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UN.VERS.TY Of IlilNOIS Ui«AtY AT URBANA-CHAMPA.GN
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I
FIELDIANA
1 1 1 ' r n ~
B°tany ^*^
Published by Field Museum of Natural History
Volume 34, No. 7 May 15, 1972
Synopsis of Hemichaena, including Berendtiella
(Scrophulariaceae)
JOHN W. THIERET
UNIVERSITY OF SOUTHWESTERN LOUISIANA, LAFAYETTE
The tribe Cheloneae sensu Wettstein (1891) of the Scrophulari-
aceae is a heterogeneous assemblage of genera whose principal com-
mon feature is the cymose disposition of the flowers. Of the various
steps that have been taken to remodel the tribe, one is of concern
here. The genera Leucocarpus, Berendtiella (Berendtia), and Hemi-
chaena possess floral characteristics that indicate their alliance with
Mimulus and so have been transferred to the Gratioleae. These
three genera are rather similar in certain vegetative features and in
inflorescence, suggesting close affinity. Recently, Dr. Louis O. Wil-
liams raised the question of possible realignment of the genera to
express their relationships better. After study of the problem I have
concluded that the complex represents but two genera: Leucocarpus
and Hemichaena (including Berendtiella). Hemichaena and Berendt-
iella are much alike in habit, habitat, vestiture, foliage, inflorescence,
flowers, fruits, and seeds, all of which convince me that the taxa are
congeneric.
Leucocarpus is distinguished from the emended Hemichaena by
its baccate — rather than capsular— fruit and by its seeds with their
distinctive intra-reticular lines. Its closest ally would appear to be
the Malaysian Cyrtandromoea (Burtt, 1965), rather than Hemichaena.
The genus Hemichaena, founded upon a Hartweg collection from
Guatemala, was established by Bentham in 1841. Twenty-seven
years later, in 1868, Asa Gray described the genus Berendtia, then
known only from Mexico. In 1889 the name Berendtia Gray was
shown by Wettstein and Harms to be a later homonym of Berendtia
Goeppert, a genus based, in 1845, on one corolla with epipetalous
stamens found in Baltic amber. Although Wettstein and Harms
Library of Congress Catalog Card Number: 71-18^078 f\ r\ -~ f\ <
OC ?5
Publication 1148 89
BIOLOGY
101 BURRILL HAH
90 FIELDIANA: BOTANY, VOLUME 34
then published the new name Berendtiella for Berendtia Gray, the
new combinations for the four species of the genus were not made
until 1955 (Thieret, 1955).
The inflorescences of Hemichaena and Berendtiella are bracteolate
cymes, those of Berendtiella being generally fewer flowered (1-5) than
those of Hemichaena. However, the cymes of Hemichaena, although
being as many as 12-flowered, are commonly only 1-5-flowered.
A common and curious feature of these plants is the presence of
what I have called "superposed" inflorescences, i.e., two inflores-
cences in each axil, one above the other. I interpret these as being
derived from superposed axillary buds. A detailed developmental
study of the inflorescence is needed to confirm or revise my interpre-
tation. The seeds, small and reticulate, are of a type common in the
tribe Gratioleae. The walls of the reticulum are thin, translucent,
and curiously radially marked.
Hemichaena and Berendtiella, native in a region whose Scrophu-
lariaceae are poorly represented in literature, have not often been
contrasted in keys. Indeed, I know of only three keys that include
both genera, those in Standley (1924), Thieret (1954), and Wettstein
(1891). Characteristics used to distinguish the genera include flower
color (Thieret, Wettstein), the length of the corolla tube in relation
to the calyx (Wettstein), position of stamen insertion in the corolla
tube (Wettstein), and depth of lobing and shape of the calyx (Stand-
ley, Thieret). Study of specimens rather than keys reveals that each
of these characteristics is a poor one, sharper on paper than in nature,
on which to base separation of the genera.
Flower color — red in Berendtiella, yellow in Hemichaena — works
well in Central America, which is why I used this feature in my key
to Central America scrophulariaceous genera. However, it is of no
use in Mexico where two of the species of Berendtiella have yellow
corollas.
According to Wettstein's key, Hemichaena is distinguished by its
corolla tube being "significantly" longer than the calyx, that of
Berendtiella little exceeding it. The tube of Hemichaena is indeed
about twice as long as the calyx. The tube of Berendtiella, however,
ranges from included or but slightly exserted to 3 or %l/2 times as
long as the calyx.
