NE boundary of Aguacatenango, Breedlove 9645 (LL); 5
km N of Ixtapa along road to Soyalo, along the Rio Laja, Breedlove 35077
(MO, TEX). Oaxaca: San Pablo Macuiltianguis, 10 km from the entrance to
la Puerta del Sol, Calzada 4975 (TEX); near Oaxaca, Rancho Benito Juarez
on road up mountains from Teotitlan de Valle, Carlson 1431 (TEX); 2kmS
of E] Punto on Hwy 175, ca. 30 km N of Oaxaca, Conrad 3145 (MO); 37.9 mi
S of bridge at Valle Nacional on Hwy 175, Croat 48130 (MO); 21 km NE of
Ixtlan de Juarez on Hwy 175, Judziewicz 3371 (WIS); El Tejocote on Rte 190,
King 6456 (MO); 20 km NE of Oaxaca on road to Ixtlan, Marcks & Marcks
1107(LL); on road from Ayutla to Zacatepec, 3 km S of turnoff to Zacatepec,
Nee & Martin 32217(TEX); between Macuiltianguis and Puerta del Sol, Or-
tega O. & Ortiz T. 1627(TEX); 20 km NE of Oaxaca, on hwy from Juarez to
Ixtlan, Rzedowski 19271 (TEX); Cuauhtlilla, 28 Nov 1895, Seler 1510 (GH);
Rancho de Calderon, 11 Feb 1895, Smith 974 (GH); 4.8 km W of Sto. Tomas
Ocotepec, Torres C. 2176 (MO); 14 km N of Diaz Ordaz, by the detour to
Cuajimoloyas, Torres C. 2817 (TEX).

Baccharis glandulifera has previously been identified as B. heterophylla, to
which it is most closely related, but the new species is particularly distinctive in
its phyllaries with dark, swollen, apical glands. Some plants of B. heterophylla
may have darkened phyllary apices but never the deep and conspicuous gland
pockets of the plants to the south. Additionally, although the measurements
are overlapping, the leaves and achenes of the two taxa are different in average
size, as noted in the following couplet.

1. Leaves (21-)30-65 mm long, (6-)8-20 mm wide; phyllaries with a prominent,
dark, glandular pocket at the abaxial apex; achenes 0.8-1.3 mm

34 PHY fT OsiwW GebA volume 69(1):32-39 July 1990

-
4 ¢
"
> Beso Zuhal
s
on eo DoD ir &
ata ale i
eee eee 8
| eS ee TO
ae ine,
- 3 =
=~ Bos &
oF Ee VG
oo; = @
as Cece. EP
Pg 0m, SG
>! 2 ~ i
an phe
2

Figure 1. Geographic distribution of Baccharis heterophylla, B. glandulifera,
B. halimifoha, and B. lancifolia.

Nesom: Two new Mexican Baccharis 35

oi Re Oe cS SS Ce ae eR eae ee eae eee nee B. glandulifera

1’ Leaves (20-)25-50 mm long, 4-12 mm wide; phyllaries sometimes darkened
at the apex but never with a gland pocket; achenes 1.3-1.6 mm
[SNE rs tee see ees eee eee came tote canine mere mere B. heterophylla

All records of Baccharis heterophylla from Oaxaca to Chiapas apparently
are referable to B. glandulifera (Figure 1). I have seen only two collections
from Oaxaca that have phyllaries with only weakly developed gland pockets:
Smith 874 and Seler 1510. Both of these are apparently from well inside the
range of the gland bearing plants, and both have larger leaves and smaller
achenes typical of those in the surrounding area. The Seler collection is a
damaged branch with a mixture of mature and immature heads.

In order to emphasize their close relationship, these two population svstems
might be regarded as conspecific and recognized at varietal rank. Nevertheless,
since they are sharply distinct morphologically and separated geographically
with no region of intergradation, the recognition of both at the rank of species
seems justified.

Also mapped on Figure 1 is Baccharts halimifolia L., which is morpholog-
ically very similar to B. heterophylla. Baccharis halimifolia typically grows in
swampy or lakeside habitats and is at the southwestern most extension of its
range in México.

II. The identity of Baccharis vacciniordes.

The population system of plants known as Baccharis vaccinioides Kunth,
which occurs from Chiapas to Guatemala, Honduras, and El Salvador, has
been accepted by most systematists, including Matuda (1957) and Nash (1976)
as a distinctive species. Apparently, however, no one has critically considered
the location of the type collection or evaluated its identity, because the col-
lection was made in the state of Hidalgo, far north of the range of the plants
it traditionally has been associated with. Kunth’s description clearly notes
that the leaves of B. vacctnioides are uninerved and entire or toothed toward
the apex of both margins, and the photograph (fiche) of the type specimen
confirms this. Thus, given the type locality and the morphology of the plant
itself, B. vacctnioides must be considered a synonym of B. conferta Kunth.
The “vaccinium like” plants, which apparently have never been given a name,
are formally provided below with a type and epithet.

Baccharis confertoides Nesom, spec. nov. TYPE: MEXICO. Chiapas:
Mpio. San Cristobal Las Casas, E side of Zonthehuitz near summit,
evergreen cloud forest, Drimys, Clethra, Quercus, and Cleyera, 2800 m,

36 PHY TOLO GTA volume 69(1):32-39 July 1990

19 Dec 1972, D.E. Breedlove 304386 (HOLOTYPE: LL!; Isotypes: CAS,
MEXU!).

Baccharis confertae Kunth similis sed foliis integris praecipue

differt.

Glabrous, glutinous shrubs or small trees 1.0-3.5 m tall. Leaves elliptic to
elliptic obovate, entire, uninerved, 10-28 mm long, 6-12 mm wide, punctate,
with minutely papillate margins. Heads 3-5 mm wide, 3-4 mm long; phyllar-
ies sometimes with a colored apical portion or rim. Hermaphroditic corollas
distinctly pubescent with short, white hairs, at least on the upper tube and
lower limb. Achenes 1.1-1.3 mm long; mature pappus 4-5 mm long.

