6 PHYTOLOGIA

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

~ Vol. 79 October 1995 No. 4

CONTENTS

Le NESOM, G.L. & TURNER, B.L., Systematics of the Sedum parvum group
(Crassulaceae) in northeastern México PNG LEXA, ois scan sare that etrectan 257
- GRAY UM,.M.H. & B.E. HAMMEL, The genus Tetranema spa wb euaataseiroay

in Costa Rica, ‘with two newsspecies es eee e os pee che in ewes ees 269

- NESOM, G.L., Key to the American genera of Asterinae (Asteraceae) ae stay 281

_ TURNER, B. ee Tridax yecorana (Asteraceae, Heliantheae) a new species from
Sonora, Te Sean ST NN tt a eGR D eee Armee cet 286

_ TURNER, B.L., Salvia booleana (Lamiaceae), a new species from northeastern
WARIS 5k Ge saa tess ca rhen sn ae soun SoC AEN Leta ig adestepaie per 289

~» TURNER, B.L., A new species of Lobelia (Campanulaceae). from Oaxaca,
1s), (25.418, 0 SRR Cue aE spn igh pc Nite ea ners Mp ayer genoa Rincuey AL Pe Cen Raa sD 293
“~*~ TURNER, B.L., po new species of .Verbesina (Asteraceae) from Oaxaca,
; RANG 0 SEER ES RAS ARBEIT SSE NS, a AE AL Roe oe Cokt OG wee wy Be eR 296
TURNER, B.L. & A.L. HEMPEL; A new species of Mentzelia (Loasaceae)
EPOMYIUE VG: OON; MERICO. 55694 aos bade Li Scr og Decina Lash ape es ata 298

_~ TURNER, B.L., A new species of Stevia (Asteraceae) from Cerro Quiexobra,
Oaxaca, Migkidbs len ae ne Ae erie Lotte 301

“ TURNER, B.L., Stellaria miahuatlana (Caryophyllaceae), a new species from
PAK SCA TER IGOR ioe 1 OS Le ae Pn PG PO ATES pote eee 303
“TURNER, B.L., A new species of Cynoglossum (Boraginaceae) from Oaxaca,
PAE NICD Ep Sis as Nien ean ng ABaRe Nad crepes UL TRAIT Soe aon 306

a TURNER, B.L:., Two new species of Ageratina (Asteraceae) from México. . 309
_ TURNER, B.L.,. A new species of Bocconia (Papaveraceae) from Oaxaca,
PRN Win itn Poke Loos sur eke y PEND Os BIG TER aay CLT ev ete Re eos 313

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Phytologia (October 1995) 79(4):257-268

SYSTEMATICS OF THE SEDUM PARVUM GROUP (CRASSULACEAE) IN
NORTHEASTERN MEXICO AND TEXAS

1Guy Nesom & 2B.L. Turner
IRte. 5, Box 298, Huntsville, Texas 77340 U.S.A.

2Department of Botany, University of Texas, Austin, Texas 78713 U.S.A.

ABSTRACT

In a systematic study of the yellow-flowered species of Sedum of
northeastern México and Texas, those traditionally interpreted as the S.
parvum Hemsl. group, we elevate or reinstate to species rank three taxa
regarded by Clausen as subspecies of S. parvum: S. catorce stat. et nom.
nov. (= S. parvum subsp. dendroides R.T. Clausen), S. nanifolium Frod. (S.
parvum subsp. nanifolium [Frod.] R.T. Clausen), and S. diminutum stat. et
comb. nov. (= S. parvum subsp. diminutum R.T. Clausen). In addition, three
new species are proposed: S. dulcinomen Nesom, S. papillicaulum
Nesom, and S. macdonaldii Nesom.

KEY WORDS: Crassulaceae, Sedum, México, systematics

This study was begun as a consequence of the observation by McDonald (1991)
that two separate species of Sedwm occur on the tops of the high peaks of southeastern
Coahuila. Robert Clausen had earlier identified and annotated nearly all of the yellow-
flowered species of Sedum in the Sierra Madre of Coahuila and Nuevo Leén as S.
parvum Hemsl. subsp. nanifolium (Frod.) R.T. Clausen, although many of the
collections cited in the present study have been made since his death. With further
study, it became apparent that only one of these high elevation taxa could be identified
as subsp. nanifolium. Several other entities related to S. parvum, at high and lower
elevations in the Sierra Madre and adjacent areas, represent undescnbed species.
Further, based on a more restncted species concept than Clausen’s, we believe that a
group of taxa described by him as subspecies of S. parvum are better treated at specific
rank. Clausen espoused, theoretically and practically, a very broad species concept
(1984, p. 9): “The concept of species is best reserved for a level of differentiation that
is major, involves many genetic characters, and is especially distinguished by
impressive biological discontinuity where two or more species occur together.” The

2}

258 PHYTOLOGIA October 1995 volume 79(4):257-268

subspecies of this group, however, are allopatnc non-intergrading taxa of northeastern
México and the morphological distinctions among them are equivalent to those of
accepted species in many genera of vanous other families.

Distinctive features of the Sedum parvum group are yellow, erect petals, yellow
anthers and ovaries, and relatively small, elongate leaves drying with a flat to slightly
concave adaxial surface (see below). Two other Mexican species clearly are members
of the same group, Sedum reptans R.T. Clausen of San Luis Potosf and Querétaro,
and S. tammaulipense Nesom of Tamaulipas. These taxa are distinctive in their
spreading petals and fully terete leaves, but in their completely prostrate habit they are
perhaps closely related to S. parvum. Sedum nutiallianum Raf., an erect annual
common in Texas and the south-central United States, also appears to be very closely
related (Nesom 1988). Clausen (1975) included the latter as the most divergent
member of subgenus Sedum sect. Lanceolata, in which he otherwise included only
species from north of México, but he noted that it is related to S. parvum “on the basis
of morphology and geography.” Finally, S. greggii Hemsl. and S. grandipetalum
Frod. of the Trans-Mexican Volcanic Range, and perhaps S. humifusum Rose and S.
cupressoides Hemsl., also appear to belong with this assemblage of taxa. According
to Praeger (1921), these would be members of sect. “Seda Genuina Koch” (= sect.
Sedum).

Other yellow-flowered species of Sedum in México may constitute more than one
natural group apart from the S. parvum group. All, however, have flat (fresh) leaves,
mostly 6-20 mm wide (much wider than the taxa treated in the present paper) and 15-
40 mm long, and the inflorescences tend to be elevated well above the normally
developed cauline leaves. Only one of them, S. palmeri, occurs in northeastern
México.

Vaniation in leaf shape and arrangement

Clausen (1978) referred to Sedum greggii and S. grandipetalum as
“heterophyllous,” in reference to the “leaves of the elongate flonferous stems [which]
are markedly different from those of the vegetative shoots or compact rosettes.”
Species treated in the present study also present a type of heterophylly, although it may
not be homologous with that of those noted above. These produce short, vegetative,
lateral branches with spreading, imbricately arranged leaves so densely packed that the
stem surface is not evident. Sometimes, however, the flonferous branches of these
also produce densely packed leaves resembling those of the lateral branches. In any
case, five essentially prostrate species covered in this study (in the key below) do not
produce this type of leaf arrangement vegetatively or otherwise but rather the leaves are
evenly and well-spaced, leaving the stem surface in view. Such species without a
marked dimorphism in leaf morphology Clausen called “homophyllous.”

In Clausen’s key to the species of the Mexican Cordilleran Plateau (1984), an early
couplet distinguished “leaves flat, 2 or more times broader than thick” (Sedum greggii)
from “leaves terete, subterete, or at least not 2 or 3 times broader than thick” (S.
parvum and S. reptans). These choices were in reference to living maternal, which
Clausen expected users of his key to have on hand (or else field notes describing the
onginal morphology), but the leaves of Sedum shrink radically upon drying and it

Nesom & Turner: Systematics of Sedum parvum group 259

becomes difficult to surmise the original shape. The relative position of the midvein in
dried leaves does provide a clue to the original shape. The key below presents choices
based on dried material, and notes on the morphology of fresh maternal are provided in
the species descriptions where this information is available. The leaves of the S.
parvum group have a strong tendency to markedly flatten when dned, even to the point
of displaying two, sharply acute, lateral margins. The leaves of S. nanifolium Frod.
and S. macdonaldii Nesom usually dry with a relatively flat adaxial surface and
convex abaxial one with rounded, poorly defined margins. These are referred to as
“half-terete” in the descriptions below. Although the leaf shape (in cross-section) is
constant within a species, it does not appear to be of cnitical importance in indicating
phylogenetic relationships, if our supposition is correct that S. nanifolium is most
closely related to S. chrysicaulum J.A. McDonald, which has markedly flattened
leaves with strongly differentiated margins.

Variation in stem and leaf surface texture

Three basic types of stem morphology in the Sedum parvum group can be
identified: 1) smooth, non-shiny, without visible cellular structure; 2) smooth, shiny,
with elongated cells evident; and 3) papillate, non-shiny, with quadrate cells evident.
The following groups of species correspond to the three stem types: 1) S. reptans, S.
nuttallianum, S. tamaulipense; 2) S. parvum, S. diminutum (R.T. Clausen) Nesom,
S. nanifolium, S. chrysicaulum, and 3) S. papillicaulum Nesom, S. macdonaldii, S.
catorce Nesom, S. dulcinomen Nesom. These three morphologies are so distinctly
different that we believe that the three species groups may represent separate phyletic
lineages, each including species both of erect and prostrate habit. It is surprising that
Clausen did not recognize the usefulness of this character among the species treated in
the present study, because he used the same character in several key couplets (Clausen
1984) to delimit groups of species unrelated to the S. parvum group.

Description of the Sedum parvum group

Unless otherwise noted, the descriptions and measurements below are taken from
dried, pressed specimens. Elaboration is found in the paragraphs above.

Erect or prostrate-decumbent perennials, annual in one species, somewhat woody
in the lower portions, completely herbaceous in one species. Stems glabrous, smooth
or papillate. Homophylious or heterophyllous, the leaves glabrous, small, 3-10 mm
long, drying flat to-half-terete. Flowers in congested, terminal cincinni, more
diffusely arranged in one species. Petals yellow, sometimes with short, red,
longitudinal stnpes, separate, each usually with a thick, raised, medial keel widened at
the apex, erect to spreading or reflexed. Stamens 10, those opposite the petals adnate
to the petal base; anthers yellow, red in one species. Carpels yellow, erect, the
follicles erect to spreading, free or sometimes basally connate, baso-ventrally gibbous,
each with numerous seeds. Seeds brown, minutely papillate, echinate in one species.
Chromosome numbers, ”=10 and 26 (known from only 2 species, see below).

260 PHYTOLOGIA October 1995 volume 79(4):257-268

KEY TO THE SPECIES OF THE SEDUM PARVUM GROUP IN
NORTHEASTERN MEXICO AND TEXAS

1. Stems minutely papillate to papillate-glandular, without a sheen................... (2)
2. Plants prostrate-decumbent from horizontal rhizomes. ......... 9. S. dulcinomen
2. Plants erect, without stolons or creeping rhiZOMES................:ssseeeeeeeeees (3)

3. Stems 10-25 cm high or long, somewhat glaucous; leaves with prominent,
wide, white margins of different texture than the blade; petals often
Spreading OL TEMexUN Ss a. re eer ee cee or eee nn eee neces sere scr or 8. S. catorce

3. Plants mostly 4-14 cm high, not glaucous; leaf margins sometimes slightly
lighter colored than the blade but then very narrow and not of different
texture) petals) (CLEC ccese tein. ot nds seed downer sccceaens neeicace spent eaeem (4)
4. Stems densely and minutely papillate, appearing somewhat stipitate-

papillate, the cellular structure not clearly perceptible; leaves flattened or
slightly convex above and beneath, the cellular structure evident only in
the striate, elongated cells at the abaxial base, the margins sharply
flattened and translucent-papillate ................... 11. S. papillicaulum
4. Stems not distinctly columnar- or stipitate-papillate but with quadrate
cells clearly perceptible; leaves half-terete, not strongly papillate but
both surfaces with clearly perceptible quadrate cells, the margins

rounded and not at all papillate............ ee 12. S. macdonaldii

1. Stems smooth, not at all papiilate or glandular-appearing, with or without a sheen.
Bet eR EE SE in ats CAR an RtsR WOT dnin sda a aorta saitwie hesctia eek Heat elena anleouee ee eee (5)
5. Plants prostrate-decumbent from horizontal rhizomes; leaves and petals without
PIOMINENE TES MALIN GS eee n seen en esas tele he eee seem ee eee eee (6)

6. Stems with a prominent sheen on the lower part, the cells elongate; petals

erect; =3-5 cm ONG: cacy... Ges. cese Meee eeeeseaas. ce otnoesnlens eas teee eee (7)

7. Rhizomes completely herbaceous; leaves 4-8 mm long, 2-4 mm wide;
flowers in compact, terminal cincinni; petals 4-5 mm long; anthers

V elonyp en swe mr, 23. sae ace ah dete oe eae 1, S. parvum

7. Rhizomes thin but noticeably woody; leaves 3.5-5.3 mm long, 1.3-1.8
mm wide; flowers arranged relatively diffusely along upper branches;
petals 3/0-3:5,mm long; anthers)red. <....-...---5-s-00-< 2. S. diminutum

6. Stems without a prominent sheen, the cellular structure not readily
apparent, petals spreading, either ca. 2.5 or 6.5-8.5 mm long............. (8)
8. Petals ca. 2.5 mm long; leaves 2.5-7.0 mm long, 0.8-1.0 mm wide....
Ua lla cia GND Rea saat ai fo hae Sina od sina oats Noten gee eenle mer 4. S. tamaulipense

8. Petals 6-8 mm long; leaves 6.5-8.5 mm long, 2.5-3.5 mm wide. .......

Shu. cidlpalleiaiae win Solent Hoe nivale Sa eeiaveislate ne alo’ RU Aabrrcdolelsigloines wees 3. S. reptans
5. Plants erect, without rhizomes; leaves and petals with or without prominent red
MAPKIN GS) ei iaidch eed docicainchce sooaeealst ds v'olye ge vkphrwiins cbhwantcd ess teheacae eate seminal (9)

9. Annual; stems without a sheen; leaves and petals without red markings.....
seanse ug oties 122 ocaancndiss aachascoccsung. dopcatadeniptncc aden S weevil Mel) Aen li
9. Perennial; stems with a prominent sheen at least on the lower part; leaves

and petals withiprominent redimarkingss «2. (01.4 ase eese essence emer (10)
10. Lower part of stems reddish-shiny; leaves 2.5-3.5 mm long, even in
length; petals'4:0-5) limi longa ih os.0-.2--ce rece seen 6. S. nanifolium

10. Lower part of stems golden-shiny; leaves of lower stems 5-6 mm long,
7-10 mm long near the inflorescence; petals 5.0-7.5 mm long .......... :
Jock vateh Siac HRee cota Rurectragaeane scabs onep qammaaen eae 7. S. chrysicaulum

Nesom & Turner: Systematics of Sedum parvum group 261

1. SEDUM PARVUM Hemsl.

Sedum parvum Hemsl., Diagn. Pl. Nov. Mex. 5O. 1880. Altamiranoa parva
(Hemsl.) Rose ex Bnitt. & Rose, Bull. New York Bot. Gard. 3:32. 1903.
Villadia parva (Hemsl.) Jacobsen, Natl. Cactus Succ. J. 13:76. 1958. TYPE:
MEXICO. San Luis Potosf: In regione San Luis Potosi, 6000-8000 ft, 1878,
Parry & Palmer 234 (HOLOTYPE: K; Isotype: GH!).

