Familia Acanthaceae en Panamá

Additions to the Acanthaceae of PanamaAuthor(s): Thomas F. Daniel and Lucinda A. McDadeSource: Annals of the Missouri Botanical Garden, Vol. 82, No. 4 (1995), pp. 542-548Published by: Missouri Botanical Garden PressStable URL: http://www.jstor.org/stable/2399835 .Accessed: 10/04/2011 20:00

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ADDITIONS TO THE Thomas F. Daniel2 and ACANTHACEAE OF PANAMA' Lucinda A. McDade:

ABSTRACT

Nine species of Acanthaceae are added to the known flora of Panama, including the newly described species Aphelandra kuna. Range extensions into Panama are reported for A. scolnikae, Justicia ephemera, Odontonema cuspidatum, 0. rubrum, Sanchezia lutea, and S. parvibracteata. Revised keys to all Panamanian species of Aphelandra, Odontonema, and Sanchezia are provided. Two cultivated species, Pachystachys lutea and Justicia betonica, are also recorded from Panama for the first time.

The treatment of Acanthaceae for the Flora of Panama (Durkee, 1978) is now 17 years old. It serves as an extremely helpful facilitator for rec- ognizing additions to the acanthaceous flora of Pan- ama. Most of the new taxa and distributional rec- ords discovered since 1978 have been in regions remote from the well-collected provinces of central Panama. Dari6n Province and the Comarca de San Blas in eastern Panama (adjacent to Colombia) and the Caribbean slope of western Panama (which is becoming increasingly accessible via new roads) have yielded most of the additions of Acanthaceae to the Panamanian flora. Further discoveries are to be expected in these regions as they become more thoroughly explored. On the other hand, because of abundant "geographic taxonomy" in the Neotropics, some species undoubtedly will be combined with others as more collecting and study make geographic patterns apparent.

In this paper, we add nine species to the Acan- thaceae known from Panama. These include the de- scription of a new species of Aphelandra; docu- mentation and discussion of six species in four genera that are newly reported from Panama; and notes on the occurrence of two species that are cultivated in Panama. We also provide revised keys to all Panamanian species of Aphelandra, Odontone- ma, and Sanchezia.

A NEW SPECIES OF APHELANDRA

About 175 species are currently recognized in the wholly neotropical genus Aphelandra (Wasshau- sen, 1975; McDade, 1984; Daniel, 1991). The up- dated checklist for Panama (D'Arcy, 1987) includ-

ed a number of nomenclatural changes and new taxa that had been recognized in the genus since Durkee's (1978) publication. D'Arcy (1987) listed a total of 19 species. McDade (1984) has argued that two of these do not actually occur in Panama. The two Panamanian specimens cited by Durkee (1978) as A. crenata Leonard (otherwise known only from South America) are readily referable to A. campanensis Durkee (Wedel 2501) and A. hartwegiana Nees (Allen 5094). These three species are part of a lineage with branched pedunculate inflorescences, obtuse bracts, falcate bracteoles, and distinctive pollen (see McDade, 1984); the Central American taxa should be viewed as provisional pending revision of the entire group. Durkee (1978) identified a number of plants from the ridges of the low, central Panamanian ranges as A. pilosa Leon- ard. Aphelandra pilosa is a South American species with profusely branching inflorescences, short bracts (5-6 mm), and relatively short corollas (ca. 60 mm). It is unlike any Aphelandra that occur in Central America. On the basis of morphology and experimental hybridizations, McDade (1984) argued that specimens treated by Durkee (1978) as A. pilosa are likely to be hybrids between A. sinclairiana Nees and A. gracilis Leonard. With the removal of A. crenata and A. pilosa and the additions (see below) of a new species and a geographic range extension, there are a total of 19 species of Aphelandra in Panama and one putative hybrid.

Aphelandra kuna T. F. Daniel & McDade, sp. nov. TYPE: Panama. Panama: El Llano-Cartf Hwy., ca. 12 km N of El Llano, 19 July 1974,

' We are grateful to S. Myers for rendering the line drawing, D. Ubick for SEM assistance, and the curators of herbaria cited in the text for loans. Daniel's fieldwork in Panama was generously facilitated by G. McPherson and H. Herrera. McDade's fieldwork was supported by the National Geographic Society (grant to W. Starnes).

