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CONTENTS

eraalacess) SORA SS is EUAN pe PMB ange MEE OE YP PRM Ob ta ge be A 405
~ HURTER, P.J.H., Two newly described cycads from Affica...............4..
s ~-REEDER, C.G. & LR. REEDER, The resurrection of a species; Muhlenbergia
Saas PMMMETILE ME BEMININOCAR Ye Oooh Sn Rane Sleek Cannas Need raee se avec seen ata ees 417
% -ENQUIST, M. & B. CROZIER, Anemone tuberosa (Ranunculaceae) in Texas. .
MD ec Okn Ci VAG Lich. iwc G wehbe ceo k caee pa be ay aca en ee he Ns the Ghee deci ey wtebee ss 428
TURNER B.L., Paronychia virginica (Caryophyllaceae), a first report of its
Manmey emeMar nIOe 1) ICKICO 08 ss i ge hes ane Bye eqn tet end covet oa 446
a TURNER, B.L., Synoptical study of Rhododon (Lamiaceae). ............-..+. 448
et _ Index to authors, VOMENC Jas xe ce iRoeiele cea ie eek ape era Uk ape ee 452
(EE GOT LAN | oh Bente S ea Ea ye Exot gt Sean Soe Rane pew ae 453
a Mua IER S AV OSTAD NE 6 585 al dope vba fac ose kr sods dk Coates ousenst aaa 510
as Soar MewOIIeWOre. NONUNE 9S. erie. ced eh ead GON gia aod boas 51}

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Phytologia (June 1995) 78(6):405-408

A NEW SPECIES AND NEW COMBINATION IN MEXICAN SEDUM
(CRASSULACEAE)

B.L. Turner

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

ABSTRACT

A new species of Sedum, S. hintoniorum B.L. Turner, is described
from southern Nuevo Leon and closely adjacent Tamaulipas, and S$.
rhodocarpum subsp. edwardsii R.T. Clausen is elevated to specific rank as S .
edwardsii (R.T. Clausen) B.L. Turner, comb nov. An account of their
relationships to closely related taxa is given, and a map showing their
distributions is presented.

KEY WORDS: Crassulaceae, Sedum, México, Nuevo Leén, Tamaulipas

Routine identification of plants from northeastern México has revealed the
following novelty.

SEDUM HINTONIORUM B.L. Turner, spec. nov. TYPE: MEXICO. Nuevo
Le6n: Mpio. Zaragoza, Cerro Viejo, 2085 m, 1 Oct 1993, Hinton et al. 23797
(HOLOTYPE: TEX!; Isotypes: GH!,MEXU!).

Sedo caduco R.T. Clausen similis sed differt caulibus non sigillatim
papillosis, foliis viridibus (vs. rubescentibus), petalis lanceolatis (vs. cucullatis
ad apicem), et antheris purpurascentibus (vs. flavidis).

Ascending to recumbent perennial herbs 15-20 cm high. Roots forming fusiform
tubers 1-2 cm long, 0.2-0.5 cm across. Lower stems suffruticose, glabrous, the
epidermal cells elongate but bulging; upper stems and branches of inflorescence similar
to the lower but upon drying seemingly winged, the uppermost portions appeanng
subpapillose. Leaves alternate, oblanceolate, those at midstem ca. 2 cm long, 0.4 cm
wide, glabrous, smooth. Flowers arranged in terminal, relatively open, paniculate
cymes 3-5 cm across and about as wide, or else arranged in leafy 1-sided flowering
branches. Sepals 5, unequal, free, ovate-lanceolate to oblanceolate, 2-4 mm long.
Petals 5(-4), 46 mm long, 1-2 mm wide, broadly lanceolate, the upper portion often
with a green or rosy mid-rib, the apices decidedly acute, reflexing at maturity.

405

406 PAY fOMO GVA June 1995 _ volume 78(6):405-408

Nectaries much-reduced, reniform, white. Stamens ca. 5 mm long, the anthers purple
or purplish. Pistils 4-S mm long, erect to somewhat spreading at maturity, the beak
ca. 1.5mm long. Seeds ovoid, ca. 0.7 mm long, brown, minutely papillose.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Nuevo Leé6n: = Mpio.
Arambern, Cerro Viejo, 2485 m, 20 Nov 1993, Hinton et al. 23981 (TEX). Mpio.
Galeana, Agua Blanca to San Miguel, 1985 m, 28 Aug 1991, Hinton et al. 21297
(TEX). Mpio. Zaragoza, Cerro Viejo, 1935 m, 6 Oct 1992, Hinton et al. 22455
(TEX). Tamaulipas: Mpio. Hidalgo, Arroyo Oscuro, 2 rd. mi NE of Paraje Los
Caballos, 1800 m, 23 Sep 1994, Nesom 7500 (TEX).

Sedum hintoniorum occurs in pine-oak woodlands from 1800-2100 m where,
according to label data, it forms colonies on bare boulders or exposed rocky places.
The specimen from Tamaulipas (growing in humid woods with Carya spp., Cornus,
Carpinus, and Dirca) differs somewhat in possessing smaller flowers (these obviously
immature).

In Clausen's 1984 account of Sedum for the Mexican Cordillera, flowering
specimens of §. hintoniorum will key to or near S$. caducum R.T. Clausen, a white-
flowered species known only from type material collected in Tamaulipas, México, near
Ciudad Victoria at 610 m. His description of the habit (stiffly erect), stems
(papillose), leaves (“prominently speckled with red and appearing reddish”),
inflorescence (2-3 parted cymes with 2-7 flowers), petals hooded at apex, basal fusion
of petals (0.8 mm), anther color (“pale yellow to pale carmine”), differs markedly
from the species descnbed here.

It is a pleasure to name this distinctive species for the remarkable G.B. Hinton
(and his extended family), the latter having assembled most of the specimens cited
herein. I am well aware that there already exists a Sedum with the name S$. hintonii
R.T. Clausen, named for the late G.B. Hinton (1882-1943), fount of this clan and
father of Jaime and grandfather of George (collectors of the present taxon). But such
an extraordinary lineage, in my opinion, is fully deserving of the apellations
concerned, as well documented in Hinton & Hinton (1995) showing their collecting
activities in northeastern México.

SEDUM EDWARDSII (R.T. Clausen) B.L. Turner, comb. & stat nov.
BASIONYM: Sedum rhodocarpum Rose subsp. edwardsii R.T. Clausen, Sedum
Mexic. Cordill., Plat. 7. 1981.

As noted by Clausen, who treated the taxon as a subspecies of Sedum
rhodocarpum, S. edwardsii is closely related to S. rhodocarpum but differs in
numerous characters, including leaf shape and vestiture; the latter character is
especially diagnostic, all of the collections listed below possessing pilose stems and
foliage, while S. rhodocarpum is completely glabrous.

The following collections of Sedum edwardsii have come to the fore since
Clausen's description of the taxon. MEXICO. Tamaulipas: Mpio. Hidalgo, Los
Caballos, 1705 m, 21 Sep 1994, Hinton et al. 25143 (TEX); El Mirador, 895 m,
Hinton et al. 25147, 11.4 mi E of Dulces Nombres, 0.3 road mi W of Paraje de Los
Caballos, 1840 m, 21 Sep 1994, Nesom 7456 (TEX).

Turner: New species of Sedum 407

NUEVO LEON

101°20'

98°|5'

Figure 1 Distribution of Sedum spp: S. caducum (solid triangle); S$. hintoniorum
(open triangles); 8. edwardsii (open circles); S. rhodocarpum (closed circles).

408 PREY DOL OIGiyA June 1995 — volume 78(6):405-408

None of these collections contained material referable to Sedum rhodocarpum,
however, Hintons’ collection 25/47 (from 857 m) differs from the other specimens
listed in reportedly having green petals. Only two plants of the latter were observed at
the locality concerned, according to the collectors.

ACKNOWLEDGMENTS

I am grateful to Guy Nesom for the Latin diagnosis, and to him and Mark Mayfield
for reviewing the manuscript.

LITERATURE CITED

Clausen, R.T. 1984. Sedum (Crassulaceae) of the Mexican Cordilleran Plateau.
Gentes Herb. 12:8-48.

Hinton, J. & G.S. Hinton. 1995. Checklist of Hinton’s collections of the flora of
south-central Nuevo Leon and adjacent Coahuila. Acta Bot. Mex. 30:41-112.

Phytologia (June 1995) 78(6):409-416

TWO NEWLY DESCRIBED CYCADS FROM AFRICA

P. J. H. Hurter

Lowveld National Botanical Garden, P. O. Box 1024, NELSPRUIT, 1200, Republic
of South Africa.

ABSTRACT

Recent intensive field study of Encephalartos in Uganda and South Africa
reveal two overlooked species. The nature of these entities, their distribution,
and conservation status is discussed.

KEY WORDS: Encephalartos, new species, Zamiaceae

Encephalartos nubimontanus P.J.H. Hurter, spec. nov. TYPE: SOUTH
AFRICA. Northern Province: 1,000 m alt., (leaf and male cone). 27 March
1995, P.J.H. Hurter 95R/1. (HOLOTY PE: PRE). (Figure 1).

E. cupidus R.A. Dyer frondes rigidae, pungentes, glaucas similis sed
plantae arborescentes; frondes recurvae et subsessiles; foliola falcata:
microsporophylla demissa; bullae labiatae.

Plant arborescent, suckering from the base. Trunk often decumbent, up to 2.5 m
long, 350-400 mm diameter. Leaf bases persistent, crown slightly tomentose.
Cataphylls velvety white, becoming sub-glabrous with age. Leaves numerous in a
dense crown, apically recurved, rigid, glaucous, 1.1-2.0 (1.4) m long, pinnae
ascendant. Petiole apparent, slightly trigonous, basally bulbous, 230 mm long,
slightly tomentose with a reddish brown collar at matunty. Rachis glabrous, round in
cross-section, becoming yellow with age. Pinnae dentate or entire, veins not raised
abaxially, margins slightly thickened, inflexed, directed toward the frond apex at an

angle of circa 45° to the rachis, opposing leaflets inflexed, set at an angle of circa 70°
to each other, oriented incubously towards each other; proximal leaflets gradually
reduced to a series of spines; median leaflets 180-250 mm long, 15-25 mm wide, outer
lamina edge falcate, narrowly elliptic, gradually acuminate apically, pungent, basally
subsessile to the rachis, apices somewhat turned toward the frond apex. Strobili
seriate, dimorphous, glabrous; scale facets smooth, light green. Microsporangiate
strobili 1-5 per trunk, subconical, 250-400 mm long, 50-90 mm in diameter, stalked
on a peduncle 30-45 mm long. Median microsporophylls slightly descending, lamina

409

410 PHY TROTuOGTA June 1995 volume 78(6):409-416

oblong, tapering to the base, 11-24 mm wide, 20-30 mm long, 5-10 mm high,
margins contracted to the pedicel, bulla with two trapezoidal lateral facets, without
median facet but well defined hump-like sagittal ndge; terminal facet a drooping lip-
like structure, often slightly crenulate. Megasporangiate strobili 1-3 per trunk, ovoid,
360-400 mm long, 180-200 mm in diameter, with peduncle 360-400 mm long, 180-
200 mm in diameter, with peduncle up to 150-310 mm long. Terminal sterile
megasporophyll bullae drawn out into ascending lip-like structures. Median
megasporophylls rhombic, 40-60 mm wide, 50-60 mm long, and 350-360 mm high.
Bulla with two smooth or slightly verrucose trapezoidal lateral facets and a single
smooth or slightly verrucose oblong median facet, terminal facet smooth, slightly
concave, 3-5 mm deep, lateral ridges slightly drawn out, sagittal crest verrucose,
drooping below the pedicel, seminal ridges smooth or slightly sagittate. Seed circa
200 per cone, sarcotesta orange-red, sclerotesta 35-38 mm long, 23-30 mm in
diameter, ellipsoidal, round and smooth.

Plants of Encephalartos nubimontanus superficially resemble those of
Encephalartos cupidus R.A. Dyer (Dyer 1971) by their stiff, pungent, glaucous
fronds. However, FE. nubimontanus can at first glance be distinguished from that
species by its arborescent habit, robust nature, and long recurved leaves.
Vegetatively, E. nubimontanus is easily distinguished from FE. cupidus by its longer,
often nearly sessile fronds. In E. cupidus the median pinnae are oblong lanceolate,
unlike the normally falcate, often marginally dentate pinnae of FE. nubimontanus (an
ecotype that has thus far been observed only in cultivation and not in habitat, produces
a lamina dentate to nearly lobed on both sides and is thus not falcate). Cones are also
distinctive. The female cones of FE. nubimontanus have sterile terminal
megasporophyll bullae that are drawn out into ascending lip like structures, which is
not the case in FE. cupidus. In the male cones of FE. nubimontanus the median
microsporophylls are slightly descending with bullae drawn out into lip-like structures,
while the median microsporophylls of E. cupidus are spreading with bullae not drawn
out.

Distribution and Habitat: At present this species is known only from one small
area in the Northern Province, at an altitude of 1000 m. Plants grow scattered in
mixed deciduous woodland, especially on cliff faces.

Material Studied: To protect plants from poachers, precise localities are not given,
and grid references are restricted to a 1:250,000 scale. 2430 Pilgrim's Rest: P.J.H.
Hurter 95R/] (leaf and male cone, PRE, holotype), 95R/2 (leaf and female cone,
PRE), 95R/3 (leaf), 95R/4 (leaf), 95R/5 (leaf and male cone), 95R/6 (leaf), 95R/7
(leaf and male cone), 95R/8 (leaf and male cone).

Encephalartos whitelockii P.J.H. Hurter, spec. nov. TYPE: UGANDA:
Mpanga River Falls, 1,200 m alt. 27 October 1994, P.J.H. Hurter 94U/3a (leaf
and median section of microstrobilus [HOLOTY PE: PRE]). (Figure 3).

E. hildebrandtii A. Br. & Bouché propter folia rigidi, dentati, pungentia
similis sed frondes planae, leviter recurvatae; foliola effusa nonsuperposita ad
apicem frondes apicem versus; microstrobilis laxus; microsporophylla patentia;
megasporophylla demissa.

Hurter: Two new African cycads 411

Plant arborescent, suckering from the base. Trunk up to 3.5 m long (rarely to 4.2
m), 350-400 mm in diameter. Leaf bases persistent. Leaves numerous in a dense
crown, apically recurved, rigid, subsessile, green, 3.1-3.4(-4.1) m long. Petiole
basally bulbous, to 130 mm long, glabrous. Rachis tomentose, becoming glabrous
with age. Pinnae dentate, veins not raised abaxially, margins slightly revolute,

spreading, opposing leaflets set at an angle of about 160° to each other, not
overlapping, but succubously oriented, proximal leaflets gradually reduced to a distinct
series of spines. Median leaflets 230-300 mm long and 20-28 mm wide, narrowly
elliptc and falcate, gradually acuminate apically, pungent and subsessile basally,
apically somewhat turned towards the frond apex. Strobili seriate, dimorphous,
glabrous with smooth scale facets, light green becoming yellow with age.
Microsporangiate strobili up to 5 per trunk, narrowly ovoid, 500 mm long and 90 mm
in diameter, stalked on a peduncle to 320 mm long. Median microsporophylls
rhombic, about 29 mm wide, 32 mm long, and 14 mm high, distinctly with two lateral
facets and one median facet, the central facet flat or slightly concave. Megasporangiate
strobili 1-3 per trunk, ovoid, 450 mm long, 350 mm in diameter, appearing sessile but
with peduncle to 100 mm long, cloaked by cataphylls in the trunk crown. Median
megasporophylls descending in appearance, rhombic, with two lateral facets and one
central facet, slightly drawn out, seminal fringe slightly verrucose, about 55 mm wide,
40 mm long, and 30 mm high with central facet a third of the horizontal diameter of the
bulla. Seed about 400 per cone, sarcotesta orange-red; sclerotesta 30-35 mm long and
25-30 mm in diameter, ellipsoidal, round and smooth.

Diagnostic features and affinities: | Encephalartos whitelockii superficially
resembles E. hildebrandtii A. Br. & Bouché (Melville 1957, 1958) because of its suff
dentate and pungent green leaves. However, even vegetatively it is easily
distinguished from this species. In Encephalartos hildebrandtii the pinnae are
ascending, succubously overlapping and falcate with the apices directed toward the
base of the leaf. In E. whitelockii the pinnae are spreading, not overlapping, and
falcate with the apices directed toward the apex of the leaf. Profound differences are
also observable between the strobili. Microstrobili of E. whitelockii are pendulous at
maturity while those of FE. hildebrandtii are erect. The microsporophylls of FE.
whitelockii are spreading while those of EF. hildebrandtii are strongly ascending. The
megasporophylls of E. hildebrandtii are spreading and the bullae are not drawn into
descending structures as with those of FE. whitelockii.

Distribution: At present this species is known only from one area along the
Mpanga River, Western Uganda (Figure 4).

Material Studied: UGANDA: Mpanga River falls, 2 km west of Lake George,
P.J.H. Hurter 94U/5, 94U/3a.

This species is named for Loran M. Whitelock, horticulturist and cycadologist,
who has made most of my recent exploration of central and east Afnca possible and
who has devoted a life time to cycads.

PHY TOLOGIA June 1995 — volume 78(6):409-416

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microsporophylls (P.J.H. Hurter 95R/5), (i) side view, (11) abaxial view, (ii) frontal
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view, (iii) frontal view; (d) median pinnae (P.J.H. Hurter 95R/1); (e) seed (P.J.H.
Hurter 95R/2); (f. i-iv) a range of pinnae shapes observed within the new species; (g)
petiole and proximal pinnae (P.J.H. Hurter 95R/1). (del. S. Burrows).

413

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414 PHY TOW OGi4 June 1995 — volume 78(6):409-416

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Figure 3. Encephalartos whitelockii (a) median microsporophyll (P.J.H. Hurter
94U/3a), (i) abaxial view, (ii) frontal view, (iil) side view; (b) median megasporophyll
(P.J.H. Hurter 94U/5) (i) adaxial view, (ii) frontal view, (ill) side view; (c) median
pinna; (d) petiole and proximal pinnae (P.J.H Hurter 94U/3a); (e) median section of
leaf (P.J.H. Hurter 94U/3a).

Hurter: Two new African cycads 415

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416 PE Y TOWwOiG vA: June 1995 volume 78(6):409-416
ACKNOWLEDGMENTS

Research in South Africa and Uganda was supported by the National Botanical
Institute and Loran M. Whitelock. Sandie Burrows is thanked for the art work in
Figures | and 3.

REFERENCES

Dyer, R.A. 1971. A further new species of cycad from the Transvaal. Bothalia
10:370-383.

Melville, R. 1957. Encephalartos in Central Africa. Kew Bull. 12:237-257.

Melville, R. 1958. Gymnospermae. in Flora of Tropical East Africa. pp 3-10.
Crown Agents, London, Great Britain.

Phytologia (June 1995) 78(6):417-427.

THE RESURRECTION OF A SPECIES; MUHLENBERGIA STRAMINEA
(GRAMINEAE)

Charlotte G. Reeder & John R. Reeder

Herbarium (ARIZ), 113 Shantz Building, University of Arizona, Tucson, Arizona
85721 USA

ABSTRACT

Muhlenbergia straminea, described by Hitchcock (1913), was recognized
as a distinct species until 1935 when he relegated it to synonymy under M.
virescens (H.B.K.) Kunth, a disposition that has been followed by all
subsequent authors. The present study, based on morphology, distribution,
and time of flowering presents strong evidence for the recognition of M.
straminea as a species distinct from M. virescens. Although quite unexpected,
we believe it is highly significant that meiosis occurs in the anthers of M.
straminea in the autumn preceding flowering the following spring.

KEY WORDS: Gramineae, Muhlenbergia straminea, M. virescens, M.
curvula, M. quadridentata, México, southwestern U.S.A.

Continuing field and herbarium studies, along with examination of type
specimens, have revealed the need to re-evaluate certain members of the Muhlenbergia
montana complex. This group includes species most easily recognized by the 3-
nerved (often 3-toothed and/or 3-awned) second glume and lower sheaths which
become flat, somewhat papery, and no longer invest the culm. A number of species,
most of which occur in México, make up this taxonomically puzzling assemblage of
both annual and perennial grasses. Of immediate concern is the perennial M.
virescens, a taxon said to range from Central México to Anzona and New Mexico,
U.S.A. The species is based on Podosaemum virescens H.B.K. (1816), which was
described from a plant collected by Humboldt & Bonpland in 1803 in the state of
Guanajuato, México: “Crescit locis asperis, excelsis regni Mexicani prope Santa Rosa
de la Sierra et Puerto de Varientos, alt. 1350 hexap. [ca. 2400 m]. Floret Septembn.”
The type specimen (P!) is fragmentary, consisting of a segment of culm with an
inflorescence partially included in the upper sheath, another short section of culm with
leaf attached, and a separate leaf. Written on the sheet is the locality, “S. Rosa.” The
description is quite ample, and along with the type specimen, gives an adequate
conception of the species. Numerous specimens (including our own) collected in the
mountains of Guanajuato can be determined as this species with little difficulty. The

417

418 PELY T'OLOGiA June 1995 volume 78(6):417-427

transfer to Muhlenbergia was made by Trinius (1824), some five years before Kunth
(1829) as cited by Hitchcock (1913, 1935, 1935a, 1951). Pnor to 1913, Trinius was
given as the authority for this transfer. It is noteworthy that Kunth (1833) recognized
the Trinius combinations.

There has been, and continues to be, confusion regarding the range of
Muhlenbergia virescens. Roemer & Schultes (1817), Hemsley (1885), and Fournier
(1886) repeat the distribution as given in H.B.K. (1816), which suggests that the
species is confined to Central México. Fournier often cited other collections but here
listed only the type number. Hemsley's work, based largely on Fournier, included no
additional information.