Sedum pososepalum Frod., Acta Horti Gotob. 10, App.: 66. 1936. TYPE:
MEXICO. San Luis Potosi: Alvarez, 13-23 Jul 1904, Palmer 25]
(HOLOTYPE: US; Isotype: GH!).

Prostrate-decumbent, completely herbaceous perennials from creeping rhizomes,
producing fibrous, adventitious roots. Stems green, smooth, with a noticeable reddish
sheen below, the cells elongated. Leaves evenly sized and spaced, green, usually
drying translucent, very flat, narrowly elliptic-oblong to narrowly oblanceolate-
oblong, 48 mm long, 2-4 mm wide. Petals erect, yellow, 4-5 mm long. Seeds
echinate.

San Luis Potosf; rock outcrops or ledges in pinyon pine, pine-oak, pine-fir woods;
2000-2700 m; June-October.

In its prostrate habit, shiny stems, and “homophyllous” leaves, Sedum parvum is
most similar to S$. diminutum. Sedum reptans, which also occurs in San Luis Potosi,
can be distinguished by its non-shiny stems, longer petals, and papillate seeds.

to

SEDUM DIMINUTUM (R.T. Clausen) Nesom, comb. ef Stat. nov.
BASIONYM: Sedum parvum Hemsl. subsp. diminutum R.T. Clausen, Bull.
Torrey Bot. Club 106:215. 1979. TYPE: MEXICO. Coahuila: Cafiada La
Hacienda, Sierra de la Madera, NW of Cuatro Cienegas, crevices of limestone in
shade of Quercus, slope on E side of Cafiada, 1860 m, 22 Jul 1978, R.T. Clausen
78-7 (HOLOTYPE: BH!; Isotypes: BH-3 sheets!).

Prostrate-decumbent perennials from slightly woody rhizomes, producing
adventitious, fibrous roots. Stems green, smooth, with a noticeably golden sheen
below, the cells elongated. Leaves evenly sized and spaced, green, very flat, narrowly
elliptic-oblong to narrowly oblanceolate-oblong, 3.7-5.5 mm long, 1.3-1.8 mm wide.
Flowers arranged rather diffusely along upper branches, not in congested cincinni.
Petals erect, yellow, 3.0-3.5 mm long. Anthers red.

Known only from the type collection. See comments following Sedum parvum.

3. SEDUM REPTANS R.T. Clausen
Sedum reptans R.T. Clausen, Bull. Torrey Bot. Club 105:222. 1978. TYPE:
MEXICO. San Luis Potosi: Sierra de San Luis Potosi, northern slope of Cerro H

Ajugon, 21° 40’ 45” N, 100° 03’ 20” W, 1720 m, 7 Sep 1977, R.T. Clausen
772,036 (HOLOTYPE: BH!).

262 PHYTOLOGIA October 1995 volume 79(4):257-268

Sedum reptans R.T. Clausen var. carinatifolium R.T. Clausen, Variation Spec.
Sedum 15, 1981. TYPE: MEXICO. Querétaro: 1 km W of Lazaro Vega, 8

km NE of Vizarron des Montes, 20° 53’ 25” N, 99° 39’ 40” W, in depressions
in limestone exposed to SE, 2250 m, 14 Apr 1980, R.T. Clausen 80-29
(HOLOTYPE: BH!; Isotype: BH!).

Completely herbaceous perennials with prostrate, creeping stems producing
fibrous adventitious roots and short, erect or decumbent, leafy stems. Cells of stems
quadrate, noticeably papillate in the youngest portions. Leaves terete to subterete to
carinate above (fresh), drying [somewhat flattened], 6.5-8.5 mm long, 1.5-2.5 mm
wide. Petals spreading, 6-8 mm long. Follicles widely spreading, ventrally gibbous,
basally connate for ca. 1/4 their length.

Eastern San Luis Potosi to northeastern Querétaro; limestone rocks; 1700-2250 m;
Apnil-September.

In its habit, leaf shape, and morphology of its stems and follicles, Sedum reptans
is similar to S. tamaulipense. Clausen (1981) described subsp. carinatum as different
from subsp. reptans “in the length (8.5 versus 6.7 mm) and length-width ratios (4
versus 2.4 of the leaves” and dorsally carinate leaves. These putative differences (we
find overlap in the leaf length) do not appear to us to justify formal nomenclature.

4. SEDUM TAMAULIPENSE Nesom

Sedum tamaulipense Nesom, Sida 13:22. 1988. TYPE: MEXICO. Tamaulipas:
Mpio. San Carlos, Sierra de San Carlos, ca. 5 mi S of San Carlos, N side of Bufa
E] Diente, 18 Jun 1987, G. Nesom 6166 (HOLOTYPE: TEX!; Isotypes: BH!,
MEXU!,UAT!,WTU}).

Perennial, prostrate herbs with numerous adventitious roots, forming mats, the
stems smooth, without a sheen. Leaves narrowly oblong, terete (fresh), drying
flattened, 2.5-7.0 mm long, 0.8-1.0 mm wide. Petals ca. 2.5 mm long, yellow,
spreading. Follicles spreading, ventrally gibbous, basally connate for 1/2-3/5 their
length.

Known only from the Sierra de San Carlos of central Tamaulipas; top of rocks and
large boulders with bryophytes, in oak woods with scattered Carya and Abies, 1100-
1250 m; May-August.

5. SEDUM NUTTALLIANUM Raf.

Sedum nuttallianum Raf., Atl. J. 1:146. 1832. TYPE: UNITED STATES.
{Oklahoma]: drainage of the Red River, Arkansas, 1819, Nuttall s.n. (NY). See
Clausen (1975) for notes on typification and synonymy.

Erect annual herbs 5-8 cm tall, from a very slender taproot, the stems smooth,
without a sheen. Leaves narrowly elliptic-oblong, mostly 3-5 mm long, terete (fresh),
drying more or less terete, all more or less similar in size and distribution. Petals
yellow, spreading, 2-4 mm long. Follicles widely spreading, ventrally gibbous.
Chromosome number, n=10 pairs.

Nesom & Turner: Systematics of Sedum parvum group 263

Texas, Oklahoma, Arkansas, Missouri; open areas in shallow soil, commonly over
granite or sandstone, usually in the area of oak or oak-juniper woods; Apnil-July.

Probably most closely related to Sedum tamaulipense, sharing with it distinctively
small petals and spreading, ventrally gibbous follicles.

6. SEDUM NANIFOLIUM Frod.

Sedum nanifolium Frod., Acta Horti Gotob. 10, App. 196. 1936. Sedum parvum
Hemsl. subsp. nanifolium (Frod.) R.T. Clausen, Bull. Torrey Bot. Club 105:223.
1978. TYPE: MEXICO. Coahuila: Chojo Grande, 27 mi SE of Saltillo, 16 Jul
1905, Palmer 722 (HOLOTYPE: UC; Isotype: GH!).

Erect perennials to 20 cm high, homophyllous or rarely producing densely leafy
shoots. Stems smooth, prominently reddish-shiny on the lower portions, cells
elongated. Leaves widely obovate to obovate-oblong, half-terete (flat adaxially), green
with prominent red dots, sometimes waxy, 2.5-3.5(-4.0) (-5.0 in Texas) mm long,
2.0-2.5 mm wide, even in length on upper and lower portions of the stem, cells
quadrate. Petals erect, 4.0-5.1 mm long, yellow with prominent, short, red,
longitudinal stripes. Chromosome number, n=26 pairs.

Widespread in eastern Chihuahua, Coahuila, central Nuevo Le6n, and apparently
rare in Brewster Co., Texas; limestone gravel, ledges, or crevices, in matorral,
chaparral, or pinyon pine woods with juniper, oak, or agave; (1200-) 1600-2300
(-2700) m; June-November.

Additional collections examined: MEXICO. Chihuahua: NW end of the Sierra

del Diablo, ca. 27° 20' N, 29 Jul 1941, Stewart 98] (GH). Coahuila: W of El Chorro
and ESE of Saltillo, 22 Jun 1978, Clausen 78,3 (BH-2 sheets); W of Chorro Grande,

25523) N, 100° 48’ W, 23 Apr 1949, Clausen 7607 (BH-2 sheets, GH); 17 mi S of
Arteaga, 18 Aug 1948, Kenoyer & Crum 2771 (GH); Del Carmen Mts., 2 Sep 1936,
Marsh 862 (TEX); Sierra de Santa Rosa, NW of Muzquiz, 25 Jul 1938, Marsh 1476
(GH,TEX); 3 mi N of Puerto Flores, 8 Nov 1957, Moran 6309 (BH); Sierra de la
Encantada, 10 km NW of Rancho Buena Vista, 5 Sep 1941, Stewart 1428 (GH,
TEX); near Linos, SE of Saltillo, Strauss s.n. (BH); Cafion de la Barrica, Sierra de la
Madera, 20 Aug 1975, Wendt 1218 (TEX); Sierra del Carmen, E of Pico de Cerda, 11
Aug 1974, Wendt 563A (LL); Sierra del Carmen, 7.9 mi N of Rancho El] Jardin on
road to Mina El Popo, 22 Sep 1973, Wendt et al. 63 (LL). Nuevo Leén: Cafiada

Zacatosa, 6 km N of La Escondida, 24° 09’ N, 99° 55’ W, 30 Aug 1977, Clausen
77,32 (BH-2 sheets); Hwy 51 between Dr. Arroyo and Galeana, 34 km S of ject with
Hwy 58 at Puerto de Pastores, 28 Jun 1978, Cochrane et al. 8459 (BH); 2 mi S of
Pablillo, 20 Jul 1958, Correll & Johnston 19889 (LL); Hwy 68, 17.7 mi S of jet of
Hwy 60 and 1.6 mi N of Puerto de Cieneguillos, 24 Sep 1973, Reveal 3409 (BH);
Hacienda Pablillo, Galeana, 8 Aug 1936, Taylor 103 (TEX)

UNITED STATES. Texas: Brewster Co.: on limestone hills in valley at the S
end of Del Nortes, Hinckley 4114 (BH); Doubtful Canyon, Del Norte Mts., Gage
Estate, 25 mi S of Alpine, 18 Sep 1947, Warnock & Hinckley 7521 (SRSC); Cox

264 PRY ROL GT: October 1995 volume 79(4):257-268

Ranch, 15 mi SE of Alpine, 21 Aug 1960, Warnock 18644 (SRSC); Doubtful
Canyon, Del Norte Mts., 25 Sep 1967, Warnock 21320 (SRSC).

7. SEDUM CHRYSICAULUM J.A. McDonald

Sedum chrysicaulum J.A. McDonald, Sida 14:315. 1991. TYPE: MEXICO. Nuevo
Leon: Mpio. Rayones, summit of Sierra La Marta, ca. 3600 m, 24 Aug 1980, I fp
A. McDonald & M. Mayfield 2556 (HOLOTYPE: TEX!).

Erect perennials 5-9 cm high. Stems smooth, prominently golden-shiny on the
lower portions, cells elongated. Leaves narrowly oblanceolate-oblong, flat, green
with prominent red dots, 2.0-2.5 mm wide, 5-6 mm long on lower stem, 7-10 mm
long on the upper portions. Petals erect, 5.0-7.5 mm long, yellow with prominent,
short longitudinal, red stripes.

Southeastern Coahuila (Sierra La Viga) and central Nuevo Leén (Sierra La Marta,
Cerro Potosf, Sierra Pefia Nevada); grassy subalpine to alpine meadows, often with
Pinus hartwegii and P. culminicola; 3400-3800 m; August-October (November).

Additional collections examined: MEXICO. Coahuila: Mpio. Arteaga, summit of
Sierra La Viga, ca. 3600 m, 24 Oct 1984, McDonald & Gomez 1157 (TEX); Sierra La
Viga, 3700 m, 22 Aug 1986, McDonald 2099 (TEX); Sierra La Marta, 22 Aug 1986,
McDonald 2136 (TEX). Nuevo Le6n: Mpio. Arambern, Cerro Viejo, 3400 m, 20
Nov 1993, Hinton et al. 23971 (TEX); Mpio. Doctor Arroyo, Sierra de Pefia Nevada,
N of Picacho de San Onofre, ca. 3400 m, 30 Nov 1984, McDonald & Gomez 1298
(TEX); Mpio. Galeana, summit or near summit of Sierra La Marta, 3600 m, 31 Aug
1980, Hinton et al. 17977 (TEX), Sierra La Marta, 3680 m, 4 Aug 1980, Hinton et al.
17919 (TEX); 25 Oct 1984, McDonald & Gomez 1242 (TEX); 22 Aug 1986,
McDonald 2136 (TEX); summit or near summit of Cerro Potosi, 23 Aug 1984, Lavin
4787 (TEX); Cerro El Potosi, 3810 m, 14 Oct 1970, Hinton et al. 17303 (TEX).

This is the most widespread of the high-elevation Sedum species of northeastern
México. It is most closely related to S. nanifolium, with which it shares an erect habit,
shiny stems with elongated cells, and the distinctive red markings in the leaves and
petals.

8. SEDUM CATORCE Nesom, nom. et stat. nov.
Sedum parvum Hemsl. subsp. dendroides R.T. Clausen, Bull. Torrey Bot. Club
105:223. 1978. TYPE: MEXICO. San Luis Potosi: 0.5 km W of Real de

Catorce, 23° 41’ 24” N, 100° 53’ 32” W, cliff of quartzite exposed to southwest,
N side of canyon, 2620 m, 23 Aug 1977, R.T. Clausen 772.028--pressed from
greenhouse-grown plants (HOLOTYPE: BH!; Isotype: BH!). Non Sedum
dendroideum DC.

Erect to semi-erect, slightly glaucous perennials 10-25 cm high, roots said to be
tuberous. Stems strongly woody, sometimes somewhat pendant from cliff sides,
reddish, minutely papillate, not at all shiny, cells quadrate. Leaves homophyllous,
lanceolate-oblong, flat but from a swollen base, (3.0-)3.5-5.0 mm long, 1.3-2.0 mm

Nesom & Turner: Systematics of Sedum parvum group 265

wide, dark green with prominent white margins, venation usually clearly discernible.
Petals erect to spreading or reflexed, 6-7 mm long, yellow.

Known only from cited collections.

9. SEDUM DULCINOMEN Neson, spec. nov. TYPE: MEXICO. Nuevo
Leon: [Mpio. Zaragoza], 2 mi E of Dulces Nombres, succulent on limestone
outcrops, 1850 m, 28 Jun 1948, F.G. Meyer & D.J. Rogers 2699 (HOLOTYPE:
BH!).

Sedo catorce Nesom, S. papillicaulo Nesom, et S. macdonaldii Nesom
caulibus papillatis similis sed distinctus habitu prostrati-decumbenti caulibus ex
rhizomatibus honzontalibus radicibus adventitiis orientibus; Sedo catorce
similis caulibus ac foliis glaucis.

Prostrate-decumbent perennials from honzontal, slightly woody rhizomes,
producing fibrous, adventitious roots, stems and leaves heavily glaucous (less so in
cultivation). Stems green, becoming reddish-tinted, but without discrete dots of red
pigment, minutely papillate with quadrate cells, arching upward or erect and ansing
from the rhizomes. Leaves heterophyllous, green, heavily glaucous (less so in
cultivation), flat, elliptic-oblong, 1.5-2.0 mm wide, 2.5-4.5 mm long. Petals erect,
yellow, 4.5-6.0 mm long.

Nuevo Leon, on the Tamaulipas border near Dulces Nombres; limestone ledges
and outcrops in pine woods; 1750-2000 m; February-June.

Additional collections examined: MEXICO. Nuevo Leén: Mpio. Zaragoza: ca.
16 km E of mine in Distnct of Dulces Nombres, Feb 1950, J.L. Edwards s.n.--
pressed from greenhouse cultivar (BH); ca. 3 km SE of Santa Teresa, “39” Jan 1980.
Clausen U2724--pressed from greenhouse cultivar (BH).

