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1 December 2004 A Revision of Capraria (Scrophulariaceae)
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Abstract

Herbarium and field studies of the chiefly neotropical genus Capraria have led to the recognition of four species. Capraria frutescens and C. mexicana are essentially endemic to Mexico. Capraria biflora is a widespread species that occurs throughout Mexico, Central and South America, the West Indies, and southern Florida. Capraria peruviana grows in northwestern South America and the Galápagos Islands. A complete account of synonymy and typification is provided, along with a key to species, scanning electron micrographs of pollen, stigmas and seeds, photographs, illustrations, and distribution maps.

Capraria L. (Latin = goat, in reference to observations that goats consume the leaves (Sprague (1921)) is a neotropical genus of suffruticose herbs and subshrubs distributed from southern Florida to central South America. Capraria is unique within the Scrophulariaceae in having alternate leaves with external punctate glands and oil-secreting cavities. The relationship of Capraria within the Scrophulariaceae has been under considerable dispute (see Generic Relations). The species, however, are easily delineated morphologically.

Linnaeus (1753) established the genus with the description of Capraria biflora L. Subsequently, 49 names were proposed in Capraria. Most of these were transferred or synonymized under other genera (e.g., Anticharis Endl., Freylinia Colla, Scop aria L., and Stemodia L.). Sprague (1921), in his treatment of Capraria, provided a list of excluded species that is included and expanded at the end of the present article.

A treatment of Capraria is pertinent at this time, considering the numerous mis- identifications of specimens in various herbaria observed by the present author and the discrepancy in the number of species recognized by various botanists: Lersten and Curtis (2001) considered Capraria to be monotypic, Mabberley (1987) recognized four species, Sprague (1921) five species, and Méndez Larios and Villaseñor Ríos (2001) six species. In this work four species of Capraria are recognized; full synonymy, illustrations and detailed exsiccate are presented for all species.

Generic Relationships

Throughout its history, the placement of Capraria has been under dispute (Table 1). Bentham (1846) placed it in subfamily (his suborder) Rhinanthoideae (with posterior corolla lobes interior in bud), tribe Sibthorpeae (with alternate leaves, and flowers solitary in the leaf axils), along with Sibthorpia L., Hornemannia Benth., Hemiphragma Wall., Camptoloma Benth. and Scoparia.

Bentham (1876) later reduced the tribe Sibthorpeae to subtribe Sibthorpieae of subfamily (his series) Rhinanthoideae, tribe Digitaleae with Hornemannia reduced to a synonym of Sibthorpia, and Camptoloma repositioned in his subtribe Digitalieae. Bentham's Digitaleae was later viewed as a heterogenous group with no clear definition by Thieret (1967).

Wettstein (1891) essentially followed Bentham's (1876) treatment, differing only in the non-recognition of subtribes for tribe Digitaleae.

Sprague (1921) retained Capraria in the tribe Digitaleae, emphasizing that Scoparia was its closest relative because both possessed leaves with punctate glands along the upper leaf surface. Sprague also noted that Hemiphragma “has relatively little in common with either” Capraria or Scoparia, suggesting that its placement near them was artificial, based largely upon a shared possession of alternate leaves.

Table 1.

Familial, subfamilial, and tribal placement of Capraria by various authors.

Thieret (1954) initially followed the treatments of Bentham (1876) and Wettstein (1891), but later transferred both Capraria and Scoparia into the subfamily Antirrhinoideae, tribe Gratioleae (Thieret, 1967). This was based, in part, on the observations of Pennell (1920, 1935), who noted that the posterior corolla lobes are external in bud in both taxa, and that they possess glandular hairs similar to other Gratioleae. Thieret also suggested that the sca- lariform-reticulate seeds (his “Bacopa” type) of Capraria and Scoparia are characteristic of many Gratioleae, and that their small 4-valved capsules are similar to those found in the genus Conobea Aubl., a native of tropical America with opposite leaves and punctate glands.

Arekal et al. (1971) related Capraria biflora to Scoparia dulcis L. on the basis of embryological data. This was later negated by Hakki (1975), who noted “no secondary haustoria developed in C. biflora as has been reported by Arekal et al. (1975) for Scoparia dulcis—the closest relative of our genus.”