Meanwhile Scribner (1882) published a detailed list of grasses collected by C.G.
Pringle in Arizona and California during the summer of 1881. Included (p. 88, 89) is:
“32. Muhlenbergia virescens, Trin., Unifl., 193; Kunth, Enum. Pl. 1, 202, et Suppl.
160; Podosaemum virescens, H B K., Nov. Gen., 1, 132; Trichochloa virescens, R.
& S., Syst., i, 389.” One finds (p. 88, 89) a rather lengthy description of a plant
collected on “Summits of the Santa Rita Mts., Arizona... . This species resembles
M. gracilis Trin., in habit, but is distinguished at once by its very long involute leaves
and light colored, more loosely flowered panicle.” [M. gracilis sensu Amer. auct. is
considered to be M. montana (Nutt.) Hitche. True M. gracilis (H.B.K.) Trinius is
not a synonym of M. montana but a related species.] Lastly this comment (p. 89):
“This is a Mexican grass, not before observed in the distributed collections made
within the limits of the United States, and perhaps now found for the first time within
our limits. It should be added that identification of Pringle's specimens with M.
virescens Trin., is based upon descriptions of that plant only.”

This report by Scribner (1882) of the occurrence of Muhlenbergia virescens in the
Southwestern United States appears to account for Vasey (1883, 1892), Beal (1896),
and subsequent authors having added Arizona and New Mexico to its range. Vasey
(1892) included a detailed description of M. virescens stating (p. 67): “. . . New
Mexico, Arizona, and Mexico.” Hitchcock (1913), in Mexican Grasses, indicated
that the range of M. virescens extended from central México into New Mexico and
Arizona. Two years later Wooton & Standley (1915), in their Flora of New Mexico
cited (p. 72) “18. Muhlenbergia virescens (H.B.K.) Kunth” [obviously following
Hitchcock 1913], quoting the type locality from H.B.K. The range is given as
“Arizona and New Mexico to Mexico.” There is no descnption; however two
collections are cited: Palmer from northwestern New Mexico, and a Bigelow
gathering from Ben More (sic!) [a peak near Santa Rita, in eastern Grant County]. A
specimen (at US) collected by Edward Palmer s.n. [no date] gives only “State of New
Mexico.”

Hitchcock (1913, p. 301) listed Muhlenbergia virescens (H.B.K.) Kunth, Rév.
Gram. 1:64. 1829. He indicated that it is based on Podosaemum virescens H.B.K..,
and quoted the type locality from the orginal description. The range, however,
includes New Mexico and Arizona. Three collections from México are listed: Rose
3527 from Sierra Madre, Zacatecas, and two Parry & Palmer numbers (920 and 928 in
1878), both from San Luis Potosi.

Reeder & Reeder: Resurrection of Muhlenbergia straminea 419

Fig. 1. Habits and spikelet details. A, Muhlenbergia
virescens, from Scribner (1897); B, M. straminea, from
Hitchcock (1951).

Fig. 2. Distributions of A, Muhlenbergia virescens and
B, M. straminea.

420

Pel VIE ONL OMGILI June 1995 volume 78(6):417-427

TABLE |

Chromosome Numbers for Muhlenbergia straminea and M. virescens (All collections
are Reeder & Reeder, with vouchers deposited at ARIZ.)

Collection
6903
6906
6907
6908
6909
6910
6911
6914
6918
6922
6924
6926
6928
6929

Collection
4750
475l

4965

tw
tw
tN
£

ww

2n
20
20+1

40

Muhlenbergia straminea
Locality

AZ: Cochise Co., Chiricahua Mts., Rustler's Park
Campground. 2530 m_ 16 Oct. 1977.

AZ: Pima Co., Santa Catalina Mts., near Loma Linda Picnic
area. 2440 m 18 Oct. 1977.

AZ: Pima Co., Santa Catalina Mts., above Geology View
Point. 2165m 18 Oct. 1977.

AZ: Graham Co., Graham [Pinaleno] Mts., near Shannon
Campground. 2774m_ 19 Oct. 1977.

AZ: Graham Co., Graham [Pinaleno] Mts., Turkey Flat.
2316 m 19 Oct. 1977.

AZ: Cochise Co., Huachuca Mts., Carr Canyon. 2195 m
21 Oct. 1977.

AZ: Cochise Co., Huachuca Mts., Carr Canyon. 2255 m
PTOct 97a:

NM: Grant Co., Cherry Creek Campground, N of Pinos
Altos. 2103 m 26 Oct. 1977.

NM: Catron Co., S of Luna. 2440 m 27 Oct. 1977.

AZ: Navajo Co., N of Indian Pines. 2250 m 27 Oct. 1977.

AZ: Coconino Co., San Francisco Mts., on Snow Bowl
Road. 2440 m 28 Oct. 1977.

AZ: Gila Co., along Old Mogollon Rim Road. 2225 m 28
Oct. 1977.

AZ: Gila Co., along Old Mogollon Rim Road. 2440 m 3
Nov. 1977.

AZ: Apache Co., SE of Big Lake Campground. 2680 m 4
Nov. 1977.

Muhlenbergia virescens
Locality

MEX: Guanajuato, 9.5 km NE of Ciudad Guanajuato. 2450
m 19'Ock 1966:

MEX: Guanajuato, 27.5 km NE of Ciudad Guanajuato.
2400 m_ 19 Oct. 1966.

MEX: San Luis Potosf, 19.5 km SW of Ciudad San Luis
Potosf. 2250 m, 15 Sept. 1967.

eee

Reeder & Reeder:

Resurrection of Muhlenbergia straminea

TABLE II

42]

Comparison of morphological features between Muhlenbergia virescens, M.
curvula, and M. straminea.

Basal sheaths and
blades
Blades

Blade width

Blade surface

Inflorescence color

Glume I

Glume II

Glume surface

Lemma

Lemma surface

Stamens

Flowering time

M. virescens

straight

involute

ca. 1.5mm

scabrous below,
hispid above

green or dark green
(1.5) 2.0--3.0 (3.5)
mm

(3.0--3.5) 4.0--4.5
(5.0): mm

glabrous, shiny,
translucent

(3.0) 3.5--4.0 mm

pubescent on lower
1/2--2/3

2.0--2.5 (3.0) mm
Sept.--Oct.

M. curvula

straight

involute

ca. 1.5mm

scabrous below,
hispid above

dark green

(2.4) 2.5--3.0 mm

4.0--4.5 (5.0) mm

glabrous,
translucent

3.5--4.0 mm

pubescent lower
2/3

ca. 2 mm

Sept.--Oct.

M. straminea

curling, like wood
shavings

flat, becoming
involute at tip

2.5--3.5 mm

scabrous below,
with tiny
spicules
between the
ribs above

yellowish or pale
green

(3.0--3.5) 4.5--
5.0 mm

(4.0) 4.5--5.5
(6.0-7.0) mm

dull, minutely
scaberulous

4--5(6) mm

pubescent on
midnb at base
and on
margins on
lower 1/2

(2.0) 2.5--3.0 mm

March--Apnl
(May)

422 PHY TOLnOG TA June 1995 volume 78(6):417-427

Recent authors, e.g. McVaugh (1983), Beetle (1987), and Allred (1993) have
continued to include the southwestern United States within the range of Muhlenbergia
virescens. McVaugh (p. 263) states “Open dry grassy places in sparse oak or oak-
pine forest, 1600--2700 m on the Central Plateau and the ranges to the west, flowering
Oct--Nov.” He includes “S w U.S.” as well as various central and western states of
México. Beetle (1987) cites “New Mexico and Anzona, south to Michoacan and
Mexico.” Allred (1993) gives only the distribution within New Mexico.

Hitchcock (1913, p. 302) described as new Muhlenbergia straminea, the type
collection, R. Endlich 1210, 10 April, 1906, from pine and oak woods, Tecorichu,
western Sierra Madre, Chihuahua, elev. 2200 m. Two other gatherings from the same
general area are cited: Endlich 1226 and 1210a, from Tierra Colorada, dated April,
1906. [Hitchcock indicated that the specimens at US are from material sent him by Dr.
I. Urban, Berlin Botanical Garden.] There is a rather lengthy description, followed
by (p. 302): “This species is distinguished by the stramineous appearance of the
glumes and foliage and by the flat, ribbon-like lower sheaths, which withdraw from
the culms and become spirally twisted or curled like shavings. The long, papery
glumes indicate an affinity with Triniochloa but it is distinguished from that genus by
the terminal awn of the lemma. It appears to be most nearly allied to M. longiglumis
[Vasey, G., Contrib. U.S. Natl. Herb. 1:283. 1893].”

Muhlenbergia straminea may somewhat resemble Triniochloa, however careful
examination reveals that the members of the latter genus are characterized by closed
sheaths, and lodicules which are often fused and lack vascular traces. Also the leaf
anatomy is pooid; in Muhlenbergia it is eragrostoid (chlondoid). Triniochloa is clearly
related to Melica and Schizachne. [See J.R. Reeder Systematic position of the genus
Triniochloa (Gramineae). Amer. J. Bot. 55:735. 1968 (abstract). ]

Muhlenbergia longiglumis is an attractive Mexican species with compressed-keeled
sheaths, and long glumes which may be anstate. Soderstrom (1967) included it as a
member of § Epicampes. Muhlenbergia straminea belongs within the § Podosaemum
as outlined by Soderstrom, and is related to the M. montana group of species. In fact
when Hitchcock treated the genus in the North American Flora (1935a), he placed M.
straminea as a synonym of M. virescens, but gave no reason for this change. Since
the original material of M. virescens was collected in central México, and the type
specimen of M. straminea came from the mountains of northern Chihuahua, it seemed

prudent to re-examine these two taxa with respect to the plants which are to be found
in Arizona and New Mexico.

Perhaps one of the most striking differences to be noted from the protologues of
these taxa is that the original material of Muhlenbergia virescens was collected in
September; that of M. straminea in early April. Anyone familiar with “M. virescens”
in the mountains of New Mexico and Arizona is aware that it blooms very early in the
spring. Another characteristic, perhaps known only to us, is that the flowers are
actually initiated in late autumn, at which tme meiosis in the microsporocytes Occurs.
We tned unsuccessfully for several years in early spring to collect young
inflorescences for cytological examination. Even though panicles had not yet emerged
from the sheath, the anthers were already fully formed and contained mature pollen
grains. At this time also, anthers have their characteristic reddish purple color, which
is retained in the fixative and in 70% alcohol in which they are stored. Eventually we
discovered that good cytological matenal can be collected in late September to early

Reeder & Reeder: Resurrection of Muhlenbergia straminea 423

November. Numerous good chromosome counts were obtained from collections
made in late autumn in various localities in Anzona and New Mexico. The possibility
that meiosis might occur in the autumn of the previous year occurred to us when we
found plants partially covered with snow in March, many of which had inflorescences
exposed, some well exserted from their sheaths! In contrast, we have observed M.
virescens in full flower from August to October, and at that ume have collected young
inflorescences in satisfactory condition for cytological examination. Although, as can
be seen in Table I, collections for chromosome counts for both M. virescens and M.
straminea were made in the month of October, there is an important difference with
respect to the time involved for the completion of flowering. As mentioned above, the
initiation of meiosis within the anthers of M. straminea takes place in late autumn, and
the plants remain dormant during the winter, blooming in early spring of the following
year. In the case of M. virescens, meiosis occurs in late summer or autumn, and the
plants bloom soon after in the same season.

In 1951 little was known regarding the flowernng habits of this southwestern
grass. In Gould (1951), C. Reeder (p. 211) made the following comment: “. . .
flowering March to June and occasionally again in the summer and fall. New Mexico
and Anzona to central Mexico. An interesting forest bunchgrass, closely related to the
widespread and vanable Muhlenbergia montana complex but rather unique in its
spring-flowering habit and conspicuous with its peculiarly coiled leaves [and sheaths]
and strikingly colored anthers.” Obviously the idea that it may flower in late summer
or fall arose from the early emergence of next-year's panicles. ARIZ has at least one

_ such specimen, L.N. Goodding M-314 (ARIZ-55284) from the Huachuca Mts.,

Cochise Co., Anzona, 5 Nov 1937. It is worth noting that specimens of this
southwestern species collected after late April ordinarily consist of empty glumes with
few or no florets. Even at this stage, the plants can usually be identified by their
characteristic glumes and curling foliage.

It is true that Muhlenbergia virescens and M. straminea share certain
characteristics, e.g., both are commonly to be found in oak or pine-oak forests in the
higher mountainous areas of México and southwestern USA (mostly between 1900
and 2800 m); a fragile hyaline ligule commonly 10 to 20 mm long; subequal long acute
or acuminate glumes, the first 1-nerved, the second 3-nerved; and narrow somewhat
contracted panicle. There are, however, a number of morphological differences which
serve to distinguish these species even on herbarium sheets (Table II).

Swallen (1950) described as new Muhlenbergia curvula, which he appears to have
compared to M. virescens based on the northern element (M. straminea). This is not
surprising since the descriptions of M. virescens in North American Flora (1935a) and
Hitchcock's Manual (1935, 1951), as well as the illustration in the latter work, are
based on the northern plants. In the Manual the figure of this species (1935, fig. 785;
1951, fig. 566) is drawn from Palmer 565 in 1890 collected at Willow Springs,
Navajo Co., Anzona. The type specimen of M. curvula came from México:
Guanajuato: “12 miles from [Cd.] Guanajuato on road to Santa Rosa,” Sept. 30, 1946
H.E. Moore Jr. 1353 (US-1963089!). Since the Moore specimen fits well within M.
virescens, we consider M. curvula Swallen and M. virescens (H.B.K.) Trinius to be
synonyms. Further justification for this change is the fact that the type collections of
both species came from essentially the same area in Guanajuato, México.

424 PHY OE OrGieA June 1995 volume 78(6):417-427

Of particular interest is Parry & Palmer 920 in 1878 (US-995807) from San Luis
Potosf area, México, elevation 6,000--8,000 ft., on which one finds in Mrs. Agnes
Chase's script: “Like HBK type of Pod. [Podosaemum] virescens.” This specimen
does, indeed, resemble the type of the Humboldt & Bonpland collection (P!), but in no
way resembles M. virescens as illustrated in Hitchcock's Manual (1935, 1951). (fig.
1,B). Scribner (1897, fig. 110, p. 128) in American Grasses-Illustrated has a sketch
of M. virescens which is quite consistent with the type, although his comment below
the figure gives: “At an altitude of 1800 to 2400 m on the mountains of Arizona and
New Mexico. Mexico. May, June.” (fig. 1,A). Although the figure fits our concept
of M. virescens, the range given for the species includes M. straminea, but he has
noted the early flowering of the southwestern material. In 1891 Vasey reported on
grasses collected by Dr. Edward Palmer in western México and Anzona. On p. 114
one finds “Muhlenbergia virescens Trin.” from “Near the summit of the mountain
Alamos.” March 26 to Apr. 8. 1890. US has four sheets of this number; all prove to
be Muhlenbergia straminea. This species has been collected more recently from about
the same locality in Sonora, México, by V.W. Steinman & M. Baker 9373 on March
11, 1993 (ARIZ). Other collections from the mountainous areas of Sonora and
Chihuahua have appeared within recent years. For the distribution of these two
species see Figure 2.

Of the new species recently described by Y. Herrera (1987, 1992) from México
only one appears to pose any problem of separation from Muhlenbergia straminea.
The type of M. durangensis came from Mpio. Mezquital, state of Durango, and like
M. straminea flowers early in the spring. However, it may be distinguished by its
very short truncate ligule (0.7--1.0 mm long) which has short acute projections from
the sheath on either side. Further, the spikelets tend to be larger (5.0--6.5--7.0 mm
long), and anthers mostly 3.0--3.5 mm long. The other species (Muhlenbergia
cualensis and M. michisensis) differ in having only one nerve in the second glume. In
both Muhlenbergia virescens and M. straminea the second glume is distinctly 3-
nerved.

In a recent paper, Herrera & Grant (1993) consider the correlation of
morphological and flavonoid data within the Muhlenbergia montana complex as
generated by the DELTA system. The discussion does not instill great confidence,
e.g. one finds the repetition of three separate collections under two different species.
Further, the inclusion of M. argentea Vasey within this complex is puzzling. Even
though Vasey (1886) in the onginal description of M. argentea stated that the upper
“empty glume” is “three-toothed at the apex,” Beal (1896, p. 232) described the
glumes of M. argentea as “subequal, linear-lanceolate, the apex more or less 2-
toothed, 1-nerved.” Hitchcock (1935a, p. 465) described the same species in some
detail, basing his descnption on the type material, which was all that was available at
the time. The glumes are said to be “about equal, thin, oblong or somewhat notched,
the midnerve extending into a mucro or short awn, the summit minutely pubescent or
ciliolate.” The glumes in specimens of M. argentea we have examined fit this
description. Whereas the second glume in M. argentea may be 3-toothed in some
plants, other characteristics do not suggest a relationship with the M. montana
complex. The sheaths invest the culms and are slightly compressed-keeled; the glumes
are l-nerved; the lemma is deeply bifid; and of special note is the fact that the caryopsis
is conspicuously flattened. Muhlenbergia argentea clearly is not a member of the M.
montana complex.

Reeder & Reeder: Resurrection of Muhlenbergia straminea 425

In summary, Muhlenbergia straminea Hitchc. is shown to be a species worthy of
recognition, and is not conspecific with M. virescens (H.B.K.) Trinius.
Muhlenbergia curvula Swallen is reduced to synonymy with M. virescens. To our
knowledge, the occurrence of meiosis in M. straminea in late autumn, preceding
flowering the following spring, is unique among members of the M. montana
complex.

ACKNOWLEDGMENTS

Our special thanks to Dra. Alicia Lourtieg of the Paris Herbarium for assistance in
making the Humboldt & Bonpland collections available for our study. Curators of the
following herbaria facilitated use of their specimens: ARIZ, ENCB, F, MO, P, RM,
TAES, and US. Thanks also to Philip Jenkins and James Rominger who reviewed the
manuscript.

LITERATURE CITED

Allred, K.W. 1993. A Field Guide to the Grasses of New Mexico. Dept. Agric.
Communications, New Mexico State Univ., Las Cruces, New Mexico. 258 pp.

Beal, W.J. 1896. Grasses of North America. Vol. 2. New York, New York. 1-viil,
1-706.

Beetle, A.A. 1987. Noteworthy grasses from Mexico. XIII. Phytologia 63(4):209-
251;

Fournier, E.P.N. 1886. Mexicanas Plantas. Pars Secunda, Gramineae. Paris,
France. pp. i-xix, 1-160.

Gould, F.W. 1951. Grasses of Southwestern United States. Univ. Anzona Biol.
Sci. Bull. 7. pp. 1-342.

Hemsley, W.B. 1885. Botany. In: Biologia Centrali-Americana. Vol. 3. London,
Great Britain. (Gramineae, pp. 474-588.)

Herrera Arrieta, Yolanda. 1987. Una nueva especie de Muhlenbergia (Gramineae) del
Estado de Durango. Phytologia 63:457-460.

Herrera Armeta, Yolanda & P.M. Peterson. 1992. Muhlenbergia cualensis and M.
michisensis (Poaceae: Eragrostideae): two new species from Mexico. Novon
2:114-118.

Herrera Arrieta, Yolanda & W.F. Grant 1993. Correlation between generated
morphological character data and flavonoid content of species in the Muhlenbergia
montana complex. Canad. J. Bot. 71:816-826.

Hitchcock, A.S. 1913. Mexican Grasses in the United States National Herbarium.
Contr. U. S. Natl. Herb. 17:181-389.

Hitchcock, A.S. 1935a. Poaceae. North American Flora 17(6):419-482.

Hitchcock, A.S. 1935. Manual of Grasses of the United States. U.S. Dept. Agric.
Misc. Publ. 200. 1040 pp.

Hitchcock, A.S. 1951. Manual of Grasses of the United States. Second Edition,
revised by Agnes Chase. U.S. Dept. Agric. Misc. Publ. 200. 1051 pp.

426 PHY TOLOGTA June 1995 volume 78(6):417-427

Humboldt, A., A. Bonpland, & C.S. Kunth. 1816. Nova Genera et Species
Plantarum. Vol. 1. Paris, France. (Gramineae, pp. 84-201.).

Kunth, C.S. 1829-1834. Révision des Graminées. Paris, France. pp. 1-666.

Kunth, C.S. 1833. Enumeratio Plantarum. Vol. 1. Agrostographia Synoptica.
Stuttgard & Tubingen, Germany. 606 pp.

Kunth, C.S. 1835. Enumeratio Plantarum. Supplementum Tomi Primi. Stuttgard &
Tiibingen, Germany. 436 pp.

McDonald, J.A. 1990. The alpine-subalpine flora of northeastern Mexico. Sida
124(1):21-28.

McVaugh, R. 1983. Flora Novo-Galiciana. Vol. 14. Gramineae. Univ. Michigan
Press, Ann Arbor, Michigan. 436 pp.

Reeder, J.R. 1967. Notes on Mexican grasses. VI. Miscellaneous chromosome
numbers. Bull. Torrey Bot. Club 94: 1-17.

Reeder, J.R. 1968. Notes on Mexican grasses, VIII. Miscellaneous chromosome
numbers-2. Bull. Torrey Bot. Club 95:69-86.

Roemer, J.J. & J.A. Schultes 1817. Systema Vegetabilium. Vol. 1. Stuttgart,
Germany. pp 1-viii, 1-642.

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

Scribner, F.L. 1897. American Grasses (Illustrated). U.S. Dept. Agric. Div.
Agrost. Bull. 7. 331 pp. [cf fig. 110].

Soderstrom, T.R. 1967. Taxonomic study of subgenus Podosemum and section
Epicampes of Muhlenbergia (Gramineae). Contr. U.S. Natl. Herb. 34(4):75-189.

Swallen, J.R. 1950. New Grasses from Mexico, Central America, and Surinam.
Contr. U.S. Natl. Herb. 29(9):395-428.