The three collections studied of Sedum dulcinomen are very similar among
themselves. The specimen collected from nature (the type), is strongly glaucous, but
the greenhouse-grown plants show clear traces of a waxy surface. Among the other
species treated in this study, only S. catorce produces a glaucous covering.

10. SEDUM PAPILLICAULUM Nesom, spec. nov. TYPE: MEXICO. Nuevo
Leén: Mpio. Zaragoza, Sierra de Pefla Nevada, Picacho San Onofre, fir and pine
forest, 3000 m, 18 Jun 1979, Hinton et al. 17551 (HOLOTYPE: TEX’).

Sedo catorce Nesom et S. macdonaldii Nesom habitu erecto et caulibus
papillatis similis sed distinctus paginis non glaucis, foliis planis, et papillis
caulinis columnanbus structuram cellulosam perspicuam carentibus.

Erect, fibrous-rooted perennials 6-25 cm high. Stems suffruticose, prominently
minutely and densely papillate, the papillae columnar and sometimes appearing
stipitate-glandular, the cellular structure not readily apparent. Leaves oblong-elliptic to
lanceolate-oblong, flat, 3-4 mm long, even in length on the upper and lower portions

266 PHYTOLOGIA October 1995 volume 79(4):257-268

of the stems, 1.5-2.2 mm wide, the cells quadrate in the distal portions, elongate in the
swollen basal portion. Petals erect, yellow, rarely with a reddish tinge, 5-7 mm long.

Nuevo Le6n; subalpine and alpine meadows of Sierra Pefia Nevada and vicinity,
usually with Pinus hartwegii or pine-fir, sometimes in oak-agave woodland; (2700-)
3000-3600 m; June-August.

Additional collections examined: MEXICO. Nuevo Leén: Mpio. Doctor Arroyo:
ridge and E side of Pefia Nevada, 5 Jul 1985, McDonald 1642 (TEX); trail from
Cafion La Tinaja to La Encantada, 4 Jul 1988, Patterson 5837 (TEX); N and NW
slope of Picacho Onofre, 10-15 Jul 1977, Wells & Nesom 369 (TEX). Mpio.
Zaragoza: Cerro El Viego, 1800 m, 7 Jul 1992, Hinton et al. 22125 (TEX); Cerro Hl
Viego, 3360 m, 6 Oct 1992, Hinton et al. 22147 (TEX); 9 km N of La Encantada,
2700 m, 25 May 1992, Herndndez et al. 2284 (TEX); 2 m NE Cerro Pefia Nevada,
2690 m, 23 Aug 1989, Nesom 7121] (TEX). Tamaulipas: 15 km NW Estanque de los
Walle, 2000 m, 25 Oct 1989, Herndndez S. 2063 (TEX).

A distinctive species restricted to the Pefia Nevada area of southeastern Nuevo
Leon but closely similar to Sedum macdonaldii, which appears to be its northern
vicariad.

11. SEDUM MACDONALDII Nesom spec. nov. TYPE: MEXICO. Nuevo
Leén: Mpio. Galeana, Sierra La Marta, S and SE sides at the top, alpine and subalpine
zone, 22 Aug 1986, Andrew McDonald 2135 (HOLOTYPE: TEX!); Isotypes:
MEXU,BH).

Sedo catorce Nesom et S. papillicaulo Nesom habitu erecto et caulibus
papillatis similis sed distinctus paginis non glaucis, foliis semi teretibus et
cellulisquadratis in lineis papillas caulinas formantibus.

Erect fibrous-rooted perennials 4-7 cm high. Stems mostly obscured by the leaves
but the surfaces low-papillate with quadrate cells in lines. Leaves half-terete, flat
above with a medial sulcus, both surfaces with quadrate cells from tip to base,
minutely striate-papillate, the cellular structure clearly perceptible. Petals yellow,
erect, 6-7 mm long.

Coahuila (Sierra Coahuil6n, Sierra La Viga), Nuevo Leén (Cerro Potosi and Sierra
La Marta); subalpine and alpine zones, often with Pinus hartwegii, Pinus culminicola,
or Pseudotsuga; 2850-3600 m; July-October.

Additional collections examined: MEXICO. Coahuila: Mpio. Arteaga, ndge and
SE side of Sierra Coahuilén, 22 Jul 1985, McDonald 1762 (TEX); summit of Sierra
La Viga, 24 Oct 1984, McDonald & Gomez 1158 (TEX). Nuevo Leén: Mpio.
Galeana: Sierra La Marta, near top, 5 Jul 1981, Hinton et al. 18310 (TEX); SE side of
Cerro Potosf, 25 Jun 1960, Beaman 3321 (GH); near top of Cerro Potosi, 3500 m, 23
May 1988, Westlund 23 (TEX).

Sedum macdonaldii apparently is most closely related to §. papillicaulum, which
differs in its flat (dried) leaves with a basal area of elongated cells and its strongly
stipitate-papillate stems, the cellular structure of which is not at all discernible.

Nesom & Turner: Systematics of Sedum parvum group 267

INCERTAE SEDIS

Sedum robertsianum E.J. Alexander, Bull. Torrey Bot. Club 63:201. 1936. Sedum
parvum Hemsl. subsp. robertsianum (E.J. Alexander) R.T. Clausen, Variation
Spec. Sedum 16. 1981. TYPE: UNITED STATES. Texas: Brewster Co.,
mountain top in shallow calcareous soil, 4000 ft, A.R. Davis s.n. (HOLOTYPE:
NY, from cultivar of Davis collection.)

Clausen (1981) could not find the type at NY and made the following comment:
“Because no type is at the New York Botanical Garden, a part of the type matenal,
made available by Mr. Alexander and cultivated and pressed at Cornell University on
July 22, 1937, may serve as the lectotype. The specimen is in the herbarium at
Cornell University.”

[It] “combines features of the other subspecies: longer leaves (8.6 mm) as in ssp.
diminutum, wider leaves (3.7 mm) as in ssp. nanifolium, longer anthers (1.1 mm) as
in ssp. dendroides, narrower nectaries (0.4 mm) as in spp. diminutum, and later
flowering (Aug.-Sept.) as in ssp. parvum. It is the most herbaceous of the five
subspecies. Otherwise, it is similar to the other subspecies in having cymes of 1-2
cincinni, yellow flowers, gibbous follicles, and fuscous, papillose seeds.” Clausen
(ms), in his forthcoming treatment of Sedum for the Flora of the Chihuahuan Desert
(Henrickson, in prep.), places S. robertsianum in synonymy under S. parvum Hemsl.

Erect, fibrous-rooted perennials. Stems [papillate?], both stems and leaves “red-
streaked and spotted.” Heterophyllous, the leaves 5-8 mm long, 3-4 mm wide,
subterete (fresh), papillate, the cells quadrate. Petals yellow, 4 mm long, spreading-
reflexed. Carpels erect, the follicles spreading, baso-ventrally gibbous.

ACKNOWLEDGMENTS

We thank the staffs of BH, GH, SRSC, and US for providing us with loans of
selected specimens. We are indebted to Mark Mayfield, Andrew McDonald, and
Gayle Tumer for reviewing the manuscript.

LITERATURE CITED

Berger, A. 1930. Crassulaceae. Jn: A. Engler & K. Prantl, Nat. Pflanzenfam. ed. 2.
18a:352-483.
Bnitton, N.L. & J.N. Rose. 1950. Sedum in N. Amer. Fl. 22:7-74.
Clausen, R.T. 1959. Sedum of the trans-Mexican volcanic belt: an exposition of
taxonomic methods. 380 pp. Ithaca, New York.
. 1975. Sedum of North America north of the Mexican Plateau. 742 pp.
Ithaca, New York.

268 PHY T OLOGIA October 1995 volume 79(4):257-268

. 1978. Sedum--seven Mexican perennial species. Bull. Torrey Bot.
Club 105:214-223.

1979. Sedum in six areas of the Mexican Cordilleran Plateau. Bull.
Torrey Bot. Club 106:216.
. 1981. Variation of species of Sedum of the Mexican Cordilleran Plateau.
Arnold Printing Corp., Ithaca, New York.
. 1984. Sedum (Crassulaceae) of the Mexican Cordilleran Plateau.
Gentes Herb. 12:8-48.
Froderstrom, H. The genus Sedum L., a systematic essay. Part 1. Acta Horti Gotob.
5 Appendix): 1-75. 1930; Part II, Ibid 6 (Appendix): 1-111. 1931: Part III. Ibid
7 (Appendix): 1-126. 1932; Part IV. Ibid 10 (Appendix): 1-262. 1935.
McDonald, J.A. 1991. Plantae alpinae novae Mexicanae: Sedum chrysicaulum
(Crassulaceae). Sida 14:315-319.
Nesom, G.L. 1988. New species of Crassulaceae from northeastern México. Sida
13:21-24.

Praeger, R.L. 1921. An account of the genus Sedum as found in cultivation. J. Royal
Hort. Soc. 46:1-314.

Phytologia (October 1995) 79(4):269-280

THE GENUS TETRANEMA (SCROPHULARIACEAE) IN COSTA RICA, WITH
TWO NEW SPECIES

Michael H. Grayum & Barry E. Hammel
Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri 63166 U.S.A.

ABSTRACT

Two new species of Tetranema (Scrophulariaceae) are described from
Costa Rica’ T. gamboanum Grayum & Hammel, known from wet forests
on both slopes, and T. floribundum Hammel & Grayum, endemic to Cerro
Turrubares in the mid-Pacific region. Both are unusual in having a long-
stemmed growth habit and red, tubular corollas presumably adapted for
pollination by hummingbirds. The new species are most similar, at least in
floral morphology, to the Mexican endemic Tetranema megaphyllum
(Brandegee) L.O. Williams. These are the first records of the genus from
south of Honduras, and increase the total number of species from four to six.

KEY WORDS: Costa Rica, Scrophulanaceae, Tetranema, systematics

A wealth of botanical material gathered during the exploration of a remote region
on the Atlantic slope of Costa Rica’s Cordillera de Talamanca in April, 1989, included
one particularly remarkable collection made by Costa Rican botanist Gerardo Herrera.
This collection was remarkable in representing a conspicuous, terrestnal herb--an
astend dicot with bright red, tubular corollas ca. 5 cm long--that we were unable to
identify even to the generic level. Though its flowers superficially resembled those of
some Acanthaceae known from the region [Odontonema tubaeforme (Bertol.) Kuntze,
Razisea spicata Oerst.], the Herrera collection was soon identified as belonging to
Scrophulariaceae. However, the combination of a caulescent, subshrubby growth
habit and axillary, long-pedunculate, bracteolate, cymose inflorescences seemed
incompatible with any known genus; indeed, we entertained the notion of establishing
a new genus to accommodate this collection and other, similar matenal that has
subsequently emerged from Costa Rica.

With respect to their shrublike habit, axillary, cymose inflorescences and red,
tubular corollas, the abovementioned Costa Rican collections suggest the genus
Russelia Jacq., of the monotypic tnbe Russeliae. Russelia differs, however, in having
septicidal capsules densely packed with hairs. The Costa Rican matenal better
concords with tnbe Cheloneae sensu Thieret (1954), charactenzed by bracteolate,

269

270 PHY DP OLOGITA October 1995 volume 79(4):269-280

cymose or racemose inflorescences and sterile posterior stamen filaments. The only
members of this group occurring naturally in the Mesoamencan region are the large
genus Penstemon Schmidel and the oligotypic Tetranema Benth. ex Lindl. and
Uroskinnera Lindl. (though none of these have been recorded from south of
Honduras). Each of these three genera includes at least one species with red, tubular,
presumably hummingbird-pollinated flowers (see, e.g., Daniel & Breedlove 1992).

Tetranema differs from Penstemon and Uroskinnera in having axillary and cymose
(rather than terminal and racemose or thrysoid) inflorescences and much reduced
sterile stamens (staminodes); it further differs from Uroskinnera in having distinct
sepals, and from Penstemon in having loculicidal capsules. In all of these cntical
details, the Costa Rican material accords perfectly with Tetranema. Moreover, the
seeds of the one Costa Rican collection examined in this regard (Figure 2) are a very
convincing match for those of Tetranema roseum (M. Martens & Galeotti) Standl. &
Steyerm., as illustrated by Beaufort-Murphy (1983: Pl. 4G) (who, unfortunately, did
not study Uroskinnera or Penstemon).

Our initial attempts to identify the Costa Rican Tetranema collections to genus level
were thwarted by our reliance on Standley & Williams’s (1973) Flora of Guatemala
Scrophulariaceae treatment. In their generic key (p. 321), the leads are inverted in the
couplet purporting to separate Tetranema from Uroskinnera and Penstemon (as pointed
out by Daniel & Breedlove 1992). Furthermore, the three Tetranema species attnibuted
to Guatemala are all quite unlike the Costa Rican matenal in being acaulescent or short-
stemmed herbs with campanulate, white or purple (fide Standley & Williams) corollas.

Tetranema has heretofore been considered a genus of four species, ranging from
southern México (Puebla) to Honduras (Méndez-Lanos & Villasefior 1995).
Tetranema roseum, the most wide-ranging species, is of modest horticultural repute as
a glasshouse plant, with at least two cultivars available commercially under the name
“Mexican foxglove” (Morrison 1981).

The Costa Rican matenal of Tetranema is here treated as comprising two species
new to science, bringing the generic total to six. Tetranema gamboanum Grayum &
Hammel is represented by the Herrera collection from the Atlantic slope and several
subsequent collections from wet-forest sites on the Pacific slope, while T.
floribundum Hammel & Grayum is known only by three collections from Cerro
Turrubares, an isolated peak in the central Pacific region.

TETRANEMA GAMBOANUM Grayum & Hammel, spec. nov. TYPE: COSTA
RICA. Puntarenas: Cant6n de Osa, Fila Costefa, cabeceras del Rio Piedras

Blancas, Cerro Anguciana, 8° 49’ 12” N, 83° 11’ 15” W, 900 m, 7 Dec 1993 (f1.,

fr.), Aguilar et al. 2700 (HOLOTYPE: INB!; Isotypes: BM!,CAS!,CR!,F!,
MEXU!,MO!,NY!,US!). Figures 1-2.

Species cum Tefranemata megaphyllo (Brandegee) L.O. Williams optime
congruens sed differt foliis apice longiacuminatis bracteis inflorescentia
brevioribus corolla longiore lobis corollae multo longioribus.

Grayum & Hammel:

Tetranema in Costa Rica 271

Figu

igure 1. Tetranema gamboanum. A.
flower (Aguilar et al. 2700); C. fruit (Hammel et al. 19429).

flowenng shoot (Aguilar et al. 2700); B.

DZ PHY TOrOGiA October 1995 volume 79(4):269-280

Figure 2. Tetranema gamboanum (Hammel et al. 19542), seed: x 150 (photo by
Betty Strack).