Chiapas (México), to Guatemala, E] Salvador, and Honduras (as noted by
Nash [1976]) (Figure 2); roadsides, slopes in oak, pine-oak, often with alder,
or evergreen cloud forests; 1600-3600 m; December- April.

Representative collections examined: MEXICO. Chiapas: Mt. Pasitar, 30
Dec 1936, Matuda 722 (LL).

GUATEMALA. Dept. Totonicapan: near San Francisco El Alto, 12 Jan
1941, Standley 83137 (LL).

EL SALVADOR. Dept. Chalatenango: near summit of Los Esesmiles, 10
Mar 1942, Tucker 986 (LL).

Baccharis conferta Kunth, Nov. Gen. & Sp. Pl. 4 {folio]:43. 1818; 4 [quarto]:55.
1820. TYPE: MEXICO. [Morelos]: near Cuernavaca, Apr [1803], Hum-
boldt & Bonpland s.n. (P fiche!, B-WILLD [photo TEX!)).

Baccharis vaccinioides Kunth, Nov. Gen. & Sp. Pl. 4 [folio]:39. 1818; 4
[quarto]:50. 1820. TYPE: MEXICO. {Hidalgo}: near Moran, May
(1803], Humboldt & Bonpland s.n. (P fiche!, photo GH!). Not B.
vaccinioides Gardn. (1845) from Brazil.

Baccharis resinosa Kunth, Nov. Gen. & Sp. Pl. 4 [folio]:41. 1818; 4
[quarto]:52. 1820. TYPE: [MEXICO]: “In America meridionali,”
no date [1803], Humboldt & Bonpland s.n. (P fiche!).

Baccharis zalapensis Kunth, Nov. Gen. & Sp. Pl. 4 [folio]:44. 1818; 4
[quarto]:56. 1820. TYPE: MEXICO. [Veracruz]: near Xalapa, Feb
[1803], Humboldt & Bonpland s.n. (P fiche!).

Baccharis congesta DC., Prodr. 5:410. 1836. TYPE: MEXICO. [Hi-
dalgo}: Real del Monte, Haenke s.n. (HOLOTYPE: G-DC fiche!
[photo TEX!]).

Baccharis orizabaensis Schultz-Bip. ez Hemsley, Biol. Centr. Amer.,
Bot. 2:130. 1881. SYNTYPES: MEXICO. [Veracruz]: Peak of
Orizaba, {no date], Linden 1138 and Liebmann 358 (K).

Nesom: Two new Mexican Baccharis 37

conferta

eee —._—_—__——. ml
150

km
25

. confertoides

® B. dioica

50
@ B.
{0}

Seis
°
PY
ee
e
e
2

)

eal

Figure 2. Geographic distribution of Baccharis conferta, B. confertoides, and
B. dioica.

38 PUHCY T OL0.GiVA volume 69(1):32-39 July 1990

San Luis Potosi, Michoacan, Guerrero, México, D.F., Hidalgo, Tlaxcala,
Morelos, Veracruz, Puebla, Oaxaca (Figure 2); clearings or woods edges, grassy
areas, often grazed or recently burned, in areas of pine-oak. pine, pine-fir, or
fir woods; (100-)1900-3250 m; February-May.

The two species are separated by the following couplet.

1. Shrubs 0.3-2.0 m tall; leaves mostly obovate-oblanceolate, with 1-2(-3)
pairs of coarse teeth near the apex; phyllaries sometimes with a dark
midvein but without a colored apical area or rim .......... B. conferta

1’ Shrubs to small trees 1.0-3.5 m tall; leaves mostly elliptic to slightly
oblanceolate, the margins entire; phyllaries often with a colored apical
POLIO FORTIN 5 ooh pn a alive aiecenP elas weyenala ehctane aie B. confertozdes

Rare individuals of Baccharis conferta produce leaves that are entire or
nearly so, but even then, a few other leaves on those plants usually have
slightly toothed margins. Even more rare are specimens on which all the
leaves are entire, such as Arsene & Nicolas 5157 (GH), collected in the vicin-
ity of Cd. Puebla. The latter has small, oblanceolate leaves and was collected
in an area from which a number of other specimens with small but typically
toothed leaves have been taken. The southeasternmost collection known of B.
conferta (Croat & Hannon 65795 |TEX], from the Uxpanapa region of eastern
Oaxaca) is morphologically very typical of the species, although at an unusu-
ally low elevation, and the distributional disjunction to the south is correlated
with an abrupt transition in morphology as well. Further study may show that
these closely related taxa are better recognized as varieties of a single species,
but such a study should necessarily also account for variation patterns in the
related B. tricuneata (L. f.) Pers. of South America. Nash (1976) has noted
that B. tricuneata, which has several varieties (Cuatrecasas 1968), might be
conspecific with B. conferta. Each of these three taxa, however, occupies a
discrete geographic region separate from the others, and it seems more reason-
able at present to recognize each as a separate species, while acknowledging
their close relationship.

Also mapped on Figures 1 and 2, respectively, are Baccharis lancifolia
Schlecht. and B. dioica Vahl, closely related species that also produce unin-
erved, elliptic to oblanceolate, and entire leaves. The former differs from both
B. conferta and B. confertoides in its much larger leaves with smooth margins
and acute apices, and the latter differs in its completely smooth leaf margins,
much shorter pappus, and its sandy, coastal habitats.

Nesom: Two new Mexican Baccharis 39

ACKNOWLEDGMENTS

I thank Drs. Billie Turner and Tom Wendt for their review and comments
on the manuscript, and GH, MO, and WIS for loans of specimens. The dis-
tribution records are from specimens on loan and from the collections at LL,

MEXU, and TEX.