Trinius, C.B. 1824. De Graminibus Unifloris et Sesquifloris. Petropoli, Russia.
319 pp. + 5 pl.

Vasey, G. 1883. The Grasses of the United States: Being a Synopsis of the Tribes

and Genera with Descriptions of the Genera and a List of the Species. U.S. Dept.
Agric. Special Report 63. 47 pp. [published 30 Aug 1883].

Vasey, G. 1886. New species of Mexican grasses. Collected by Dr. Ed. Palmer, in
S.W. Chihuahua, in 1885. Bull. Torrey Bot. Club 13:229-232.

Vasey, G.R. 1892. Monograph of the Grasses of the United States and British

America. Contr. U.S. Natl. Herb. 3(1):1-89. [published 25 Feb 1892].
Wooton, E.O. & P.C. Standley. 1915. Flora of New Mexico. Contr. U.S. Natl.

Herb. 19:1-794. [published 24 Jun 1915]. Reprinted 1972. (Poaceae, pp. 43-_

109.).

Note: The chromosome number of Reeder 4965 from San Luis Potosi was |

determined to be 2n = 40. It is of interest that in this instance meiosis is somewhat

regular with 4 or 5 quadrivalents clearly visible, suggesting that it may be an

autotetraploid.

A better understanding of specific features, and the separation of Muhlenbergia

straminea, reveal that three of our collections for which chromosome counts were —

reported earlier as M. virescens in fact represent the closely related M. quadridentata
(H.B.K.) Trinius. The specific collections are 4664 and 4475 from Durango (Reeder

Reeder & Reeder: Resurrection of Muhlenbergia straminea 427

1967) and 4703 from Zacatecas (Reeder 1968). Muhlenbergia quadridentata may be
distinguished by its shorter ligule which tends to be firmer near the base, frequently
with vascular traces evident; blades, which are flat becoming involute, are strongly
ribbed with tiny spicules on the ribs; and glumes that are dull (scaberulous and may
have a few hairs at the base).

A few reports of Muhlenbergia virescens in the literature need to be clarified as
well. McVaugh (1983) cited Reeder & Reeder 6437 from Monte Escobida, Zacatecas,
as M. virescens. Re-examination of the specimen reveals it to be M. quadridentata, the
lemma of which is pubescent on the lower 1/2 to 2/3. This type of pubescence is the
principal character used by H.B.K. (1816) to segregate Podosaemum gracilis H.B.K.
There is little else to distinguish these plants from their close relative, M.
quadridentata.

McDonald (1990) listed Muhlenbergia virescens in an enumeration of species from
alpine-subalpine mountains of northeastern México. He indicated that the grasses
were identified by us. Another check, along with the separation of M. straminea and
M. virescens, shows that indeed McDonald 2045 from the Pefia Nevada, Tamaulipas,
is true M. virescens, whereas McDonald 2100 from Sierra La Vega, Coahuila, is M.
quadridentata.

Phytologia (June 1995) 78(6):428-445S.

ANEMONE TUBEROSA (RANUNCULACEAE) IN TEXAS

Marshall Enquist
11511 Metric Blvd. #1033, Austin, Texas 78758 U.S.A.
&
Bonnie Crozier

3411 Spanish Oak Drive, Austin, Texas 78731 U.S.A.

ABSTRACT

Four species of Anemone, all with tuberous rootstocks, occur in Texas:
Anemone caroliniana Walter; A. berlandieri Pritzel; A. edwardsiana Tharp; and
A. tuberosa Rydberg. The recently described species A. okennonii Keener &
Dutton is treated as within A. tuberosa where it is described anew as var.
texana Enquist & Crozier, var. nov. A key and distribution maps are
provided for the Texas taxa of Anemone.

KEY WORDS: Ranunculaceae, Anemone, Texas

INTRODUCTION

Anemone okennonii Keener & Dutton (1994) was recently described as a new
species endemic to central Texas. Its authors noted (p. 192) it is “taxonomically
closest to A. edwardsiana” but added the caveat that “whether these two species are
closest phylogenetically remains in question.” They observed that the geographic
range of the new species overlaps with A. berlandieri Pritzel but appears to lie isolated
between A. edwardsiana Tharp and A. tuberosa Rydberg. Based on our own field and
herbarium experience, however, we believe that the relationship of A. okennonii lies
with A. tuberosa. Earlier botanists, including Donovan Correll, Cyrus L. Lundell,
Marshall Johnston, Billie Turner, and Barton Warnock, correctly identified material
referable to A. okennonii as belonging to A. tuberosa (s.l.). In addition, we find
several misidentified collections that were later correctly annotated as A. tuberosa by
Marshall Johnston and Lloyd Shinners. In this paper we have attempted to assess the
status of A. okennonii, especially as regards its relationship to A. tuberosa and A.
edwardsiana. This was accomplished by the examination of numerous populations in

428

Enquist & Crozier: Anemone tuberosa in Texas 429

the field as well as a review of herbarium maternal so as to determine the biological
status of each.

MATERIALS AND METHODS

The authors examined fifteen populations of Anemone tuberosa in Crockett,
Edwards, Kimble, Sutton, and Val Verde counties, from which more than 300
individuals from seven populations (see Table 1) were studied by Crozier for
characters of the sepals and achenes. Including our own collections, we studied
approximately 140 sheets of A. tuberosa from México, Arizona, New Mexico, and
Texas.

DISCUSSION

Keener & Dutton (1994), relying upon Joseph & Heimberger (1966), assert that
the range of Anemone tuberosa extends only into “extreme western Texas (El Paso
Co.)”. However, since the Joseph & Heimberger paper there has been a gradual
published acknowledgment that A. tuberosa extends much farther east. Warnock
(1970) identified A. tuberosa as present in Big Bend National Park in Brewster
County, Texas. Correll & Johnston (1970) did not orginally include A. tuberosa in
the flora of Texas, but by 1976 Marshall Johnston had annotated numerous sheets in
the University of Texas collection as A. tuberosa |Warnock T205, Warnock 442,
Smith M338, Correll, Correll, & Benson 30844; Correll & Correll 30884, McVaugh
7729; Tharp 1800; Correll & Flyr 38359; Rose-Innes & Warnock 565; Warnock 5199,
Turner & Warnock 273; Lundell & Lundell 10176; and Worthington 4229). Johnston
(1988 & 1990), in reference to A. tuberosa, stated, “The occurrence of this species in
Western Texas has been confirmed, including the occurrence in Big Bend National
Park and other southern trans-Pecos areas east to the Pecos River.” This located the
species in Texas as far east as Pecos, Terrell, and Val Verde counties.

Dr. James Hennckson has placed a rough draft of the unpublished “Chihuahuan
Desert Flora” in the University of Texas herbarium for corrections and annotations. In
its present form, the draft gives the following range information for A. luberosa .. .
“Very local in trans-Pecos Texas. (El Paso, Culberson, Brewster, Pecos, Val Verde
Cos.); rare in Coah. (Tule Canyon near the Rio Grande), mostly below 1000 meters;
Baja C., Calif., Son., Ariz., N.M., Tex., Coah., N.L., early spring, Feb.-Mar.”

Recent collections by the authors, in 1995, indicate that Anemone tuberosa Rydb.
is not limited to trans-Pecos Texas (which is considered to be that part of Texas west
of the Pecos River) but also occurs widely in the Edwards Plateau east of the Pecos
River in Crockett, Val Verde, Sutton, Kimble, Edwards, and Uvalde counties (see
Map 4).

In large part, Keener & Dutton differentiate Anemone okennonii by contrasting it
with A. edwardsiana. We believe that such a comparison is irrelevant. Many

430 PHYTOLOGIA June 1995 volume 78(6):428-445

characters of A. okennonii that constitute “differences” when compared to A.
edwardsiana become similarities when compared to A. tuberosa (see Keener & Dutton
1994, Table 1, p. 194-195). Of the “differences” that remain, most are the result of
incomplete or inaccurate information regarding one or both of the subject species.

Keener & Dutton provided a chart (p. 201) summarizing eleven differences
“between A. okennonii and other Texas species of Anemone with tuberous rootstocks.
....” Four of the eleven contrasted characters, including leaf division, number of
flowers per stem, time of flowering of central and lateral flowers, and sepal number
were admitted to be like A. tuberosa. One other character, the linear divisions of the
bracts, was said to be like A. berlandieri, but this would apply also to A. edwardsiana
or the eastern elements of A. tuberosa. This leaves six characters that were said to be
unlike those of the other Texas species of Anemone. Of these six, we find two,
flower size and sepal shape, to be of little use due to the broadly overlapping ranges of
these characters in the Texas species. Reddish-glandular leaf margins and tips are not
distinctive, but are common to all the Texas species of Anemone, particularly in the

colder months. Likewise, yellow anthers are common to all the Texas species. We |

believe that attempts to distinguish subtle shades of yellow are too subjective to be of
any value. The two remaining characters, style color and leaf dissection, are also
problematic. The lavender-tinted styles of A. okennonii were said to be distinctive,
but elsewhere in their paper (p. 196) Keener & Dutton stated that the styles of A.
tuberosa are lavender. Additionally, Keener & Dutton reported that the styles of A.
caroliniana and A. berlandieri are greenish with lavender tips. We believe the last
remaining “distinctive” character of A. okennonii, the “typically 3-ternate” character of
its leaves, results from a misinterpretation of leaf morphology. Despite the assertion
that the leaves of A. okennonii are “typically 3-ternate”, we find no instance in which
this 1s the case, that is, thrice-ternate leaves for this taxon are not only not typical, but
are also apparently non-existent. The leaves of A. tuberosa in Texas are typically once
to twice-ternate (see Figure 1), as are the leaves of A. tuberosa in New Mexico,
Arizona, Utah, and California. We carefully searched our collections for any leaf that
might be considered 3-ternate. The most extreme example of leaf dissection we could
find was from a depauperate population infected with black rust found 1.35 miles
south of the Llano River on highway 385 in Kimble County. This leaf is illustrated
with a line drawing in Figure 1 - H. Close examination of the actual leaf reveals that
the margins of the central subleaflet of the central leaflet are continuous, i.e., there is
no formation of the petiolules that would be necessary to describe this as a further
ternate division. The leaf is smaller than normal, with a reduced surface area, and 1s
probably more deeply incised than normal due to disease stress. None of the other

populations we found, which were all healthy, displayed central subleaflets so deeply
incised.

Alternatively, we note that the leaves of Anemone tuberosa often have lateral
leaflets that are asymmetnically ternately divided (see Figure 1, A-G). The two leaves
shown in the type drawing of A. okennonii (Keener & Dutton 1994, p. 199)
apparently are drawn from a collection in which the positions of the central leaflet and
a lateral leaflet are transposed, that is, the apparent central leaflets are actually lateral
leaflets. This may have contributed to the confusion over the number of ternate
divisions.

Sy = 3

Enquist & Crozier: Anemone tuberosa in Texas 431

TABLE 1 — VARIATION OF SEPAL AND ACHENE CHARACTERS WITHIN AND BETWEEN
POPULATIONS OF ANEMONE TUBEROSA VAR. TEXANA.

Population _sepal sepal sepal achene achene style
number length width length width length
(mm) (mm) (mm) (mm) (mm)
Edwards Co. (Hwy. 377 NE of Carta Valley]
(N=31)
average - - - 2.7 2.4 1.0
std - - - 03 03 0.1
range - - - 2.0-3.1 2.0-3.0 0.7-1.2
Val Verde Co. [Hwy. 189 NE of Juno]
(N=41) (N=35)
average 8.0 144 43 2.9 2.8 1.0
std 05 2.2 0.9 0.2 05 0.1
range 7-9 10.0-19.5 3.06.0 2.5-3.1 2.0-3.2 1.0-1.1
Val Verde Co. [Hwy. 277 N of Del Rio]
(N=23)
average - - = 32 3.1 11
std - - - 03 0.2 0.1
range - - - 3.0-4.0 2.8-3.8 0.9-1.3
Crockett Co. [Hwy. 2083 28.7 miles N of Pandale]
(N=25) (N=39)
average 85 12.3 3.7 3.0 3.0 1.0
std 13 24 1.0 0.1 0.1 0.1
range 7-12 5.0-18.0 2565 2.7-3.2 2.5-3.2 0.9-1.2
Crockett Co. [Hwy. 2083 18.2 miles N of Pandale]
(N=40)
average - - - 28 23 1.0
std - - - 03 0.9 0.1
range . - - 2.1-3.3 2.1-3.1 0.9-1.2
Kimble Co. [Hwy. 385 1.3 miles S of Llano R.]
| (N=37)
average - - - 2.7 25 1.0
std - - - 03 05 0.1
range - - - 2.0-3.0 1.8-3.2 0.7-1.2
Kimble Co. [Hwy. 385 3.2 miles S of Llano R.]
(N=21)
average - - - 3.0 3.0 11
std - - - 0.1 02 03

range - - - 2.8-3.2 2.5-3.4 0.9-2.2

432

PHY TOLOGEA

June 1995

volume 78(6):428-445

TABLE 2 - COMPARISON OF ANEMONE TUBEROSA AND ANEMONE EDWARDSIANA.

TUBEROUS ROOTSTOCKS
length
width
BASAL LEAVES
dissection
petiole length

HEIGHT AT ANTHESIS

PEDUNCLE
pubescence below bracts
number of flowers
time of flowering

BRACT
shape

SEPALS
number
color
shape

length x width

FRUITING HEAD
shape

length
width

ACHENE
pubescence

width

STYLE
color
length

HABITAT

A. tuberosa
var. tuberosa

15-2 cm
1.0 cm

1-2 ternate
4-10 cm

1-3dm

glabrous to glabrescent
1-3
central before lateral

similar to basal leaves
but shorter petiole &
longer lobes and teeth

6-13
white to pink
linear-oblong

20.0 x 3.0 mm
cylindrical to
ellipsoidal
1.5-3.0 cm
1.0-1.5 cm
villous
2.14.0 mm
violet

1.5mm

high dry rocky slopes

A. tuberosa
var. texana

1-3 cm
(0.7)1(1.2) cm

1-2 ternate
(3)4-12(16) cm

1-3(4) dm

glabrous to glabrescent
1-3
central before lateral

linear lobes

6-12

white to pink
linear-oblong or
oblanceolate
5.0-19.5 x 2.0-6.5 mm

cylindrical to
ellipsoidal
1.5-3.5(4) cm
0.8-1.0(1.2) cm

villous
2.2-3.5 mm
violet
0.8-1.6 mm

rocky slopes and
grassy flats

A. edwardsiana

1-1.5(2) cm
0.5-1(1.2) cm

ternate
(3)4-12(20) cm

1-4(4.5) dm

glabrous to glabresceni
1-3
central before lateral

linear lobes

7-13

white
linear-oblong or
oblanceolate }
5.0-22.0 x 1.5-6.2 mm

cylindrical to
ellipsoidal
1-15 cm
0.4-0.9 cm

sparse with hairs on
margin and base
2.2-3.0 mm

yellow-green to green |
0.5-1.4 mm

shaded canyons to
open, rocky slopes

Enquist & Crozier: Anemone tuberosa in Texas 433

SS be
Zs 2 ro
sts : ~~ | i Z

LL) H. Texas, Kimble Co., On 385, 1.35 mi. S

434

PHYTOLOGIA June 1995 volume 78(6):428-445

FIG. 2— BRACT VARIATION IN ANEMONE TUBEROSA. A. Mexico, Nuevo Leon, Villaldama, 15 Mar
1983, Cowan & Nixon 3809 (TEX-LL) B. Arizona, Pinal Co., 1 Mar 1968, Keil, Pinkava, & Lahto 10790
(TEX-LL ) C. Arizona, Pima Co., Ajo Mts., 4 Apr 1948, Fouts 368 (NMU) D. Arizona, Santa Cruz Co., 1
Mar 1986, Ferguson S.N. (TEX-LL) E. New Mexico, Dona Ana Co., Las Cruces, ridge N of Bishop's Cap,
16 Apr 1995, M. Enquist & B Crozier 2799 (TEX-LL) F. Texas, El Paso Co., Franklin Mts., 1 Apr 1979,
Worthington 4229 (TEX-LL) G. Texas, Pecos Co., NE of Ft. Stockton, 15 Apr 1941, Lundell 10176 (TEX-
LL) H. Texas, Crockett Co, 24 mi N of Pandale, 12 Mar 1995, M. Enquist & B. Crozier 2734 (TEX-LL) I &
J. Texas , Val Verde Co., On 277, 1.1 mi S of int of 277 & 377, 17 Apr 1995, M. Enquist & B. Crozier 2802
(TEX-LL) K. Texas, Kimble Co., 3.2 mi. S of Llano River on 385, 17 Apr 1995, M. Enquist & B. Crozier ”
2804 (TEX-LL) L. Texas, Kimble Co., On 385, 1.35 mi. S of Llano River, 9 Apr 1995, Enquist 2790 (TEX-
LL).

Enquist & Crozier: Anemone tuberosa in Texas 435

FIG. 3 — LEAF AND BRACT VARIATION IN ANEMONE EDWARDSIANA. A. Texas, Kinney Co., On
674, 0.7 mi S of Edwards/Kinney line, 2 Apr 1995, M. Enquist 2770 (TEX-LL) B. Texas, Val Verde Co., On
1024, 6.8 mi E of int. of 1024 & Pandale Road, 5 Mar 1995, M. Enquist & B. Crozier 2731 (TEX-LL) C.
Texas, Crockett Co., Ft. Lancaster, on 290 at roadside park at top of Lancaster Hill, 26 Feb 1995, M.
Enquist 2709 (TEX-LL) D & E. Texas, Edwards Co., Towards Barksdale, on 55, 14 mi S of int. of 55 & 377
in Rocksprings, 4 Mar 1995, M. Enquist & B. Crozier 2723 (TEX-LL) F. Texas, Travis Co., Austin, 1 Mar
1937, B. C. Tharp 43999 (TEX-LL) G. Texas, Kinney Co., On 674, 0.7 mi S of Edwards/ Kinney line, M.
Enquist 2770 (TEX-LL) H. Texas, Crockett Co., Ft. Lancaster, on 290 at roadside park at top of Lancaster
Hill, 26 Feb 1995, M. Enquist 2709 (TEX-LL) L Texas, Edwards Co., Long roadcut on 2523, 1.7 mi N of
Edwards/ Kinney line, 25 Mar 1995,'M. Enquist, B. Crozier, & B.L. Turner 2755 (TEX-LL) J. Texas,
Medina co., On 1283, E of Medina Lake, 0.2 to 0.3 mi SE of int. turning off to Mico, 25 Feb 1995, M.
_ | Enquist 2704 (TEX-LL) K. Texas, Medina Co., On 12853, E of Medina Lake, 0.4 mi SE of int. turning off to
Mico, 4 Mar 1995, M. Enquist & B. Crozier 2716 (TEX-LL) L & M. Texas, Travis co., Paisano Ranch, 21
Feb 1995, B. Crozier 118 (TEX-LL).

436 PHY TOLOGTA June 1995

a —— ae
bee a me, ees

a i | Px

MAP2 - Range of Anemone berlandieri in Texas.

volume 78(6):428-445

nquist & Crozier: Anemone tuberosa in Texas 437

i 1 aa] ' ' ! é Ye
lia Ce tk. SS es ee
5 rs yen mal Ait 1 a ot oy ‘

I
!
|
I
i

MAP 4 - Range of Anemone tuberosa in Texas. Solid dots = A. tuberosa var. texana. Open circles = A.
tuberosa var. tuberosa.

438 PHYTOLOGIA — June 1995 _ volume 78(6):428-445

We believe that once these putative “differences” are discounted, a comparison of
leaf characters, achene size and pubescence, and style color indicates that the type of
Anemone okennonii clearly fits within A. tuberosa (see Table 2).

The obvious differences between Anemone tuberosa and A. edwardsiana (compare —
Figure | & Figure 2 with Figure 3) may be easily contrasted in the field, where the
two are often found in either closely adjacent or intermingled populations. We found —
the two species growing together in two populations in Val Verde County and two |
populations in Crockett County. Our collections indicate A. edwardsiana ranges from
the eastern edge of the Edwards Plateau west to the valley of the Pecos River (see Map |
3). We have seen one sheet that appears to be a mixed collection of A. tuberosa and
A. edwardsiana from Brewster County. Although we consider it likely that A. —
edwardsiana is also continuously distributed from the Pecos River west to Brewster |
County, we were not able to confirm its range west of the Pecos because that area, |
roughly to Sierra Blanca, was suffering a severe four year drought at the time we made —
our collections.