Grayum & Hammel: Tetranema in Costa Rica 273

Erect, decumbent-based herbs 1-2+ m tall. Internodes to at least 11 cm long,
strigulose when young. Petioles obsolete to ca. 1 cm long, strigulose, canaliculate
above, the margins ciliate proximally, the hairs extending in a line across the node.
Leaves 14-31 x 5-11 cm, elliptical to oblanceolate or spatulate, long-acuminate at

apex, attenuate to the base (where decurrent onto the petiole), the margins + coarsely
serrate, glabrous above or with few, distant hairs along the midnb and major veins
(especially proximally), pubescent along the veins below, midrib often falcate, pnmary
lateral veins ca. 9-13 per side, prominulous on both sides when dry. Inflorescences
axillary, cymose; peduncle 9-24 cm long, divergent, green, quadrangular with the
angles narrowly winged. Flowers ca. 2-12 per inflorescence, bracteate, the bracts
0.5-2.0 mm long, subulate to narrowly triangular, ciliate on margins; pedicels ca. 9-11
mm long at anthesis, to ca. 20 mm in fruit, glabrous; calyx 5-merous, divided nearly

to base, the lobes ca. 3-5 mm long at anthesis (to ca. 6 mm in fruit), + narrowly to

broadly ovate, imbricate, + cornute apically, ciliate on margins; corolla ca. 4.9-5.5 cm
long, scarlet, tubular, slightly curved (convexly) upward and gradually expanded
distally, glabrous throughout or (Herrera & Chacon 2644) with flat hairs at the mouth
and onto the lower lobes, the lobes 4, 11-13 x 3.0-3.5 mm, imbnicate, lanceolate, the
3 lower ones obtuse to rounded apically and spreading-reflexed, the upper one
emarginate and slightly wider; fertile stamens 4, exserted from the throat (but not
exceeding the upper corolla lobe); filaments attached at base of corolla tube, glabrous,

+ dilated toward base; anther sacs 0.8-1.0 mm long, confluent apically and becoming
divergent, glabrous; staminode ca. 1.5-2.0 mm long; ovary 3-4 mm long, narrowly
ovoid, glabrous; style exserted, glabrous; stigma clavate to funnelform, hollow, the
nm papillose; fruit a loculicidal capsule, ca. 6-9 mm long, subglobose-apiculate; seeds
ca. 0.6-0.7 x 0.45 mm, oblong, amber to black, densely foveolate.

Additional specimens examined: COSTA RICA. Limon: Cordillera de
Talamanca, entre Cerro Muchilla y Cerro Avioneta, cabeceras de Rio Suruy, Fila
Matama, 9° 47’ 25” N, 83° 06’ 30” W, 550 m, 17 Apr 1989 (fl.), Herrera & Chacdén
2644 (BM,CR,INB,MEXU,MO,USJ). Puntarenas: Cantén de Osa, upper head

waters of Rio Piedras Blancas, W slopes of Cerro Anguciana, Fila Cruces, 8° 49’ 12”

N, 83° 11’ 09” W, 950-1,150 m, 10 Dec 1993 (f1.), Grayum 10663 (CAS,BM,CR,
F,INB,MEXU,MO); same locality, 7 Dec 1993 (fl., fr.), Hammel et al. 19200 (CR,
INB,F,MO). San José: Canton de Pérez Zeledén, Fila Costefia, Fila Tinamastes, por

la carretera entre Dominical y San Isidro, 9° 18’ 43” N, 83° 46’ 19” W, 950 m, 3 Feb
1994 (fl., fr.), Hammel et al. 19429 (COL,CR,INB,MICH,MO,TEX); same locality,
28 Mar 1994 (f1., fr.), Hammel et al. 19542 (CR,INB,MO; live at MO).

Tetranema gamboanum is endemic to Costa Rica, where it is known by a single
collection from the Atantc slope of the Cordillera de Talamanca (Fila de Matama) at
550 m elevation, and from two widely separated sites in the Pacific Fila Costefia at ca.
900-1,000 m (Figure 3). All of these stations appear to lie in the Premontane Rain
Forest Life Zone of the Holdridge system (cf. Tosi 1969). Flowering matenal of T.
gamboanum has been collected from December through Apnil.

As mentioned previously, Costa Rican matenal of Tetranema does not concord
with any of the three species treated in the Flora of Guatemala (Standley & Williams

274 PHYTOLOGIA October 1995 volume 79(4):269-280

1973). It does, however, compare reasonably well with the Chiapan endemic 7.
megaphyllum (Brandegee) L.O. Williams, at least in terms of gross floral
morphology. The original description of Allophyton megaphyllum Brandegee (1914)
specified tubular, red corollas (“Corollae tubus cylindraceus . . . Corollae coccineae”’)
that “resemble those of Russelia,” and subsequent descriptions agree on this point.
This is the only Tetranema species from north of Costa Rica that has tubular corollas,
though those of T. evolutum Donn. Sm. may be red (fide Méndez-Lanos & Villasenor
1995; Standley & Williams described them as “bright purple”).

Tetranema gamboanum would seem to differ from T. megaphyllum in comprising
taller (1-2+ m), coarser plants. Although the specimens of the latter species studied by
Brandegee (1914) were “not complete enough to give the size of the plant,” the leaves
were said to be “crowded,” suggesting that the plants may have been short-stemmed.
Pennell (1925), the first to ally the “most remarkable” Allophyton megaphyllum with
Tetranema (using the name Allophyton Brandegee for the entire group), stated that “all
the species of Allophyton have short stems,” more specifically, “1 dm long or less.”
Pennell cited three duplicates of a topotype collection (Purpus 7921) not cited by
Brandegee. Méndez-Larios & Villasefior (1995), citing three additional collections not
seen by previous authors, characterize T. megaphyllum as “la especie con desarrollo
vegetativo mds vigoroso”; nevertheless, they descnbe it as having “tallos muy
reducidos,” 25-40 cm tall.

Although we have been unable to obtain the holotype of Tetranema megaphyllum
on loan, we have studied an isotype (Purpus 6855 [NY]), as well as the NY duplicate
of the topotype cited by Pennell (1925). While neither of these specimens bears label
data indicating either the habit of the plants or the color of the corollas, the following
differences from T. gamboanum are manifest: T. megaphyllum has inflorescence
bracts to ca. 10 mm long and corollas ca. 2.5-3.6 cm long with rounded, apparently
forward-directed lobes ca. 2-4 mm long; T. gamboanum, on the other hand, has
inflorescence bracts to ca. 2 mm long and corollas ca. 5 cm or more long with
elongate, spreading-reflexed lobes ca. 11-13 mm long. These observations are
corroborated by Méndez-Larios & Villasefior’s (1995) description of T. megaphyllum.

The occurrence of Tetranema gamboanum on both the Atlantic and Pacific slopes
has innumerable precedents in the Costa Rican flora. The lone collection from the
Atlantic slope (Herrera & Chacoén 2644) is essentially a perfect match for the Pacific
material, except for the unusual corolla hairs noted in the description. Whether or not
these hairs are characteristic of Atlantic populations, and thus potentially indicative of
infraspecific rank, cannot be decided without additional material.

Tetranema gamboanum is probably more widespread in Costa Rica than our
scattered records indicate; it may also yet be found in Panama. Although it is locally
more or less abundant, none of the three known stations lies within a protected area.
This appears to be a species of relatively undisturbed habitats.

We take great pleasure in dedicating this new species to William Gamboa Elizondo
(1958- ) of Las Mellizas de Coto Brus, Costa Rica, who has_ participated
enthusiastically in virtually every major botanical expedition into the Cordillera de
Talamanca since 1983 as cook, porter, scout, negotiator, and occasional collector.

Grayum & Hammel: Tetranema in Costa Rica 275

86.00 85.00 84.00 83.00

11.00

10.00

et @ Tetranema floribundum

m@ Tetranema gamboanum

8.00

Figure 3. Distnbution of Tetranema in Costa Rica (500 m contour is indicated).

276 PHYTOLOGIA October 1995 volume 79(4):269-280

TETRANEMA FLORIBUNDUM Hammel & Grayum, spec. nov. TYPE:
COSTA RICA. San José: lado N de Cerro Turrubares, al S de San Rafael por

Quebrada Pital, 9° 48’ 05” N, 84° 27’ 52” W, 1,200-1,300 m, 5 Jan 1996 (f1.,
fr.), Hammel, Jiménez, & Morales 20068 (HOLOTYPE: INB!; Isotypes:
BM!,CR!,F!,MO!). Figure 4.

Species ex affinitate Tetranematis megaphylli (Brandegee) L.O. Williams et
T. gamboani Grayum & Hammel, ab utroque inflorescentiis omnibus (8-)14-
30-floris tubo corollae intus ventraliter in longitudinem pubescenti distincta.

Erect, decumbent-based herbs (0.35-)0.80-2.00 m tall, often rooting at decumbent
nodes. Internodes to at least 5 cm long, densely matted-, arachnoid-, or woolly-
pubescent when young. Petioles essentially obsolete, the often undulate margin of the
leaf blade reaching nearly to the node. Leaves 21.0-23.5 x 9-13 cm, broadly elliptic
to oblanceolate or spatulate, rounded, abruptly acute or short-acuminate at apex, acute
to mostly concavely and abruptly attenuate to the base, the margins coarsely serrate to
undulate-toothed, glabrous above except on the midnb at the very base, strigulose on
the midrib and main veins below and minutely scaly (and thus shiny, when dry)
throughout the abaxial leaf surface, midrib occasionally falcate, primary lateral veins 8-
10(-11) per side, prominent below. Inflorescences axillary, cymose; peduncle 13-23
cm long, purple, quadrangular with the angles narrowly winged. Flowers ca. (8-)14-
30 per inflorescence, bracteate, the bracts 1-5 mm long, narrowly tnangular, ciliate
(often only at base) on margin; pedicels ca. 10 mm long at anthesis, to ca. 20 mm in
fruit, glabrous; calyx 5-merous, divided nearly to base, the lobes 2-3 mm long at

anthesis (to 4 mm in fruit), broadly ovate, + cormute apically, ciliate on margins;
corolla 2.6-3.5 cm long, red, tubular, gradually slightly curved (convexly) upward
and expanded distally, glabrous externally, internally pubescent with a narrow band of
flat, yellow hairs (to ca. 1 mm long) on the ventral surface of the tube from near the
base to the mouth and often all along the median lower lobe, the lobes 4, ca. 13 x 2.5-

5.5 mm, + lanceolate, the 3 lower ones rounded apically and spreading-reflexed, the
upper one emarginate and slightly wider; fertile stamens 4, exserted from the throat
(but held just below the upper corolla lobe and not exceeding it); filaments attached at
the base of the corolla tube; anther sacs 0.8-0.9 mm long, confluent apically (where
attached to the filament), divergent at dehiscence (full length) and then broadly elliptic,
glabrous; staminode ca. 0.5 mm long; ovary ca. 3.5 mm long, narrowly ovoid,
glabrous; style exserted (with the stamens), glabrous; stigma clavate, hollow; fruit to
ca. 8 mm (immature), ovoid.

Additional specimens examined: COSTA RICA. San José: Canton de
Turrubares, Z. P. Cerros de Turrubares, Potenciana arnba, cerca del Cerro
Turrubares, 9° 48’ 00” N, 84° 27’ 10” W, 1,600 m, 4 Mar 1993 (f1., fr.), Jiménez et
al. 1155 (BM,CR,INB,MO); Z. P. Cerros de Turrubares, Cerros de Puriscal, sector
San Rafael, Sitio Cerro Pelén, 09° 49’ 00” N, 84° 28’ 50” W, 1,200 m, 6 Dec 1991
(fl.), Zuiniga 599 (INB).

277

Tetranema in Costa Rica

Grayum & Hammel:

rh

as

Figure 4. Tetranema floribundum. A. flowering shoot (Jiménez et al. 1155), B.

corolla (Hammel et al. 20068); C. fruit (Jiménez et aa T1S5):

278 Det ye 1 OULO GION October 1995 volume 79(4):269-280

Tetranema floribundum is endemic to Costa Rica, where it is known only from
Cerro Turrubares, an isolated peak in the central Pacific region, at 1,200-1,600 m
elevation (Figure 3). This region apparently corresponds to the Lower Montane Rain
Forest Life Zone of the Holdridge system (cf. Tosi 1969). The three flowering
collections of T. floribundum are from December, January, and March.

The specific epithet of Tetranema floribundum reflects the fact that it has more
flowers (ca. 14-30) per inflorescence, on average, than any other known Tetranema
species (JT. roseum may have as many as 20). It differs additionally from T.
gamboanum and T. megaphyllum, the only other species with red, tubular corollas, in
having the corolla tube internally pubescent along the ventral surface. The corollar
pubescence of the sole Atlantic slope collection of 7. gamboanum, discussed
previously, does not extend into the tube. Tetranema floribundum is further
distinguished from 7. gamboanum in having (as T. megaphyllum) merely acute to
short-acuminate (rather than long-acuminate) leaf apices and smaller corollas, and from
T. megaphyllum in having (as T. gamboanum) longer stems and generally larger
corollas with relatively and absolutely much longer, spreading-reflexed lobes.

It may seem unusual that Tetranema populations on Cerro Turrubares, in the
central Pacific region of Costa Rica, should differ specifically from populations in the
southern Pacific region, while the latter populations should be conspecific with
matenal from the Atlantc slope (as discussed under T. gamboanum). Cerro
Turrubares, however, is relatively high and quite isolated, and is known to harbor
other endemic plant species (cf. Burger & Jiménez 1994). Tetranema floribundum
occurs at slightly higher elevations and, ostensibly, in a different life zone than T.
gamboanum.

Tetranema floribundum should presently be considered an endangered species,
since it is known from just a few populations in a site that has already been seriously
degraded by human activity. Two of the three collections were made within a
protected area (Zona Protectora Cerro de Turrubares), but from a region dominated by
pastures.

Both of the new Costa Rican Tetranema species described herein will come out to
T. megaphyllum in the key of Méndez-Larios & Villasefior (1995). The distinguishing
characteristics of these three species may be summarized as follows:

1. Corolla lobes ca. 2-4 mm long, < 1/5 the total corolla length, apparently directed
forward; floral bracts ca. 6-10 mm long; stems ca. 0.25-0.40 m tall; leaf apex acute
to short-acuminate; inflorescence 3-10-flowered; corolla ca. 2.5-3.6 cm long,

glabrous throughoutsChiapas®....ce.-ce:ccesnecdee catestee ceases T. megaphyllum.
1’ Corolla lobes ca. 11-13 mm long, > 1/5 the total corolla length, spreading-reflexed;
floral bracts 0.5-5.0 mm long; stems (0.35-)0.80-2.00 m tall; Costa Rica. ....... (2)

2. Inflorescence many- (14-30-) flowered, the peduncle purple; corolla 2.6-3.5
cm long, pubescent within in a band of flat, yellow hairs along the entire
ventral surface and onto the lower lobe; leaf apex rounded to short-acuminate;
CERrOnl UNTUDANES earth ee ciiccest een sae wats eae ane tae T. floribundum

Grayum & Hammel: Tetranema in Costa Rica 279

2' Inflorescence few- (2-12-) flowered, the peduncle green; corolla ca. 4.9-5.5 cm
long, glabrous throughout or (rarely) pubescent on lower lobe and at mouth;
leaf apex long-acuminate; Fila Costena and Atlantic slope of Cordillera de
SUNTAN Celeron cua ruins obs cae mere aires wa ep one eM ateee aac ua nels Peed T. gamboanum

The recent discovery of Tetranema in Costa Rica is surprising, especially since
both species comprise shrubby, understory plants with large, vividly scarlet corollas.
Though the distribution of the genus in Costa Rica appears spotty, 7. gamboanum, at
least, may be locally abundant. At the Tinamastes site, a sizeable population occurs
right at the roadside along a moderately well-botanized route (San Isidro de EJ General
to Dominical).

It is likely that earlier Costa Rican collections of Tetranema, not seen by us, will
yet be discovered filed as undetermined, or misdetermined, in some of the many
scattered herbaria housing Costa Rican matenal. As in the case of Ticodendron
(Ticodendraceae), another conspicuous Central American plant described only
recently, the belated recognition of Tetranema in Costa Rica is “perhaps explainable by
the fact that although it looks very much like something well known [e.g., an
Acanthaceae, Scutellaria, or Russelia], it really is something different” (Hammel &
Burger 1991: 92).