2 Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. 3Department of Ecology and Evolutionary Biology and Herbarium, University of Arizona, Tucson, Arizona 85721,

U.S.A.

ANN. MISSOURI BOT. GARD. 82: 542-548. 1995.

Volume 82, Number 4 Daniel & McDade 543 1995 Acanthaceae of Panama

R. Dressler 4667 (holotype, US; isotype, DUKE). Figure 1.

Herbae perennes usque ad 1.3 m altae. Folia opposita, petiolata, laminae ellipticae vel obovato-ellipticae, 125- 240 mm longae, 48-91 mm latae, 2.2-2.9-plo longiores quam latiores. Spicae dense bracteatae, 60-100 mm longae. Bracteae subroseae, ovatae vel ligulatae, erectae, (26-)30-49 mm longae, 10-17 mm latae, margine den- tatae dentibus utroque latere 3-4. Corolla atropurpurea, 43-45 mm longa, extus dense sericea. Stamina inclusa, 4.5-5.5 mm longa, thecae 2.8-3.5 mm longae, apice pubescentes.

Monocaulous perennial herbs to 1.3 m tall. Young stems subquadrate (often somewhat collapsed in dried specimens) or flattened, sparsely pubescent with appressed eglandular trichomes to 1 mm long or nearly glabrate. Leaves opposite, pet- iolate, petioles to 50 mm long, blades elliptic to obovate-elliptic, 125-240 mm long, 48-91 mm wide, 2.2-2.9 times longer than wide, gradually attenuate to decurrent along petiole at base, acuminate at apex, the surfaces sparsely pubescent with appressed eglandular trichomes or becoming glabrate on adaxial surface. Inflorescence a terminal, sessile to short-pedunculate (peduncles to 5 mm long), densely bracteate spike, 60-100 mm long (excluding flowers), 32-45 mm wide (excluding flowers) near midspike, rachis pubescent with an- trorsely appressed eglandular trichomes 0.4-1 mm long (sericeous). Bracts pinkish green to pinkish purple, ovate to strap-shaped, erect, (26-)30-49 mm long, 10-17 mm wide, attenuate to long-attenuate at apex, abaxial surface sericeous with eglandular trichomes, lacking nectariferous glands, margin conspicuously dentate (except for proximalmost bracts) with 3-4 teeth per side, the teeth 1.5-7 mm long. Bracteoles lance-subulate, strongly condupii- cate, 19-31 mm long, 1.6-3 mm wide, pubescence similar to bracts. Calyx 5-lobed, 8-10 mm long, lobes lanceolate to subulate, unequal in size (i.e., one conspicuously longer and wider than others, the remaining lobes subequal to unequal in size), 6-9.7 mm long, 0.4-1 mm wide, abaxial surface sparsely pubescent to nearly glabrous. Corolla dark purplish with white on lower lip, 43-45 mm long, externally densely sericeous (especially the tube), upper lip 8-10 mm long, bilobed with rounded to acute lobes 3.5-5.5 mm long, 1.5-2.5 mm wide, lower lip 11-12 mm long, deeply trilobed, lateral lobes linear-elliptic, 8.5-11.5 mm long, 2.5-4 mm wide, lower central lobe elliptic, 9-12 mm long, 4.2-7.2 mm wide. Stamens inserted in distal 1/3 of corolla tube, included, 4.5-5.5 mm long, thecae 2.8-3.5 mm long, apically pubescent; staminode 1 mm long. Style 32 mm long, stigma asymmetrically

funnelform, 1 mm long. Capsule and seeds not seen.

Distribution and habitat. Known only from the vicinity of the type locality in northeastern Panama (Panama and Comarca de San Blas), in the Serrania de San Blas; plants occur along streams in a region of tropical wet forest at elevations from 300 to 400 m.

Phenology. Flowering: June-July, September, November.

Paratypes. PANAMA. Panama: El Llano-Carti Rd., 10-12 km from El Llano, 12 Nov. 1974, P Maas & R. Dressler 1706 (U, US). Comarca de San Blas: El Llano- Cartif Rd., 18 km from Interamerican Hwy., 9'19'N, 78055'W, 7 Sep. 1984, G. de Nevers 3876 (CAS); Km 16.7 of El Llano-Carti Rd., 9'19'N, 78055'W, 16 June 1985, G. de Nevers 5906 (DUKE).