The position of insertion of the stamens ranges from about one-
fourth way up the corolla tube in Hemichaena to about one-third to
one-half way up in Berendtiella. In an occasional flower of Hemi-
chaena, the lower pair of stamens may be adnate even a bit more than
FIG. 1. Calyces of Hemichaena. A, H. spinulosa, shallowly lobed calyx, X9,
Johnston & Muller 1142 (GH); B, H. spinulosa, deeply lobed calyx, X9, Pringle
2663 (F); C, H. coulteri, X8.5, Moore 5431 (GH); D, H.frulicosa, shallowly lobed
calyx, X3.5, Steyermark 34958 (F); E, H. fruticosa, deeply lobed calyx, X3.5,
Standley 83910 (F).
91
92 FIELDIANA: BOTANY, VOLUME 34
one-fourth way up the tube, approaching but not quite reaching the
one-third level of some Berendtiella. The differences in stamen inser-
tion, then, appear to be ones of degree only.
In the words of Standley (1924), the calyx of Hemichaena is
"capanulate, 5-lobate" and that of Berendtiella is "tubular-campanu-
late, 5-dentate." Study of the calyces of Hemichaena and Berendt-
iella reveals these to be rather tenuous distinctions. In Hemichaena,
the calyx ranges from campanulate to tubular; in Berendtiella, from
narrowly tubular-campanulate to tubular. In Hemichaena, the per-
centage of total length of the calyx that is contributed by the lobes
ranges from about 35 to 53 per cent; in Berendtiella it ranges from
about 14 to 30 per cent (fig. 1). These differences are also ones of
degree only.
Thus, the characteristics that have been used to maintain Hemi-
chaena and Berendtiella as separate genera do not hold up under care-
ful examination. I have been unable to find others to replace them.
The conclusion that Hemichaena and Berendtiella are one seems rea-
sonable.
In addition to H. spinulosa, shown in Figure 2, only two other
species of the genus are illustrated in the literature: H. fruticosa, in
Curtis' 's Botanical Magazine 101: Tab. 6164 (1875); and H. fruticosa
and H. rugosa in the forthcoming treatment of the Scrophulariaceae
in Standley and Williams' "Flora of Guatemala." Documented dis-
tribution of the five species of Hemichaena is shown in Figures 3 and
4.
The specimens on which this study was based were received on
loan from Field Museum of Natural History (F), United States
National Museum (US), and Arnold Arboretum (A) and Gray Her-
barium (GH) of Harvard University.
Hemichaena Bentham, PI. Hartw. 78. 1841. Berendtia Gray,
Proc. Am. Acad. 7: 379. 1868, non Goeppert, 1845 (type species: B.
ghiesbrechtii Gray) . Berendtiella Wettst. et Harms in Engl. et Prantl, I
Pflanzenf., Gesamtregister zum II. bis IV. Teil: 459. 1899.
North American herbs or shrubs. Leaves opposite or fascicled; inflorescences
axillary, cymose. bracteolate, 1-12-flowered; flowers zygomorphic; calyx campan-
ulate to tubular, angled or prismatic, 5-toothed to 5-lobed; corolla 2-lipped, yellow,
orange, or red, tube equalling or exceeding the calyx, lobes spreading, rounded,
equalling or shorter than the tube the two posterior free or united nearly to apex;
stamens 4, didynamous, adnate to the corolla tube, included to exserted, the
thecae at length divergent; stigmas 2, plate-like; fruit a loculicidal, many-seeded
capsule; seeds reticulate.
FIG. 2. Hemichaena spinulosa. Flowering branch, Pringle 11645 (F). The ver-
tical line represents 5 cm.
93
94 FIELDIANA: BOTANY, VOLUME 34
Type species. — H. fruticosa Bentham.
Leaves clasping; stamens included .2. H. fruticosa.
Leaves not clasping; stamens included or exserted.
Corolla yellow.
Corolla tube included in or but slightly exserted from the calyx;
peduncles 1-5 flowered I. H. coulter i.
Corolla tube 1 V^ to 2 times as long as the calyx; peduncles 1-flowered.
5. H. spinulosa.
Corolla orange to red.
Plant vernicose, glabrous 3. H. letigata.
Plant glandular pubescent 4. H. rugosa.