Of the nine collections of Anemone okennonii cited in the protologue, we interpret
seven of them, all made in the 1940's, as typical A. tuberosa. The remaining two —
collections were made by Mr. O'Kennon in 1992 and 1993. Since the type description
and illustration of A. okennonii do not match the appearance of A. tuberosa, we tned
to visit the O'Kennon collection sites to determine if these populations were somehow
different. The type collection (as published) is said to be from Gillespie County, “high
on a shallow roadcut in sandy loam, FM 783, 2 mi S of Doss, 22 Apr 1993”. At the
stated locality, a closely grazed hillside, we found an abundance of A. berlandieri but
nothing resembling A. okennonii or A. tuberosa. The other collection (as published) |
is said to be from Kimble County, “Hwy 385, 2.5 mi S of Llano River, 9 Mar 1992”. |
At this locality, a gently rolling plain in red sandy loam, we again found an abundance |
of A. berlandieri, but nothing resembling A. okennonii or A. tuberosa. Repeat visits |
and careful searches of these localities on an almost weekly basis from mid February
through mid Apnmil yielded only A. berlandieri. |

|
Examination of the holotype (BRIT) and the Kimble County collection (BRIT) |
offered little additional information about the exact locality of Anemone okennonii, but
we did note that the type specimen we were shown was heavily infected with black
rust. The label for the type locality reads. . . “Gillespie Co.: high on shallow roadcut
in sandy loam, along FM 783 at Edwards Creek, 2 miles S of Doss, Robert J.
O'Kennon 11390, 22 Apr 1993.” We found such a roadcut 2.3 miles south of Doss at
Edwards Creek but again found nothing resembling A. tuberosa or the plant described
as A. okennonii.
|

The specimen label for the only other cited collection was similarly brief, reading:
“Kimble Co., Hwy 385 2.5 miles So. Llano River. High on shallow roadcut in rocky
limestone, Robert J. O'Kennon 8813, 9 March 1992.” The label data is ambiguous.
Although there is a South Llano River, Highway 385 does not cross it. The North and
South forks of the Llano River unite in Junction, with the river then flowing northeast
towards Highway 385. We conclude that the label intends to indicate a locality 2.5
miles south of the Llano River on Highway 385, as stated in Keener & Dutton (1994).
As we have said, this location is a gently rolling plain of red sandy loam, with no
roadcut in sight. Here, we did find Ferocactus setispinus Engelm., which was listed

|

}
t
|

Enquist & Crozier: Anemone tuberosa in Texas 439

as a plant associate of Anemone okennonii, but we found no evidence of the latter
_ species. Searching for roadcuts in limestone, we found one at 1.35 miles south of the
_ Liano River with a small population of A. tuberosa. About 80% of the plants in this
_ population were heavily infected with black rust, as was the type specimen said to be
from Gillespie County. We also found an uninfected population of A. tuberosa at
another roadcut 3.2 miles south of the Llano River.

We find the label data for the O'Kennon localities, including the type locality, to be
inaccurate, confused, and incorrect as to mileages. We also believe there is a strong
possibility that the locality of the diseased Gillespie County type specimen is actually
the same as that of the diseased population 1.35 miles south of the Llano River in
Kimble County.

In this paper, we treat Anemone okennonii as a synonym of A. tuberosa, but

_ because plants resembling A. okennonii cannot be found at the type locality as stated in

the label by Mr. O'Kennon for this taxon, and because the infected plants of the type

collection seem likely to have been collected in Kimble County rather than Gillespie

County, the problematic and ambiguous typification of this taxon cannot serve to
establish its identity.

Because we regard Anemone okennonii as within A. tuberosa but as a distinctive
eastemmost segment of it, we treat the former at varietal rank. Recognition that A.
_ tuberosa extends from El Paso County to central Texas amounts to a 400 mile range
extension eastward for the species. The plants of the new vaniety differ slightly from
typical A. tuberosa, and we have modified the description of the species to reflect their
- inclusion.

| ANEMONE TUBEROSA Rydberg, Bull. Torrey Bot. Club 29:151-152. 1902.

Anemone sphenophylla sensu Britton, Ann. New York Acad. Sci. 6:220. 1891.
(but see discussion)

Robust, apically villous herbs; stems simple below, usually branched above
-involucre, 1-4 dm tall at anthesis, from brownish, oblong-obovate tuberous rootstocks
1-3 cm long and 1 cm thick. Radical leaves several, 1-2 ternate; petioles 4-12 cm
long. Leaflets 1-2 cm wide, subglabrous, variously parted or cleft, with cuneate
obovate segments having acute tips, sessile to petiolulate. Involucral bracts 3, similar
to radical leaves in dissection or dissimilar with elongate lobes, reduced, short-
petiolate, thinly pilose, positioned above middle of scape at anthesis; secondary
branches one or more, 2-bracteate. Flowers 1-several per scape; lateral flowering
branches maturing later than central flower: scape villous above involucre,
subglabrous below. Sepals 6-13, white to pink, linear-oblong, 5-19 mm long, 2.5-
6.5 mm wide, pubescent on the outer side; anthers yellowish-brown, ca. 1 mm long.
Fruiting heads cylindrical to ellipsoidal, 1.5-4.0 cm long and 1.0-1.5 cm broad;
achenes flat, orbicular, 2-4 mm wide, densely villous; styles reddish-purple to
lavender, + or - erect, filiform, 1/3 to 1/2 length of achenes; 2n = 16 (Joseph &
Heimberger 1966).

| February to May. High dry rocky slopes and grassy flats, southeastern California,
southern Nevada, and southwestern Utah, southeastward through Anzona and New

440 PHY TOL GGA June 1995 volume 78(6):428-445

Mexico to central Texas; south into Baja California, Coahuila, and Nuevo Leo6n in
México. [description adapted from Keener & Dutton 1994].

ANEMONE TUBEROSA Rydberg var. TUBEROSA
Specimens examined:

MEXICO. Baja California: On bare rocky E slope of Cerro Matomi, 4 May 1973,
R. Moran 20797 (TEX-LL).

U.S.A. Arizona: Gila Co., Beside Beeline hwy, 14 mi S of Payson, 2 Apr 1960,
N.H. Russell 11352 (BRIT). Maricopa Co., Desert slopes by Apache Trail, 2 mi W
of Canyon Lake, 28 Feb 1960, N.H. Russell 11123 (BRIT); E side of Canyon Lake,
28 Feb 1960, N.H. Russell 11159 (BRIT); Lake Pleasant Regional Park, 6 Apr
1966, Moore, Pinkava, & Keil 152 (TEX-LL). Pima Co., Ajo Mts., 4 Apr 1948,
Fouts 368 (NMC); Hills W of Tucson, 17 Feb 1935, F. Shreve 13163 (BRIT);
Rocky hillside near Colossal Cave, Rincon Mountains, 25 Mar 1946, C. Van Cleve 5
(TEX-LL). Pinal Co., 1 mi N of Peppersauce Canyon, 1 Mar 1968, Keil, Pinkava,
& Lahto 10790 (TEX-LL). Santa Cruz Co., Summit of Fraguita Peak, Cobre Ridge,
Coronado Natl Forest, 1 Mar 1986, G.M. Ferguson s.n. (TEX-LL). New Mexico:
Dona Ana Co., Bishop's Cap, 23 Mar 1935, Hershey s.n.(NMC); Tortugas Mts., 14
Mar 1941, No collector, s.n. (NMC); West of Organ Mts, 21 Mar 1941, No

collector, s.n. (NMC); Little Mt., mesa west of Organ Mts., 7 Mar 1900, E.O. |

Wooton s.n. (NMC); Ridge on N side of Bishop's Cap, SE of Las Cruces, 16 Apr

1995, M. Enquist & B. Crozier 2799 (TEX-LL,TAES). Luna Co., Florida Mts., 7 |

Mar 1897, C.L. Herrick 309 (NMC); Hermanas, 18 Mar 1940, A.L. Hershey s.n.

(NMC). Sierra Co., 1 mi W of Hillsboro, 30 Apr 1904, O.B. Metcalfe 1558 (NMC). |
Texas: El Paso Co., E lower slopes of Mt. Franklin, 19 Apr 1952, Barton Warnock |
10409 (SRSC); E slopes of Fusselman Canyon, Franklin Mts., 25 Apr 1973, Barton —
Warnock 23688 (SRSC); Slopes below Cottonwood Spring above Tom Mays —
Memorial Park, W side of Franklin Mts., 6 Apr 1970, D.S. Correll & D. Flyr 38359 |

(BRIT,TEX-LL); Franklin Mts. along Trans-Mountain Road near Fusselman Canyon
flood control dam, 1 Apr 1979, R.D. Worthington 4229 (TEX-LL). Winkler Co., 5
mi E of Kermit, 24 Apr 1975, Barton Warnock 23875 (SRSC).

Since the 1930's, many collections of Anemone tuberosa have been made in the
western half of Texas. Its variation has not gone unnoticed. As a result,
identifications have largely been divided between A. tuberosa and A. sphenophylla

Poepp. We believe taxonomists have divided their identifications between these two |

names due to the character of the involucral bracts in most of the Texas plants.

Rydberg (1902) described the leaves and involucral bracts of Anemone tuberosa as
follows “. . . basal leaves with petioles about 5 cm. long, twice termate; divisions
rhombic-cuneate in outline, ternately cleft and toothed; teeth oblong-ovate:

involucral leaves similar to the basal ones, but short-petioled and with longer lobes and :
teeth. . .”. He cited specimens from Anzona, New Mexico, Utah, and California |

which presumably fit his description of a “leafy-bracted” Anemone.

Although specimens with involucral bracts fitting the typical elements of Anemone
tuberosa can be found in extreme West Texas (El Paso County), other plants from

|
|

- rootstocks; radical leaves petiolate, 1-2 ternate, acute-tipped, 2-4 cleft, cuneate at the

base; peduncle typically branched above the involucra] bracts, appressed pubescent
above the bracts and glabrate below: involucral bracts sessile, palmatifid into linear or
oblong acute segments; flowers (1) 2 (3) per peduncle, the first peduncle naked,

Enquist & Crozier: Anemone tuberosa in Texas 441

there east to Edwards County have involucral bracts unlike the type description. These
plants have bracts with elongate, linear lobes and segments, usually with entire
margins but sometimes few-toothed (see Figure 2).

Faced with this variation from the typical elements, many taxonomists in Texas
have relied on the interpretation of Britton (1891), who expanded the name Anemone
sphenophylla to include this taxon of the southwestern U.S. Britton described the
leaves and involucral bracts of A. sphenophylla as follows: “Radical leaves slender-
petioled ternately divided, the divisions obovate, obtuse, cuneate at the base, variously
lobed and cleft; leaves of the involucre short-petioled or sessile by a narrowed base,
palmatifid into linear or oblong acute segments .. .”. This description would seem to
better describe the linear-lobed bracts common to A. tuberosa in Texas and is the
probable reason for the numerous label and annotation identifications of this taxon as
A. sphenophylla. (We have seen no specimens of A. sphenophylla from Chile and do
not address the question of whether this name has priority over the name A. tuberosa.

_ We note the disjunct occurrence of A. multifida Poir. in Chile [Lourteig 1951] and

would not be surprised at the disjunct occurrence there of the taxon under discussion
in this paper).

Rydberg's description of the bracts of Anemone tuberosa was brief and somewhat
ambiguous, describing the involucral leaves as similar to the basal ones “but short-

_ petioled and with longer lobes and teeth. . .”.. When Rydberg described the bracts as

being similar to the radical leaves, he may have had in mind bracts such as those in

Figure 2 B, E, & F, as well as those with “longer lobes and teeth” exemplified by

Figure 2 C & D. We believe the bracts illustrated in Figure 2 C & D represent an

_ intermediate morphology between the western and eastern elements of A. tuberosa. In

_acomparison of specific individual plants, the difference between Figure 2 D (Santa
Cruz Co., Arizona) and Figure 2 K (Kimble Co., Texas) is one of degree only.

We feel it is useful to contrast the western and eastern populations of Anemone

_ tuberosa; those in Arizona are predominantly “leafy bracted” while those in most of the

Texas range have bracts that are not “leafy bracted” but instead are predominantly

_elongately lobed. We propose that the name Anemone luberosa var. texana be applied

to those plants with elongately lobed bracts that are dissimilar to the radical leaves.

ANEMONE TUBEROSA Rydberg var. TEXANA Enquist & Crozier, var. nov.
TYPE: U.S.A. Texas: Val Verde Co., On Highway 277 north of Del Rio, 1.1
miles south of intersection of 277 & 377, abundant in bar ditch on east side of
road, 25 Mar 1995, M. Enquist, B. Crozier, & B.L. Turner 2757 (HOLOTYPE:
TEX-LL; Isotypes: TAES, BRIT)

Anemone okennonii Keener & Dutton, Sida 16(1):191-202. 1994.

Anemone tuberosae Rydb. var. tuberosae similis sed segmentis ultimis
bracteorum involucralium linearibus integribusque differt.

Slender perennial herb, 4-30 cm tall at anthesis, from oblong or clavate tuberous

442 PHY DOLOG WA June 1995 volume 78(6):428-445 |

subsequent ones involucellate; sepals 6-12, oblong-oval obtuse, often with retuse tips; |

pink, white, or white with pink streaks on the outer side, 0.5-1.9 cm long, pubescent
on the outer side; fruiting head ovoid to cylindncal, 1.5-3.5 cm long; achenes
tomentose, styles violet-purple, filiform, about 1/3 the length of the achene.

February to April. Rocky slopes to grassy flats, predominantly over limestone. In
Texas, from El Paso County east to Kimble County; also in northern México.

Specimens examined:
MEXICO. Coahuila: Rio Grande, Tule Canyon, on Coahuila side above Upper

Madison Falls, 10 Apr 1973, M.C. Johnston, T.L. Wendt, & F. Chiang C. 10614
(TEX-LL). Nuevo Leén: Monterrey, 9 Mar 1923, B.C. Tharp 1800 (TEX-LL); On

“M” ridge side above San Pedro, Monterrey, 5 Feb 1961, R. F. Smith M338 (TEX-
LL); Rocky hillside 3.7 mi W of Bustamante, 4 Feb 1983, C. Cowan 3782 (TEX- |
LL); Villaldama, Mountains E of state highway 34, 1.4 km N of junction to |

Bustamante, 15 Mar 1983, C. Cowan 3809 (TEX-LL)

U.S.A. New Mexico: Eddy Co., Carlsbad Caverns Natl. Park, 19 Apr 1977, |

T.L. Burgess 4476 (TEX-LL). Texas: Brewster Co., E slope of Pulliam above
Moss Well at Big Bend National Park, 21 Mar 1967, Barton Warnock 21142
(SRSC); Glass Mts., hills in back of Jim Nichols ranch house, 9 Mar 1947, Barton
Warnock 47007 (SRSC); Alpine, 20 Mar 1937, Barton Warnock T205 (SRSC,
TEX-LL); Pine Mt., about 12 mi E of Alpine, 19 Mar 1938, B. Peyton & Barton

Warnock T439 (SRSC); Dog Canyon, Big Bend Natl Park, 8 Apr 1958, Barton |
Warnock & M. Johnston 15986 (SRSC); Lowest foothill slopes, Glass Mts., 21 Mar —

1941, Reg. Rose-Innes & Barton Warnock 565 (BRIT,TEX-LL); Chisos Mts., 15
Mar 1941, Barton Warnock 442 (TEX-LL). Crockett Co., 14.8 mi W of Ozona, 14
Mar 1949, B.L. Turner & Barton Warnock 273 (SRSC,BRIT,TEX-LL); Hwy 290, 1
mi E of Pecos River bridge, 9 Apr 1964, J. Read 572 (SRSC); On 290, at roadside
park at top of Lancaster Hill, Ft. Lancaster, 26 Feb 1995, 50 to 500 yds S of park

under top of west-facing rim, 26 Feb 1995, M. Enquist 2710 (TEX-LL,BRIT, |
TAES); On 290, at roadside park at top of Lancaster Hill, Ft. Lancaster, 5 Mar 1995, |

M. Enquist & B. Crozier 2733 (TEX-LL); W side of Pandale Road, 24.4 mi N of int.
of 1024 & Pandale Road, 12 Mar 1995, M. Enquist & B. Crozier 2734 (TEX-LL,
BRIT,TAES,MO,SRSC). Culberson Co., Victonio Canyon, E margin of Sierra
Diablo, 18 Apr 1973, M.C. Johnston, T.L. Wendt, & F. Chiang 10686 (TEX-LL).
Edwards Co., Rock knob 14.2 mi NE of int. of 277 & 377, 4 Mar 1995, M. Enquist
& B. Crozier 2724 (TEX-LL,BRIT,.TAES,MO,SRSC,NMC); On 2523, 1.7 mi N of
Edwards/Kinney Co. line at long roadcut, 25 Mar 1995, M. Enquist, B. Crozier, &
B.L. Turner 2750 (TEX-LL,BRIT,TAES). Kimble Co., On 385, 1.35 mi S of Llano
River, E side of road on low roadcut through nodular limestone, 2 Apr 1995, M.
Enquist 2776 (TEX-LL,BRIT,TAES,MO,SRSC,NMC); On 385, E side of road on
low roadcut through nodular limestone, plants heavily attacked by black rust, 9 Apr
1995, M. Enquist 2790 (TEX-LL,BRIT,TAES,MO,SRSC,NMC); On 385, low
roadcut to drainage, 3.2 mi S of Llano River, 17 Apr 1995, M. Enquist & B. Crozier
2804 (TEX-LL,BRIT,TAES,MO,SRSC,NMC); On 377, S of Junction, 0.3 - 0.4 mi
N of first crossing of the Llano River, W side of curve, 2 Apr 1995, M. Enquist 2780
(TEX-LL,BRIT,TAES,MO,SRSC,NMC). Pecos Co., Along Sanderson hwy 10-18
mi out of Ft. Stockton, 5 Apr 1953, J. Scuddy 133 (SRSC); Along hwy 20 mi NE of
Ft. Stockton toward McCamey, 27 Apr 1947, Barton Warnock 5199 (SRSC,BRIT,

Enquist & Crozier: Anemone tuberosa in Texas 443

TEX-LL); N side of Sierra Madera range, 24 mi S of Ft. Stockton, 12 Apr 1947,
Barton Warnock 5064 (SRSC); 13.5 mi E of Ft. Stockton along hwy 290, 17 Apr
1965, D.S. Correll & H. B. Correll 30884 (TEX-LL); On mesa N of Tunis Spring,
about 19 mi E of Ft. Stockton, 5 Apr 1965, D.S. Correll, H. B. Correll, & L. Benson
30844 (TEX-LL); Off US 67, E of Ft. Stockton, 15 Apr 1941, C.L. Lundell & A.A.
Lundell 10176 (TEX-LL). Sutton Co., on 189, SW comer of county, 0.8 mi NE of
int. of 189 & Co. Rd. 410, 12 Mar 1995, M. Enquist & B. Crozier 2741 (TEX-LL,
BRIT,TAES); On 277, 4.3 mi S of int. of 277 & Loop 476 in Sonora, 19 Mar 1995,
M. Enquist & B. Crozier 2746 (TEX-LL,BRIT,TAES). Terrell Co., 13 mi W of
Sanderson, 28 Mar 1947, Barton Warnock 47083 (SRSC); 4.2 mi W of Sanderson,
14 Mar 1949, B.L. Turner & Barton Warnock 319 (SRSC,BRIT); 31 mi S of
Sheffield, 14 Mar 1949, B.L. Turner & Barton Warnock 307 (SRSC,BRIT); SmiE
_ of Dryden on US 90, 30 Mar 1950, H.L. Surrat 179 (SR); 8 mi E of Dryden, 28 Mar
_ 1947, Barton Warnock 47107 (SRSC). Uvalde Co., Uvalde, common on sandy hills
and prairies, 28 Feb 1919, H.C. Hanson s.n. (TEX-LL). Val Verde Co., 20 mi N of
_ Langtry, 30 Mar 1947, Barton Warnock & G.W. Brown 4730] (SRSC); Hills above
dam at foot of Devils Lake, about 20 mi NNW of Del Rio, 31 Mar 1947, R. McVaugh
7729 (BRIT,TEX-LL); 1.1 mi S of int. of 277 & 377, 25 Mar 1995, M. Enquist, B.
Crozier, & B.L. Turner 2757 (TEX-LL,BRIT,TAES,MO); 1.1 mi S of int. of 277&
_ 377, 17 Apr 1995, M. Enquist & B. Crozier 2802 (TEX-LL,BRIT,TAES); Low,
_ curving roadcut on 1024, 6.8 mi E of int. of 1024 and Pandale Rd in Pandale, 5 Mar
_ 1995, M. Enquist & B. Crozier 2730 (TEX-LL,BRIT,TAES,MO,SRSC,NMC);

_ County road to Juno at its int. with 1024, 12 Mar 1995, M. Enquist & B. Crozier

, 2738 (TEX-LL); Loma Allta, 0.2 - 0.3 mi N of Loma Alta on E side of 277, 4 Mar

| 1995, M. Enquist & B. Crozier 2727 (TEX-LL,BRIT,TAES,SRSC,NMC): E side of
_ Pandale Road, 4.8 mi S of Pecos River crossing in narrow strip of grass along road,
| 12 Mar 1995, M. Enquist & B. Crozier 2737 (TEX-LL); On 189, 0.8 mi S of
| Buckley's Crossing, 19 Mar 1995, M. Enquist & B. Crozier 2747 (TEX-LL,BRIT,
| TAES,MO,SRSC,NMC); On 1024 to Pandale ca. 3 mi N of Junction with US 90 in
_ Comstock, 14 Mar 1985, B. Ertter & K.A. Bear 5601] (TEX-LL).

pi.

KEY TO TEXAS SPECIES OF ANEMONE
(adapted from Keener & Dutton 1994)

Plant spreading by rhizomes or stolons, involucre below middle of scape at
anthesis; scape nearly glabrous below involucre; involucral bracts similar to at least
some of the radical leaves; styles as long or longer than the ovoid achenes. .........
Bete ae Sane ce oe cc Mack. Les een ees. sae. 1. A. caroliniana
Plant without rhizomes or stolons, involucre above middle of scape at anthesis;
scape glabrous to pubescent below involucre, involucral bracts dissimilar or
similar to radical leaves; styles less than half as long as the orbicular, flat achenes..

eet aicieisAsnehincieeNcwclsaiine dnavst sone vasvachaadecutcseaudds 145 a ee ee: (2)
2. Scape simple, bearing one flower; involucral bracts distinctly dissimilar to
radical leaves; scape densely pubescent below involucre. .....2. A. berlandieri

2. Scape usually branched, bearing two or more flowers per stem; involucral
bracts similar or dissimilar to radical leaves: scape nearly glabrous to glabrous
SIS VCC VS | RR ea» eee eae ae mare mance Ta NR Te (3)

444 PH Yor @iTe O1GirA June 1995 volume 78(6):428-445

3. Leaves 1 (-2) ternate; flowers white; translucent styles colorless to yellow-

PLCEMM ee eee tees Oa onchtnce eee users serine cab aVenet sec 3. A. edwardsiana

3. Leaves (1-) 2 ternate; flowers white to pink; translucent styles pale lavender

topunplish=red yess eect sence ce term. fee woo nate see eeteminee 4. A. tuberosa
CONCLUSION

We conclude that the label data on type material of the O'Kennon collections are
inaccurate and probably reflect confusion as to where the type specimens of Anemone
okennonii were actually collected. Because of this, and because we recognize the
taxon at a lower rank, we here redescribe it with a different name and better
typification. Although we are aware that the International Code of Botanical
Nomenclature recommends the retention of any name which might be replaced in a
change of rank, we believe the broader systematic community is better served by the
nomenclature provided here as to the geography to which it is largely confined, its
inextricable relationship with A. tuberosa, and by the precise typification.