LITERATURE CITED

Cuatrecasas, J. 1968. Notas adicionales, taxonomicas y corologicas, sobre

Baccharits. Rev. Acad. Colomb. Cienc. 13:201-226.

Matuda, E. 1957. El género Baccharis en México. Anales Inst. Biol. Univ.
Nac. México 28:143-174.

Nash, D.L. 1976. Baccharisin Flora of Guatemala. Fieldiana: Bot. 24(12):143-
148.

Nesom, G.L. 1990. Infrageneric taxonomy of North and Central American
Baccharis (Asteraceae: Astereae). Phytologia 69(1):40-46.

Phytologia (July 1990) 68(6):40-46.

INFRAGENERIC TAXONOMY OF NORTH AND CENTRAL AMERICAN
BACCHARIS (ASTERACEAE: ASTEREAE)

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

The 43 Baccharis species of the United States, México, and Central
America are placed into six sections, based on morphology: sect. Bac-
charts (13 species), sect. Sergilae (3 species), sect. Glandulocarpae
sect. nov. (10 species), sect. Aristidentes sect. nov. (12 species), sect.
Baccharidastrum (1 species), and sect. Molinae (4 species). Taxa with
paleate receptacles, previously placed in sect. Trinervatae. are here in-
cluded with the epaleate taxa of sect. Molinae. Long disjunctions be-
tween North and South America are noted among the species of sect.
Sergtlae and sect. Baccharidastrum.

KEY WORDS: Baccharis, taxonomy, Astereae, Asteraceae, North

America, South America.

The genus Baccharis is one of the largest in the Astereae, with approxi-
mately 450-500 species, all of which are native to the New World. It is most
highly speciose in South America, where about 90% of the species occur and
where a great amount of morphological diversity is found. Additional taxa
are endemic to the West Indies. Cuatrecasas (1967) outlined the composi-
tion of sections named up to that time, and clarified the typification for many
of them. Recent taxonomic treatments are available for species of a num-
ber of regions of South America, e.g., Colombia (Cuatrecasas 1969), central
Argentina (Espinar 1973), and Brazil (Barroso 1976). The author of each
of these presented a summary of the relationships among the species in the
geographic area under consideration. Espinar, in particular, provided signifi-
cant information regarding morphological variation among the species groups.
Zdero, et al. (1986) studied the chemistry of nine Argentinian species, and in
a broader view, identified eleven sections in the genus based on the distribu-
tion of secondary chemical compounds. An overview of various aspects of the
taxonomy and biology of the genus has recently been published (Boldt 1989),
but a modern taxonomic summary of the entire genus is lacking.

40

Nesom: North and Central American Baccharis 4]

During the preparation of a taxonomic treatment of the Baccharis species of
México, it has become clear that they can be placed into relatively few natural
groups, based on morphology. The following sectional synopsis accounts for
all known North and Central American species: only four species occur in
the United States that are not also found in México, and all of the Central
American species also occur in México. About fifteen other sections have been
identified among the South American species.

Baccharis L., Sp. Pl. 860. 1753. Type species: Baccharis halimzfolia L.. typ.
cons. prop. (Hellwig 1989).

1. Section Baccharis. Type species: Baccharis halimifolia L., typ. cons. prop.
(Hellwig 1989).

Sect. Cunezfoliae DC., Prodr. 5:405. 1836. Type species: Baccharis
cuneifolia (Lam.) DC.

Sect. Involucratae Heering in Reiche, Fl. Chile 4:17. 1903. Lectotype
species (Cuatrecasas 1967): Baccharis conferta Kunth, Nov. Gen.
& Sp. Pl. 4 [folio]:43. 1818; 4 [quarto):55. 1820. TYPE: MEXICO.
[Morelos]: near Cuernavaca, Apr {1803], Humboldt & Bonpland s.n.
(P fiche!, B-WILLD [photo TEX!]).

Sect. Glomeruliflorae Heering, Jahrb. Hamburg. Wiss. Anst. 21:32.
1904. Type species: Baccharis glomeruliflora Pers.

Glabrous shrubs or small trees: leaves cuneate to obovate oblong, punc-
tate, not papillate, glutinous, uni- or trinerved, entire or with a few coarse,
blunt teeth near the apex; heads sessile to subsessile in terminal or axil-
lary glomerules, sometimes racemoid; receptacles epaleate; achenes glabrous,
1.0-1.8(-2.0) mm long, with 8-11 thin nerves; pappus bristles in 2(-3) series.
Widespread in North and South America, apparently including many more
species, and perhaps more sectional synonyms as well, than those listed be-
low. Baccharis angustifolia and B. glomeruliflora, both of the southeastern
United States, do not occur in México.

Species included: Baccharis angustifolia Michaux, B. conferta Kunth, B.
confertoides Nesom, B. dioica Vahl, B. emory: A. Gray, B. glandulifera Nesom,
B. glomeruliflora Pers., B. halimifolia L., B. heterophylla Kunth, B. lancifolia
Schlecht., B. neglecta Britt., B. pilularis DC., and B. salicina Torrey & A.
Gray.

Additional representative species of South America: Baccharis caespitosa
(Ruiz & Pavon) Pers., B. chilco Kunth, B. macrantha Kunth, B. petiolata DC.,
B. spicata (Lam.) Baill., B. tricuneata (L. f.) Pers.. and B. tridentata Vahl.

Cuatrecasas (1967) chose Baccharis tridentata Vahl as the lectotype of sect.
Cylindricae Heering, highly polyphyletic as constituted by Heering, with the

42 PH YHOO GIA volume 69(1):40-46 July 1990

intention of synonymizing this name with that of sect. Baccharis. Espinar
(1973) disagreed with this lectotypification and suggested that B. santiagensis
Heering was a better choice. As defined by Espinar, the two sections appar-
ently are not particularly closely related.