Two of the known collections of Aphelandra kuna were identified as A. tridentata Hemsl., a species known only from Costa Rica that bears a superficial resemblance. Using keys to species of Aphelandra in southern Central America (Durkee, 1978, 1986), one might mistake A. kuna for either A. tridentata or A. aurantiaca (Scheidw.) Lindl. It differs from the former by its pinkish (vs. green) bracts with 3-4 (vs. 1-2) teeth per side, 19-31 (vs. 11-12) mm long bracteoles that are strongly (vs. slightly, if at all) conduplicate, purplish (vs. red) corollas that are externally densely sericeous (vs. pubescent with flexuose trichomes) and have a deeply (vs. shallowly) trilobed lower lip with lobes 8.5-11.5 (vs. up to 1) mm long, included (vs. exserted from mouth of corolla) stamens with smooth (vs. papillose) filaments, and thecae 2.8- 3.5 (vs. 5.2-6.5) mm long. Aphelandra tridentata is presently known only from central and northern Costa Rica. Aphelandra kuna can be distinguished from the wide-ranging A. aurantiaca by its eglandular (vs. glandular) pubescent bracts with 3-4 (vs. 5-15) marginal teeth per side, eglandular (vs. glandular) bracteoles that vary from 19 to 31 (vs. 7-15) mm in length, purplish (vs. orangish or red) corollas that vary from 43 to 45 (vs. 50-63) mm in length, included (vs. exserted from mouth of corolla) stamens, and thecae 2.8-3.5 (vs. 4-5) mm long. Aphelandra kuna might also be mistaken for A. dolichantha Donn. Sm. (cf. "dolicantha" in Durkee, 1978, and D'Arcy, 1987), an unusual species that also occurs near the crest of the El Llano-Cartf Road. Aphelandra dolichantha can be readily distinguished by its entire bracts subtend- ing flowers with minute calyx lobes and white corollas. Further elucidation of relatives within the genus must await a detailed subgeneric treatment

544 Annals of the Missouri Botanical Garden

1 mm

FIGURE 1. Aphelandra kuna T. F. Daniel & McDade. A. Habit (Dressier 4667). B. Bract (Maas & Dressier 1706). C. Bracteole (Maas & Dressier 1706). D. Base of one bracteole and calyx (Maas & Dressier 1706). E. Apex of bract and corolla (Dressier 4667). F. Anthers (Dressier 4667). G. Distal portion of style (Dressier 4667). Drawn by Sheva Myers.


of Aphelandra. None of the South American species examined by us or treated by Wasshausen (1975) is likely to be confused with A. kuna. A revised key to all of the Panamanian species is provided below.

Pollen of Aphelandra is generally 3-colpate (Wasshausen, 1975; McDade, 1984; Daniel, 1991). Pollen of A. kuna (i.e., Maas & Dressler 1706) was observed with a scanning electron microscope. All pollen was either partially collapsed or otherwise degraded. Although the number and type of aper- tures from this collection could not be determined, the exine surface is reticulate as in some other species of the genus.

The specific epithet is derived from the Kuna people who live in the region where this species occurs and who are making concerted efforts to preserve native plant communities in the Serranfa de San Blas.

SPECIES NEWLY REPORTED FROM PANAMA

APHELANDRA SCOLNIKAE LEONARD

This species was described by Leonard (1953) from plants collected at 1700-2500 m elevation in

several passes in the mountains surrounding Me- dellfn, Antioquia Province, Colombia. Until re- cently, the species was known only from this region. A collection from Panama (Dari6n: Cerro Mali, ca. 22 km E of Pucuro, 1250-1500 m, 20-26 Oct. 1987, G. de Nevers et al. 8475, CAS, MO) extends the range of this unusual species considerably. Aphelandra scolnikae is a member of the A. pulcherrima (Jacq.) Kunth complex (McDade, 1984); within that group, it is a member of the lineage marked by patches of numerous small (less than 0.1 mm in diameter) extrafloral nectaries on the floral bracts. Plants of A. scolnikae can be distinguished from others in this lineage by their suffru- tescent to treelet habit (vs. monocaulous herbs), relatively small leaves (up to about 22 cm long vs. up to 40 cm long in other species), and densely pubescent (vs. glabrous to puberulent) floral bracts with comparatively few extrafloral nectaries per glandular patch (mean of about 50 vs. more than 100 in other species).