ACKNOWLEDGMENTS

The manuscript was cniically reviewed by Thomas B. Croat and Gordon
McPherson. We are also grateful to Silvia Troyo for the excellent line drawings; to
William C. Burger and Betty Strack for arranging and executing (respectively) the
SEM micrograph (Figure 2); to Jacqueline Kallunki for expediting the delivery of
important specimens from NY; and to Quirico Jiménez, for leading the second author
to the type locality of Tetranema floribundum. Field work was supported by National
Geographic Society grants 3317-86 and 4682-91 to the first author. Publication was
supported by National Science Foundation grant DEB-9300814 to both authors.

LITERATURE CITED

Beaufort-Murphy, H.T. 1983. The seed surface morphology of the Gesneriaceae
utilizing the scanning electron microscope and a new system for diagnosing seed
morphology. Selbyana 6:220-422.

Brandegee, T.S. 1914. Plantae mexicanae purpusianae, VI. Univ. Calif. Publ. Bot.
6:51-77.

Burger, W. & Q. Jiménez. 1994. A new species of Psychotria subgenus Psychotria
(Rubiaceae) from Costa Rica. Novon 4:206-208.

Daniel, T.F. & D.E. Breedlove 1992. A new species of Uroskinnera
(Scrophulanaceae) from southern Mexico. Madronio 39:13 1-136.

280 PHY TOLOGGIA October 1995 volume 79(4):269-280

Hammel, B. & W.C. Burger. 1991. Neither oak nor alder, but nearly: the history of
Ticodendraceae. Ann. Missoun Bot. Gard. 78:89-95.

Méndez-Larios, I. & J.L. Villasefior. 1995. Revisién taxondmica del género
Tetranema (Scrophulanaceae). Acta Bot. Mex. 32:53-68.

Morrison, P. 1981. Tetranema roseum (Mexican Foxglove) formerly Allophyton
mexicanum. Light Gard. 18:180-181.

Pennell, F.W. 1925. The genus Allophyton of southern Mexico and Guatemala
Proc. Acad. Nat. Sci. Philadelphia 77:269-272.

Standley, P.C. & L.O. Williams. 1973. Scrophulariaceae. Jn, P.C. Standley, L.O.
Williams, & D.N. Gibson (editors), Flora of Guatemala. Fieldiana, Bot.
24(9):3 19-416.

Thieret, J.W. 1954. The tribes and genera of Central American Scrophulariaceae.
Ceiba 4: 164-184.

Tosi, J.A., Jr. 1969. Mapa ecoldgico, Republica de Costa Rica: Segiin la
clasificacion de zonas de vida del mundo de L. R. Holdridge. San José, Costa
Rica: Centro Cientffico Tropical.

Phytologia (October 1995) 79(4):281-285

KEY TO THE AMERICAN GENERA OF ASTERINAE (ASTERACEAE)

Guy L. Nesom

Texas Regional Institute for Environmental Studies, Sam Houston State University,
Huntsville, Texas 77341 U.S.A.

ABSTRACT

An artificial key is provided for identification of Aster sensu stricto and the
fourteen genera that have been recently proposed to encompass the ca. 180
New World species segregated from Aster: Almutaster, Ampelaster,
Canadanthus, Chloracantha, Doellingeria, Eucephalus, Eurybia, lonactis,
Oclemena, Oreostemma, Psilactis, Sericocarpus, Symphyotrichum, and
Tonestus. Aster sensu stricto is represented by only a single species native to
the New World, A. alpinus. Also included in the key are Aster tataricus,
naturalized in eastern North Amenica, and the distinct genus Boltonia, which is
often associated with a group of Old World Aster.

KEY WORDS: Aster, Asteraceae, Asterinae, New World, systematics

In a systematic review of the genus Aster as it has been broadly conceived in recent
treatments, it was proposed that the ca. 180 American species of this alliance be
divided among a number of segregates (Nesom 1994). In this view, only a single
species of Aster sensu stncto occurs natively outside of the Old World: A. alpinus
grows in northern Eurasia and across Beringia into Alaska and southward along the
Rocky Mountain cordillera as far as Colorado. Aster tataricus, which is native to
northeast Asia, is naturalized in the eastern United States; as noted in the review, this
species probably should be placed in a genus separate from Aster sensu stricto. Only
Doellingeria among the American segregate genera also has species in the Old World.

Several of the genera included here (particularly Tonestus, Ionactis, Boltonia, and
Chloracantha) are ambiguous in their relative positions among other potentially related
genera (Nesom 1994). Tonestus kingii is the only species of that genus that has been
treated within Aster, and Tonestus may be more closely related to the Solidagininae
than to genera it is associated with among segregates of Aster. Jonactis has been
hypothesized to be related to Eucephalus and to the goldenasters, but it differs from
both in a number of cntical morphological features. Boltonia is isolated among
Amenican genera associated with Aster; it has long been considered to be closely
related to the Asian genus Kalimeris (an Aster segregate), but morphological features

281

282 PHY TOE OGHA October 1995 volume 79(4):281-285

in the key below suggest that it may be closer to the South American subtribe
Brachycominae. Chloracantha also appears to be phyletically isolated although it is
similar to Boltonia in some features, particularly habit. Other North American species
previously treated within Aster have recently been repositioned in Erigeron and
Machaeranthera, and several South American species of Aster sensu lato have recently
been dispersed among phyletically diverse genera.

The recognition of the genera segregated from Aster apportions the morphological
variation into reasonably discrete entities, but apparent parallelisms create practical
difficulties in the definition of some genera. The generic placement of certain species
(particularly within Eurybia) will be problematic because of distinctive morphological
specializations. These problems are discussed in detail elsewhere (Nesom 1994) and
reflected in the artificial key provided here. In any case, the key should serve at least
as a starting point for those who elect to use this taxonomic system or something
similar to it. Construction of keys and the identification of genera and species groups
will be considerably easier on a regional basis, just as it has been for Aster sensu lato.
Detailed descriptions of these genera, species groups, and problematic species are
found in the Aster review (Nesom 1994), as are authonties for all names used in the
present report.

In previous keys and discussions, | have used the terms “ligule” and “achene” in
reference to the expanded portion of the pistillate corollas and the fruit of Astereae.
Those terms are replaced here by “lamina” and “cypsela,” in acknowledgment of their
more technical correctness and their ineluctable fate in forthcoming application.

KEY TO THE AMERICAN GENERA OF ASTERINAE

1. Cypselas strongly flattened with lateral wings; pappus of two lateral awns (or
thickened bristles) and a series of short, highly reduced, awns or scales; disc
corollas with tube 0.2-0.5 mm long and abruptly expanded into the limb, the veins
accompanied: Dy OFAN GE TESINGAUCES. o.6o osc cemiicens yajomatytic soe mmcers sea ieee Boltonia

1. Cypselas flat to terete, without wings; pappus of barbellate bristles disc corollas
with a longer tube, abruptly or gradually opening into the limb, the veins without
Orange resin/ducts (except in'Chloracantha) 5.0. o:a(2.00 serdeinsne esis tone e eee (2)
2. Stems suffrutescent, usually sparsely to densely thomy, sometimes unarmed in

var. spinosa; leaves deciduous by anthesis; heads terminal on wiry, green
stems, arranged in a diffuse capitulescence; resting axillary buds with bud
SCANS Sioa Rear naetiance Castmn aegceacetcasaer stratal ooaeant aero ea eeger aah Chloracantha
2. Stems usually herbaceous, suffrutescent in a few species, never thorny; at least
the cauline leaves persistent and present at flowering (the stems of Oreostemma
scapose); heads variously arranged but not on wiry green stems in a diffuse

capitulescence; resting buds'not formed... 727..-s.ceacawsnos-05- toe arene eee (3)
3. Plants arising from long or short rhizomes and fibrous roots, not strongly woody
atthe! DASE. oc ccncumrcmsin nse eemesiet oe oe aeGiee sian tee cae eER CEE Hohe eae eeen eee eer (9)

3. Plants arising from a distinct taproot or thick, woody, mostly erect caudex
DIAM GIES ances cen cosmoen Sanna Ga Cane eR CEC Rink. «OM RERT hoe AEE EDGE ER Reece (4)

Nesom: Key to Amencan Asterinae 283

4. Plants perennial, usually ansing from a thick taproot or thick caudex branches.

4. Plants annual, usually arising from a slender taprool...................::::0006- (5)

5. Heads and upper stems stipitate-glandular; ray cypselas epappose.. Psilactis, in part
5. Plants completely eglandular; ray cypselas pappose (Symphyotrichum, in part). .(6)
6. Phyllaries evenly herbaceous and of subequal length; pistillate flowers in 2-4
series in a broad outer zone, the lamina absent or rudimentary to filiform and

short; disc (staminate) flowers fewer than the pistillate; pappus bnstles in 2
semes,/all of equal length: ..:.00.25..-cn.wa2- Symphyotrichum sect. Conyzopsis

6. Phyllaries with a green, rhombic apical patch, basally indurate, graduated in
length (imbneate); pistillate flowers in 1(-2) series, the lamina prominent or
strongly reduced; disc flowers more numerous than the ray; pappus bnistles of
equamiensthiand in a Single Scenes. achat Ree deo eee eee ee see anes,

50, AEE Lee REPRE S ORCC CCCEECE EERECERE Symphyotrichum sect. Oxytripolium, in part

7. Stems scapose, eglandular or minutely granular-glandular near the apex; heads
solitary; plants arising from a thick taproot or sometimes a short rhizome............
Beret Os ene eet Laat, oii tar acai Maa Ua ntrte hindi ak RI eee oe SiS Oreostemma

7. Stems with well-developed cauline leaves, eglandular or densely glandular; heads
solitary or few and loosely associated in a corymbiform capitulescence; plants
ansing from a thick taproot or thick, woody caudex branches. ..................... (8)

8. Stems and leaves eglandular or with short-stipitate glands; leaves 1-nerved,
congested on the stems; phyllanes stiff, evidently indurate-thickened, distinctly
keeled; rays mostly blue to purple; disc cypselas commonly 2-nerved, ray
cypselas usually 3-4 nerved; carpopodium oblique; pappus with an outer series
eiebnsties muchyshorter thanithe nner is: ate .o2-se-sckeeesee ees ene Tonactis

8. Stems and leaves usually with long-stipitate glands (eglandular in some
species); leaves with at least the secondary veins evident, not crowded on the

stems; outer phyllaries loose, foliaceous, rays yellow, white, or absent;
cypselas mostly 5-8-nerved; carpopodium a symmetrical ning at nght angles to

the long axis of the cypsela; pappus of (1-)2 series of bristles of equal length,

Faleuymwithea: SNODErOUIeR SEMES I. eee ee doe, Pee Tonestus
Semienyilaies, without-avgreen apical! patchascas....aao.c5. cnn seus. deseaes: clesteemencs => (14)
9. Phyllaries with a distinct, green apical patch or zone, the lower portion of the

POR MRAT VALINE. hs betas «SSE Es octet cd Pinte ati Ea pa (10)

10. Capitulescence diffuse or the heads terminally clustered but not in a distinctly
corymboid association; apical patch of phyllanes rhombic, sharply delimited at
the base and basally acute or attenuate, basally truncate in some species;
pappus bristles apically attenuate, in a single series........................205. (12)

10. Capitulescence corymboid or reduced to glomerate clusters; apical patch of
phyllaries basally truncate, sometimes not sharply delimited; pappus bnstles
apically dilated, in (1-)2-3 senes of equal or subequal length................. (11)

11. Heads pedicellate, mostly distinct (subsessile in Eurybia compacta); leaves
stipitate-glandular in a few species, otherwise eglandular; disc corollas yellowish;
style branch appendages spreading hairy from base to tip (closely papillate in a few
species); rays blue and strongly coiling, or white and non-coiling in sect. Biotia,
cypselas narrowly cylindric, glabrous to moderately strigose................. Eurybia

11. Heads sessile or subsessile in glomerate clusters; leaves sessile- or punctate-
glandular; disc corollas white; style branch appendages closely papillate; rays
white, not coiling; cypselas turbinate, stm gose-sericeous................: Sericocarpus

284 PHY TOLOG TTA October 1995 volume 79(4):281-285

L2e Ray CypselasiepappOse scare -c amcp oes acts e eee oe dere amie sete Psilactis, in part
12s RAY (CY PSClAS PAP POSC senescent apis cn anv come scm ceajee as- avon cetoreee hehe eener (13)
13. Plants trailing or climbing (not twining) vines. ................-20.0eee eee Ampelaster

13. Plants mostly erect, sometimes leaning but never trailing or even subscandent.....
Je aces anata adnate mee Os mimo eae nce G-Lisjecossis satstse sine Symphyotrichum, in part

14. Leaves all cauline, glabrous, linear with 3 parallel veins; pappus of a single
series of equal-length, apically attenuate bristles; involucre glandular. ...........

HEARS See ae a acc dtarbre aioe apace Clas aol isiejaeloeelaleps lotta ae telys eatathc are slemine 6 oe neo Almutaster

14. Leaves various but not as above; pappus bnistles in (1-)2-3 series of equal
length, apically dilated or attenuate; involucre glandular or eglandular. ..... (15)

15. Plants monocephalous; phyllaries evenly herbaceous, in 2(-3) series of subequal
length; cypselas obovate, 2-nerved and flattened, usually sessile-glandular near the

apex; pappus often with an evident short, outer series. .................. Aster alpinus
15. Plants with two or usually more heads, or if monocephalous then without the
above® combination Of {feataress 6.5.0 daccsudensdeoceeosments ownscees ee cee (16)

16. Leaves neither clasping nor subclasping; phyllaries usually strongly graduated
in length, not foliaceous; stems, leaves, and phyllaries eglandular or sometimes
sessile-glandular but without stipitate glands. .......................0ceseeeee es (18)

16. Leaves clasping or subclasping; phyllaries subequal in length, at least those of
the outer series foliaceous; stems, leaves, and phyllanes with stipitate glands. .
baci enlace ak cs SaUNE LS dooce aan ttie Sadatttatrea asbestos ence te ee 1

17. Outer phyllaries foliaceous, the inner usually with a green apical patch or zone;
basal leaves usually the largest, persistent; cypselas cylindric; pappus bnistles
usually dilatediat thelapex (nse ..ocesaftcataccmenans cece osene Eurybia sect. Herrickia

17. Outer phyllanes similar to the inner, herbaceous from base to apex; lowermost
cauline leaves greatly reduced in size (scale-like) and not persistent; cypselas
flattened; pappus bnistles apically attenuate. ...................2.0c0eee ees Canadanthus

18. Phyllaries herbaceous, 1-nerved, with a green band along the midvein from
base to tip, often purple-margined; basal leaves the largest, persistent; cypselas
LETELE crea oa a EU os via scisBure ease sutaeaerstlamacweubaeseatnee flog Aster tataricus

18. Phyllaries usually somewhat indurate at least near the base, with 1 or more
nerves, never with a medial green band; lowermost cauline leaves greatly
reduced in size (scale-like); cypselas terete to flattened. ....................... (19)

19. Heads mostly solitary or sometimes few and in a loosely corymboid
capitulescence; leaves thickened and stiff, l-nerved, congested on the stems

(internodes abbreviated); disc cypselas commonly 2-nerved, ray cypselas usually

3-4-nenved:, carpopodiumm oblique rtic....tusceotecs sce .ceeven -debeeccreneeeaceee Tonactis

19. Heads in a distinctly corymboid capitulescence; leaves relatively thin and
flexuous, spaced along the stem with internodes prominent, venation with at least
the secondary nerves evident; all cypselas 4-9 nerved; carpopodium at nght angles

tothe: long-axis:ofitheteypselas, esac} a essa aenas aed eset ntok oe one eee Eee (20)

20. Leaves usually sessile-glandular on the lower surface; collecting appendages
of the disc style branches spreading-hairy from base to tip; cypselas densely
sessile-glandular; pappus bnstles apically attenuate or (in Oclemena reticulata)
slightly dilatediat the apercc.vs,..0..cqns norsk eee coats teeter see teal eens Oclemena

20. Leaves not sessile-glandular, rarely short-stipitate glandular; collecting
appendages of the disc style branches closely papillate at least in the distal
portion; cypselas eglandular; pappus bnistles usually prominently dilated at the
AD OX Go cane sins cnn aaa ie oa eee eee ove aS aNd ara Te It (21)

Nesom: Key to Amencan Asterinae 285

21. Cypselas terete or subterete, with (4-)5-9 evenly spaced, orange-resinous nerves,
at maturity about the same length as the phyllanes; phyllaries oblong, not keeled,
each with a midvein and 1-2 lateral pairs of nerves; eastern North America and
BEIMCANLEDH ASIA: coef =o. on chm snag) aatinemadtnags* senmnete anise ae Nee entre? Doellingeria

21. Cypselas distinctly flattened, with a pair of lateral nerves and sometimes 1-2
whitish, subepidermal nerves on each face, shorter than the phyllanes at matunty;
phyllaries ovate to ovate-oblong, keeled, 1-nerved; western North America.........
I sa ios. occas cass «sone sagem aaageca ss poaceesoes snag dnauanaes eae asec Eucephalus

LITERATURE CITED

Nesom, G.L. 1994. Taxonomic overview of Aster sensu lato (Asteraceae: Astereae),
emphasizing the New World species. Phytologia 77: 141-297.