2. Sect. Sergilae DC., Prodr. 5:424. 1836. Lectotype species (Cuatrecasas
1967): Baccharis scoparia (L.) Pers.

Sect. Aphyllae Baker, Fl. Bras. 6(3):45. 1884. Type species: Baccharis
aphylla DC.

Glabrous shrubs; leaves linear to linear oblanceolate, punctate, not papil-
late, glutinous, uninerved, entire; heads mostly solitary, sometimes in few
headed, terminal glomerules; receptacles epaleate; achenes glabrous, 1.0-1.8
mm long, with 8-11 thin nerves; pappus bristles in 2(-3) series. Southwestern
United States and adjacent México, South America.

Species included: Baccharis sarothroides A. Gray, B. sergiloides A. Gray,
and B. vanessae Beauchamp.

Additional representative species of South America: Baccharis genistifolia
DC., and B. notosergila Griseb.

These species are remarkable in their broomlike habit with narrow leaves
and essentially solitary heads, but they appear to be closely related to sect.
Baccharts on the basis of other features. I originally thought that sect. Baccha-
ris would be paraphyletic without the inclusion of at least the North American
sergiloid taxa but the resemblance between the North and South American
species with this morphology is so close that they should be considered mono-
phyletic, at least as a null hypothesis. The chemical data of Zdero, et al. (1986)
also suggest that they are closely related.

3. Sect. Aristidentes Nesom, sect. nov. Type species: Baccharis multiflora
Kunth, Nov. Gen. & Sp. Pl. 4 [folio|:46. 1818; 4 [quarto]:59. 1820.
TYPE: MEXICO. (Edo. México]: Tianguillo, (Sep-Oct, 1803], Humboldt
& Bonpland |4372| (P fiche!). Kunth also cited a collection from near
Toluca but only the plant from Tianguillo is represented on the fiche
showing the P specimens.

Capitulis pedicellatis in capitulescentiis corymboideis et foliis
papillati punctatis serraturis brevi-aristatis diagnoscenda.

Glabrous to puberulent shrubs or small trees; leaves oval to linear lanceo-
late, uni- or trinerved, often serrate with numerous, shallow teeth with aris-
tate apices, punctate, each punctation usually with a minute but definitely
extruded papilla, glutinous or not; heads pedicellate, in a corymboid capit-
ulescence; phyllaries fringed ciliate; receptacles epaleate; achenes glabrous,

Nesom: North and Central American Baccharis 43

(1-)2-3 mm long, with 5-6(-8) relatively thick ribs; pappus bristles in (1-)2-3
series. Southwestern United States. México. and Central America. Baccharis
plummerae A. Gray is known only from California.

Species included: Baccharis bigelovw A. Gray, B. havardu A. Gray, B. mez-
icana J. Cuatrecasas, B. multiflora Kunth, B. palmer Greenm., B. potosina A.
Gray, B. plummerae, B. serraefolia DC., B. sordescens DC., B. sulcata DC.,
B. thestordes Kunth, and B. zamorensis Rzedowski.

These taxa are somewhat similar to those of sect. Molinae but different
in their leaves with papillate punctations and aristate serrate margins, their
tendency to produce a vestiture of puberulent trichomes, and their pappus
bristles mostly in 2-3 series. A study of the Baccharis theszordes-bigelovu-
sulcata complex is in progress (Nesom, in prep.).

4. Sect. Glandulocarpae Nesom, sect. nov. Type species: Baccharis wrightu
A. Gray,. Pl... Wright., 1:101., 1852.. TYPE:. UNITED. STATES. Texas:
Jeff Davis Co., valley of the Limpia, Aug [1849], C. Wright s.n. (HOLO-
LYPE:.GH).

Capitulis pedicellatis in capitulescentiis corymboideis et acheniis

grandibus trichomatibus papillati-glandulosis imprimis diagnoscen-
da.

Glabrous to minutely hispidulous shrubs or small trees; leaves mostly lin-
ear lanceolate, uninerved, entire or with a few shallow, blunt, subapical teeth,
punctate, not papillate, usually glutinous; heads solitary or in racemoid panic-
ulate to corymboid capitulescences; receptacles epaleate; achenes (1.5-)2.0-4.5
mm long, with 5-6(-10) relatively thick ribs, sparsely to densely invested with
thick, viscid, multicellular, often recurved or slightly coiled trichomes; pappus
bristles in 3-5 series. Apparently restricted to México and the United States.

Species included: Baccharis brachyphylla A. Gray, B. erosoricola Rze-
dowski, B. macrocephala Schultz-Bip. er Greenm., B. occidentalis S.F. Blake,
B. pteronioides DC., B. pyramidata (B.L. Robins. & Greenm.) Rzedowski, B.
ramiflora A. Gray, B. squarrosa Kunth, B. terana A. Gray, and B. wrightu A.
Gray.

This group of species is highly diverse vegetatively. Baccharis occiden-
talis and B. squarrosa produce stems with solitary heads and greatly reduced
leaves. Other species produce solitary heads on densely leafy stems or heads
in racemoid paniculate capitulescences. The large and distinctly pubescent
achenes are found in all of the species. Baccharis pyramidata is tentatively
included here, largely on the basis of its similarity in habit and capitulescence
to B. pteronioides, but its stiffly strigose achenes and peculiar ericoid leaf
morphology are anomalous among the Mexican species. The species of the
South American sect. Discolores DC. are similar in leaf morphology but have
a different capitulescence as well as details of the phyllaries and achenes.

44 PA YEO 10.G..A volume 69(1):40-46 July 1990

5. Sect. Baccharzdastrum (Cabrera) Nesom, Phytologia 65:170. 1988. BA-
SIONYM: Baccharidastrum Cabrera, Not. Mus. La Plata Bot. 2:175.

1937. Type species: Conyza triplinervia Less. (= Baccharis vulnearza

Baker).