1. Hemichaena coulter! (Gray) Thieret, comb. nov.Berendtia
coulteri Gray, Proc. Am. Acad. 7: 380. 1868 (Holotype: Mexico,
Coulter 1335 [GH]). Berendtiella coulteri (Gray) Thieret, Ceiba 4:
305. 1955.
Erect shrubs to 2 m., glandular pubescent; leaves opposite, elliptic, 2-5.8 cm.
long, 0.6-3.1 cm. wide, the margins revolute, entire to obscurely toothed or undu-
late, the base acute to rounded, the apex acute to obtuse, the petioles 1-2 mm.
long, obscure, much flattened at base, the base persisting as a scale after the fall of
the blade; inflorescences solitary or superposed, axillary to leaves; peduncles 1-5-
flowered, to 15 mm. long, primary bracteoles linear to narrowly obovate, to 6 mm.
long and 1.5 mm. wide, in 1-flowered inflorescences inserted below the middle of
the peduncle, sometimes lacking, secondary bracteoles lacking; pedicels to 5 mm.
long; calyx tubular to narrowly tubular campanulate, 5-8 mm. long, 5-toothed to
5-lobed, the teeth or lobes triangular, 0.5-2 mm. long; corolla yellow, 1.3-1.5 cm.
long, its tube included in or but slightly exceeding the calyx; stamens exserted,
anthers 1.6-1.8 mm. long; capsule ovate in lateral view, 7-8 mm. long.
Dry, rocky places in mountains at 5,000-6,000 ft. elevation,
Hidalgo (fig. 3).
2. Hemichaena fruticosa Benth. PI. Hartw. 78. 1841 (Holo-
type: Guatemala, Quetzaltenango, Hartweg [K; not seen]). Leuco-
carpus fruticosus (Benth.) Benth. in DC. Prodr. 10: 336. 1846.
Erect herbaceous, suffrutescent, or shrubby plants to 2 m., simple or branched,
glandular pubescent; leaves opposite, sessile, ovate or elliptic, sometimes narrowly
so, 4-16 cm. long, 1.5-5.5 cm. wide, the margins slightly revolute, coarsely to finely
toothed, usually less so or even entire toward the base, the base slightly clasping to
cordate-amplexicaul, sometimes slightly pandurate, the apex acute to acuminate;
inflorescences solitary or superposed, axillary to leaves; peduncles 1-12-flowered,
to 4 cm. long, primary bracteoles narrowly ovate, obovate, or elliptic, to 22 mm.
long and 7 mm. wide, in 1-flowered inflorescences inserted near the middle of the
peduncle, secondary bracteoles similar but smaller, elliptic to subulate, to 10 mm.
long and 4 mm. wide, sometimes lacking; pedicels to 20 mm. long; calyx tubular to
campanulate, 14-17 mm. long, 5-lobed, the lobes elongate triangular, to 8 mm.
-C
1
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CO
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96 FIELDIANA: BOTANY, VOLUME 34
long; corolla yellow, 2.5-3.2 cm. long, its tube 2 times as long as the calyx; stamens
included, anthers 3 mm. long; capsule ovate to elliptic in lateral view, 12-15 mm.
long.
Forests, open slopes, cliff faces, river banks, sandbars, rocky
places, thickets, and roadsides in mountains at 3,000-11,500 ft. ele-
vation, Oaxaca and Chiapas, Guatemala, Costa Rica (fig. 3).
3. Hemichaena levigata (Robins, et Greenm.) Thieret, comb,
nov. Berendtia levigata Robins, et Greenm. Proc. Am. Acad. 32: 39.
1897 (Holotype: Puebla, calcareous ledges near Tehuacan, 24 Decem-
ber 1895, Pringle 6294 [US; isotypes at A, F, GH, US]). Berendtiella
levigata (Robins, et Greenm.) Thieret, Ceiba 4: 305. 1955.