ACKNOWLEDGMENTS

We would like to thank Mike Powell and Sharon Yarborough of SRSC, Barney |

Lipscomb and Fiona Norris of BRIT, and Richard Spellenberg of NMC.

We thank Sara Hoot, to whom we have forwarded material for DNA work, for her |

comments.

We also thank Guy Nesom for the Latin diagnosis.

LITERATURE CITED

Britton, N.L. 1891. The American species of the genus Anemone and the genera |

which have been referred to it. Ann. New York Acad. Sci. 6:215-238.

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

Texas Research Foundation, Renner, Texas.

Hoot, S.B., A.A. Reznicek, & J. Palmer. 1994. Phylogenetic relationships in |
Anemone (Ranunculaceae) based on morphology and chloroplast DNA. Syst. |

Bot. 19:169-200.

Johnston, M.C. 1990. The Vascular Plants of Texas: A List Updating the Manual of |

the Vascular Plants of Texas. published by the author, Austin, Texas.

Henrickson, J. & M.C. Johnston. A flora of the Chihuahuan desert region.

Unpublished manuscript.

Enquist & Crozier: Anemone tuberosa in Texas 445

Joseph, C. & M. Heimberger. 1966. Cytotaxonomic studies on New World species
of Anemone (Section Eriocephalus) with tuberous rootstocks. Canad. J. Bot.
44:899-928.

Keener, C.S. & B.E. Dutton. 1994. A new species of Anemone (Ranunculaceae)
from central Texas, Sida 16(1):191-202.

Lourteig, A. 1951. Ranunculaceas de Sudamerica templada. Darwiniana 9:397-608.

Rydberg, P.A. 1902. Studies on the Rocky Mountain flora -- VII. Bull. Torrey Bot.
Club 29: 145-160.

Warnock, B.H. 1970. Wildflowers of the Big Bend Country, Texas. Sul Ross State
University, Alpine, Texas.

Phytologia (June 1995) 78(6):446-447.

PARONYCHIA VIRGINICA (CARYOPHYLLACEAE), A FIRST REPORT OF ITS
OCCURRENCE IN MEXICO

B.L. Turner

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

ABSTRACT

Paronychia virginica Spreng. is widespread in the eastern U.S.A. but has
not previously been reported for México, although it is relatively common in
central Texas. Recent collections by D.H. Riskind from the Serranias del
Burro, Mpio. Villa Acufia, Coahuila, México has extended its distribution
about 300 kilometers southwestwards from its previously known range. A |
map showing its distributions in Texas and México is provided.

KEY WORDS: Caryophyllaceae, Paronychia, Texas, México

Routine identification of plants from northern México has revealed the occurrence |
of Paronychia virginica Spreng. in México, where it has not heretofore been reported.
All of the collections (3 sheets) were made by Mr. David H. Riskind, in the Serranias |
del Burro, Mpio. de Villa Acufia, Rancho el Bonito (ca. 29° 01’ 30” N, 102° 07’ 30”
W), as follows:

1. Canyon El Toro, 18 Sep 1977, R. 2142 (TEX).

2. Head of Canyon El Bonito, plateau of open pine-oak woodland, 2300 m, 20
Sep 1977, R. 2263 (TEX).

3. Open slope between canyons E! Bonito and Pantera, 20 Sep 1977, R. 2286
(TEX).

Figure 1 shows the distribution of Paronychia virginica in Texas and México. |
This is based upon collections from Texas as reported in Turner (1983. Phytologia
54:9-23.) and upon collections assembled at LL, TEX since that time.

447

{ RSeiAiis -—
iggy

Fig. 1. Distribution of Paronychia vi

Phytologia (June 1995) 78(6):448-451.

SYNOPTICAL STUDY OF RHODODON (LAMIACEAE)

B.L. Turner

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

ABSTRACT

Rhododon, endemic to Texas, is treated as having two species: R. ciliatus
(Benth.) Epling, and R. angulatus (Tharp) B.L. Turner, comb. nov. The
latter taxon was originally positioned in the genus Stachydeoma by Tharp in
1945 after the erection of Rhododon by Epling in 1939. Tharp was unaware
of Epling's establishment of Rhododon, typical elements of which had
previously resided in Stachydeoma. A key to the two species of Rhododon is
provided, along with complete synonymies and maps showing the distribution
of each.

KEY WORDS: Lamiaceae, Rhododon, Stachydeoma, Hedeoma, Texas

Rhododon was established as _a monotypic genus by Epling in 1939. The type’
species, R. ciliatus (Benth.) Epling, was originally described by Bentham as Keithia
ciliata Benth., based upon material collected in Texas by Berlandier in 1828. Bentham
(Nov. 1848) subsequently transferred the species to Hedeoma where it was retained as
A. ciliata (Benth.) Benth. ex DC., in spite of the existence of H. ciliata Nutt. (Apr.
1848), which occasioned the replacement name, H. texana Cory, in 1936. Earlier,
however, Small in 1903 had transferred Keithia ciliata into his newly erected
Stachydeoma, which apparently contained two disparate elements, neither of which
was selected by Small to typify his genus. Epling (1939) corrected this oversight with
the erection of Rhododon, typifying Stachydeoma with S. graveolens (Chapm.)
Small, a Floridean endemic.

Tharp (1945), when undertaking his study of Stachydeoma for Texas, was}
unaware of Epling's Rhododon and maintained three taxa within Stachydeoma, S.}
ciliata and two newly described species, S. angulata Tharp and S. duvalii Tharp.}
Subsequent workers have more or less ignored Rhododon, although Irving (19860) |
excluded R. ciliata from his concept of Hedeoma, in effect accepting Epling's
Rhododon, but treating this as_ monotypic, in spite of Tharp's two newly described
species, both of which Irving thought synonymous with R. ciliatus.

Turner Study of Rhododon 449

I have gone over the Rhododon problem in some detail, studying specimens of the
genus both in herbaria and in the field. My treatment recognizes Rhododon as distinct
from Stachydeoma, accepting two species, R. ciliatus (including S. duvalii) and R.
angulatus, the former occupying deep sandy soils in east central Texas, the latter
occurring in coastal relic dunes in southern Texas, as shown in Figure 1.

A key to the species follows, along with brief descriptions and’ complete
synonymies.

1. Midstem leaves broadly ovate to ovate-elliptic, mostly 1.5-2.0 times as long as
| wide; calyx lobes merely ciliate, their surfaces essentially glabrous, the marginal
hairs mostly 0.5 mm long or less; flowers arranged in interrupted spikes; coastal
felle Gunes i southernmost Texas. ................--ssessseosenceaassasss- R. angulatus
_1. Midstem leaves elliptic to oblanceolate, mostly 2.0-3.5 times as long as wide; calyx
| lobes pubescent both along their margin and upon their surfaces, the hairs, at least

some or most of them 1-2 mm long; flowers usually arranged in dense spikes,

iaieny insereupted: east-central Texas. ...0........s00-d:scoceennee-desecesacs R. ciliatus

.
|
.

RHODODON ANGULATUS (Tharp) B.L. Turner, comb. nov. BASIONYM:
__ Stachydeoma angulata Tharp, Brittonia 5:304. 1945. TYPE: U.S.A. Texas:
Aransas Co., Rockport, 2 Jul 1939, B.C. Tharp 4399] (HOLOTYPE: TEX!).

Tharp (1945) provided an adequate description of this species, along with a
photograph of the holotype. He also constructed a key to the several taxa recognized,
but emphasized in this several characters which I have not drawn upon.

Rhododon angulatus is a rarely encountered taxon. I visited the type locality in
May of 1994 to ascertain its relative commonness in the area concerned. As indicated
in the specimens cited below, I first collected the species in June of 1964, 5 miles
north of Aransas Pass, growing upon large stabilized sand dunes along the east side of
‘State highway 35. At that time I encountered only two plants in the immediate vicinity,
both growing upon the dunes concerned. My revisit to this site in 1994 was most
disappointing, for all of the prominent dunes along the roadway had been leveled for
highway expansion and commercial development.

I did, however, locate Rhododon angulatus in similar dune sands along the
roadway that circles the Aransas County airport, about 1 km west of the earlier site.
None of the plants was in flower at the time, but I counted several hundred or more
Over an acre or less, nearly all in deep sandy soils among and upon seemingly
stabilized dune sands. Since the relic dunes in this region harbor a number of
interesting plants, especially Rhododon angulatus, some effort should be made by
conservancy agencies to protect at least a few such areas.

ADDITIONAL SPECIMENS EXAMINED: U.S.A. Texas: Aransas Co., “On
sandy bank south of Aransas County Airport,” 8 Jul 1957, Correll & Johnston 17613

(LL); S mi N of Aransas Pass, roadside, growing in live oak mott, 16 Jun 1964,
Turner 5030 (TEX).

volume 78(6):448-45]

June 1995

PHY TOLOGIA

450

| | pe
Bilge eee [
| | a ae
..—- = A
lite f
See e774
re AS
AG ani)
Ne

@ Rhododon ciliatus

@ Rhododon angulatus

per map, R. ciliatus; lower map,

Distribution of Rhododon spp. Up

Fig. 1.
R. angulatus (based upon records at BRIT, LL

>; LEX):

Turner Study of Rhododon 451

RHODODON CILIATIS (Benth.) Epling, Rep. Spec. Nov. Beih. 115:14. 1939.
=Hedeoma ciliata (Benth.) Benth. in DC., Prodr. 12:245. Nov 1848. [not H.
ciliata Nutt.]. = Stachydeoma ciliata (Benth.) Small, Fl. S.E. U.S. 1041. 1903,

1937. =Hedeoma texana Cory, Rhodora 38:405. 1936. TYPE: U.S.A. Texas:

w/o county, 1828, Berlandier s.n. (HOLOTYPE: K).

Stachydeoma duvalii Tharp, Brittonia 5:306. 1945. TYPE: U.S.A. Texas:
Austin Co., 4 mi NW of Bellville, 22 Jun 1923, B.C. Tharp 43992
(HOLOTYPE: TEX!).

This taxon has been adequately described by Tharp (1945). He separated from
this, however, material collected in Austin County having interrupted spikes (the latter
character approaching that of Rhododon angulatus), calling this Stachydeoma duvalii.
Except for the interrupted spikes, nearly all of the characters of the latter are those of
Rhododon ciliatus and | have little hesitancy in treating these as synonymous.

Rhododon ciliatus is linearly widespread and abundant in eastcentral Texas, where
it is largely restricted to the deep white sandy soils of the Carnzo formation (cf,
McBryde 1933), although occasional populations occur elsewhere, as shown in Figure
I.

LITERATURE CITED

Irving, R. 1980. The systematics of Hedeoma (Labiateae). Sida 8:218-304.

Tharp, B. 1945. Noteworthy plants of Texas - III. Stachydeoma as represented in
Texas. Brittonia 5:304-307.

McBryde, J. 1933. The vegetation and habitat factors of the Carrizo sands. Ecol.
Monographs 3:247-297.

Phytologia (June 1995) 78(6):452

INDEX TO AUTHORS, VOLUME 78

Adams, R.P. 134
Anderson, L.C. 246
Affolter, JM. 127
Aruna, V. 357
Babu, G.J. 357
Baird, G.I. 61
Benitez de Rojas, C. 353
Bridges, E.L. 246
Constance, L. 127
Crozier, B. 428
Debreezy, Z. 217
Douglas, P.P. 249
Egger, M. 256
Enquist, M. 428
Gandhi, K.N. 1
Grant, J.R. 119
Hays, J. 264

He, Ping 277, 282
Hurter, P.J.H. 409
Jones, S.D. 244
Kartes7- 320 1
King, R.M. 124, 381
Klackenberg, J. 189
Kumar, K.V. 260
Lafernére, J.E. 214
Landry, P. 287

Li, Hong 277

de la Luz Arreguin-Sanchez, A. 361

452

MacRoberts, B.R.

MacRoberts, M.H.

Martinez, M. 353

Meagher, W.L. 317

Morales, J.F. 192, 195, 197

Naidu, K.C. 357

Nesom: G:L> Gly 13:13

Orzell, S.L. 246

Palacios-Chavez, R. 361

Pushpalatha, V. 357 |

Quiréz-Garcia, D.L. 361

FRACZ ANE 27,

Rao, B.H. 260

Reeder, €.G. 417

Reeder, J.R. 417

Robinson, H. 124, 381, 384

Shaw, R.B. 249

Turner, B.L. 28, 36, 39, 199, 204,
209, 211, 285, 400, 405, 446,
448

Warnock, M.J. 73, 102

Wipff, J.K. 244

18, 291, 402
18, 291, 402

Phytologia (June 1995) 78(6):453-509.

INDEX TO TAXA, VOLUME 78

New taxa described in this volume are indicated by bold face type.

A

Abies 113, 217-220, 222-229, 231,
232, 242
bracteata 218
concolor 217-219, 224, 232
var. lowiana 219, 232
durangensis 218, 224, 225, 228
var. coahuilensis 218
firma 218
grandis 219, 224, 232
guatemalensis 217-219, 222,
226,227, 229, 231
var. longibracteata 218,
29) 227, 229, 231

var. lacanensis 218, 219,

231
hickelii 218, 226, 227
hidalgensis 218-220, 228,
229

453

Abies (cont.)
lowiana 217-219, 224, 231, 232
var. viridula 218, 231,
232
magnifica 218
neodurangensis 218, 219,
223, 228, 229
oaxacana 218
religiosa 222, 231
vejarii 218, 222
var. mexicana 218
zapotekensis 218, 219, 225,
228, 229
Abutilon 328, 331, 347
simulans 328, 331, 347
Acacia’ 237; 326, .328-332;/335,
346, 351, 358
arabica 358
farnesiana 328-332, 346

leucophloea 358

454 PHY TOLOGIA

Acacia (cont.)
schaffneri 326, 328, 330, 332,
335; 346; S51
sundra 358
Acalypha 331
Acanthaceae 21, 268, 336, 343
Acer 233, 247
glabrum 233
rubrum 247
Aceretes 296
angustifolia 296
Adenostemma 125
Adiantaceae 268, 342, 343
Adiantum 241
Adina 359
cordifolia 359
Aegle 359
marmelos 359
Aeonium 1,5, 6, 13, 16
haworthii 1, 5,6, 13
Agalinis 23, 273
fasciculata 23
skinneriana 23
tenuifolia 273

Agarista 247

June 1995 volume 78(6):453-509

Agarista (cont.)
populifolia 247
Agave 34, 165, 203, 337, 350
filifera 350
lechugilla 203
Agavaceae 2]
Ageratella 204-208
microphylla 204-208
var. microphylla 204-208
var. palmeri 204-207
var. seemannil 204, 205
var. sonorana 204-206,
207
palmeri 206
Ageratina 131
Ageratum 204, 205, 328, 332, 333,
343
corymbosum 328, 332, 333,
343
microphyllum 204, 205
Agrostis 10, 11, 244, 274
aspera 10, 11, 244
clandestina 244
composita 10, 11, 244

elliottiana 274

|

Agrostis (cont.)
hyemalis 274
var. hyemalis 274
perennans 274
Albizzia 358
lebbeck 358
odoratissima 358
Alcantarea 119, 120, 122, 123
brasiliana 120
duarteana 120
extensa 120
farneyi 120
geniculata 120
imperialis 120
nahoumii 120
odorata 120
regina 120

vinicolor 120

Aletris 402

aurea 402
Alismataceae 123
Allium 274
canadense 274
var. canadense 274

var. mobilense 274

Index to taxa volume 78

Allomiinae 124, 125
Alnus 223, 225, 227
firmifolia 223, 225
glabrata 227
Alomiinae 381
Alopecurus 265, 274
carolinianus 265, 274
Alstonia 192-194
longifolia 192-194
macrantha 192, 193
pittiert 192, 194
Amaranthaceae 341, 343
Amaranthus 330, 331, 341, 343
hybridus 330, 331, 341, 343
Ambrosia 21, 269, 332, 343
artemisiifolia 21, 269
bidentata 269
cordifolia 332, 343
Amsonia 295,311
ludoviciana 295,311
Anacardiaceae 21, 269, 343, 357
Anacardium 357
occidentale 357
Andropogon 22, 274, 350

gerardii 22,274

456 PHY TOLOGTA

Andropogon gerardii (cont.)
var. gerardii 274
glomeratus 22
tenarius 22
virginicus 22
Anemone 23, 428-441, 443, 444,
445
berlandieri 428, 430, 434, 438,
443
caroliniana 23, 428, 430, 434,
443
edwardsiana 428-430, 433,
435, 437, 438, 444
sect. Eriocephalus 445
multifida 441
Okennonii 428-430, 438, 439,
441, 444

sphenophylla 439-44]

tuberosa 428-432, 435-441, 444

var. texana 428, 435, 436,
441
var. tuberosa 435, 440, 441
Angelica 127
Angiospermae 343, 380

Anisacanthus 328, 334, 343

June 1995 volume 78(6):453-509

Antsacanthus (cont.)
quadrifidus 328, 334, 343
Annonaceae 358
Anoda 335, 347
cristata 335, 347
Anogeissus 358
acuminata 358
latifolia 358
Antennaria 269
plantaginifolia 269
Aphanactis 131
Apiaceae 21, 127
Apioideae 127
Apocyanaceae 21, 192, 195, 197,
311,336, 343; 358
Apocynoideae 197
Apocynales 191
Apocynum 21
cannabinum 21
Aquifoliaceae 21
Arbutus 227, 233, 235, 241
arizonica 235
glandulosa 233,235
xalapensis 227, 241

Archibaccharis 131

_.. aa

Index to taxa volume 78 457

Arecaceae 358 Asclepias (cont.)
Arenaria 270 viridis 21
serpyllifolia 270 Aspidium 1, 6,7
Argemone 342, 348 alpestre 1,6,7
ochroleuca 342, 348 Asplenium 268
Aristida 22, 268, 274 platyneuron 268
dichotoma 274 var. platyneuron 268
var. dichotoma 274 Aster 21, 265, 269, 275, 331, 343
longespica 274 dumosus 21
var. longespica 274 linariifolius 269
oligantha 22 forma linariifolius 269
purpurascens 274 var. linariifolius 269
Artemisia 65 oolentangiensis 21
Artocarpus 359 patens 21
integrifolia 359 pilosus 269
Asanthus 125, 126 sericeus 21,269
Asclepiadaceae 21, 189, 191, 343 var. microphyllus 21
Asclepiadeae 190 forma sericeus 269
Asclepias 21, 296, 308, 329, 330, subulatus 343
343 Asteraceae 1, 2, 21, 26, 36, 39, 61,
hirtella 308 65, 124-126, 153, 187, 188,
linaria 329, 330, 343 204, 208, 211, 246, 248, 267,
longifolia 308 269, 3105, 313; 318,320, 336;
stenophylla 296 337, 340-345, 352, 381, 383,

viridiflora 21, 296 384, 399-401

458 PHY TOLOGIA

Astereae 61, 63, 65, 341
Astragalus 296
crassicarpus 296
var. trichocalyx 296
Alasites 187
Athenaea 354
peruviana 354
Athyrium 1, 6,7
alpestre 1,6,7
distentifolium 6
felix-femina 6
Avicenniaceae 72, 152, 316, 404
Ayapana 125
Azadirachta 358

indica 358

B

Baccharis 328, 331, 334, 335, 343
multiflora 328, 343
salicifolia 328, 331, 334, 335,
343
Bacopa 340, 349
procumbens 340, 349
Bahia 343

schaffneri 343

June 1995 volume 78(6):453-509

Bambusa_ 350
Baptisia 270
alba 270
Barroetea 124-126
brevipes 124
glutinosa 124, 126
laxiflora 125
setosa 125
sonorana 125
subuligera 125
Bartonia 312
texana 312
Bassia 359
latifolia 359
Bauhinia 358
purpurea 358
racemosa 358
Berberidaceae 214-216
Berberis 214, 215, 387
chochoco 214, 215
tinctoria 215
Berchemia 23,24
scandens 23,24
Bidens 331, 341, 343, 344

angustissima 343

Index to taxa volume 78 459

Bidens (cont.)

ferulifolia 343

odorata 341, 343

pilosa 341, 344
Bignoniaceae 336, 345, 358
Boehmeria 247

cylindrica 247
Boerhavia 348

coccinea 348
Bombacaceae 358
Bombax 358

malabaricum 358
Boraginaceae 39, 58, 60, 260
Boraginoideae 60
Borassus 358

flabellifer 358

Borreria 349, 361-364, 368, 369,

378, 379
laevis 361-364, 368, 369
verticillata 349, 361-364, 368,
369
Boswellia 358
Serrata 358
Botrychium 1,9

pumicola 1,9

Bouchea 331, 342, 349
prismatica 331, 342, 349
var. brevirostra 342, 349
Bouteloua 350
curtipendula 350
gracilis 350
repens 350
Bouvardia 328, 330, 333, 349,
361-365, 368, 370, 371, 378,
379
subgen. Bouvardia 379
subgen. Bouvardiastrum 379
subgen. Bouvardioides 379
longiflora 361, 362, 364, 368,
370; 379
multiflora 328, 349, 361, 362,
36413685370; 379
obovata 361-364, 368, 379
terniflora 361, 362
lernifolia 328, 333, 349, 365,
368, 371, 379
Brachiaria 341, 350
meziana 341, 350
Brachyactis 275
Brassicaceae 1, 3, 254, 269, 341,