Glabrous, perennial herbs; leaves nearly linear to broadly lanceolate, strongly
trinerved, shallowly serrate with numerous, nonaristate teeth, punctate, usu-
ally glutinous; heads pedicellate, in a tightly compact, corymboid capitules-
cence; receptacles epaleate: achenes densely and minutely hispidulous, 0.7-1.0
mm long, with 4(-6) thin ribs; pappus bristles in a single series. South Amer-
ica, with a single species in California and Baja California.

Species included: Baccharis douglasu DC.

Additional species of South America: Baccharis breviseta DC., B. ptngraea
DC., and B. vulnearna Baker.

Section Baccharidastrum previously included one dioecious species ( Bac-
charts pingraea) and two monoecious ones (Nesom 1988). Baccharis douglasi,
which is here added to the section, also is dioecious. I[t is so similar to some
forms of B. pingraea that the two must be considered extremely closely re-
lated if not conspecific. Baccharis pingraea, however, is highly variable and a
more detailed study is needed before an understanding of the overall pattern of
variation can be reached. The plants of these species are recognized by their
trinerved. closely serrulate leaves. small, minutely hispidulous achenes, and
uniseriate pappus. These species were included in sect. Molinae by Espinar
(1973), but because of their extremely distinctive achenial vestiture, I think
they are best regarded as a separate group.

6. Sect. Molinae (Ruiz & Pavon) Pers., Syn. Pl. 2:424. 1807. BASIONYM:
Molina Ruiz & Pavon, Prodr. 111, t. 24. 1794. Type species: Baccharis
latufolia (Ruiz & Pavon) Pers.

Sect. Trinervatae DC., Prodr. 5:399. 1836. Type species: Baccharis
trinervis Pers.

Sect. Corymbosae Heering in Reiche, Fl. Chile 4:5. 1903. Lectotype
species (Cuatrecasas 1967), Baccharis marginalis DC.

Shrubs, small trees, sometimes sprawling or subscandent, glabrous or less
commonly puberulent; leaves linear lanceolate to broadly ovate lanceolate or
elliptic, trinerved, shallowly serrate with numerous, nonaristate teeth, less
commonly entire, punctate, papillate, usually glutinous; heads pedicellate, in
a corymboid capitulescence; phyllaries ovate and usually distinctly yellow-
ish; pistillate receptacles epaleate or paleate; achenes glabrous, less commonly
sparsely strigose, 1.0-1.8 mm long, with 4-6(-8) thin nerves; pappus bristles
in a single series. México and Central America to South America; Baccharis

Nesom: North and Central American Baccharis 45

salicifolia, the most widespread species in the genus, extends from the south-
western United States to the southern tip of South America.

Species included (epaleate): Baccharis monoica Nesom, B. salicifolia (Ruiz
& Pavon) Pers. (= B. glutinosa Pers.); (paleate): B. pedunculata (Mill.) Cabr-
era, and B. trinervis Pers. (including B. rhezzordes Kunth).

Additional representative species of South America (epaleate): Baccha-
ris prunifolia Kunth; (paleate): B. brachylaenoides DC., and B. cotinifoha
(Willd.) Urban (distinct from B. pedunculata, in contrast to the view of Cua-
trecasas {1968]).

These species with paleate and epaleate receptacles have not previously
been regarded as closely related, but I can find no other differences among
what is otherwise a group of species with an easily recognizable set of mor-
phological similarities. The species with paleate receptacles were included by
Cabrera (1955) in a broad and highly heterogeneous genus Pszla, united only
by the paleate receptacles of the pistillate heads. More recently, Cuatrecasas
(1982) considered these all to be species of Baccharvs, formally dividing them
into three different sections: sect. Trinervatae, sect. Psila (Phil.) J. Cuatre-
casas, and sect. Pseudobaccharis (Cabrera) J. Cuatrecasas. A situation anal-
ogous to that in sect. Molinae is found in the “paleate” species of the genus
Heterothalamus and a number of the “epaleate” species included in sect. Pseu-
dobaccharis, where no other difference in morphology can be found to separate
them.

ACKNOWLEDGMENTS

I thank Drs. Billie Turner and Tom Wendt for their review and comments
on the manuscript.

LITERATURE CITED

Barroso, G. 1976. Compositaei-Subtribo Baccharidinae Hoffman Estudo das
especies ocorrentes do Brasil. Revista Rodriguesia 40:7-273.

Boldt, P.E. 1989. Baccharis, (Asteraceae), a review of its taxonomy, phyto-
chemistry, ecology, economic status, natural enemies and the potential
for its biological control in the United States. Texas Agric. Exp. Station,
College Station, Texas.

Cabrera, A.L. 1955. La identidad del género Psila Philippi. Bol. Soc. Ar-
gentina Bot. 5:209-211.

46 PHY BODO Gara volume 69(1):40-46 July 1990

Cuatrecasas, J. 1967. Revision de las especies Colombianas del género Bac-

charts. Rev. Acad. Colomb. Cienc. 13(49):5-102.

. 1968. Notas adicionales, taxonomicas y corologicas, sobre Baccha-

ris. Rev. Acad. Colomb. Cienc. 13(50):201-226.

. 1969. Prima Flora Colombiana. 3. Compositae - Astereae. Webbia
24:1-335.

. 1982. Miscellaneous notes on neotropical flora, XV. New taxa in
the Astereae. Phytologia 52:166-177.

Espinar, L. 1973. Las especies de Baccharis (Compositae) de Argentina
central. Bol. Acad. Nac. Cienc. Argentina 50:175-305.

Hellwig, F. 1989. Proposal to conserve 8933 Baccharis L. (Asteraceae) with
a conserved type. Taxon 38:513-515.

Matuda, E. 1957. El género Baccharis en México. An. Inst. Biol. Mex.
28:143-174.