Erect shrubs to 0.9 mm., glabrous, vernicose especially in the younger parts;
leaves opposite or fascicled, sometimes clustered on spur shoots, elliptic, ovate, or
rhombic, sometimes narrowly so, 1.5-5 cm. long, 0.4-2.2 cm. wide, the margins
usually revolute, entire to distally several toothed, the teeth widely spaced, tha
base acute to attenuate, the apex acute to broadly acute, petioles lacking or up to
8 mm. long, winged distally; inflorescences solitary, axillary to leaves; peduncles
1-flowered, to 20 mm. long, bracteoles inserted at, above, or below the middle of
the peduncle, often leaflike, linear to narrowly elliptic or ovate, to 14 mm. long and
3.8 mm. wide, sometimes with rudimentary buds in their axils; calyx tubular to
narrowly tubular campanulate, 9-15 mm. long, 5-toothed to 5-lobed, the teeth or
lobes triangular to subulate-tipped triangular, 2.5-3.5 mm. long; corolla reddish
orange, 2.5-3.5 cm. long, its tube 3 times as long as the calyx; stamens barely or
not exserted, anthers 2-2.5 mm. long; capsule narrowly elliptic or ovate in lateral
view, 9-11 mm. long.
Ledges and rocky soil in mountains at 5,000-6,000 ft. elevation,
Puebla and Veracruz (fig. 3).
4. Hemichaena rugosa (Benth. in DC.) Thieret, comb. nov.
Diplacus rugosus Benth. in DC. Prodr. 10: 368. 1846 (Holotype:
Chiapas, Linden 201 [K; not seen]). Berendtia rugosa (Benth. in
DC.) Gray, Proc. Am. Acad. 7: 380. 1868. B. ghiesbrechtii Gray,
Proc. Am. Acad. 7: 380. 1868 (Holotype: Chiapas, Ghiesbrecht &
Berendt 134 [GH]). Berendtiella rugosa (Benth. in DC.) Thieret,
Ceiba 4: 305. 1955.
Erect, arching, or sometimes pendent shrubs to 4 m., glandular pubescent;
leaves opposite or fascicled, ovate or elliptic, sometimes narrowly so, 2-8 cm. long,
0.7-3.5 cm. wide, the margins revolute, coarsely to finely many to few toothed, the
base cuneate to attenuate, the apex acute to rounded, the petioles 1-7 mm. long,
obscure, winged, much flattened at base, the base persisting as a scale after the fall
of the blade; inflorescences solitary or superposed, axillary to leaves or to indurate-
based bracts of compact shoots; peduncles 1-3-flowered, to 15 mm. long, primary
bracteoles subulate or linear to narrowly elliptic or obovate, to 10 mm. long and 4
mm. wide, in 1 flowered inflorescences inserted below the middle of the peduncle,
97
98 FIELDIANA: BOTANY, VOLUME 34
secondary bracteoles subulate, to 2 mm. long, sometimes lacking; pedicels to 12
mm. long; calyx tubular to narrowly tubular campanulate, 8-14 mm. long, 5-
toothed, the teeth subulate to subulate-tipped triangular, 1-2 mm. long; corolla
orange to red, 2.5-3.5 cm. long, its tube 2.5-3.5 times as long as the calyx; stamens
exserted, anthers 1.5-2 mm. long; capsule ovate in lateral view, 8-10 mm. long.
Rocky slopes, cliff faces, roadbanks, and forests in mountains at
4,600-10,000 ft. elevation, Chiapas to Honduras (fig. 4).
5. Hemichaena spinulosa (S. Wats.) Thieret, comb. nov.
Berendtia spinulosa S. Wats. Proc. Am. Acad. 25: 159. 1890 (Holo-
type: Nuevo Leon, dry limestone cliffs of the Sierra Madre near
Monterey, 27 June 1888, Pringle 1952 [US; isotypes at USJ). Be-
rendtiella spinulosa (S. Wats.) Thieret, Cieba 4: 305. 1955.
Erect, pendent, or trailing shrubs to 0.9 m., glandular pubescent; leaves oppo-
site or fascicled, elliptic, ovate, or obovate; sometimes narrowly so, 0.5-3.2 cm.
long, 0.1-1.1 cm. wide the margins revolute, entire to several toothed, the teeth
widely spaced, the base acute to attenuate, the apex acute to rounded, the petioles
1-2 mm. long, obscure, much flattened at base, the base persisting as a scale after
the fall of the blade; inflorescences solitary or very rarely superposed, axillary to
leaves; peduncles 1-flowered, to 10 mm. long, sometimes persisting and "thornlike"
on dead branches, bracteoles inserted at or above the middle of the peduncle, sub-
ulate to elliptic, to 2 mm. long and 0.7 mm. wide, sometimes lacking; calyx tubular
to narrowly tubular campanulate, 5-6.5 mm. long, 5-toothed to 5-lobed, the teeth
or lobes very broadly depressed triangular to triangular, 0.5-1.7 mm. long; corolla
yellow, 1.3-1.8 cm. long, its tube 1.5-2 times as long as the calyx; stamens exserted,
anthers 0.7-1 mm. long; capsule narrowly ovate in lateral view, 4.5-6 mm. long.
