LIBRARY OF THE UNIVERSITY OF ILLINOIS AT URBANA-CHAMPAIGN 580.5 F6 33-35 BIOLOGY The person charging this material is re- sponsible for its return to the library from which it was withdrawn on or before the Latest Date stamped below. Theft , and underlining of books or. r^son, octlon and may result In dismissal from ec UN.VERS.TY Of IlilNOIS Ui«AtY AT URBANA-CHAMPA.GN L161— O-1096 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 Q CO C3 95 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 UNIVERSITY OF ILLINOIS URBANA 580 5FB C001 FIELDIANA. 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