Phytologia (October 1995) 79(4):286-288.

TRIDAX YECORANA (ASTERACEAE, HELIANTHEAE) A NEW SPECIES
FROM SONORA, MEXICO

B.L. Turner

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

ABSTRACT

Tridax yecorana B.L. Turner, spec. nov., is described and illustrated.
tv is an annual herb known only from type material collected near Yecora,
Sonora, and is related to T. erecta. It differs from the latter in numerous
characters which are discussed in the text.

KEY WORDS: Asteraceae, Heliantheae, Tridax, México, Sonora, systematics

Routine identification of Mexican Asteraceae has revealed the following novelty.

TRIDAX YECORANA B.L. Turner, spec. nov., Figure 1. TYPE: MEXICO.
Sonora: Arroyo El Otro Lado, Mesa El Otro Lado, 1-2 km NNE of Yecora on old

road to Maycoba, pine-oak forest, 28° 23’ 49” N, 108° 54’ 48” W, 1520 m, 7 Sep
1995, T.R. Van Devender 95-836 (with A.L. Reina G., D.A. Yetman, and M.E.
Fishbein) (HOLOTYPE: TEX).

Similis T. erectae A. Gray sed foliis lineanbus-lanceolatis (vice foliorum
ovatorum), glaberis aut sparsim glanduliferis-pubescentibus (vice
hispidissimorum), involucns campanulatis (vice urceolatorum) glaberisque
(vice pubescentium), acheniis mgide pubescentibus (vice molliter
pubescentium), et pappis 1-2 mm longis (vice 2.5-5.0 mm).

Annual herbs 7-20 cm high. Stems mostly unbranched, sparsely pubescent with
glandular tichomes 0.5-1.0 mm long. Leaves linear-lanceolate, mostly 1-2 mm wide.
Heads single on peduncles, 4-15 cm long, pubescent like the stems. Involucres
campanulate, 4-6 mm high, 4-9 mm wide (pressed); bracts 3-4 senate, broadly
elliptical to oblanceolate, glabrous, the apices broadly rounded, scarious. Receptacles
conical, 2-3 mm across, 2.0-2.5 mm high; bracts scarious, persistent, oblanceolate to
linear-oblanceolate, variously 2-3 cleft at their apices. Ray florets pistillate, fertile;

286

Tumer. New Tridax from Sonora

/cm

Figure 1. Tridax yecorana, from holotype; left, a single head; nght, a disk achene.

288 PHY TOLOGIA October 1995 volume 79(4):286-288

corollas yellow; tube ca. 2 mm long, densely pilose; ligules mostly 4-5 mm long, 3-4
mm wide. Disk florets 10-25; corollas yellow, ca. 3 mm long, the tubes ca. 0.8 mm
long, densely pilose; throat ca. 2 mm long, gradually arnpliate upwards, the 5 lobes
markedly nervate. Anthers yellow, their apices trianguloid, keeled inwardly. Achenes
of disk and ray florets similar, obpyramidal, ca. 2 mm long, 0.8 mm wide, densely
pubescent with stiff ascending hairs 0.5-1.0 mm long; pappus of 20 or more short
plumose scales 1-2 mm long.

Tridax yecorana, in habit, superficially resembles T. coronopifolia H.B.K. but is
clearly most closely related to T. erecta A. Gray, differing from the latter in having
linear, nearly glabrous leaves, campanulate completely glabrous involucres, ray florets
with densely villous tubes, and achenes with stiffer hairs and shorter pappus scales.

Tridax erecta (including the recently descnbed T. durangensis A. Garcia Arévalo,
which appears to be but a form of that species) has ovate, coarsely pubescent leaves,
involucres urceolate with loose outer bracts and coarsely pubescent inner bracts, and
more softly pubescent achenes with longer pappus scales.

According to label data on the type sheet, Tridax yecorana is a “Locally very
common annual.”

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscript.

Phytologia (October 1995) 79(4):289-292.

SALVIA BOOLEANA (LAMIACEAE), A NEW SPECIES FROM
NORTHEASTERN MEXICO

B.L. Turner

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

ABSTRACT

Salvia booleana B.L. Turner spec. nov., is described and illustrated. It
belongs to the sect. Fulgentes, a small group with about eight species, all
having large red flowers (corollas mostly 3-5 cm long), where it relates to S.
fulgens Cav. It is distinguished from the latter by numerous characters
including habit, leaf shape, bract size, vestiture and distribution.

KEY WORDS: Lamiaceae, Salvia, sect. Fulgentes, México, Nuevo Leon,
San Luis Potosi, systematics

Routine identification of Mexican plants has revealed the following novelty.

SALVIA BOOLEANA B.L. Turer, spec. nov. Figure 1. TYPE: MEXICO. San
Luis Potosi: Mpio. Charcas, Charcas, “on wetbank of Arroyo”, Jul-Aug 1934,
C.L. Lundell 5470 (HOLOTYPE: LL!, Isotype: TEX!).

Similis Salviae fulgenti Cav. (Salvia fulgens) sed differt laminis foliorum
subdeltatis, basibus foliorum plerumque cordatis, et caulibus_ valde
glandulosis-pubescentibus, indumento 0.6-1.0 mm alto.

Perennial herbs 60-100 cm high. Stems densely glandular-hirsute, the vestiture
0.6-1.0 mm high. Midstem leaves 4-7 cm long, 2.5-4.0 cm wide; petioles 1.5-3.0 cm
long; blades cordate-deltoid to subdeltoid, about as wide as long, mostly subcordate at
base, pubescent like the stems, margins crenulodentate, the apices mostly obtuse.
Floral bracts ovate, soon deciduous, the upper immature bracts 8-10 mm long, 2-4
mm wide, the apices gradually acuminate. Flowers (2-)4-6 to a node. Calyces mostly
11-15 mm long, glandular-pubescent; upper lobes 3-4 mm long, 9-ribbed. Corollas
red to orangish-red, 3.0-4.2 cm long; upper lips 12-15 mm long; lower lips 10-12 mm
long. Stamens attached near the onfice, the anthers mostly loosely exserted somewhat
beyond the upper lip, rarely not, ca. 2 mm long, attached near the base (1/4 the
anthers’ length). Styles pubescent, the upper branches 2-3 times as long as the lower.
Nutlets linear-ovoid, ca. 4 mm long, 1.5 mm wide, veinous, glabrous.

2) 32)

Poy TOLOGcilA

October 1995 volume 79(4):289-292

Tumer: New Salvia from Nuevo Leon 291

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Nuevo Leon: Mpio.
Arambern, N of Arambern, 995 m, 16 Jun 1990, Hinton et al. 20340 (TEX); N of
Arambern, 970 m, 1 Sep 1990, Hinton et al. 25019 (TEX); Sierra Vieja, 12.2 mi
along dirt road turnoff to Ejido Capadero, just N of Dr. Arroyo, 6900 ft., “In dry
stream bed”, 20 Oct 1984, Saunders-Scherrer 13476 (TEX).

Salvia booleana belongs to the sect. Fulgentes of Salvia, sensu Epling (1939).
The nomenclatural history of this section is discussed in some detail by Ramamoorthy
(1987), but no recent taxonomic study of the taxon is available, in spite of its array of
attractive large red-flowered species.

Epling (1939) recognized (and keyed) six species as occurring in the section,
adding an additional species with the description of Salvia sharpii Epling & Mathias in
1957, which is probably a weakly differentiated populational element of S.
microphylla H.B.K. The present addition brings this total to eight, and additional
species are certain to follow as Mexico becomes more thoroughly collected.

Type matenal of Salvia booleana was apparently included by Epling (1939) in his
concept of S. fulgens, but with the comment, “Lundell’s specimen from Charcas,
while similar in flowers to the southern forms is markedly glandular with short-deltoid
leaves.” Which is certainly true; indeed, all of the specimens cited above possess such
leaves and, combined with their relatively small calyces and much-reduced floral
bracts, mark the plants concerned as very distinctive, certainly deserving of specific
rank as morphologically defined by Epling and yet others.

Salvia booleana reportedly occurs along dry washes in relative xeric habitats from
800 to 2000 m; S. fulgens is a taller plant with much larger leaves occurring in mostly
moist montane habitats above 2000 m (distributed from southern San Luis Potosi
southwards to the states of Puebla and Morelos).

It is a pleasure to name this taxon for George Boole Hinton (great grandson of the
late renown Mexican collector, George Boole Hinton), frequent companion on field
forays with Jaime and Jorge Hinton, son and grandson, respectively of the pnmal sire,
G.B. Hinton. A photograph of this young Hinton can be found in Turner (1996). My
principal reason for selection of the epithet concerned is to establish a familial record of
sorts: five names from a male lineage representing four generations, all included in the
same genus. These include:

Salvia hintonii Epling - named for G.B. Hinton, the father.

Salvia jacobi Epling - for James Hinton, the son (pers. comm., James Hinton)
Salvia jaimehintoniana Ramamoorthy - honoring James Hinton, the son.

Salvia jorgehintoniana Ramamoorthy - honoring George Hinton, the grandson.
Salvia booleana B.L. Turner - honoring George Boole Hinton, the great grandson.

NE el rial dl

And this does not include Salvia leninae Epling, named for a remarkable pack
animal of the Hinton’s, a mule named Lenina. Salvia, with SOO or more species, can
comfortably ingest such effrontery. What I like about the eponyms concerned is that
most of the species (all except S. jacobi and S. hintonii) occur in the state of Nuevo
Le6n, and the surviving kin of G.B. Hinton, all residing in Nuevo Leon on their

292 PHY POLO GEA October 1995 volume 79(4):289-292

Rancho Aguililla, are now surrounded by flonstic “headstones” that will extend far
beyond their natural lives. I like that kind of perpetuity for such dedicated workers!

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Piero Delprete
for reviewing the manuscnpt

LITERATURE CITED

Epling, C. 1939. A revision of Salvia, subgenus Calosphace, Fedde Repert. Sp.
Nov. Beih. 110:1-388.

Ramamoorthy, T.P.. 1987. Typifications in Salvia (Lamiaceae). Taxon 33:322-324.

Turmer, B.L. 1996. Sedum booleanum (Crassulaceae), a new red-flowered species
from Nuevo Leon, México. Phytologia 79:31-34.

Phytologia (October 1995) 79(4):293-295S.

A NEW SPECIES OF LOBELIA (CAMPANULACEAE) FROM OAXACA,
MEXICO

B.L. Turner

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

ABSTRACT

Lobelia hintoniorum B.L. Tumer, spec. nov. from _ Distnto
Miahuatlan, Oaxaca, is described and illustrated. It belongs to the sect.
Hemipogon, subsect. Leiospermae, where it relates to L. occidentalis
McVaugh. It differs from the latter in possessing very large dark blue corollas
and nonhispidulous anthers.

KEY WORDS: Campanulaceae, Lobelia, México, Oaxaca, systematics

Routine identification of Mexican lobelioids has revealed the following novelty.

LOBELIA HINTONIORUM B.L. Turner, spec. nov. Figure 1. TYPE:
MEXICO. Oaxaca: Distnto Miahuatlan, S side of Cerro Quiexobra, 1-3 km NE
of La Cieneguilla on road to summit, in damp ravines below understory of pine-
oak forests, 2900 m, 2 Oct 1990, Andrew McDonald 2982 (HOLOTYPE: TEX).

Similis L. occidentali McVaugh & Huft sed foliis midcaulis majoribus, ([6-
]12-15 cm longis vice 4-10 cm longis), pedunculis valde majonbus (5-6 cm
longis vice 2.5-4.0 cm longis), tubis corollarum longionbus (12-15 mm longis
vice 7-9 mm longis), et sacculis supens antherarum glabns (vice sacculorum
hispidorum).

Weakly ascending or procumbent herbs to 60 cm high ansing from slender
rhizomes, forming colonies. Midstems 1-3 mm across, glabrous. Midstem leaves
glabrous, mostly linear to linear-lanceolate, gradually reduced upwards, (5-)6-15 cm
long, 0.3-0.7 cm wide, remotely denticulate. Inflorescence of (2-)5-25 flowers, when
numerous the latter disposed in a secund fashion. Bracts linear, mostly 1/2 as long as
the pedicels, or more. Pedicels of mature flowers mostly upwardly arcuate, 2-6 cm
long. Ovary ca. 1/3 to 1/2 inferior, the calyx cup ca. 2 mm high, glabrous, the lobes

293

volume 79(4):293-295

October 1995

PHYTOLOGIA

294

, »S y=

Figure 1. Lobelia hintoniorum, from holotype.

Turner: New Lobelia from Oaxaca 295

linear-lanceolate, 4-6 mm long, reflexing with age. Corollas dark blue, the tubes 12-
16 mm long, not fenestrate, the dorsal slit 9-11 mm deep; upper two lobes linear-
lanceolate, 6-8 mm long; lower 3 lobes neatly elliptical, 7-10 mm long, 2.5-4.0 mm
wide. Filaments ca. 10 mm long, united for ca. 4 mm apically; anthers 3-4 mm long,
the lower 2 tufted, otherwise glabrous. Fruits not available.

ADDITIONAL SPECIMENS EXAMINED: MEXICO. Oaxaca: Distrito
Miahuatlan, Quiexobra, 2920 m, 14 Oct 1995, Hinton et al. 26104 (TEX); Siete
Ocotes, 2950 m, 20 Oct 1995, Hinton et al. 26256 (TEX); Siete Ocotes, 2880 m,
Hinton et al. 26265 (TEX).

Lobelia hintoniorum clearly belongs to the sect. Hemipogon subsect. Leiospermae
(sensu Wimmer 1953) where it relates to L. occidentalis McVaugh and L. dielsiana
Wimmer. McVaugh (1975) provided a detailed key to both of these taxa. In this, L.
hintoniorum, because of its very large corollas, will key to L. sublibera S. Wats., a
very distinctive species confined to northeastern México (Nuevo Leon and
Tamaulipas). Lobelia hintoniorum has the habit, leaves, and general inflorescence of
L. occidentalis, but differs in the characters called to the fore in my diagnosis.