The following key distinguishes all of the species of Aphelandra now known to occur in Panama:

la. Floral bracts marginally toothed. 2a. Floral bracts with paired lateral patches of nectariferous glands.

3a. Floral bracts apically obtuse, 14-20 mm wide, usually entire, rarely with 2-3 pairs of minute marginal teeth each less than 1 mm long; corolla 65-71 mm long -A. sinclairiana Nees

3b. Floral bracts apically attenuate, 6-9 mm wide, with 2-4 pairs of well-developed marginal teeth each 1-2 mm long. 4a. Corolla 36-41 mm long, thecae 3-4 mm long; plants of lowland deciduous forests and edges

--- -- - -- - - -- - -- - - -- - -- - - -- - -- - -- - - -- - -- - - -- - -- - - -- - -- - -- - - -- - -- - - -- - -- - - -- - -- - AA . sc a b ra (V ah l) S m . 4b. Corolla 55-70 mm long; thecae 4-5 mm long; plants of cloud forests - A. panamensis McDade

2b. Floral bracts lacking nectariferous glands. 5a. Floral bracts less than 15 mm long; corollas less than 20 mm long.

6a. Mature cauline leaves less than 10 cm long; floral bracts imbricate, ovate, with 2-3 well-developed teeth on each side of margin -A. seibertii Leonard

6b. Mature cauline leaves greater than 10 cm long; floral bracts lax, lanceolate, teeth minute (occasionally lacking). 7a. Leaves glabrous to sparsely pubescent, apically acute; corolla red or pink A. arnoldii Mildbr. 7b. Leaves hispid, with trichomes exceeding 1 mm long, usually slightly attenuate apically;

corolla white to purplish -A. tonduzii Leonard 5b. Floral bracts greater than 20 mm long; corolla greater than 40 mm long.

8a. Floral bracts pubescent with glandular trichomes, marginal teeth 5-15 per side; bracteoles glandular, 7-15 mm long; corolla orangish or red, 50-63 mm long; stamens exserted from mouth of corolla, thecae 4-5 mm long -A. aurantiaca (Scheidw.) Lindl.

8b. Floral bracts pubescent with eglandular trichomes, marginal teeth 3-4 per side; bracteoles eglandular, 19-31 mm long; corolla purplish, 43-45 mm long; stamens included in corolla tube, thecae 2.8-3.5 mm long -A. kuna T. F. Daniel & McDade

lb. Floral bracts entire. 9a. Floral bracts with paired lateral patches of nectariferous glands.

10a. Nectariferous glands numerous (at least 50, usually more than 100) per patch and minute (each ca. 0.1 mm diam.). lla. Floral bracts more than 30 mm long, apically rounded and reflexed; calyx segments much

shorter than floral bracts; capsules ca. 35 mm long -A. darienensis Wassh. lib. Floral bracts less than 20 mm long, apically obtuse to acute to acuminate, not reflexed; calyx

segments equaling or longer than floral bracts; capsules to 30 mm long. 12a. Floral bracts distantly spaced along rachis, lax -A. laxa Durkee 12b. Floral bracts closely adjacent or imbricate, rachis not visible during anthesis.


13a. Floral bracts relatively narrow, width no more than 2/3 of length. 14a. Plants rarely taller than 1.5 m; corolla 55-60 mm long -- A. lingua-bovis Leonard 14b. Plants taller than 1.5 m when reproductive; corolla longer than 60 mm.

15a. Spikes sessile, strongly tetrangular; floral bracts sparsely and minutely puberulous -A. leonardii McDade

15b. Spikes pedunculate, nearly terete in cross section; floral bracts conspicuously pubescent -A. scolnikae Leonard

13b. Floral bracts relatively broad, width exceeding 3/4 of length. 16a. Bracteoles strongly falcate, 7-10 mm long, 3-5 mm wide; corolla usually

yellow (rarely orange); capsules 28-31 mm long -A. hartwegiana Nees 16b. Bracteoles slightly falcate, 11-13 mm long, 2-3 mm wide; corolla orange;

capsules 20-24 mm long -A. campanensis Durkee lOb. Nectariferous glands 10 or fewer per patch, each > 0.2 mm diam.