342, 345

460 PHY TOLOGIA

Braya 1,3, 14
glabella 1,3
var. glabella 3
var. purpurescens 3
purpurascens 1,3
Brickellia 124-126, 328-330, 332-
334, 344, 381
subgen. Phanerostylis 124, 126
problematica 124
secundiflora 328, 334, 344
sonorana 124-126
veroniciflora 329, 330, 332,
333, 344
Brickelliastrum 124-126
fendleri 125
nesomii 125
villarrealii 125, 126
Bridelia 358
retusa 358
Bromeliaceae 119, 120, 123, 350
Bromus 331, 350
Buchanania 357
lanzan 357
Buddleia 223, 328, 331, 345

cordata 223, 328, 331, 345

June 1995 volume 78(6):453-S509

Buddeleiaceae 336, 345
Bulbostylis 273

capillaris 273
Bumelia 272

lanuginosa 272
Burmannia 296

biflora 296
Bursera 328, 333, 345

fagaroides 328, 333, 345

var. fagaroides 328, 333,
345

Burseraceae 336, 345, 358
Butea 358

frandosa 358

Cc

Cacalia 21, 169, 246
plantaginea 21
spathulata 169
suaveolens 246

Cactaceae 269, 337, 345

Caesalpinaceae 269, 358

Caesalpinoideae 340

Calliandra 328, 346

grandiflora 328, 346

Index to taxa volume 78 461

Callicarpa 23 Cardiospermum 331, 349
americana 23 halicacabum 331, 349
Callitrichaceae 269 Carea 358
Callitriche 269 arborea 358
heterophylla 269 Garex 21) 247. 249-252) 254.255:
var. heterophylla 269 2S 2972313
Calopogon 296, 306 bushii_ 273
barbatus 296, 306 chapmanii 247
pallidus 296 cherokeensis 21
Camassia 274, 296 flaccosperma 21, 273
scilloides 274, 296 var. glaucodea 273
forma scilloides 274 hyalina 297, 313
Campanulaceae 21, 336, 346 inversa 249-252, 254
Canthium 359 sect. Inversa 249
dicoccum 359 leptalea 247
Capparaceae 336, 346 meadii 21, 297
Caprifoliaceae 21, 269 microdonta 297
Cardamine 1, 3-5, 16, 269 tenax 297
californica 1, 3-5 subgen. Vignea 249
var. californica 4 wahuensis 250
var. integrifolia 4,5 subsp. rubiginosa 250
concatenata 269 subsp. wahuensis 250
integrifolia 1, 3-5 Carpinus 247, 406
parviflora 269 caroliniana 247

var. arenicola 269 Carya 271, 328, 347, 406

462 PHY TOLOGITA

Carya (cont.)
illinoinensis 328, 347
texana 271
Caryophyllaceae 199, 270, 346,
446
Caryota 358
urens 358
Casearia 358
elliptica 358
Cassia 22,358
fasciculata 22
fistula 358
siamea 358
Castilleja 70, 131-133, 256-259,
286, 329, 330, 333, 349
altorum 258
sect. Castilleja 131, 133, 256,
258, 259
subgen. Castilleja 258, 259
conzattii 132, 133
clenodonta 256, 258
sect. Euchroma 131, 133, 258
filiflora 258
integrifolia 131

macrostigma 258

June 1995 volume 78(6):453-S09

Castilleja (cont.)
nervata 131, 132
nivibractea 131, 132, 133
ornata 258
pallescens 70
var. inverta 70
parviflora 70
var. albida 70
var. oreopola 70
quiexobrensis 131, 132
scorzonerifolia 131-133
spiranthoides 256-258
tenuiflora 133, 259, 329, 330,
333, 349
tolucensis 131, 133
zempoaltepetlensis 131, 133
Castillejinae 259
Ceanothus 1, 11, 23, 297
americanus 23
greggi 1,11
subsp. franklinii 1, 11
var. franklinii 11
herbacea 297
Ceiba 358

pentandra 358

Celtis 273
tenuifolia 273
var. tenuifolia 273
Centrosema 22
virginianum 22
Cephalanthus 272
occidentalis 272
Cerastium 270
brachypetalum 270
Cercis 269
canadensis 269
Cestrum 223, 233
fasciculatum 223
nocturnum 233
purpureum 223 ~
Chamaelirium 297
luteum 297
Chamaesyce 22
cordifolia 22
Chaptalia 153-188
albicans 154, 156, 157, 167,
174, 185, 188
sect. Archichaptalia 155

carduacea 163

Index to taxa volume 78 463

Chaptalia (cont.)

crispula 167

dentata 161, 188

diversifolia 163

erosa 163

estribensis 153, 155-157,
160, 161, 171, 178

exscapa 154

fallax 167

graminifolia 168

hintonti 155-157, 161, 171, 179

hololeuca 156, 157, 164, 166,
174, 184

integrifolia 157, 187

leiocarpa 167

leonina 164

sect. Leria 154-156

sect. Lieberkuhnia 154-156

sect. Loxodon 154, 156

lyrata 153, 169, 187

lyratifolia 155-157, 159-161,
he a

madrensis 153, 155, 157,

158,159,166; 170, 176

sect. Chaptalia 153, 155, 159, majuscula 163

169

464

Chaptalia (cont.)

sect. Microchaptalia 155, 156
nutans 153, 154, 156, 157,
161, 162, 164-167, 169,
173, 181, 187
var. leiocarpa 167
var. fexana 163, 164

obovata 167

petrophila 153, 154, 164, 165,

169, 182, 187
piloselloides 154
pringlei 153, 161, 171, 180
sect. Pseudotrichocline 155
runcinata 154, 156, 167, 168,

172, 186

var. graminifolia 168

var. runcinata 168
semifloscularis 158
spathulata 153, 154, 165, 169,

173

subcordata 163

texana S45 156) 157 159 Nel;

163-166, 169, 173, 182
tomentosa 153, 155, 157-159,
7, UO, WAS

PHY TOLCGIA June 1995

Chaptalia (cont.)

volume 78(6):453-S09

transiliens 156, 157, 165, 166,

172, 183
Chasmanthium 22

sessiliflorum 22

Cheilanthes 268, 297, 333,

342, 343
beitelii 342
bonariensis 342
incana 342
lanosa 268, 297
lendigera 342
myriophylla 333, 343
sinuata 343
Chiranthodendron 231
pentadactyon 231
Chloraxylon 359
swietenia 359
Chloris 313, 341, 350
subdolichostachya 313
virgata 341, 350
Chresta 384
pinnatifida 384
Chrysodendron 214, 215

tinctorium 214, 215

338,

Index to taxa volume 78 465

Chrysodendron tinctorium (cont.) Citharexylum 328, 330, 349

var. latifolium 214, 215 lycioides 328, 330, 349
var. longifolium 214, 215 Cleistanthus 358

var. oblongifolium 214, 215 collinus 358

Chrysothamnus 61-65
albidus 63
consimilis 64
eremobius 63, 65
sect. Graminei 63
gramineus 63
sect. Nauseosi 61, 62, 64
nauseosus 64
var. artus 64
var. oreophilus 64
oreophilus 64
var. artus 64
var. oreophilus 64
sect. Punctati 61, 62, 64
spathulata 63
vaseyi 63
Cinchoneae 378
Cinchoninae 378
Cinchonoideae
Cirsium 247

muticum 247

patulus 358
Clethra 227
mexicana 227
Clusia 387
Clusiaceae 21, 270
Coffeoideae 378
Comarostaphylis 227, 241
arguta 227,241
conzattii 227, 241
Combretaceae 358
Commelina 313, 350
benghalensis 313
dianthifolia 350

erecta 350

Commelinaceae 273, 311, 337, 350

Compositae 38, 65, 124, 187, 188,

ZIT. 599
Condylopodium 381-383

fuliginosum 381

killipti_ 381, 382

Coniferae 217

466 PEPY OwOiGivA June 1995 volume 78(6):453-509

Convolvulaceae 342, 346 Cosmos 344

Conyza 334, 341, 344, 385 bipinnatus 344
divaricata 385 Crassula 241, 249, 254
sophiifolia 334, 341, 344 sieberiana 249, 254

Cordia 260-263 Crassulaceae 1, 5, 16, 254 337;
alba 260-263 346, 405, 408
dichotoma 260-263 Crataegus 19, 23, 24
domestica 260-263 crus-galli 23
evolutior 260-263 marshallii_ 23
macleodii 260-263 spathulataa 23
monoica 260-263 Critoniopsis 384, 386-390
sebestena 260-263 choquetangensis 386, 387
wallichii 260-263 gynoxiifolia 387

Coreopsis 21 jJaramilloi 388
lanceolata 21 lindenii 390

Comaceae 21, 270 oolepis 389

Cornus NON Zl 245 225) 227,233, peruviana 387
247, 270, 301, 406 thomasii 388
disciflora 225, 227 zarucchii 389, 390
drummondii 19,21, 24 Crossothamnus 381-383
florida 21, 270, 301 gentryi 381, 382, 383
foemina 247 killipiti 381, 382, 383
sericea 233 pascoanus 381-383

Coryphantha 339, 345 weberbaueri 381, 383

elephantidens 345 Crotalaria 341, 346

Index to taxa volume 78 467

Croton (cont.)
pumila 341, 346
Croton 270, 328, 346
adspersus 328, 346
capitatus 270
var. capitatus 270
Crotonopsis 267, 270
elliptica 267, 270
grandiflora 267

Cruciferae 14, 16

Crusea 227, 361-363, 365, 368,

371-373, 378, 379
coccinea 227, 361-363, 365,
368, 371

diversifolia 361, 362, 365, 368,

BIZ
longiflora 361, 362, 365, 368,
Sie; 3715
Cucurbitaceae 336, 346
Cuphea 271, 332, 347
viscosissima 271

wrightii 332, 347

Cupressaceae 21, 134, 149, 150,

268

Cupressus 222, 231, 240

Cupressus (cont.)
arizonica 240
lusitanica 222,231

Cyclanthera 335, 346
dissecta 335, 346

Cyperaceae 21, 123, 249, 255, 273,
310, 350

Cyperus’ 265;,'273, 298, 310; 334;
335,338, 350
acuminatus 265, 273
aristatus 273
calderoniae 350
esculentus 335, 350
grayioides 298, 310
niger 334, 335, 350
retroflexus 310
spectabilis 350
virens 334, 335, 350

Cypripedium 298
kentuckiense 298

Cystopteris 268

lennesseensis 268

D

Dahlia 231

468 PHY TOLOGITA June 1995 volume 78(6):453-509

Dahlia (cont.) Delonix (cont.)
excelsa 231 regia 358
Dalbergia 358 Delphinastrum 95, 99
latifolia 358 leucophaeum 99
Dalea 22, 328-330, 340, 346, 347 menziesii 95
bicolor 328, 330, 346 Delphinium 23, 73-118

var. bicolor 346
candida 22
lutea 329, 347
prostrata 340, 347
purpurea 22
Danthonia 250, 254, 274
gracilis 250
pilosa 254
spicata 274
var. spicata 274
unarede 250
Dasyanthina 397
Dasylirion 34, 241
lucidum 241
Datura 341, 349
stramonium 341, 349
Decachaeta 205
seemannit 205

Delonix 358

alabamicum 93, 102
alpestre 79
andersonii 83, 103
andesicola 79, 80, 103
antoninum 88
azureum 105
bakeri 93
barbeyi 79, 103
basalticum 88, 91
bicolor 87-91, 112
subsp. bicolor 89-91
subsp. calcicola 89,90,
91
forma devriesti 89
forma helleri 112
forma montanense 89
var. montanense 89, 90
var. nuttallii 98

sect. Bicoloria 87

Index to taxa volume 78

Delphinium (cont.)

species group Bicoloria 87
subsect. Bicoloria 75, 77, 87,

88, 92 .
bicornutum 80, 103

subsp. bicornutum 103

subsp. oaxacanum 103
brachycentrum 79, 104
calcar-equitis 80, 104
californicum 79, 80, 104

subsp. californicum 104
cardinale 80,81, 104, 118
species group Caroliniana 85
subsect. Caroliniana 85
carolinianum 23, 85, 86, 104,

106, 107, 118

subsp. carolinianum 104

subsp. penardii 108

subsp. vimineum 85, 106,

107

subsp. virescens 108
chilliwacense 95
columbianum 98
cyanoreios 108, 112

forma multiplex 112

469

Delphinium (cont.)

decorum 88, 93, 108, 110
subsp. fracyi 108

sect. Delphinastrum 73

subgen. Delphinium 73

subsect. Depauperata 76, 84

depauperatum 8A, 85, 88, 108

distichum 84, 108

subsect. Echinata 77, 85-87

series Echinatae 86, 100

subsect. Elata 73, 76, 78

elatum 78, 104, 109

subsect. Exaltata 76, 77-81

exaltatum 78,79, 98, 109
nuttallii 98

flexuosum 116

geraniifolium 81, 109

geyeri 78, 81,91, 109, 110

glareosum 88, 90,91

glaucescens 80

glaucum 79, 80, 110

gracilentum 94, 113

sect. Grumosa 92

subsect. Grumosa 75, 77, 88,

92

470 PHY TOLO@GTA

Delphinium (cont.)

gypsophilum 83, 110, 118
subsp. gypsophilum 110
subsp. parviflorum 110

hansenii 86, 87, 110, 111
subsp. ewanianum 110
subsp. hansenii 111
subsp. kernense 111

hesperium 87, 111
subsp. cuyamacae 111
var. cuyamacae 111
subsp. hesperium 111
forma pallescens 111
subsp. pallescens 87, 111

hutchinsoniae 87

inopinum 82, 113

leucophaeum 99

lineapetalum 83

luteum 88, 111, 117

madrense 86

menziesii 92-98, 111, 112
var. levicaule 112
subsp. menziesii 95, 96,

111

var. menziesii 111

June 1995 volume 78(6):453-S09

Delphinium menziesii (cont.)
var. ochroleucum 98
subsp. pallidum 95, 96,
1 |
var. pavonaceum 111
subsp. pyramidalis 95
var. pyramidale 95
multiplex 81, 82, 112
subsect. Multiplex 76, 78, 81,
82, 84
nelsonii 112
newtonianum 93
novomexicanum 79, 80, 112
nudicaule 88, 93, 112
nuttallianum 89, 93, 94, 97,
2S
var. pilosa 89
nuttallii 92, 93, 95-99, 112
var. leucophaeum 99
subsp. nuttallii 98, 99, 112
subsp. ochroleucum 98,
99, 112
var. ochroleucum 112
x occidentale 79, 110

oreganum 95

Index to taxa volume 78 471

Delphinium (cont.)

parishii 82, 83, 113
subsp. pallidum 113
subsp. parishii 113

parryi 82,83, 113
subsp. blochmaniae 113
var. blochmanae 113
subsp. maritimum 113
subsp. parryi 113
subsp. purpureum 113

patens 88, 93, 94, 113
subsp. hepaticoideum 113
subsp. patens 113

pauperculum 95

x pavonaceum 92, 96

pedatisectum 80, 113

series Pelligerae 100

penardii 85, 108

polycladon 82, 114

purpusti 82, 83, 114

ramosum 79,80, 114

recurvatum 83, 114

robustum 79, 80

sapellonis 79

Scaposum 82, 83, 114

Delphinium (cont.)

scopulorum 80,81, 115
stachydeum 81
subscandens 80, 115
subsect. Subscaposa 75, 77, 78,
82, 83, 100
“tribus” Subscaposa 82
sutherlandii 93, 94,97, 115
tenuisectum 80
treleasei 94, 115
tricorne 93-95, 116
subsp. menziesii 95
var. menziesii 95S
trolliifolium 88, 91-93, 96, 116
tuberosum 95
uliginosum 84
umbraculorum 83, 116
valens 79
variegatum 87, 116
subsp. variegatum 116
virescens 85, 108
subsect. Virescens 75, 77, 84,
85, 86
viride 80

viridescens 82

472 PHY TOLOGTA

Delphinium (cont.)
willametense 99
subsect. Wislizenana 75, 78,
80, 81
wislizeni 80, 116, 117
wootonit 86, 117
xantholeucum 83, 117
Dentaria 1, 3-5, 14
californica 3-5
var. integrifolia 4,5
integrifolia 3-5
var. californica 4,5
Deprea 354
Desmanthus 22
illinoensis 22
Desmodium 22, 328, 330-332
Deutzia 277-283
subsect. Cymosae 277
sect. Deutzia 277
discolor 282, 283
var. bicruristyli 282, 283
var. discolor 282
multiradiata 277-281
series Multiradiatae 277

Dichanthelium 22

June 1995 volume 78(6):453-S509

Dichanthelium (cont.)
aciculare 22
acuminatum 22
dichotomum 22
Dichelachne 250
crinita 250
Dichondra 330, 333, 334, 346
argentea 330, 333, 334, 346
Dicliptera 343
peduncularis 343
Dicotyledonae 336, 343
Dicrastylidaceae 72, 172, 316, 404
Didymaea 361, 362, 365, 366, 368,
373, 378
alsinoides 361, 362, 365, 368,
373
floribunda 361, 362, 366, 368,
373
Digitaria 274
cognata 274
var. cognata 274
Diodia 267, 272
teres 267, 272
Diospyros 19, 21, 24, 270, 358

melanoxylon 358

Index to taxa volume 78 473

Diospyros (cont.)
montana 358
sylvatica 358
virginiana 19, 21, 24, 270
forma platycarpa 270
var. platycarpa 270
Dipladenia 197
boliviensis 197
Dipterocarpaceae 358
Dirca 406
Dodecatheon 298
meadia 298
Dodonaea 249, 254
Draba 269, 298
brachycarpa 269
cuneifolia 298
Drosera 402
capillaris 402
Drymaria 199-203, 330, 333, 338,
346
arenarioides 346
ser. Arenarioides 199, 201, 203
axillaris 199, 201
barkleyi 203

coahuilana 199, 200-202

Drymaria (cont.)
elata 201
laxiflora 333, 346
lyropetala 199-202
var. coahuilana 199, 200
var. lyropetala 199-201
ser. Lyropetala 199, 201
pattersonii 199, 201, 203
pratheri 199, 200-202
subumbellata 201
suffruticosa 201
xerophylla 330, 346
Dryopteridaceae 1, 6, 268
Dryopteris 247, 268
ludoviciana 247
marginalis 268
Dyschoriste 343
Dyscritogyne 125, 126
Dyssodia 213, 344
papposa 344
porophylla 344

var. cancellata 344

E

Ebenaceae 21, 270, 358

474 PHY TOLOGTA

Echeandia 350
mexicana 350
Echinocactus 318, 345
grusonii 318, 345
Echinochloa 341, 350
Ehrharta 254
stipoides 254
Elaphoglossum 241
Eleocharis 335, 350
macrostachya 335, 350
montevidensis 335, 350
Encephalartos 409-416
cupidus 409, 410
hildebrandtii 410, 411
nubimontanus 409, 410,

412, 413

whitelockii 410, 411, 414,

415
Eragrostideae 425
Eragrostis 22, 254

brownei 254

spectabilis 22
Eremanthus 384, 390

hatschbachii 390

pinnatifidus 384

June 1995 volume 78(6):453-S09

Eremanthus (cont.)
rondoniensis 390
Ericaceae 270
Ericameria 61-65
sect. Asiris 62
crispa 64
sect. Ericameria 61, 62
lignumviridis 61, 64
sect. Macronema 61, 62, 64
nauseosa 61-64
var. arta 64
var. mohavensis 63
var. oreophila 61, 64
paniculata 61, 62
parryi 61
leretifolia 61, 62
Erigeron 21, 131, 233
strigosus 21
tenuis 21
Eniocaulaceae 72, 172, 311, 316,
404
Eriocaulon 298, 398, 402
decangulare 402
lexense 298

Eriochloa 335, 350

Eriochloa (cont.)
acuminata 335, 350
Eriogonum 299
longifolium 299
multiflorum 299
Erodium 335, 347
cicutarium 335, 347
| Eruca 341, 345
sativa 341, 345
Eryngium 21
yuccifolium 21
Erythrina 328
coralloides 328
Erythronium 299
rostratum 299
Eulophia 304
ecristata 304
Eucalyptus 250
Eulophia 312
cristata 312
Enterolobium 358
samman 358
Eucalyptus 359
glabulus 359
Eupatorieae 124-126, 204,
341, 381, 383

Index to taxa volume 78 475

340,

Eupatorium 21, 124, 125, 328,
329, 333, 334, 338, 344
collinum 328, 344
espinosarum 329, 333, 344

var. espinosarum 344
pycnocephalum 328, 334, 344
serotinum 21

Euphorbia 22, 233, 270, 299, 330-
332, 335, 338, 340, 341, 346
bicolor 299
corollata 22,270
dentata 332, 346
graminea 335, 346
heterophylla 331, 335, 346
indivisa 340, 341, 346
maculata 346

Euphorbiaceae 22, 270, 341, 346,
358

Eutetras 329, 333, 344
pringlei 329, 333, 344

Evolvulus 299, 342, 346
alsinoides 342, 346
sericeus 299

Eysenhardtia 328, 347

polystachya 328, 347

476 Pipe MOO GIA June 1995 volume 78(6):453-509

F Flyriella 124
Fabaceae 71720) 2700 337) Forestiera 328, 330, 331, 348
340-342. 346. 347. 358 durangensis 328, 330, 331, 348

Fagaceae 270 Forsteronia 195, 196

Ferocactus 320, 332, 333, 339, floribunda 196
345. 438 monteverdensis 195, 196
histrix 333, 345 peninsularis 196
ia 200. 2
latispinus 332, 345 Fouqueria 200, 203

setispinus 438 splendens 203

Feronia 359 Fragaria 233
elephantum 359 Fraxinus 271, 328, 333, 348
Festuca 250 americana 271

‘o}
novae-zealandiae 250 uhdei 328, 333, 348

Ficus 359

bengalensis 359

religiosa 359 G
Fimbristylis 273 Galactia 22, 347

autumnalis 273 brachystachys 347
Flacourtiaceae 358 volubilis 22
Flaveria 400, 401 Galieae 378

angustifolia 400, 401 Galinsoga 332, 341, 344

kochiana 400, 401 parviflora 332, 341, 344
Flaveriinae 401 Galium 23> 233, 3315 33353425
Florestina 331,341, 344 361-363, 366-368, 374, 375,

pedata 331, 341, 344 378, 379

Index to taxa volume 78

Galium (cont.)
aschenbornii 361-363, 365,
) 368, 374, 378
mexicanum 331, 333, 349, 361,
362, 366, 368, 374
subsp. mexicanum 361,
362, 366, 374
pilosum 23
praetermisum 363
Seatonii 361-363, 366, 368, 374
sphagnophilum 361-363, 366,
368, 375
var. mazocarpum 361, 362,
366
trifidum 361, 362, 365, 368,
375
var. pacificum 361, 362,
3655375
uncinulatum 361, 362, 367,
368, 375
Gardenia 359
latifolia 359
resinifera 359
Garrya 225
laurifolia 225

Garuga 358
pinnata 358
Gaudichaudia 347
cynanchoides 347
Gaultheria 227, 241
acuminata 227
hirtiflora 227, 241
Gaura 22, 334, 348
coccinea 334, 348
Gelsemium 22
sempervirens 22
Gentianaceae 22
Geraniaceae 22, 336, 347
Geranium 22

carolinianum 22

477

Gerbera 153,155, 157, 158, 162.