It is a pleasure to name this taxon for the Hinton family, who collected three of the
only four collections known to me. Label data on the Hinton matenal report the
species to form scattered but common procumbent plants or colonies to 60 cm high.
Hinton 26104 is a depauperate plant with relatively small leaves, but its flowers are
typical of the taxon concerned.

The type of Lobelia hintoniorum was obtained by Andrew McDonald in 1990
(from among whose many collections I named Lobelia macdonaldii B.L. Turner), but
this collection remained unnamed awaiting additional material. The several Hinton
specimens cited above leave little doubt that the taxon is quite distinct and undescribed.

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscnipt. Ms. Mana Thompson provided the
illustration.

LITERATURE CITED

McVaugh, M. & M.J. Huft. 1975. Rediscovery of Lobelia dielsiana Wimmer, and a
related species new to science. Contr. Univ. Mich. Herb. 11:65-68.

Turner, B.L. 1992. A new species of Lobelia (Campanulaceae) from Oaxaca,
México. Phytologia 72:34-36.

Wimmer, F.E. 1953. Lobelia, in Pflanzenreich IV. 276b (Heft 107): 408-695.

Phytologia (October 1995) 79(4):296-297

A NEW SPECIES OF VERBESINA (ASTERACEAE) FROM OAXACA, MEXICO

B.L. Turner

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

ABSTRACT

Verbesina miahuatlana B.L. Tumer spec. nov., is described and
illustrated from Distrito Miahuatlién, Oaxaca. It is known only from two
collections, both obtained in pine-oak forests between 2700-2825 m. It
belongs to the Verbesina virgata complex (ca. eight species) but can be
distinguished from all of these by its much larger coarsely serrate leaves and
loosely corymbose paniculate capitulescence.

KEY WORDS: Asteraceae, Verbesina, México, Oaxaca, systematics

Routine identification of Mexican Asteraceae has revealed the following novelty.

VERBESINA MIAHUATLANA B.L. Turner, spec. nov. TYPE: MEXICO.
Oaxaca: Distrito Miahuatlin, Xianaguilla, 2700 m, oak and pine forests, 21 Oct
1995, Hinton et al. 26294 (HOLOTYPE: TEX).

Similis V. virgatae sed foliis lationbus (3-9 cm latis vice 1.5-2.5 cm latis),
cum marginibus valde serratis, et capitulis parvioribus, dispositis in paniculis
rotundatis et corymbosis, pedunculis ultimis gracilibus et flexuosis (vice
crassorum et rigide erectorum).

Shrub to 2.5 m high. Stems sparsely strigose, narrowly corky winged for 1-3 cm
below each node. Larger leaves alternate, 9-24 cm long, 3-8 cm wide; petioles 5-20
mm long; blades pinnately nervate, broadly ovate to elliptic, gradually tapenng upon
the petioles, sparsely strigose above and below, especially along the major veins, the
margins irregularly serrate. Heads numerous, arranged in terminal corymbose
panicles, scarcely exceeding the leaves, the ultimate peduncles mostly 5-15 mm long.
Involucres broadly campanulate, 4-5 mm high, 6-8 mm wide (pressed), bracts 2-4
senate, narrowly ovate, subgraduate, black, the apices acute. Receptacle ca. 2 mm
across, | mm high, the chaff shorter than the subtended florets, their apices abruptly
acute. Ray florets 5-8, pistillate fertile; ligules yellow, 6-9 mm long, 2-3 mm wide,
4-6 nervate, their apices with 2-3 shallow lobes; tubes ca. 1.5 mm long, pubescent.

296

Turner: New Verbesina from Oaxaca 297

Disk florets 30-40 (est.); corollas yellow, ca. 3 mm long, the tube ca. 0.75 mm long,
pubescent; lobes glabrous, ca. 0.7 mm long. Anthers brown. Achenes ca. 2 mm
long, the faces sparsely strigose, the margins ciliate; pappus of 2 subequal persistent
awns ca. 2 mm long.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Oaxaca. Distnto
Miahuatldn, Siete Ocotes to Xianaguilla, 2825 m, 21 Oct 1995, Hinton et al. 26277
(TEX).

The present novelty is closely related to a group of species centering about the
widespread Verbesina virgata. The distnbution of this complex is shown in more
detail by Turner (1992). Verbesina miahuatlana differs from these in possessing
broader leaves, more numerous heads arranged in rounded corymbose panicles, and
having black, broadly campanulate involucres, among yet other characters.

The holotype represents a lush collection with very large leaves, while the
additional collection has much smaller, less serrate leaves, but in all other characters
the two plants are alike and unquestionably belong to the same species.

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Justin
Williams for reviewing the paper.

LITERATURE CITED

Turner, B.L. 1992. Two new species of Verbesina (Asteraceae) from southern
México. Phytologia 72: 109-114.

Phytologia (October 1995) 79(4):298-300.

A NEW SPECIES OF MENTZELIA (LOASACEAE) FROM NUEVO LEON,
MEXICO

B.L. Turner & Alice L. Hempel

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

ABSTRACT

Mentzelia hintoniorum B.L. Tumer & Hempel, spec. nov, is
described and illustrated. It is known only from gypseous soils near San
Roberto, Mpio. Galeana, Nuevo Leén. The taxon belongs to the sect.
Bartonia and is seemingly most closely related to M. mexicana but is
distinguished from that species by numerous features including habit,
vestiture, and flower size.

KEY WORDS: Loasaceae, Menizelia, México, Nuevo Leon, systematics

Routine identification of Mexican plants has revealed the following novelty.

MENTZELIA HINTONIORUM B.L. Turmer & Hempel, spec. nov. Figure 1.
TYPE: MEXICO. Nuevo Leén: Mpio. Galeana, San Roberto to “Y,” (24° 41’

55" N, 100° 10’ 34” W) 2015 m, gypsum hillside, 5 Sep 1995, Hinton et al.
25495 (HOLOTYPE: TEX).

Similis M. mexicanae M.J. Thompson et Zavort. sed caulibus rectis, non
ramosis infra, em corona radicum lignearum exonentibus, et floribus
majoribus, petalis plerumque 20-22 mm longis (vice 10-15 mm longis),
staminibus exterioribus ca. 13 mm longis (vice ca. 9 mm longis).

Simple-stemmed (or sparsely branched following injury) perennial herbs ca. 30 cm
high, ansing from the crown of woody roots. Stems straight, not at all fractiflex, ca.
3 mm across at midstem, moderately pubescent with stiff, multiseptate, glochidiate
hairs, forming a vestiture ca. 0.5 mm high. Leaves linear-oblanceolate, not clearly
petiolate, gradually reduced upwards, those at midstem mostly 3-4 cm long, 4-7 mm
wide, pubescent like the stems, but sparsely so, and the surfaces mostly glabrous, the
margins with 3-7 shallow lobes. Flowers 1-3, terminal. Calyx cup at anthesis 3-5
mm high; lobes lanceolate, ca. 12 mm long, 2.5 mm wide at base, fused below for

298

Tumer: New Menizelia from Nuevo Leén

Figure 1. Mentzelia hintoniorum, from holotype

300 PHY POLMOGTA October 1995 volume 79(4):298-300

1.5-2.0 mm, pubescent like the stems. Petals 10, yellow, 20-22 mm long, ca. 5 mm
wide, gradually tapered from above into a narrow claw ca. 8 mm long. Stamens
numerous, 10-13 mm long, the outermost anthers borne on narrow filaments.
Capsules 20-25 mm long, 8-10 mm wide (pressed); lobes 4-6 mm long. Seeds white,
smooth, 2.5-3.0 mm long, ca. 2 mm wide; wings ca. 0.5 mm wide.

Menizelia hintoniorum is closely related to M. mexicana Thompson & Zabort. of
the sect. Bartonia (cf. Thompson & Powell 1981). Itis readily distinguished from M.
mexicana by its unbranched straight stems which arise from the crown of woody tap
roots (vs. much-branched stems from tough but scarcely woody tap roots), more
prominent stem-hairs, the vestiture ca. 0.5 mm high, lacking an understory of minute
hairs (vs. vestiture ca. 0.25 mm high and minutely pubescent beneath), and much
larger petals (20-22 mm long vs. 10-15 mm long).

Thompson & Powell (1981) provided a detailed account of Mentzelia mexicana
and closely related taxa, mapping the distribution of each taxon. None of these was
shown to occur in Nuevo Le6én. Menizelia hintoniorum occurs in a region of Nuevo
Leén (near San Roberto) where numerous gypseous endemics occur, the present
apparently being yet another.

It is a pleasure to honor the remarkable Hinton clan with this rare novelty, the
collectors noting the taxon to be represented by only “a few plants.” at the locality

concerned, which is very near the type locality of the localized Arenaria hintoniorum
B.L. Turner.

ACKNOWLEDGMENTS

We are grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the paper. Maria Thompson provided the illustration.

LITERATURE CITED

Thompson, H.J. & A.M. Powell. 1981. Loasaceae of the Chihuahuan Desert
Region. Phytologia 49: 16-32.

Phytologia (October 1995) 79(4):301-302.

A NEW SPECIES OF STEVIA (ASTERACEAE) FROM CERRO QUIEXOBRA,
OAXACA, MEXICO

B.L. Turner

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

ABSTRACT

Stevia quiexobra B.L. Turner, spec. nov. is described from Cerro
Quiexobra, Oaxaca, México, where it occurs in pine-fir forests at ca. 3400 m.

KEY WORDS: Asteraceae, Stevia, México, Oaxaca, systematics

Stevia is represented in México by numerous species, most of these treated by
Grashoff (1972). Since the latter’s treatment, numerous additional species have been
added, the most recent being those of Turner (1995) and Yahara & Soejima (1995). I
add here a newly discovered taxon from Cerro Quiexobra, Oaxaca.

STEVIA QUIEXOBRA B.L. Turner spec. nov. TYPE: MEXICO. Oaxaca:
Distnito Miahuatlin, Cerro Quiexobra, 3385 m, “steep fir and pine woods”, 15 Oct
1995, Hinton et al. 26141 (TEX).

Similis S. perfoliatae Crong. sed foliis non perfoliatis et achenibus
exaristalis.

Perennial rhizomatous herbs 20-30 cm high. Stems with a dense vestiture of
glandular-capitate tnchomes about 0.5 m high. Leaves mostly opposite (except for 3-5
uppermost leaves), gradually reduced upwards. Midstem leaves ovate to ovate-
elliptic, sessile or nearly so, widest at or about the middle, 3-4 cm long, 1.0-1.8 cm
wide, with 3 principal nerves arising from above the base, glandular-punctate on both
surfaces, glandular pubescent like the stems, the margins weakly crenate. Heads
arranged in bracteate congested glomerules ca. 1.5 cm high, 1.5 cm across.
Subtending bracts glandular pubescent, similar to the involucral bracts. Involucres ca.
7 mm high, sparsely glandular pubescent to glabrous. Corolla tubes ca. 5S mm long,
sparsely pubescent; lobes 1.5-2.0 mm long, sparsely pubescent on the outer surfaces.
Achenes (immature) all alike, ca. 4.5 mm long, glabrous except for a few hispid hairs
near the apices; pappus a crown of short scales ca. 0.75 mm high.

301

302 PHY TOLOGIA October 1995 volume 79(4):301-302

This taxon is known only by the type; label data note it to occur as “thin colonies
0.3 m high.” Because of its broad sessile glandular pubescent leaves, S. quiexobra is
readily distinguished from most other Mexican taxa. It is seemingly most closely
related to S. perfoliata Cronq., but lacks the perfoliate leaves and anstate achenes of
that species.

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the paper.

LITERATURE CITED

Grashoff, J.L. 1972. A systematic study of the North and Central Amencan species
of Stevia. Doctoral Dissertation. The University of Texas, Austin, Texas.

Turner, B.L. 1995. Stevia calzadana (Asteraceae) a new species from Oaxaca,
México. Phytologia 79:5-7.

Yahara, T. & A. Soejima. 1995. A new species of Stevia from México. Phytologia
79:35-37.

Phytologia (October 1995) 79(4):303-30S.

STELLARIA MIAHUATLANA (CARYOPHYLLACEAE), A NEW SPECIES
FROM OAXACA, MEXICO

B.L. Turner

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

ABSTRACT

Stellaria miahuatlana B.L. Turmer, spec. nov., is described from
Distnto Miahuatldn, Oaxaca, México. It is closely related to S. irazuensis but
differs in its 5-parted calyx, larger corollas and much larger leaves.

KEY WORDS: Caryophyllaceae, Stellaria, México, Oaxaca, systematics

Routine identification of Mexican plants has revealed the following novelty.

STELLARIA MIAHUATLANA B.L. Tumer, spec. nov. Figure 1. TYPE:
MEXICO. Oaxaca: Distnto Miahuatlan, above Xianaguilla, 2510 m, “mixed

woods of oak, pine, arbutus..Common”, 24 Oct 1995, Hinton et al. 26426
(TEX).

Similis S. irazuensis Donn. Sm. sed calycibus cum 5 lobis (vice 4),
_ corollis majoribus, ca. 9 mm longis (vice 3-6 mm longis), et foliis majoribus
cum laminis 30-40 mm latis (vice 10-20 mm).

Sprawling perennial (?) herbs to0.4 m high. Younger stems mostly pilose; older
stems glabrate and shiny, the internodes mostly 2-3 times as long as the leaves.
Stipules absent. Midstem leaves (4-)5-6 cm long; petioles 1.0-2.5 cm long, pilose;
blades cordate, 3.0-4.5 cm long, 3.0-3.5 cm wide, more or less glabrous on both
surfaces, the margins and veins sparsely pilose. Flowers 5-10, mostly axillary in
bracteate dichasial cymes, rarely solitary. Pedicels mostly 1.5-2.0 cm long, densely
glandular-pilose. Sepals 5, ovate-lanceolate, 4-5 mm long, ca. 1.5 mm_ wide,
sparsely pilose below, the margins white-scanious. Petals 5, white, ca. 9 mm long,
deeply cleft for 4-5 mm, the lobes linear to linear-oblanceolate, weakly nervate, if at
all. Stamens 10, ca. 4 mm long, the anthers white. Style branches 3, ca. 4 mm long,
free to the base. Capsules (immature) ca. 4.5 mm long, the young seeds numerous
and peripherally ornate with bulging cells.

303

304 PHYTOLOGIA October 1995
Aa
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Figure 1. Stellaria miahuatlana, from holotype.

volume 79(4):303-305

Turner: New Stellaria from Sonora 305

This taxon, because of its inflorescence, glandular-villous pedicels and markedly
cordate leaves, appears to be closely related to Stellaria irazuensis Donn. Sm. a species
of Central Amenca (Guatemala to Panama), nicely illustrated by Duke (1961) in his
treatment of Stellaria for Panama. Stellaria miahuatlana is readily distinguished from
S. irazuensis in having larger more broadly cordate blades (30-35 mm wide vs. 5-15
mm wide) mostly 5 sepals (vs. 4 sepals), and larger petals.

ACKNOWLEDGMENTS

I am grateful to Gayle Turner for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscript.

LITERATURE CITED

Duke, J.A. 1961. Stellaria, in Flora of Panama. Ann. Missouri Bot. Gard. 48:438-
442.

Phytologia (October 1995) 79(4):306-308.

A NEW SPECIES OF CYNOGLOSSUM (BORAGINACEAE) FROM OAXACA,
MEXICO

B.L. Turner

Department of Botany, University of Texas, Austin, Texas 78713 OBS A

ABSTRACT

Cynoglossum hintoniorum B.L. Turner, spec. nov., is described and
illustrated from high elevational regions on and about Cerro Quiexobra,
Oaxaca. It is closely related to C. amabile, but differs markedly from that
species in possessing mericarps with relatively few smooth elongate spines,
otherwise they appear very similar.