17a. Floral bracts green to occasionally dull brown-orange, narrowly ovate to ovate, to 7 mm wide. 18a. Leaves glabrous; floral bracts 5-8 mm long, not imbricate, visible internodes 7-10 mm

long -A. gracilis Leonard 18b. Leaves pubescent with trichomes densest on veins; floral bracts longer than 8 mm, slightly

to closely imbricate, internodes not visible during anthesis. 19a. Trichomes of distal stems erect; floral bracts slightly imbricate, apically obtuse;

corolla pubescent with trichomes of tube exceeding 0.5 mm long ---- - -- - ---- ---- - -- - -- -- --- -- ---- ---- --- - -- --- -- --- --- ----- ---A . g racilis X sin clairian a

19b. Trichomes of distal stems appressed; floral bracts imbricate, apically acute; corolla minutely puberulent with trichomes of tube less than 0.2 mm long ------------------------------------------------------------A . golfod izlcensis M cD ade

17b. Floral bracts orange, broadly ovate, 8-20 mm wide. 20a. Distal stems, leaves, and bracts sparsely pubescent; floral bracts 10-12 mm long, 8-10

mm wide; corolla 57-62 mm long -A. terryae Standl. 20b. Distal stems, leaves, and bracts pilose; floral bracts 15-20 mm long, 14-20 mm wide;

corolla 64-71 mm long -A. sinclairiana Nees 9b. Floral bracts lacking nectariferous glands.

21a. Stems and leaves somewhat fleshy; floral bracts longer than 30 mm; calyx reduced to minute, hyaline lobes ca. 1 mm long; corolla longer than 30 mm, white -A. dolichantha Donn. Sm.

21b. Stems and leaves not fleshy; floral bracts less than 10 mm long; calyx well developed, lobes ca. 5 mm long; corolla less than 20 mm long, red, pink, purplish, or white. 22a. Leaves glabrous to sparsely pubescent, apically acute; corolla red or pink - A. arnoldii Mildbr. 22b. Leaves hispid with trichomes greater than 1 mm long, usually slightly attenuate apically;

corolla white to purplish -A. tonduzii Leonard

JUSTICIA EPHEMERA LEONARD

Justicia is both the largest genus of Acanthaceae and the most species-rich in Panama with 23 species (Daniel & Wasshausen, 1990). With more than 70 species of Justicia known from neighboring Co- lombia and given the still rudimentary knowledge of the eastern Panamanian flora, additional range extensions and new species are to be expected. A recent Panamanian collection of J. ephemera (Da- rie6n: Rfo Pirre, near El Real, 1-2 km upstream of water intake station for town of El Real, Feb. 1985, L. McDade 687, DUKE, MO, PMA) represents just such an extension. Justicia ephemera was previously known from northwestern Colombia (near Turbo in the department of Antioquia, Leonard, 1958). The Panamanian collection is not distinguishable from this species and shares with it a number of distinctive features: pandurate leaves with auricu- late bases, bifariously pubescent stems with retrorse trichomes, densely flowered verticillate inflorescences, and short white corollas. This combination of character states is not shared by any other species of Justicia in Panama.

ODONTONEMA CUSPIDATUM (NEES) KUNTZE

Here and below, we bring the treatment of Pan- amanian Odontonema into accord with those of Baum (1982) and Daniel (1995), and we provide a key to the species. Durkee (1978) recognized five species of Odontonema in Panama. Three of these, O. flagellum (Oerst.) Kuntze, 0. longifolium (Oerst.) Kuntze, and 0. strictum (Nees) Kuntze, were included in the synonymy of 0. tubaeforme (Bertol.) Kuntze by Baum (1982, as "tubiforme") and were similarly treated by D'Arcy (1987). Odontonema callistachyum (Schltdl. & Cham.) Kuntze, a species restricted in distribution to Mexico, Belize, and Guatemala, is neither known from Panama nor synonymous with 0. tubaeforme (cf. D'Arcy, 1987). Odontonema cuspidatum has been reported from Mexico and the West Indies (Baum, 1982; Daniel, 1995) and is much cultivated throughout tropical regions. A collection from central Panama (Panama: Cerro Azul, near Goofy Lake, 17 July 1962, J. Dwyer 2076B, MO, WIS) represents this species and the first report


of its occurrence in Panama. Collection notes in- dicate that the plant grew in a thicket. It is not known whether the occurrence of 0. cuspidatum at this locality represents an indigenous population, naturalized plants, or a remnant from cultivation. The specimen was annotated by Durkee and cited in the Flora of Panama (Durkee, 1978) as 0. callistachyum.