167-169, 188
albicans 167
bicolor 168
leiocarpa 167
lyrata 169
nutans 162
spathulata 169
tomentosa 157

walteri 157

478 PHY TLOLOGIA

Glandularia 23
canadensis 23
Gnaphaliothamnus 131
Gnaphalium 21, 233, 331, 333, 344
arizonicum 344
obtusifolium 21
Gomphrena 330, 332, 339, 341,
343
decumbens 330, 332, 339, 341,
343
Gonolobus 343
uniflorus 343
Gramineae 16, 245, 417, 425, 426
Gratiola 265, 273
neglecta 265, 273
Grewia 359
disperina 359
flavescens 359
tiliaefolia 359
Grosvenoria 125
Guilleminea 340, 343
densa 340, 343
Gymnospermae 242, 380, 416
Gymnosperma 344

glutinosum 344

June 1995 volume 78(6):453-S09

H

Habenaria 311
integra 311
Halimodendron 1,7
halodendron \,7
Haplopappus 62-65
sect. Asiris 65
lignumviridis 64
sect. Macronema 65
Hardwickia 358
binata 358
Hasteola 246-248
robertiorum 246-248
suaveolens 246, 248
Hastingia 312
Hedeoma 448, 451
ciliata 448, 451
texana 448, 451
Hedyotideae 378
Hedyotis 23, 272, 300, 361-363,
367, 368,375; 378
cervanlesti 361-363, 367, 368,
395
crassifolia 272

nigricans 23

Index to taxa volume 78 479

Hedyotis (cont.)
purpurea 300
Heimia 347
salicifolia 347
Helenieae 400
Helenium 335, 344

mexicanum 335, 344

Heliantheae 38, 208, 310, 313,

340, 341
Helianthus 21

hirsutus 21
Heliopsis 269, 344

annua 344

helianthoides 269

var. occidentalis 269

Heliotropium 299

tenellum 299
Hesperodoria 62, 63
Heteropogon 350

contortus 350
Heterosperma 330, 332, 341, 344

pinnatum 330, 332, 341, 344
Heterotheca 344

inuloides 344

Heuchera 241, 265, 273

Heuchera (cont.)

x hirsuticaulis 265, 273
Hexalectris 299

spicata 299
Hieracium 131, 161, 169, 187, 269

gronovii 269

pusillum 161

Stipatum 169, 187
Hinterhuberineae 62
Holarrhena 358

antidysenterica 358
Hololepis 384, 390, 391

hatschbachii 390

pedunculata 391
Holopetalea 359

integrifolia 359
Houstonia 300

purpurea 300
Huperzia 1,8

occidentalis 1,8
Hydrangea 225

seemannii 225
Hydrangeaceae 277, 280-282
Hydrodyssodia 211, 213

Stevensit 213

480 PHY T OLO@OGIA

Hydropectis 211-213
aquatica 211-213
estradii 211-213
stevensii 211-213

Hymenoxys 1, 2
acaulis 2

var. epunctata 2
var. nana 2

Hypericum 21, 267, 270
gentianoides 267, 270
hypericoides 21
punctatum 270

Hypoxis 274
hirsuta 274

forma vilosissima 274

|

Ilex 21, 24
decidua 21, 24
vomitoria 21

Indigofera 347
miniata 347

Inuleae 341

Iochroma 354

suffruticosa 354

June 1995 volume 78(6):453-S09

Ipomoea 330, 331, 335, 340, 346
longifolia 330, 340, 346
painteri 335, 346
pubescens 330, 331, 346
purpurea 346

Iridaceae 22, 273, 350

Tresine 328, 343

cassiniaeformis 328, 343

J

Jatropha 329, 330, 332, 346
dioica 329, 330, 332, 346
Juglandaceae 271, 336, 347
Juncaceae 274
Juncus 274
interior 274
Juniperus 21, 24, 102, 115, 117,
134-139, 141-150, 233, 267,
268
australis 145
barbadensis 134-136, 138, 139,
141, 144, 145, 147-149
var. barbadensis 142, 144
var. lucayana 134, 142,

144, 145, 147, 148

Index to taxa volume 78 481

Juniperus barbadensis (cont.)

var. urbaniana 144
bermudiana 134-136, 138, 139,
141-143, 147-149
blancoi 146
sect. Caryocedrus 134
chinensis 142
comitana 146, 149
communis 146
davurica 142
deppeana 233
var. robusta 233
ekmanii 134-139, 141, 144
gamboana 146, 149
gracilior 134-139, 142-144,
147, 148
var. ekmanii 134, 142,
144, 147, 148
var. gracilior 143, 147
var. urbaniana 134, 142,
144, 147, 148
horizontalis 146
sect. Juniperus 134, 146
lucayana 134-136, 138, 139,

141, 145, 148, 149

Juniperus (cont.)

oppositifolia 143
phoenicea 146
sabina 146
sect. Sabina 134, 135, 142,
146, 150
saxicola 134-136, 138, 139,
141, 142, 147-149
scopulorum 146, 149
silicicola 135, 149
squamata 142
standleyi 146, 149
urbaniana 134-139, 141, 144
virginiana 21, 24, 134-136,
138, 139, 141, 143-146,
148-150, 267, 268
var. australis 145
var. barbadensis 144
var. bermudiana 143
var. humilis 145
var. montana 146
var. prostrata 146
var. scopulorum 146
var. silicicola 135, 136,

138, 139, 146, 148

482 PHY TOLOGRA

Juniperus virginiana (cont.)
var. virginiana 136, 139,
268
Justicia 343

furcata 343

K

Keithia 448
ciliata 448
Kigelia 358
pinnata 358
Koeleria 300
macrantha 300
Krigia 269
dandelion 269
virginica 269
Kydia 358

calycina 358

1B
Labiatae 451
Lachnocaulon 300, 311

digynum 300

Lactuceae 341

June 1995 volume 78(6):453-509

Lamiaceae 22, 39, 209, 210, 347,
352, 448
Lamourouxia 330, 349
rhinanthifolia 330, 349
Lannea 357
coramandelica 357
Lantana 328, 331, 349
camara 328, 331, 349
Larnax 353-356
hawkesii_ 354
hunzikeriana 353, 355
peruviana 354, 356
sachapapa 354
steyermarkii 354
subtriflora 354
suffruticosa 354
Larrea 200, 203
tridentata 203
Lasiarrhenum 39, 40, 60
Lauraceae 14, 336, 347
Lecythidaceae 358
Leersia 265, 274
oryzoides 265, 274
virginica 274

Leguminosae 39

Index to taxa volume 78 483

Lieberkuhna 154, 169 Lepidaploa (cont.)
bracteata 154 pseudomuricata 395, 396
spathulata 169 sanmartinensis 396, 397
Leibnitzia 153, 154, 156, 168, 169, Lepidium 249, 254, 341, 345
188 hyssopifolium 249, 254
anandrina 169 virginicum 341, 345
lyrata 153, 169 Leria 154, 162, 167, 169
seemannii 153 albicans 167
Leonotis 333, 334, 347 leiocarpa 167
nepetifolia 333, 334, 347 lyrata 162
Leontodon 167 nutans 154, 162, 167
tomentosum 167 var. leiocarpa 167
Lepidaploa 384, 385, 391-397 spathulata 169
almasensis 385 Lespedeza 22
araujoa 391 Lessingianthus 384, 385, 397
bahiana 392 hatschbachii 397
beckii 394, 395, 397 Leucospora 273
chiriquiensis 385 multifida 273
davidsmithii 392, 393 Leucothoe 247
jJuruenensis 393 racemosa 247
krukovii 394 Liatris 21, 300
lewisii 394, 395 pycnostachya 2)
muricata 396 squarrosa 2)
nitens 394 squarrulosa 21

pseudaurea 385 tenuis 300

484 PHY TOL GGIA

Liliaceae 274, 337, 350
Liliopsida 273
Lilium 300
michauxii 300
Linaceae 27]
Linum 271
medium 271
var. texanm 271
Lipocarpha 273
micrantha 273
Lithospermeae 60
Lithospermum 40, 51, 57, 60, 300
caroliniense 300
molle 51
virginianum 57
Litsea 225, 241
glaucescens 225, 241
Loasaceae 336, 347
Lobelia 21, 346
appendiculata 21, 346
fenestralis 346
Loeselia 328, 330-331, 333, 348
mexicana 328, 330-331, 333,
348

Loganiaceae 22, 358

June 1995 volume 78(6):453-S09

Lonicera 233, 269
flava 269
involucrata 233
pilosa 233
Loranthaceae 336, 347
Loxodon 154, 167
brevipes 154
longipes 167
Ludwigia 271, 334, 335, 340, 348
alternifolia 27)
peploides 334, 335, 340, 348
Lupinus 132, 233
Luzula 233, 274
bulbosa 274
Lycopodiaceae 1, 8, 402, 403
Lycopodiella 300, 402
appressa 402
cernua 300, 402
Lycopodiophyta 317, 336
Lycopodium 8, 300, 313, 402, 403
cernuum 300, 313, 402, 403
lucidulum 8
forma occidentale 8
Lyonia 301

mariana 301

Lythraceae 271, 347
Lythrum 265, 271
alatum 265, 271

M

Machaeranthera 341,344
gymnocephala 344
pinnatifida 341, 344

Macromeria 39, 40, 60

Macroptilium 347
atropurpureum 347
gibbosifolium 347

Magnolia 247
virginiana 247

Magnoliophyta 268, 317

Magnoliopsida 268

Mahonia 214-216, 231, 233

chochoco 214,215
eutriphylla 233
lanceolata 231
repens 233
var. pumila 233
linctoria 214,215
Maianthemum 301

racemosum 301

Index to taxa volume 78 485

Malaxis 301
unifolia 301
Malpighiaceae 336, 347
Malvaceae 340, 341, 347, 348,
351,358
Malvastrum 347
coromandelianum 347
Mammiillaria 320, 338, 339, 345
magnimamma 339, 345
rettigiana 339, 345
uncinata 339, 345
wildii 339, 345
zephyranthoides 339, 345
Mandevilla 197, 343
boliviensis 197
cereola \97
foliosa 343
pittieri 197
Manfreda 21
virginica 21
Mangifera 357
indica 357
Marsilea 334, 335, 343
mollis 334, 335, 343

Marsileaceae 336, 343

486 PESY TOLEOGIA

Martyniaceae 336, 342, 348
Maurandya 349
barclaiana 349
Mayaca 301
aubletii 301
fluviatilis 301
Melampodium 332, 335, 341, 344
glabrum 335, 344
longifolium 332, 344
sericeum 341, 344
Melastomaceae 358
Melia 358
azadirach 358
Meliaceae 358
Melica 422
Melilotus 335, 341, 347
officinalis 335, 341, 347
Memecylon 358
umbellatum 358
Menabea 189-19]
venenata 189-191
Menizelia 330, 331, 347
hirsuta 347
hispida 330, 331, 347

Metrosideros 249, 254

June 1995 volume 78(6):453-S09

Miconia 387
Mikania 247
cordifolia 247
Milla 350
biflora 350
Millingtonia 358
hortensis 358
Mimosa 318, 328-330, 347
aculeaticarpa 318, 328-330, 347
var. biuncifera 318, 328-
330, 347
monancistra 328, 347
Mimosaceae 271, 358
Mimosoideae 340
Mimulus 334, 340, 349
glabratus 334, 340, 349
Mimusops 359
hexandra 359
Minasia 384, 397, 398
splettiae 397
Mirabilis 342, 348
Jalapa 348
nyclaginea 342, 348
Mitragyna 359

parvifolia 359

Index to taxa volume 78 487

Monocotyledonae 336, 350 Muhlenbergia (cont.)
Monotropa 301 mexicana 274
hypopithys 301 michisensis 424, 425
Montanoa 328-331, 333, 344 montana 417, 418, 422-425
leucantha 328, 329, 344 sect. Podosaemum 422
var. arborescens 328, 329, subgen. Podosemum 426
344 quadridentata 417, 426, 427
tomentosa 328, 330, 331, 333, robusta 350
344 straminea 417, 419-427
Moraceae 336, 348, 359 virescens 417-427
Morinda 359 Murraya 359
tinctoria 359 exotica 359
Morus 328, 333, 348 Mutisieae 153, 187, 188, 341
celtidifolia 328, 333, 348 Myoporum 254
Muhlenbergia 22, 244, 274, 350, Myrica 22, 247
417-427 cerifera 22,247
argentea 424 Mynicaceae 22
capillaris 22 Myrtaceae 359
clandestina 244 Myrtillocactus 320, 330-333, 339,
cualensis 424, 425 345
curvula 417, 421, 423, 425 geometrizans 320, 330-333, 345

durangensis 424
sect. Epicampes 422, 426 N

gracilis 418 Nama 131

i is 422
eee Nemastylis 350

488 PHY TOEGGIA

Nemastylis (cont.)
tenuis 350
Neptunia 22
lutea 22
Nicotiana 328, 341, 349
glauca 328, 341, 349
tabacum 349
Nothoscordum 274
bivalve 274
Nuttallanthus 273
canadensis 273
Nyctaginaceae 342, 348
Nyctanthaceae 72, 152, 316, 404
Nyctocereus 320, 339, 345

serpentinus 320, 345

O

Obolaria 301
virginica 301
Ochna 359
squarrosa 359
Ochnaceae 359
Oenothera 271,348
linifolia 271
rosea 348

June 1995 volume 78(6):453-S09

Oleaceae 271, 348
Onagraceae 22, 271, 348
Onosmodium 39-60
alabamense 57
bejariense 39-47, 49, 51-53, 55,
56, 58
var. bejariense 39, 41-44,
46, 47, 49, 52
var. hispidissimum 39-
42, 44, 45-47, 49,
52553
subsp. occidentale 43
var. occidentale 39, 41-
44, 46, 47, 49, 52
subsp. subsetosum 41
var. subsetosum 39, 42,
44, 47, 49, 52, 55
carolinianum 46, 51, 52
var. molle 46
var. mollis 51
dodrantale 39, 40, 47, 50, 53,
56
floridanum 57
helleri 39, 40, 48, 51, 54

hispidum 57

Index to taxa volume 78 489

Onosmodium (cont.)
hispidissimum 43, 45, 52
var. macrospermum 45
longistylum 57
molle 39, 40, 42, 43, 51-53,
55, 58, 60
subsp. bejariense 43
var. bejariense 43, 51
subsp. hispidissimum 45
var. hispidissimum 45
subsp. occidentale 46
var. occidentale 46
subsp. subsetosum 47
var. subsetosum 47, 51
oaxacanum 39, 40, 50, 53
occidentale 43, 46, 52
subsetosum 47
unicum 39, 40, 50, 56
virginianum 39, 40, 46, 52, 57-
59
var. hirsutum 46, 57,58
Ophioglossaceae 1, 9
Ophioglossum 1,9
azoricum 1,9

Opiliaceae 261

Oplismenus 247
setarius 247
Opuntia 200, 269, 320, 330-333,
338, 339, 345, 351
amarilla 345
cochinera 345
durangensis 320, 345
fuliginosa 320, 345
humifusa 269
hyptiacantha 345
imbricata 320, 330-333, 345
incarnadilla 320, 345
joconostle 345
lasiacantha 345
leucotricha 320,345
lindheimeri 345
var. cuija 345
matudae 345
pachona 320, 345
subgen. Platyopuntia 331
robusta 320, 330, 332, 345
var. robusta 320
stenopetala 320, 345
streptacantha 320, 345

tomentosa 320, 345

490 PHY TOEQGIP,

Orchidaceae 1, 9, 14, 274, 311,312
Oreopanax 231
xalapense 231
Orobanche 301
uniflora 301
Oroxylum 358
indicum 358
Osmunda 247, 402
cinnamomea 247
regalis 402
Oxalidaceae 22, 271, 342, 348
Oxalis 22, 271, 330, 331, 342, 348
corniculata 22, 330, 331, 342,
348
decaphylla 348
lunulata 348
violacea 271
Oxydon 154, 167, 168

bicolor 154, 167, 168

P

Palhinaea 402, 403
cernua 402, 403
Panic 2A 2055 274 27>.

302

June 1995 volume 78(6):453-509

Panicum (cont.)
acuminatum 274
var. acuminatum 274
anceps 22
commutatum 247
depauperatum 274
flexile 274, 302
philadelphicum 274
var. philadelphicum 274
virgatum 274
Papaveraceae 336, 342, 348
Papilionoideae 340
Paronychia 270, 302, 446, 447
drummondii 302
fastigiata 270
var. paleacea 270
virginica 446, 447
Parthenium 341, 344
bipinnatifidum 341, 344
Parthenocissus 334, 349
quinquefolia 334, 349
Paspalum 22
Passiflora 22
lutea 22

Passifloraceae 22

Pectis 340, 341, 344
prostrata 340, 341, 344
Pediculareae 259
Pediomelum 302
hypogaeum 302
Pellaea 343
ternifolia 343
Pennellia 341, 345
longifolia 341, 345
Penstemon 23, 132, 273, 302
murrayanus 302
pallidus 273
tubaeflorus 23
Peperomia 334, 348

campylotropa 334, 348

Index to taxa volume 78

Pernettya (cont.)
ciliata 241
Pervillea 189-191
decaryi 190, 191
tomentosa 190
venenata 191
Petasites 154
hybridus 154
Petradoria 62, 63, 65
discoidea 63
Phacelia 302
strictiflora 302
Phanerostylis 124
glutinosa 124

Philadelphus 231

Perdicium 154, 157, 158, 169, 187 mexicanus 231

lyratum 169

piloselloides 154

semiflosculare 157, 158, 187

Pericalia 334, 344
sessilifolia 334, 344
Periplocaceae 190, 191
Periplocoideae 190
Perittostema 40

Pernettya 241

Phlebodium 333, 340, 343

areolatum 333, 340, 343
Phlox 23,271

pilosa 23, 271

subsp. ozarkana 271

Phoebe 328, 334, 347

arsenei 328, 334, 347
Phoenix 358

sylvestris 358

49]

492 PHY T'OEOGTA

Phyla 349
nodiflora 349
Phyllanthus 358
emblica 358
Physalis 354
subtriflora 354
Phytolacca 348
icosandra 348
Phytolaccaceae 336, 348
Picea 220
engelmanii 220
pungens 220
Pinaceae 22, 242, 287
Pinaropappus 344
roseus 344
Pinophyta 268
Pintse22 NOs OA TLAS ose
209 LDS 223% 230433.
235-239, 241-243, 287-290
albicaulis 288
apulcensis 223, 235
attenuata 237
ayacahuite 233
var. brachyptera 233

subgen. Balfouria 288

June 1995

Pinus (cont.)
balfouriana 288
sect. Cembra 290
coulteri 288
durangensis 233, 235, 236
echinata 22
engelmannii 235, 236
estevezii 217, 235
subgen. Gerardia 287, 288
gerardiana 288
herrerai 235, 236
lawsonii 217-219, 230, 237-
239. 242
var. gracilis 218, 219,
230, 238
leiophylla 235-237, 288
var. chihuahuana 235, 236
sect. Leiophylla 290
lumholtzit_ 235, 236
maximinot 235, 236
michoacana 217, 235, 237
montezumae 219
Montezumae group 217, 219
nelsonit 288

nubicola 217, 235

volume 78(6):453-509

Index to taxa volume 78

Pinus (cont.)
oaxacana 217-219, 230, 237,
‘241, 243
var. diversiformis 218,
230, 237
oocarpa 209, 236
sect. Parrya 287, 290
patula 223
pinea 288
sect. Pinea 290
subgen. Pinea 287, 288
subgen. Pinus 288
pseudostrobus 223, 241, 243
var. apulcensis 223
var. megacarpa 24]
var. oaxacana 243
Pseudostrobus group 217, 219
sect. Quinquefoliis 290
subgen. Sabinia 288
sabiniana 288, 289
Strobus 288
subgen. Strobus 288
sylvestris 288
subgen. Tamaulipasa 287, 288

teocote 223, 239, 241

Pinus (cont.)
lorreyana 287, 288
yecorensis 218, 219, 230,
233; 2355236
var. sinaloensis 218, 236
Piperaceae 336, 348
Piqueria 329, 330, 334, 341, 344
trinervia 329, 330, 334, 341,
344
Pisoniella 329, 348
arborescens 329, 348
Pithecelobium 358
dulce 358
Plantaginaceae 27]
Plantago 233,271
aristata 27\
pusilla 27)
var. pusilla 271
virginica 271
Platanthera 1,9, 10, 14, 302, 311
blephariglottis 302
grandiflora 9
incana 302
integra 311

x keenanii 1,9, 10

493

494 PHY TOLOGLIA

Platanthera (cont.)
lacera 9, 14
zothecina 1, 10
Platypetalum 1,3
purpurascens 1,3
Plectrornis 89
bicolor 89
var. montanense 89
Ioan PBs) PSO
caespitosa 250
Poaceae 1, 10, 22, 244, 267, 274,
2D 2 1 S564 Is 3425350"
351, 425
Podocarpus 218
sect. Eupodocarpus 218
rostratus 218
Podosaemum 417, 418, 424, 427
gracilis 427
virescens 417, 418, 424
Polanisia 303, 346
erosa 303
uniglandulosa 346
Polemoniaceae 23, 271, 336, 348
Polyalthia 357

cerasoides 357

June 1995 volume 78(6):453-S09

Polygala 23, 272, 303
sanguinea 272
forma sanguinea 272
verticillata 23, 303
Polygalaceae 23, 272
Polygonaceae 272, 341, 348
Polygonella 303, 311
americana 303
polygama 303, 311
Polygonum 272, 334, 335, 341,
348
lapathifolium 334, 335, 341,
348
mexicanum 335, 348
lenue 272
Polypodiaceae 336, 343
Polypodiophyta 317, 336
Polypodium 6, 233
rhaetium 6
Polytaenia 303
nuttallii 303
Polytrichum 241
Pongamia 358
glabra 358
Populus 233