KEY WORDS: Boraginaceae, Cynoglossum, México, Oaxaca, systematics

Identifications of collections from Cerro Quiexobra, Oaxaca, and immediate
environs has revealed the following novelty.

CYNOGLOSSUM HINTONIORUM B.L. Turner, spec. nov. GEE:
MEXICO. Oaxaca: Distrito Miahuatldn, Cerro Quiexobra, 3145 m, 19 Oct 1995,
Hinton et al. 26206 (HOLOTYPE: TEX).

Similis C. amabili Stapf & Drumm. sed menicarpiis cum solum 10-15
spinis elongatis laevibusque (vice spinarum multarum, brevium, et
muricatarum).

Erect perennial herbs 20-60 cm high, arising from stout ligneous taproots. Basal
leaves mostly 10-18 cm long, 1.5-3.0 cm wide; petioles 3-6 cm long; blades narrowly
elliptic, widest at or near the middle, pinnately veined, moderately pilose above and
below, strigose along the major veins, the surfaces minutely atomiferous-glandular,
the margins entire. Midstem leaves 5-10 cm long, 1-3 cm wide, the petioles winged
throughout, tapered upon by the blades. Flowers terminal, arranged in scorpioid-
racemic inflorescences 10-20 cm long, the pedicels 2-5 mm long, recurved in fruit.
Sepals ovate-lanceolate, ca. 3 mm long, strigose externally, free to the base or nearly
so. Corollas blue, 8-10 mm across, the throat nearly closed by hispidulous bilobate
appendages. Stamens 5, nearly sessile, the anthers ca. 1 mm long, not excurrent.

306

Turner: New Cynoglossum from Oaxaca 307

Figure 1. Mericarps of Cynoglossum amabile (lower left, Webster 11327 [TEX]) and
C. hintoniorum (upper night, from holotype).

308 PHY TOLOGIA October 1995 volume 79(4):306-308

Style ca. 3 mm long, the stigmatic surface more or less peltate. Mericarps (3 of them),
each with 10-15 long flattened smooth spines, 3-4 mm long, their apices with 2-4
hooked hairs, 1 of the mericarps tending to abort, nearly rugose, not at all spinose or
very weakly so.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Oaxaca: Distnto
Miahuatlén, Xianaguilla, 2715 m, oak and pine forest, 13 Oct 1995, Hinton et al.
26063 (TEX).

This taxon has most of the characters of Cynoglossum amabile Stapf & Drumm.,
except for the markedly different fruits, as shown in Figure 1. Examination of 30 or
more sheets of C. amabile (LL, TEX) from both México and Central America revealed
no fruits remotely approaching those of C. hintoniorum.

Mexico is now known to have four species of Cynoglossum: C. amabile, C.
henricksonii Higgins (=C. erectum Higgins 1976, not C. erectum Sweigg ex Schrank
1822), C. hintoniorum, and C. pringlei Greenm. Cynoglossum amabile is said to be
native to China, being introduced into México and elsewhere in Central and South
Amenca (cf. Nash & Moreno 1981, who provided an excellent illustration). Brand
(1921), however, does not note a New World distribution in his treatment.
Apparently C. amabile is used as a folk medicinal, having largely spread throughout
the tropical and subtropical regions of the New World over the past 50 years (it was
not described as new to science until 1906). Gibson (1970) thought the plant to be
largely cultivated for omamental purposes in Guatemala, the very adherent seeds
readily dispersed by mammals, including man. Finally, it should be noted that C.
hintoniorum may be a stabilized or populational fruit-form of C. amabile; if so, it is a
remarkable populational variant, especially since it occurs at two distant locales in
Miahuatldn at very high elevations (2715-3145 m) in regions relatively remote from
human population centers.

ACKNOWLEDGMENTS

I am grateful to Gayle Turner for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscript. Marcia Thompson provided the illustration.

LITERATURE CITED

Brand, A. 1921. Cynoglossum, in Pflanzenreich IV (252):114-153.

Gibson, D. 1970. Cynoglossum, in Flora of Guatemala. Fieldiana: Bot. 24:133-
134.

Nash, L. & N. Moreno. 1981. Cynoglossum, in Flora de Veracruz. 18:52-55.

Phytologia (October 1995) 79(4):309-312.

TWO NEW SPECIES OF AGERATINA (ASTERACEAE) FROM MEXICO

B.L. Turner

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

ABSTRACT

Two new species of Ageratina are described from México: A.
ayerscottiana B.L. Turner, from the vicinity of Basaseachi, Chihuahua; and
A. miahuatlana from Oaxaca. The former belongs to the subgenus
Neogreenella and relates to A. petiolaris, the latter belongs to the subgenus
Ageratina and relates to A. viscosissima. A map showing the distnbution of
A. ayerscottiana and A. petiolaris is provided.

KEY WORDS: Asteraceae, Eupatoneae, Ageratina, Mexico, Chihuahua,
Oaxaca, systematics

The genus Ageratina is a segregate from Eupatorium (s.l.). It is a large highly
variable complex in Mexico, 110 or more species currently recognized (cf. Tumer &
Nesom 1993). The present account, along with others described since the 1993
survey, adds two additional species, bringing to ca. 125 the number currently
recognized for México (Tumer 1996).

AGERATINA AYERSCOTTIANA B.L. Turner, spec. nov. TYPE: MEXICO.
Chihuahua: | mi N. of Maguarachi, ca. 22 mi S of junction with Basaseachi-San
Juanito road, “steep S-facing cliff in drainage,” ca. 6000 ft, May 1984, T.J. Ayers
399, with R. Scott (HOLOTYPE: TEX!)

Similis A. petiolari (DC.) R.M. King & H. Rob. sed foliis parvioribus
cum venatione valde elevata et sine trichomatibus glandulosis.

Suffruticose herbs or shrublets. Young stems densely hirsute with white
eglandular hairs. Leaves opposite throughout; uppermost leaves thick and strongly
venose beneath; petioles 10-15 mm long; blades neatly cordate, 2-3 cm long, 2-3 cm
wide, 3-5 nervate from the base, densely hirsute above and below with eglandular
hairs, the surfaces densely atomiferous-glandular, the margins crenulate. Heads
terminal, arranged 30-100 in rounded corymbose capitulescences, the ultimate
peduncles mostly 5-15 mm long. Involucres campanulate, 5-6 mm high, ca. 10 mm

309

310 PHYTOLOGIA October 1995 volume 79(4):309-3 12

wide (pressed); bracts linear-lanceolate in ca. 2 series, pubescent with eglandular
hairs, the surfaces atomiferous-glandular. Receptacles convex, ca. 4 mm across, 1.5
mm high, glabrous. Disk florets 50 or more (est.); corollas white, 4-5 mm long,
glabrous; tubes ca. 2 mm long; lobes ca. 0.5 mm long, atomiferous-glandular, but
without hairs. Achenes ca. 3 mm long, hispidulous; the pappus of ca. 20 barbellate
bristles 5 mm long in a single senes.

ADDITIONAL SPECIMEN EXAMINED: MEXICO. Chihuahua: just E of
Maguarachi on road between Basaseachi and San Juanito, headwaters of the Rio
Oteros, “steep sided mountain slopes in narrow arroyo,” 17 May 1984, Lavin 5427
(TEX), with R. Scott et al.

This taxon belongs to the subgenus Neogreenella (sensu King & Robinson 1987),
superficially resembling Ageratina petiolaris (DC.) King & H. Rob. It is amply
distinct from the latter by a number of characters, most notably through the absence of
glandular trichomes, and by the seemingly smaller, thicker more venous leaves. I
retained such plants under my concept of A. petiolaris for several years, but closer
inspection has suggested that these are deserving of specific status. The distributional
relationship of A. ayerscottiana and A. petiolaris is shown in Figure 1.

It is a pleasure to name this isolated species in honor of Dr. Tina Ayers and her
husband Dr. Randy Scott, both having participated in the collection of the only two
specimens known to me. Tina and Randy obtained their doctorates under my
direction, and are currently located at Northern Anzona University, Flagstaff, Anzona.
Their wedded name also appears on one other Mexican species, Wedelia ayerscottiana
B.L. Turner.

AGERATINA MIAHUATLANA B.L. Turner, spec. nov. TYPE: MEXICO.
Oaxaca: Distrito Miahuatlén, Quiexobra, 3050 m, 22 Oct 1995, Hinton et al.
26304 (HOLOTYPE: TEX!).

Similis A. viscosissimae (Rolfe) R.M. King & H. Rob. sed involucns
majoribus (10-12 mm altis vice 6-8 mm altis) et setis papporum pluribus (ca.
30 vice 10-15).

Suffruticose herbs or shrublets 0.5-1.2 m high. Midstems 3-5 mm across,
densely pubescent with a vestiture of glandular trichomes ca. 0.25 mm high. Leaves
opposite throughout, but occasionally the uppermost alternate; those at midstem mostly
cordate; petioles 2-3 cm long; blades 5-7 cm long, 4-7 cm wide, thin, 3-nervate from
the base, moderately to sparsely pubescent above and below, the margins
crenulodentate. Heads arranged in relatively loose terminal cymes, the ultimate
peduncles mostly 1-3 cm long, pubescent like the stems. Involucres campanulate, 11-
12 mm high; bracts linear-lanceolate, 2-3 senate, subequal, glandular-pubescent, the
apices narrowly acute. Florets 20-30 per head (est.); corollas white, 6-7 mm long,
glabrous except for the sparsely pilose lobes. Achenes (immature) ca. 3 mm long,
hispidulous; pappus of ca. 30 readily deciduous white bristles ca. 6 mm long.

ADDITIONAL COLLECTIONS EXAMINED: MEXICO. Oaxaca: Distrito
Miahuatlan, Xianaguilla, 2715 m, 13 Oct 1995, Hinton et al 26062 (TEX); Siete
Ocotes, 2950 m, 20 Oct 1995, Hinton et al. 26258 (TEX).

Tumer: Two new Ageratina from México 311

Figure 1. Distribution of Ageratina petiolaris (closed circles) and A. ayerscottiana
(open circle). Based upon specimens at LL, TEX.

312 PHY OOIG PA October 1995 volume 79(4):309-3 12

Ageratina miahuatlana relates to a group of species with large heads and glandular -
pubescent foliage centering about A. viscosissima (Rolfe) King & H. Rob. The latter
occurs in northwesten México and belongs to the subgenus Ageratina (sensu King &
Robinson 1987). It differs from the latter in having leaves with shorter petioles and
larger heads, the involucres 10-12 mm long (vs. 6-8 mm long), and pappus of more
numerous bnistles (ca. 30 vs. 10-15).

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnoses, and to her and Justin
Williams for reviewing the paper.

LITERATURE CITED

King, R.M. & H. Robinson. 1987. The genera of the Eupatorieae (Asteraceae).
Monographs Syst. Bot. Missoun Bot. Gard. 22:1-581.

Turner, B.L. 1996. Asteraceae of Mexico vol. 1 (of a contemplated 10 vol. account):
in prep.

Turner, B.L. & G. Nesom. 1993. Biogeography diversity, and endangered or
threatened status of Mexican Asteraceae, in Biological Diversity of Mexico [T.P.
Ramamoorthy et al., eds.] Oxford Univ. Press, Oxford.

Phytologia (October 1995) 79(4):313-316.

A NEW SPECIES OF BOCCONIA (PAPAVERACEAE) FROM OAXACA,
MEXICO

B.L. Turner

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

ABSTRACT

Bocconia hintoniorum B.L. Turner, spec. nov., is described and
illustrated from Cerro Quiexobra, Distnto Miahuatldn, Oaxaca. It is a small
tree 3-5 m high having undivided, thick coriaceous leaves, and flowers with 7-
8 anthers. It is closely related to the more southern B. gracilis, differing from
the latter in having smaller, thicker leaves with minutely crenulodentate
margins and fewer anthers.

KEY WORDS: Papaveraceae, Bocconia, México, Oaxaca, systematics

Routine identification of Mexican plants has revealed the following novelty.

BOCCONIA HINTONIORUM B.L. Turner, spec. nov. Figures 1-2. TYPE:
MEXICO. Oaxaca: Distrito Miahuatldn, Cerro Quiexobra, 3070 m, 19 Oct 1995,
Hinton et al. 26227 (HOLOTYPE: TEX).

Similis Bocconiae gracili Hutch. sed foliis crassioribus glabrisque,
marginibus uniformiter minuteque crenulatis-dentatis, et antheris 7-8 (vice ca.
12).

Small tree 3-5 m high. Young stems densely hirsute. Leaves'12-13 cm long, 2-3
cm wide, pubescent at the base like the stem, often winged throughout by the
gradually tapering blades, the latter narrowly elliptic to elliptic-oblanceolate, pinnately
nervate, the margins minutely crenulodentate for about 2/3 of their length. Flowers
arranged in terminal panicles ca. 30 cm long, 10 cm across, the pedicels mostly 4-10
mm long, glabrous. Sepals 9-11 mm long, 2.5-3.0 mm wide, the apices abruptly
constricted forming a lanceolate extension ca. 2 mm long. Petals absent. Stamens 7
or 8. Fruits on recurved pedicels at matunty, glaucous-black, glabrous. Seeds ovoid,
ca. 4mm long, 3 mm across, the caruncle broadly conical, ca. 2 mm long.

313

314 PHYTOLOGIA October 1995 volume 79(4):3 13-316

Figure 1. Leaves of Bocconia hintoniorum: left side (lower surface); nght side (upper
surface); circular inset (undersurface, showing detail); from holotype.

Turer: New Bocconia from México 315

r Wann rm eT me mney

aT. LTO Y) Sas whe
a ee am ‘ er

Figure 2. Flower of Bocconia hintoniorum with one of the two sepals removed (from
holotype).

316 PHY TOLOGTA October 1995 volume 79(4):3 13-316

This newly described taxon first came to my attention in the fall of 1980 while on a
Bocconia collecting expedition with Ms. Joan Johnson (accompanied by Dr. David
Northington and Dr. Wayne Elisens). Ms. Johnson was in the early stages of a
doctoral systematic study of Bocconia, having borrowed a wide range of matenal from
various institutions so as to prepare herself for the field tnp concerned. We collected
the commonly occurring bocconias throughout most of México (mainly B. frutescens
L., including B. latisepala S. Wats.), but were startled to find small populations of the
presently described species along highway 175 in the vicinity of Miahuatlan, Oaxaca.
Unfortunately, Ms. Johnson abandoned her doctoral program and failed to preserve
the various collections made during this sojourn. She also left me, her major
professor, with a large set of Bocconia specimens to annotate and return to vanous
institutions, none of these representing the species described herein. Thus my delight
to find among Hinton’s numerous collections from Cerro Quiexobra, newly assembled
specimens that might serve as type maternal for this long-remembered but unnamed
taxon.

Bocconia hintoniorum will key to B. integrifolia Kunth in Standley’s (1922) Trees
and Shrubs of Mexico. The latter, however, is typified by Peruvian material and, as
noted by Hutchinson (1920) in his account of the genus, is restricted to South
America. Although the matenal of B. hintoniorum will key to B. integrifolia in the
treatment of Hutchinson, it is seemingly more closely related to the Central American
B. gracilis Hutch., with which it is compared here.

It is a pleasure to name this attractive new species for the Hinton family, whose
collections in México are becoming increasingly legendary.

ACKNOWLEDGMENTS

I am grateful to Gayle Tumer for the Latin diagnosis, and to her and Ted
Delevoryas for reviewing the manuscript.

LITERATURE CITED

Hutchinson, J. 1920. Bocconia and Macleaya. Kew Bull. 1920:275-282.
Standley, P.C. 1922. Bocconia, in Trees and Shrubs of Mexico. Contr. U.S. Natl.
Herb. 23:299-301.

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