ODONTONEMA RUBRUM (VAHL) KUNTZE

Baum (1982) cited collections of 0. rubrum from Panama, Colombia, and Venezuela. We have stud- ied these and other specimens of 0. rubrum from Panama and concur that this species is distinct from other Panamanian Odontonema. However, as noted by Baum (1982: 73), plants from the western portion of the range of 0. rubrum (i.e., eastern Pan- ama and northwestern Colombia) are rather differ- ent from plants from eastern Colombia and Vene-

la. Plants often decumbent; mature cauline leaves less than 50 mm long -0. microphyllum Durkee lb. Plants erect; mature cauline leaves greater than 50 mm long.

2a. Dichasia sessile, at least some (often most) whorled at inflorescence nodes; rachis pubescent with flexuose to retrorse to appressed trichomes 0.2-1 mm long, trichomes usually concentrated in 2 or more lines ---------------------------------------------------------------------------O- 0. tubaeforme (Bertol.) Kuntze

2b. Dichasia pedunculate, alternate or opposite at inflorescence nodes; rachis evenly pubescent with erect to flexuose to antrorse trichomes 0.05-0.3(-0.5) mm long. 3a. Corolla externally pubescent with eglandular trichomes, tube abruptly expanded near the midpoint

into a prominent throat, bent at point of expansion, upper lip 6-13 mm long; pollen 3-aperturate --- -- ------------------------------------------------------------------------ 0. rubrum (Vahl) Kuntze

3b. Corolla externally glabrous, tube barely or gradually expanded distally into a t distinct throat, straight, upper lip 2-5 mm long; pollen 4-aperturate -0. cuspidatum (Nees) Kuntze

SANCHEZIA LUTEA LEONARD

Leonard & Smith (1964) recognized 59 species of Sanchezia. Most of these are endemic to Peru. The only species previously reported from Panama is S. pennellii Leonard (Durkee, 1978; D'Arcy, 1987). Recent collections from eastern Panama have resulted in the addition of two other species of Sanchezia to the flora of Panama. A collection of Sanchezia lutea (Darie6n: Rio Pirre, trail up river from house of Bartolo, 16 Mar. 1973, H. Kennedy 2883, TEX, US) was so annotated by Durkee in 1975, but neither the name nor the collection were subsequently listed by him (Durkee, 1978). An- other collection from the same general region (Da- rien: along Rfo Pirre just above Choc6 village of Pijibasal, 23 Feb. 1985, L. McDade 804, DUKE, MO) also represents this species. Sanchezia lutea was heretofore known only from various localities in Colombia (Leonard & Smith, 1964). The dis-

zuela. Our understanding of this species (and of other taxa with ranges including both Panama and Colombia) will benefit greatly from additional collecting in the poorly known areas of eastern Pan- ama and adjacent Colombia.

Odontonema rubrum has been collected in the Canal Area, Col6n, Darien, Panama, and the Co- marca de San Blas. Representative specimens of 0. rubrum from Panama include:

Canal Area: drowned forest of Q. Ancha, 18 Dec. 1934, C. Dodge & J. Steyermark 17043 (MO). Colon: Rfo Boquer6n, near Peluca, 27 Jan. 1973, R. Dressler 4263 (MO). Darien: between upper Rfo Membrillo and Camp 7 on road to San Blas, J. Duke 10915 (MO, US). Panama: 25 km NE of Cerro Azul on Rfo Pedras, 25 Nov. 1974, S. Mori & J. Kallunki 3471 (MO). Comarca de San Blas: lower Rfo Ailigandi, 17 Jan. 1967, J. Duke 9321 (MO).

Species of Odontonema in Panama can be distinguished by the following key:

tinctions among the species of Sanchezia now known from Panama are discussed below.

SANCHEZIA PARVIBRACTEATA SPRAGUE & HUTCH.

This species was described from plants cultivated at the Royal Botanic Gardens at Kew with- out reference to its place of origin. It is widely cultivated in the tropics and under glass in temperate regions. Among the specimens cited by Leonard & Smith (1964), all appear to represent cultivated material except for a collection (Cua- trecasas 11231, US) from "banks of the Rio Gua- mu6s" from the Intendencia de Putumayo in Co- lombia. A recent Panamanian collection of S. parvibracteata (Darien: Rfo Cocalito, 6 Feb. 1982, C. Whitefoord & A. Eddy 60, BM), presumably from an indigenous population, was also collected on a river bank.