Nesom, G.L. 1988. Baccharis sect. Baccharidastrum (Compositae: Aster-
eae), including two monoecious and one dioecious species. Phytologia

65:169-173.

Zdero, C., F. Bohlmann, R. King, & H. Robinson. 1986. Diterpene glycosides
and other constituents from Argentinian Baccharis species. Phytochem-
istry 25:2841-2855.

Phytologia (July 1990) 68(6):47-50.

CAREX FISSA, SECTION MULTIFLORAE (CYPERACEAE), NEW TO
TEXAS

Stanley D. Jones!, Gretchen D. Jones? & J.K. Wipff'

1$.M. Tracy Herbarium, Department of Range Science, Texas A&M
University, College Station, Texas 77802 U.S.A.

*Department of Biology, Texas A&M University, College Station, Texas
¥TS02°U. SrA.

ABSTRACT

Carez fissa Mackenzie, Section Multiflorae (Cyperaceae), previously
unreported in Texas, which resembles C. annectens (Bickn.) Bickn.
has been found in several Texas counties: Austin, Colorado, Franklin,”
Hopkins, Galveston, and Raines.

KEY WORDS: Carez fissa, Carez section Multiflorae, Cyperaceae,
Texas.

Carez section Multiflorae Kunth is represented by thirteen species in the
temperate and warmer parts of North America; by a number of species in
eastern Asia, and by two in South America (Mackenzie 1931). Four species of
section Multiflorae are now found in Texas: Carez annectens (Bickn.) Bickn.,
C. fissa Mackenzie, C. triangularis Boeck., and C. vulpinozdea Michaux. Carez
annectens and C. triangularis were treated by Correll & Johnston (1970) under
C. vulpinoidea. Correll & Johnston (1970), Correll & Correll (1972), Gould
(1975), Mahler (1988), nor Johnston (1988) have listed Carez fissa as occurring
in Texas.

Carez fissa, a southeastern species, was first described from Sapulpa, in
eastern Oklahoma (Mackenzie 1931) where it was believed to be an endemic.
Since Hermann (1965) described Carez fissa var. aristata Hermann from north
central Florida, Carez fissa has been found throughout the Gulf States. The
habitat in Texas is similar to what Godfrey & Wooten (1979) described for
Carez fissa var. aristata in north and north central Florida; wet woodlands,
pine flatwoods, wet clearings, and ditches. This treatment does not recognize
varieties. However, if the reader is interested, Hermann (1965) provides a key
to separate varieties as seen by him. The following key will help differentiate
the closely related taxa of section Multiflorae found in Texas.

47

48 Pa YoD OD O.G 5 A volume 69(1):47-50 July 1990

KEY TO TEXAS SECTION MULTIFLORAE

1. Most leaves equal to or exceeding the culms; perigynia 1.0-1.8 mm broad,
narrowly ovate; beak of the perigynium tapering from the body, beak
1/2 as long as the body to as long as body ............. C. vulpinoidea

1’ Most leaves shorter than the culms; perigynia 1.6-3.0 mm broad, narrowly
ovate to broadly ovate, orbicular to reniform; beak of the perigynium
tapering or arising abruptly from the body, beak to 1/2 the length of
the bodys e260 10.2. Be a eee 2

2. Perigynia very conspicuous in the infructescence; perigynia 2.4-3.0
mm broad, orbicular to reniform, often broader than long, with
red glandular dots; beak of perigynium arising abruptly from the
body; perigynial scales not conspicuous; apex of membranous in-
ner band of the upper leaves somewhat thickened, prolonged and
usually sharply rounded 2... c00.9. A Sees C. triangularis

2’ Perigynia conspicuous or not in the infructescence; perigynia 1.6-
2.7 mm broad, narrowly ovate to ovate orbicular, rarely broader
than long, with or without red glandular dots; beak of perigynium
tapering or arising abruptly from the body; perigynial scales usually
conspicuous; apex of membranous inner band of the upper leaves
somewhat thickened or not, prolonged and slightly
rounded. 2)... j2sicgaww'satve dis cae oeee't ee 3

3. Base of culms 2.5-3.0(-4.5) mm thick; rhizomes not elongate;
perigynia 1.6-2.4 mm broad, occasionally with red glandular
dots apically; beak of perigynium usually arising abruptly from
body, (infrequently tapering); apex of the membranous inner
band of the upper leaves somewhat thickened ...C. annectens

3’ Base of culms (3.0-)3.5-6.0 mm thick; rhizomes elongate; perig-
ynia 2.0-2.7 mm broad, never with red glandular dots; beak of
perigynium usually tapering from body, (infrequently arising
abruptly); apex of the membranous inner band of the upper
leaves friable, not thickened .....7. .:.teeemeneeereraee C. fissa

Specimens collected: UNITED STATES. Texas: Franklin Co., 9.2 miles
(14.7 kilometers) N on Hwy 37 from its jct. with Hwy 196, N of Mt. Vernon,
15 May 1989, S. & G. Jones 2852 & Tim Powell (ASTC, MICH, TAES). It was
frequent in open wet areas adjacent to White Oak Creek. Associated species:
Carez amphibola Steud., C. lupulina Muhl., C. crus-corvi Kunze, C. arkansana
Bailey, C. annectens, C. typhina Michaux, C. bushtz Mackenzie, C. reniformis
(Bailey) Small, Iris brevicaulis Rafinesque, Danthonia spicata (L.) Beauv.,
Lolium perenne L., Stipa leucotricha Trin. & Rupr., Asclepias amplezicaulis

Jones, et al.: Carer fissa, new to Texas 49

Small, and A. viridis Walt. Galveston Co., Pelican Island, 4.2 miles (6.7
kilometers) E on main road through Pelican Island from its jct. with Port Ind.
Blvd., 08 May 1989, S. Jones 2718 & J. Wipff (ASTC, MICH, SMU, SRSU,
SWT, TAES, WARM). It was occasional in an open hydric roadside ditch.
Associated species: Carex triangularis, C. brittonzana Bailey, C. cherokeensis
Schwein., C. crus-corvi, C. leavenworthii Dew., C. amphibola, Eleocharis albida
Torrey, Scirpus maritimus L., Asclepias perennis Walt., Spartina spartinae
(Trin.) Hitchc., Setaria geniculata (Lam.) Beauv., and Tamarzz sp. Raines
Co., South side of Lake Fork Reservoir at the lake’s crossing of FR 2946,
13 May 1989, S. & G. Jones 2889 (ASTC, MICH, TAES). It was frequent
around the mesic to hydric lakeside in red clayey soil. Associated species
Carer crus-corvi, C. hyalinolepis Steud., Carer flaccosperma Dew., Scirpus
pendulus, Eleocharis sp., and Juncus spp.