Ledges, crevices, and talus slopes in mountains at 2,000-3,000 ft.
elevation, Coahuila to Nuevo Leon-Chihuahua border region (fig. 4).
REFERENCES
BURTT, B. L.
1965. The transfer of Cyrtandromoea from Gesneriaceae to Scrophulariaceae
with notes on the classification of that family. Bull. Bot. Surv. India 7: 73-88.
STANDLEY, P. C.
1924. Trees and shrubs of Mexico (Passifloraceae-Scrophulariaceae). Contr.
U. S. Nat. Herb. 23: 849-1312.
THIERET, J. W.
1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4:
164-184.
1955. The status of Berendtia A. Gray. Ceiba 4: 304-305.
WETTSTEIN, R. VON
1891. Scrophulariaceae. In Engler, A., and Prantl, K. Die Natur. Pflanzenfam.
IV. 3b: 39-107.
THIERET: SYNOPSIS OF HEMICHAENA 99
LIST OF EXSICCATAE
Explanation of citations: name of collector, collection number (number of the
species as it appears in this work).
Breedlove 7160 (4), 8001 (4), 8568 (2), 11682 (4), 13766 (4); Carlson 3597 (2);
Coulter 1335 (1); Cruz 133 (2); Eggler 494 (2); Ghiesbrecht & Berendt 134 (4);
Ghiesbrecht 504 (4) ; Heyde & Lux 3113 (4) ; Hunnewell 17240 (4), 17243 (2) ; Jimenez
M. 893 (2), 1432 (2); Johnston 1334 (2), 9140 (5); Johnston & Mutter 1142 (5);
Kellerman 5938 (2); Leavenworth 161 (5); Liebmann 9466 (3); Marsh 1969 (5);
Matuda S-44 (2), 131 (2), 16262 (2), 17674 (2); Melhust & Goodman 3586 (2);
Molina R. 21189 (4); Molina R., Burger, & Wallenta 16421 (2), 16423 (4), 16552 (4);
Moore 5431 (1); Muller 2873 (5), 3078 (5); Nelson 2501 (2), 3775 (2), 3788 (2);
Purpus 461 (3), 6883 (2), 7287 (2), 10692 (3); Pringle 1952 (5), 2663 (5), 6294 (3),
9386 (3), 11645 (5); Relso 4220 (3); Rose & Hay 5849 (3); Rose, Painter, & Rose
9890 (3); Rzedowski 25460 (1); Sesse, Mocino, Castillo, & Maldonado 2581 (3);
Shannon 289 (4), 339 (2); Skutch 1134 (4), 1172 (2), 1201 (2), 1527 (2), 2996 (2);
Smith, Peterson, & Tejeda 3791 (3); Smith & Tejeda 4406 (3); Soils 346 (2); Stand-
ley 41880 (2), 42227 (2), 42645 (2), 58345 (2), 65341 (2), 65633 (4), 65660 (4), 65667
(4), 65812 (2), 80569 (2), 81394 (4), 81399 (4), 81986 (2), 82178 (4), 82188 (4),
82215 (4), 82552 (4), 83380 (2), 83910 (2); Standley & Williams 4539 (4); Stewart
1092 (5); Steyermark 34958 (2), 35955 (2), 36308 (2), 42951 (4), 47486 (2), 50350
(4), 50820 (4), 52036 (4); Stork 1596 (2); Valerio 1208 (2); Williams 16102 (2),
19688 (2); Williams, Jimenez M., & Williams 24180 (2); Williams & Merrill 15705
(4); Williams & Molina R. 11207 (4), 11503 (4), 12000 (4), 14994 (4), 23259 (4);
Williams, Molina R., & Williams 21809 (4), 21828 (2), 22123 (2), 22506 (4), 23115
(2), 25664 (2), 26071 (2), 26196 (2); Williams, Molina R., Williams, & Gibson
28305 (2); Williams & Williams 21725 (4).
f
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