Populus (cont.)

tremuloides 233
Portulaca 272, 349

oleracea 272
Portulacaceae 271, 337, 349
Potentilla 233, 285, 286

heterosepala 286

macdonaldii 285, 286

sect. Multifidae 286

sect. Multijugae 285, 286

richardii 285, 286
Prenanthes 303

barbata 303
Prionosciadium 127-129

acuminatum 128

madrense 127

mexicanum 127

pringlei 127

thapsoides 127

turneri 127, 128, 129
Priva 349

grandiflora 349
Proboscidea 342, 348

louisianica 342, 348

subsp. fragrans 342, 348

Index to taxa volume 78 495

Prosopsis 1, 7, 8, 108, 328, 330-
332, 347
glandulosa 8
var. lorreyana 8
laevigata 328, 330-332, 347
odorata 1,7,8
Prunella 22
vulgaris 22
Prunus: 195234235. 241, 272329:
330, 349
hortulana 272
mexicana 272
microphylla 329, 330, 349
serotina 235, 241
subsp. capuli 235, 241
umbellata 23
Pseudotsuga 217, 218, 220, 233,
239-24]
brevifolia 240
flahaultii 220, 241
guinieri 220, 241
macrolepis 220, 241
menziesii 218, 220, 233, 239-
241

var. oaxacana 218, 239

496 PHY TOEOGIA

Pseudotsuga (cont.)
rehderi 220, 241
wilsoniana 240

Psidium 359
guajava 359

Psittacanthus 335, 347
calyculatus 335, 347

Psychotriinae 378

Ptelea 223, 328, 349
trifoliata 223, 328, 349

Pteridophyta 268, 380

Pterocarpus 358
marsupium 358

Pteroglossaspis 304
ecristata 304

Purshia 51
mollis 51

Pycnanthemum 22
albescens 22

Pyrrhopappus 21

carolinianus 21

June 1995

Q

Quechualia 397

fulta 397

Quercus 34, 103, 104, 114,

volume 78(6):453-509

iS?

147, 160)°223; 225) 227 e2sa-

233, 235-237, 241, 247, 264,

265, 267, 270

alba 270

calophylla 241
candicans 233, 235, 241
castanea 24)
coccolobaefolia 235
crassifolia 227, 241
laurifolia 247

laurina 223, 227, 241
magnoliifolia 225, 235, 236
marilandica 265, 270
stellata 270

var. stellata 270

Index to taxa volume 78 497

R

Ranunculaceae 23, 73, 100-102,
117, 118, 272, 310, 428, 444,
445

Ranunculus 233, 272
fascicularis 272
harveyi 272

forma harveyi 272

Ratibida 21
pinnata 21

Rauvolfia 192-194
longifolia 192, 193
stenophylla 194

Relbunium 361-363, 367, 368,
376, 378, 379
microphyllum 361-363, 367,

368, 376

mmaminaceae * 1, Ll, 14,.«23)..272,
359

Rhamnus 235, 272
betulaefolia 235
caroliniana 272

Rhapidophyllum 247
hystrix 247

Rhododon 448-45]

Rhodon (cont.)
angulatus 448, 449-45]
ciliatus 448-45]
Rhus 19, 21, 269
copallina 19, 21, 269
glabra 269
Rhynchelytrum 318, 330-333, 336,
341, 351
repens 318, 330-333, 336, 341,
351
Rhynchosia 22
Rhynchospora 247, 273, 304, 402
globularis 273
var. recognila 273
miliacea 247
oligantha 304, 402
Ribes 233
madrensis 233
Richardia 361-363, 367, 368, 376-
378
scabra 361-363, 367, 368, 376
tricocca 361, 3C2, 367, 368,
S77
subsp. lefracocca 361, 362,

367, 368, 377

498 PHY TOLOGIA

Ricinus 329, 341, 346
communis 329, 341, 346
Robinia 7
halodendron 7
Rosa 272
carolina 272
Setigera 272
forma setigera 272
var. setigera 272
var. tomentosa 272
Rosaceae 23, 272, 285, 336, 349
Rotala 265, 271
ramosior 265, 271
Rubiaceae 23, 272, 349, 359, 361-
363, 378-380
Rubieae 378
Rubioideae 378
RUDUSE2S a2 S352 12:
flagellaris 272
invisus 272
Rudbeckia 21, 269, 304, 309, 310
hirta 21
laciniata 304
missouriensis 269

scabrifolia 304, 309

June 1995 volume 78(6):453-509

Ruellia 21, 268
humilis 268
pedunculata 21
Rumex 272, 335, 341, 348
acetosella 272
crispus 335, 341, 348
Rutaceae 336, 349, 359

S

Sabal 247
palmetio 247
Sabatia 22,304
campestris 22
macrophylla 304
Sabazia 131
Sabina 143, 144
barbadensis 144
bermudiana 143
Saccopetalum 358
tomentosum 358
Salicaceae 15, 336, 349
Salix 70, 233, 247, 320, 328, 334,
335, 349
bonplandiana 320, 328, 334,
335, 349

Index to taxa volume 78 499

Salix (cont.)
floridana 247
gooddingii 233
serissima 70
Salvia 22, 329, 330, 331, 347
azurea 22
hirsuta 330, 347
lyrata 22
nana 347
polystachya 330, 331, 347
reflexa 347
Sambucus 223
mexicana 223
Sanvitalia 330, 332, 340, 341, 344
procumbens 330, 332, 340,
341, 344
Sapindaceae 349, 359
Sapindus 359
emarginatus 359
Sapotaceae 272, 359
Sarracenia 1, 12, 13, 16, 17, 402
alata 402
gibbosa 12, 13
heterophylla 13

purpurea 1, 12, 16, 17

Sarracenia purpurea (cont.)
var. burkii 13, 16
subsp. gibbosa 12
var. heterophylla 13
subsp. purpurea 12, 13
var. purpurea 13
var. ripicola 13
var. stolonifera 13
var. lerrae-novae 12, 13
subsp. venosa 12, 13, 16
venosa 12, 13
Sarraceniaceae 1, 12, 16, 17
Saxifragaceae 15, 273
Schinus 320, 328, 330-335, 343
molle 320, 328, 330-335, 343
Schizachne 422
Schizachyrium 22,267, 274
scoparium 22, 267, 274
lenerum 22
Schkuhria 335, 344
pinnata 335, 344
var. virgala 335, 344
Schleichera 359
oleosa 359

Schoenolirion 305, 312

500 PHY TOLOGIA

Schoenolirion (cont.)
croceum 305
wrightii 305, 312
Schrankia 22, 271
microphylla 22
nuttallii_ 271
Scleria 21
oligantha 21
Scrophulaniaceae 23, 28, 35, 131,
1335425653259) (27135 286; 313%
349
Scutellaria 22
elliptica 22
Secamone 189-191
Secamoneae 189, 190
Secamonoideae 191
Sedum 241, 333, 334, 346, 405-
408
caducum 405-407
ebracteatum 333, 334, 346
edwardsii 405, 406, 407
hintonii 406
hintoniorum 405-407
rhodocarpum 405-408

subsp. edwardsii 405, 306

June 1995 volume 78(6):453-S509

Selaginella 305, 319, 333, 342
arenicola 305
subsp. riddellii 305
lepidophylla 319, 333, 342
Selaginellaceae 336, 342
Semecarpus 357
anacardium 357
Sempervivum 1, 5,6
haworthii 1, 5,6
Senecio 1,2, 131,227, 2335241
andrieuxit 227
angularis 241
barba-johannis 241
bicolor 2
subsp. cineraria 2
cineraria 1,2
Senecioneae 34]
Senna 328, 347
polyantha 328, 347
Setaria 22, 331, 341, 351
geniculata 22, 331, 341, 351
grisebachii 35)
Seymeria 28-35
bipinnatisecta 30

coahuilana 30

Index to taxa volume 78 SO]
Seymeria (cont.)

Silphium (cont.)

cualana 28-30, 31 integrifolium 21

decurva 29 laciniatum 21

deflexa 28-31 Simsia 341, 344

falcata 28, 29, 32 amplexicaulis 341, 344

var. falcata 32 Sisymbrieae 16

gypsophila 28, 29, 32-35
integrifolia 28, 29, 31, 33, 34
laciniata 29, 34

pailana 28, 29, 33

pennellii 28-30, 34

scabra 30

sinaloana 30

tamaulipana 28, 29, 31-35

virgata 29

Sherardia 361, 362, 368, 377, 378

arvensis 361, 362, 368, 377,
378

Shorea 358

talura 358

Sida 341, 348

abutifolia 341, 348

Silene 267, 269

regia 267, 269

Silphium 21

Sisyrinchium 22, 273, 350
campestre 273
forma campestre 273
langloisii 22
ltenuifolium 350
Smilacina 301
racemosa 301
Solanaceae 341, 342, 349, 353, 356
Solanoideae 353
Solanum 231, 330, 331, 338, 341,
349
cervantesii 231
dulcamaroides 341, 349
eleagnifolium 341, 349
nigrescens 331, 341, 349

rostratum 349

Solidaginineae 62

Solidago 21, 269, 305

auriculata 305

502 PHY Ow O'!IG A June 1995 volume 78(6):453-SO09

Solidago (cont.) Sporobolus asper (cont.)
nemoralis 269 var. clandestinus 244
nitida 21 aspera 1, 10, 11, 16
radula 21 var. aspera 10
ulmifolia 269 var. drummondii 1, 11

Sonchus 341, 344 var. macer 1, |1
oleraceus 341, 344 canovirens 244

Soymida 358 clandestinus 244, 274
febrifuga 358 var. canovirens 244

Spermacoceae 378 compositus 1, 10, 11, 244

Sphaeralcea 330, 331, 340, 341, var. clandestinus 244
348 var. drummondii 1, 11
angustifolia 330, 331, 341, 348 var. macer 1, 11
hastulata 340, 348 drummondii 11

Sphagnum 402 macer \\

Sphenopholis 22, 274 ozarkanus 267, 274, 305
obtusata 22,274 vaginiflorus 274, 275

Spiranthes 274 Stachydeoma 448, 449, 451
tuberosa 274 angulata 448, 449

Spirolobium 1, 7,8 ciliata 448, 451
odoratum 1,7,8 duvalii 448, 449, 451

Sporobolus 1, 10, 11, 16, 244, graveolens 448
245, 267, 268, 274, 275, 305 Stachys 209, 210
asper 244, 245, 274 coccinea 209, 210

var. canovirens 244 manantlanensis 209, 210

Index to taxa volume 78 503

Stachys (cont.) Strebulus (cont.)
pacifica 209, 210 aspera 359
Stenanthium 305 Streptanthus 306
gramineum 305 hyacinthoides 306
Stenocactus 320, 339, 345 Strychnos 358
Stenocereus 320, 339, 345 nux-vomica 358
dumortieri 320, 345 potatorum 358
Stenotus 62, 63 Stylisma 306
Sterculia 359 aquatica 306
urens 359 Stylosanthes 270
Sterculiaceae 359 biflora 270
Stevia 206, 328, 330, 333, 334, Styphelia 249, 254
344 Symphoremaceae 72, 152, 316, 404
bisecta 206 Symphoricarpos 233, 269
micrantha 330, 334, 344 orbiculatus 269
ovata 334, 344 roundifolius 233
Serrata 328, 330, 333, 344 Syngonium 223
var. serrata 328, 333, 344 podophyllum 223
Steviopsis 124-126 Syzygium 359
nesomii 124, 125 cumini 359

Stilbaceae 72, 152,316, 404

Stilpnopappus 384, 398, 399 ap

pantanalensis 398, 399 Taenidia 306

trichospiroides 399 j
piroides 3 integerrima 306

Strebulus 359 Tageteae 211, 213, 341

504 PHY TOLOGIA

Tagetes’ (213330."332, 333, 335,
341, 344
lunulata 330, 332, 333, 335,
341, 344
Talinopsis 349
frutescens 349
Talinum 272, 305, 306, 349
calycinum 272
napiforme 349
paniculatum 349
parviflorum 305, 306
Tamarindus 358
indica 358
Taraxacum 331, 344
officinale 331, 344
Tecoma 329, 345
stans 329, 345
Tectona 359
grandis 359
Tephrosia 22, 270
virginiana 22, 270
Terminalia 358
arjuna 358
bellerica 358

catappa 358

June 1995 volume 78(6):453-509

Terminalia (cont.)
chebula 358
tomentosa 358

Ternstroemia 223
pringlei 223

Tetragonotheca 306, 313
ludoviciana 306

Tetramerium 331, 343
nervosum 331, 343

Tetraneuris 1,2
acaulis 1,2

var. epunctata 1, 2
var. nana |, 2
epunctata 2

Thalictrum 233

Thecophyllum 122
werckleanum 122

Thelypteris 247
palustris 247

Thespesia 358
populnea 358

Thrysanthemum 350
macrophyllum 350

Thuidium 241

Thyrsanthema 154, 155, 158, 162,

167-169, 187

Thyrsanthema (cont)
hybrida 155
lyrata 169
nutans 154, 162
runcinata 168
semifloscularis 158
spathulata 169
tomentosa 167

Tilia 247
caroliniana 247

Tiliaceae 359

Tillandsia 331, 333, 350
recurvata 331, 333, 350
usneoides 350

Tillandsioideae 119

Tithonia 341, 344
tubiformis 341, 344

Tonduzia 192-194
longifolia 192, 193
longipedunculata 194
macrantha 192, 193
parvifolia 194
pittieri 192, 194
stenophylla 194

Toxicodendron 334, 335, 343

Index to taxa volume 78

Toxicodendron (cont.)
radicans 334, 335, 343

Toxocarpus 189-191
decaryi 189-191
kleinii 189

tomentosus 189, 190

Tradescantia 273, 306, 311, 350

crassifolia 350
ohiensis 273
reverchonii 306
Tragia 270, 346
betonicifolia 270
nepetifolia 346
Trichochloa 418
virescens 418
Trichocline 153, 188
Tridax 332, 344
coronopifolia 332, 344
palmeri 344
Tridens 22,274
flavus 22,274
var. flavus 274
Trifolium 267, 270

reflexum 267, 270

var. reflexum 267, 270

506 PHY TOLOGTIA

Trigonospermum 36-38
annuum 36-38
hintoniorum 36-38
Stevensii 36-38

Triniochloa 422

Triphora 307
trianthophora 307

Tripsacum 351

Trixis 328, 329, 333, 334, 344
mexicana 328, 329, 333, 334,

344

var. auriculata 329, 333,
334, 344

var. mexicana 333, 344

Tussilago 154, 157, 162, 167, 169
albicans 167
bicolor 167
exscapa 154
hybridus 154
integrifolia 157
lyrata 162, 169
nutans 154, 162
tomentosum 167

vaccina 162

June 1995 volume 78(6):453-5S09

U

Ulmaceae 273, 359
Ulmus 273
rubra 273
Umbelliferae 130
Utricularia 307
juncea 307
purpurea 307
Uvularia 307

sessilifolia 307

V

Vaccinium 241, 265, 267, 270
arboreum 265, 267, 270
confertum 241

Valeriana 233

Vanclevea 63, 65

Veratrum 233

Verbena 23, 330, 332, 342, 349
bipinnatifida 332, 342, 349
carolina 330, 349
halei 23

Verbenaceae 23, 72, 152, 316, 342,

349, 359, 404

Index to taxa volume 78 S07

Verbesina 131, 237, 328, 334, 344 Viburnum 19, 21, 269
serrata 328, 344 dentatum 19, 21
sphaerocephala 334, 344 rufidulum 269

var. sphaerocephala 344 Viguiera 331-333, 341, 344, 345

Vernonanthura 384-386 dentata 341, 344
diffusa 385 var. canescens 344
divaricata 385 linearis 331-333, 345
lindbergii 385 Vilfa 10, 11, 244
lipeoensis 385 aspera 10, 11
nebularum 385 clandestina 244
pseudonudiflora 385 drummondii 11
schulziana 386 macra 11
vinhae 386 Villadia 346

Vernonia 237, 385, 386, 389, 399 parviflora 346
almasensis 385 Viola 23, 233, 273, 306
chiriquiensis 385 lanceolata 306
sect. Eremosis 389, 399 pedata 273
lindbergii 385 forma pedata 273
lipeoensis 385 rafinesquii 273
nebularum 385 walteri 23
pseudaurea 385 Violaceae 23, 273
pseudo-nudiflora 385 Vitaceae 23, 336, 349
schulziana 386 Vitis 23
vinhae 386 mustangensis 23

Vernonieae 384, 399 Vriesea 119-123

508 PHY TOWOGTA June 1995 volume 78(6):453-509

Vriesea (cont.)

breedloveana 119, 121, 122 Vulpia 267, 274
burgeri 119, 121 octoflora 267, 274
chiapensis 119, 121 var. glauca 274

cornus-cervii 119, 121, 122
dodsonii 121 W

greenbergii 121 Wedelia 208

macrantha 119, 121 Weinmannia 387

macrochlamys 121 Werauhia 119-123

nutans 121 breedloveana 122

; 5) oy)
ovandensis 119, 121, 122 burgeri 119,121

s 5
patzeltii 119, 121 chiapensis 121

subsp. panamaensis 119, a
jeay cornus-cervil 122

121 greenbergii 119, 121

var. panamaensis 121 sect. Jutleya 120

‘ 5
pectinata 121 macrantha 119, 121

aD)
pycnantha 122 nephrolepis 122

rugosa 121 :
§ ovandensis 122

j 2)
subgen. Vriesea 120 patzeltii 121
Meee ee subsp. panamaensis 121

iphi 3) 9%)
sect. Xiphion 120, 122 pectinata 119, 121

Werauhia (cont.)
pycnantha 119, 121,122
sect. Werauhia 120
werckleana 119, 121, 122
Wrightia 358

tincotoria 358

X

Xanthium 341, 345
strumarium 341, 345
Xanthorhiza 307, 310
simplicissima 307, 310
Xylea 358
xylocarpa 358
Xyris 307-309, 402
drummondii 307, 309
scabrifolia 307

Stricta 308

i'd

Yucca 34, 200

Index to taxa volume 78

Z

Zaluzania 328, 330-332, 345
augusta 328, 330-332, 345
Zamiaceae 409
Zephyranthes 350
Zigadenus 308
densus 308
Zinnia 330-332, 341, 345
peruviana 330-332, 341, 345
Zornia 308, 340, 347
bracteata 308
thymifolia 340, 347
Zyzyphus 359

frinervi 359

509

Phytologia (June 1995) 78(6):510.

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INDEX TO REVIEWERS, VOLUME 78

The editor expresses his most sincere appreciation to the following individuals.
These are persons who have reviewed papers that were submitted for publication in
volume 78 of Phytologia. Without the willingness and diligence of these reviewers,
the task of the editor would be much more difficult, and the quality of the papers
would be lessened. To each of you, I offer my most sincere thanks.

Michael J. Warnock, Editor.

Almeda, F. Holmes, W.C.
Anderson, C. latte, JL.
Ashton, P.S. Hyatt, P.
Barkley, T.M. Jenkins, P.
Brown, L.E. Karis, P.O.
Carr B: Kral, R.
Cayouette, J. Jihua, Xiong
Chiang, F. Jiménez, Q.
Cody, W.J. Jones, G.D.
Davila, P.D. Jones, S.
Delavoryas, T. Larke, J.
Delprete, P. Laven, R.D.
Doolittle, W. Lellinger, D.B.
Ertter, B. Lelong, M.G.
Fernandez N., R. Lowrey, T.
Folsom, J. MacRoberts, D.T.
Fox, W.E. Marroquin, J.S.
Funk, V. Mason, C.
Grayum, M. Mayfield, M.
Hammel, B. Moore, M.O.
Herbst, D. Nesom, G.L.
Hedenas, L. Prather, A.

S11

$12 PHY TOLOGIA June 1995
Pruski, J.

Rao, K.G. Raja
Rao, P.N.
Reveal, J.L.
Reznicek, A.A.
Rodrfguez J., C.
Rominger, J.
Rundell, J.R.
Rzedowski, J.
Smith, L.
Stevens, P.
Stewart, R.
Strother, J.
Thomas, R.D.
Turner, B.L.
Wetherwax, M.
Whitelock, L.M.
Xiong, J.H.
Yatskievych, G.C.
Zamora, N.

volume 78(6):511-512

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Code of Botanical Nomenclature) citation of sources of epithets and Br
combinations.

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