Additional specimens: UNITED STATES. Texas: Austin Co., 8 April 1939,
B. Tharp 43279 (TEX). Colorado Co., 03 May 1948, E. Whitehouse 19871
(SMU). Gregg Co., 20 April, 1988, E. Nizon 16568 (ASTC, TAES). Hopkins
Co., 08 June 1953, L. Shinners 15087 (SMU); 05 June 1989, A.A. Reznicek
8484 & R.F.C. Naczi (MICH, TAES).

A careful examination of Carer section Multiflorae specimens in other
herbaria should yield additional Texas specimens.

ACKNOWLEDGMENTS

We thank David Castaner (WARM) for verification of SMU material. We
thank Tony Reznicek (MICH) for verification and determination of the senior
author’s collections. We also thank David Castaner and Tony Reznicek for
helpful suggestions concerning this manuscript.

LITERATURE CITED

Correll, D.S. & H.B. Correll. 1972. Aquatic and Wetland Plants of South-
western United States. Stock #5501-0177. Environmental Protection
Agency Research and Monitoring, Washington, D.C.

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

Godfrey, R.K. & J.W. Wooten. 1979. Aquatic and Wetland Plants of South-
eastern United States. Univ. of Georgia Press, Athens, GA.

50 PHY TOLOGIA volume 69(1):47-50 July 1990

Gould, F.W. 1975. Tezas Plants - a checklist and ecological summary. Texas
Agric. Exp. Sta. Bull. MP-585.

Hermann, F.J. 1965. An eastern var. of Carez fissa (Multiflorae). Rhodora
67:198.

Johnston, M.C. 1988. The Vascular Plants of Tezas. A list up-dating the
manual of the vascular plants of Texas. Marshall C. Johnston, Austin,

aXe

Mackenzie, K.K. 1931. North American Flora (Poales) (Cyperaceae). 18:64.
The New York Botanical Garden, N.Y.

Mahler, W.F. 1988. Shinners’ Manual of the North Central Tezas Flora.
S.M.U. Herbarium, Dallas, TX.

A

Phytologia (July 1990) 68(6):51-55.

TAXONOMIC OVERVIEW OF THE SENECIO FLACCIDUS COMPLEX IN
NORTH AMERICA, INCLUDING S. DOUGLASII

B.L. Turner! & T.M. Barkley?
‘Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

*Division of Biology, Kansas State University, Manhattan, Kansas 66506
U.S.A.

ABSTRACT

The earliest specific name for the widespread North American taxon
long known as Senecio douglasi: DC. is shown to be S. flaccidus Less.
This has occasioned two new infraspecific combinations in the complex,
including: Senecio flaccidus Less. var. douglasii (DC.) B. Turner &
T. Barkley, comb. nov., and S. flaccidus Less. var. monoensis (E.
Greene) B. Turner & T. Barkley, comb. nov. A key to the taxa and a
map showing their distribution in Mexico is provided.

KEY WORDS: Senecio, Asteraceae, Senecioneae, taxonomy, nomen-
clature, North America.

Preparation of a treatment of Senecio (sensu lato) for México (Barkley
& Turner, in prep.) has revealed that 5. douglasa DC., long recognized as
the name for a common, widespread, highly variable, perennial subshrub of
the western United States (Barkley 1978) and México, must be replaced by
the earlier name, S. flaccidus Less., which is typified by material originally
collected along the semi-arid western slopes of Cofre de Perote, Veracruz,
Mexico (near the village of Perote).

Material from the type locality and throughout most of northern México is
essentially indistinguishable from what Barkley (1978) has treated as Senecio
douglasz var. longilobus (Benth.) Benson. The latter taxon is known to inter-
grade with S. douglasz var. douglasii in northern México and adjacent Arizona,
and both of these appear to intergrade with S. douglasii var. monoensis (E.
Greene) Jepson over at least part of their distribution. In order to provide
correct names for members of the “S. douglasiv’ complex, we have provided
here an overview of the S. flaccidus complex as it appears in México. Except
where noted, types for most of the names referred to in the present paper are

51

52 Pay T'OOLKOOG VA volume 69(1):51-55 July 1990

accounted for by Barkley (1978). The distributional map (Figure 1) is based
upon a wide range of material available on loan to the junior author and a
large assortment of collections (LL, TEX) available to the senior author.

Senecio flaccidus Less., Linnaea 5:161. 1830.

Perennial subshrubs or rarely appearing annual, 3-12 dm high, persistently
tomentose to variously glabrate; stems several, mostly branched in the upper
third, arching upward from a taprooted, woody base; leaves about equally
distributed along the stem, linear to narrowly filiform, or deeply pinnatifid
into long, narrow segments, 2-10+ cm long overall, sometimes with fascicles
of smaller leaves in the axils of the principal leaves; inflorescence a series of
corymbiform or subcorymbiform cymes, each with 3-10(-20+) cylindrical or
campanulate heads; involucral bracts ca. 21 or 13, 5-8+ mm long, greenish
or stramineous at the tip, not black tipped; calyculate bracts prominent and
up to 1/2 as long as the principal bracts, or reduced and inconspicuous, or
sometimes absent; ray florets ca. 13 or 8, the ligule 10-15(-20) mm long, yel-
low or sometimes light yellow to ochroleucous; achenes canescent hirtellous;
chromosome number, n = 20 pairs.