The Panamanian species of Sanchezia can be distinguished by the following key:


la. Bracts 30-60(-75) mm long, 20-50 mm wide, the pair at a node fused from base for up to 35 mm toward apex thereby forming cuplike involucres along the rachis; bracteoles and calyx concealed by bracts; corolla externally glabrous - ------- -- - S. pennelli

lb. Bracts 6-20 mm long, 1.8-16 mm wide, free to base, not forming cuplike involucres; bracteoles and calyx surpassing bracts, clearly visible; corolla externally pubescent distally. 2a. Bracts acute at apex; thecae 5-7 mm long --S. parvibracteata 2b. Bracts gradually acuminate at apex; thecae 3.5-4.5 mm long ----- - S. lutea

CULTIVATED SPECIES

In treating the Acanthaceae of Panama, Durkee (1978) made an admirable attempt to include cultivated species as well as native and naturalized taxa. Numerous additional cultivated Acanthaceae are likely to be encountered in Panama beyond the two additions noted below.

PACHYSTACHYS LUTEA NEES

This species, a native of Peru, is widely cultivated for ornament in the American tropics and in temperate regions under glass. The shrubs are rec- ognizable by their erect, quadrangular spikes of large (up to 25 mm long), bright yellow bracts sub- tending white flowers. In Panama, plants occur in cultivation outdoors (Canal Zone: western edge of Panama City near Smithsonian Tropical Research Institute, 22 Feb. 1988, T Daniel et al. 5455, CAS). There are no reports of these plants becoming naturalized in Panama.

JUSTICIA BETONICA L.

This species is a native of the Indian subconti- nent and eastern tropical Africa. Justicia betonica is a shrub distinguishable from all other species of Justicia in Panama by its long (up to 16 cm), lax spikes with white to pinkish (with dark purple markings) corollas subtended by relatively large (ca. 10 mm long) bracts that are white with dark green venation. It is sometimes cultivated as a hedge or garden plant in warm tropical regions, and although not listed in Hortus Third (Bailey Horto- rium, 1976), is cultivated in sheltered locations in

Florida and under glass elsewhere in the United States. In Panama, plants are cultivated in yards (Coclh: La Mesa along road between El Valle de Anton and Cerro Gaital, 24 Feb. 1988, F Almeda et al. 5927, CAS). It is known to become naturalized in some tropical regions (e.g., Hawai'i) and may do so in Panama.

Literature Cited

Bailey Hortorium. 1976. Hortus Third. MacMillan, New York.

Baum, V. M. 1982. A revision of the genus Odontonema (Acanthaceae). M.Sci. Thesis, University of Maryland.

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. 1995. A revision of Odontonema in Mexico. Contr. Univ. Michigan Herb. 20: 147-171.

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Article Contentsp. [542]p. 543p. 544p. 545p. 546p. 547p. 548

Issue Table of ContentsThe American Historical Review, Vol. 109, No. 5 (December 2004)Volume Information [pp. ]Front Matter [pp. ]The Biology and Systematics of Fuchsia in the South Pacific [pp. 473-516]Pollination Biology of Lapeirousia Subgenus Lapeirousia (Iridaceae) in Southern Africa; Floral Divergence and Adaptation for Long-Tongued Fly Pollination [pp. 517-534]Review of the Genus Strephonema (Combretaceae) [pp. 535-541]Additions to the Acanthaceae of Panama [pp. 542-548]Revision Taxonomica del Genero Alexa Moq. (Fabaceae, Sophoreae) [pp. 549-569]A Study of the Vegetation and Floristic Affinity of the Limestone Forests in Southern and Southwestern China [pp. 570-580]Chromosomal Cytology and Evolution in Eupatorieae (Asteraceae) [pp. 581-592]Numeros Cromosomicos de Algunas Especies Mexicanas de Aristida (Poaceae: Aristideae) [pp. 593-595]Chromosome Counts of Compositae from Ecuador and Venezuela [pp. 596-602]Wood and Bark Anatomy of Nothotsuga (Pinaceae) [pp. 603-609]Back Matter [pp. ]

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Familia Acanthaceae en Panamá