A widespread, common and complicated entity that has long gone under
the name Senecio douglasz in the U.S.A. It is well adapted to sites with con-
tinual, mild disturbance. Three regional intergrading varieties are recognized,
but subsequent studies may alter this concept of the species.

Key to Senecio flaccidus Varieties in México

1. Heads subcylindrical when young, principal involucral bracts ca. 13 or 21,
5-8+ mm long, calyculate bracts absent or inconspicuous; herbage to-
mentose with long, lanate hairs, sometimes unevenly glabrate; northern
México, south to Puebla and Veracruz ................-- var. flaccidus

1’ Heads large and campanulate, principal involucral bracts ca. 21, 7-10+ mm
long, calyculate bracts usually prominent and up to 1/2 as long as the
Principal DrActs b.. .és 2:20 are o,0.0 » 4 2,016 6\n'yb\o.0 ei eee 2

2. Herbage glabrous or nearly so at maturity; Baja California,
Sonora’ Sa. ee. Pe ee var. monoensis
2’ Herbage tomentose with persistent short, grayish hairs, sometimes

unevenly tomentose; northern Sonora .............. var. douglas

Senecio flaccidus Less. var. douglasii (DC.) B. Turner & T. Barkley, comb.
nov. BASIONYM: Senecio douglasii DC., Prodr. 6:429. 1837.

53

Overview of Senecio flaccidus complex

Turner & Barkley:

‘OOIXAJA| Ul SNPIOORIJ OLNaUaS JO UOTINGNSIq “| “Sty

SISUDOUOW “IRA @
SNploor[y “IBA O

useysnop “wAV
snplooryj
OIOANAS

54 Pay TD GUL OG TLA volume 69(1):51-55 July 1990

Senecio douglasu DC. var. tularensis Munz, El] Aliso 4:99. 1958.

Known in México from only a few questionable collections in northern
Sonora (extending into México from a much wider distribution in the U.S.A.),
open sandy or rocky sites in desert hills, mostly below 1500 m, July-October.

Senecio flaccidus Less. var. flaccidus TYPE: MEXICO. Veracruz: “In llanos
de Perote,” Sep 1819, Schiede & Deppe s.n. (HOLOTYPE: B?, not

examined; drawing in GH!).

Senecio douglastt DC. var. jamesii (Torrey & A. Gray) Ediger ex Correll
& Johnston, Man. Vasc. Pl. Tez. 1712. 1970.

Senecio douglasii DC. var. longilobus (Benth.) Benson, Amer. J. Bot.
30:631. 1943. BASIONYM: Senecio longilobus Benth., Pl. Hartw.
18. 1839.

Senecio orthophyllus E. Greene, Leafl. Bot. Observ. Crit. 1:221. 1906.
Senecio filifolius Nutt., Trans. Amer. Phil. Soc. II. 7:414. 1841.
Senecio reguomontanus DC., Prodr. 6:429. 1838.

Scattered through the Central Plateau of northern México southwards to
Puebla and adjacent Veracruz (Figure 1), occurring mostly in open sandy or
rocky floodplains, creek beds, roadsides, and similar, mildly disturbed places
in open desert regions, chiefly below 2000 m, also adjacent U.S.A.; flowering
the year around.

The typical var. flaccidus passes into the other two varieties in the U.S.A.,
where their distributions overlap. Occasional specimens of both var. monoen-
sis and var. douglasw from México approach var. flaccidus, although neither
of the former are clearly sympatric with the latter in México. Var. flacczdus
apparently intergrades to some extent with Senecio stoechadiformis in México
along the eastern edges of the Sierra Madre Occidental. Additional study may
indicate that var. flaccidus, as conceived here, incorporates two or more mor-
phological phases, based on head size, distribution and duration of pubescence,
and gross aspect. Whether or not these phases warrant taxonomic recognition
is yet to be determined.

The type specimen of Senecio flaccidus was apparently at Berlin and it has
not been examined. However, a pen and ink tracing of the type is among the
Klatt materials at GH. This drawing, plus the excellent original description and
comparison with more recent collections, leaves no doubt as to the application
of the name.

Senecio flaccidus Less. var. monoensis (E. Greene) B. Turner & T. Barkley,
comb. nov. BASIONYM: Senecio monoensis E. Greene, Leafl. Bot. Ob-
serv. Crit. 1:221. 1906. Senecio douglasiti DC. var. monoensis (E.
Greene) Jepson, Man. Fl. Pl. Calif. 1149. 1925.

Turner & Barkley: Overview of Senecio flaccidus complex 55

Senecio filicifoltus Greenm., Ann. Missouri Bot. Gard. 1:774. 1914.
Senecio lathyroides E. Greene, Leafl. Bot. Observ. Crit. 2:21. 1909.

This variety occurs in northwestern Mexico (Figure 1) and adjacent U.S.A..,
where it is frequent in open, rocky or sandy sites, especially on alluvial fans
and floodplains in the low desert, in México mostly below 800 m; flowering all
seasons.

ACKNOWLEDGMENTS

We are grateful to Guy Nesom and Andrew McDonald for reviewing the
manuscript.

LITERATURE CITED

Barkley, T.M. 1978. Senecio, in N. Amer. Fl., Ser. II. 10:50-139.

Phytologia (July 1990) 69(1):56.

PUBLICATION DATES FOR VOLUME 68

68(1) mailed 2 February 1990
68(2) mailed 6 March 1990
68(3) mailed 3 April 1990
68(4) mailed 26 April 1990
68(5) mailed 6 June 1990
68(6) mailed 2 July 1990

56

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