Niezgoda and Tomb (1975) showed that the pollen of Capraria is spherical and 3-colpate, as opposed to 4-colpate pollen found in most Scrophulariaceae. They also found 3-colpate pollen in Leucophyllum Bonpl., Eremogeton Standi. & L. O. Williams (tribe Leucophylleae), and some My-oporaceae. Due to the similarity in pollen structure, Niezgoda and Tomb suggested the placement of Capraria close to the Leucophylleae, which they favored placing as a subfamily of the Myoporaceae. Argue (1980), however, observed 3-colpate pollen in Mimulus L. (sections Mimulus and Erythranthe), Penstemon Schmidel (tribe Cheloneae), Celsia L. (tribe Verbasceae), and selected genera of tribe Gratioleae {Lancea Hook. f. & Thomson, Artanema D. Don, and Conobea). Indeed his work vitiates the dramatic placement of Capraria in the Myoporaceae, suggesting instead its retention in the Scrophulariaceae tribe Gratioleae near Conobea, as proposed by Thieret (1967).

Henrickson and Flyr (1985) discussed in detail the systematic position of Eremogeton and Leucophyllum, concluding that both genera were more closely related to members of the traditional Scrophulariaceae and not to the Myoporaceae. However, they did not study Capraria, nor suggest it as a possible relative.

Barringer (1993) did not recognize subfamilies for the Scrophulariaceae and placed Capraria in his newly described monotypic tribe Caprarieae, suggesting no close relatives for it.

Table 2.

Morphological variation among the four species of Capraria.

Lersten and Curtis (2001) noted that Capraria and Leucophyllum are the only genera of Scrophulariaceae with true secretory oil cavities but noted that these two genera differ in the number and arragement of these cavities (see Morphology section). They did not propose a systematic placement of Capraria; instead, they cited Raman (1991) who, based on a study of trichomes in the Scophulariaceae, proposed no “taxonomic conclusion” for Capraria.

Recently, Olmstead (2004) placed Capraria in the tribe Leucophylleae with Eremogeton and Leucophyllum.

The systematic position of Capraria within the Scrophulariaceae appears unresolved. I am inclined to recognize Capraria in the monotypic Caprarieae as suggested by Barringer (1993). Nevertheless, I acknowledge that Capraria shares many characters with various members of the tribes Gratioleae and Leucophylleae.

Morphology

Methods: The morphological study is based on field observations by the author in Mexico and the examination of approximately 1600 herbarium specimens. Specimens were examined at or borrowed from the following institutions: BM (2 types), F (330 specimens), FLAS (40), K (3 types), MA (1 type), MO (274), NY (445), P (2 types), TEX-LL (96), US (354). There are at least 16 morphological characters (Table 2) that serve to distinguish Capraria from other genera and/or the four species from one another as noted below.

For Scanning Electron Microscopy (SEM) studies, the following procedures were followed. Pollen, stigma and seeds of the four species were collected from herbarium specimens housed at TEX-LL and SHST and the sheets annotated to that effect. These were mounted onto SEM stubs with conductive graphite tape and placed in a silica dessicator for three days. After dehydration the speciemens were coated with gold using a LADD sputter coater. Observations of the pollen and stigmas were made with a Vega Tescan 5130 (Department of Biological Sciences, Sam Houston State University) and photographed digitally. Observations of the seeds were made with a Phillips 515 scanning electron microscope (Cell Research Center, University of Texas) and photographed using Polaroid type 55 positive/negative film.

Leaves: The leaves of Capraria are alternate, simple and dentate. They vary in shape from lanceolate to spatulate. Three leaf characters in combination distinguish Capraria from other genera of Scrophulariaceae: 1) alternate leaves; 2) punctate glands; 3) oil-secreting cavities. Alternate leaves occur throughout the Scrophulariaceae, as do punctate glands. The possession of oil cavities is the only one of these three characters that does not appear to occur randomly throughout the family. In the Scrophulariaceae, only Leucophyllum shares true secretory oil cavities with Capraria. Despite the fact that Capraria and Leucophyllum share alternate leaves and oil cavities, Leucophyllum lacks punctate glands along the upper leaf surface and has a different number and arrangement of the oil cavities. Capraria has numerous small oil cavities scattered throughout the leaves, while Leucophyllum has two large oil cavities paired at the apex of the leaves (Lersten and Curtis, 2001).

Corollas: There is a wide range of variation among the corollas of the species of Capraria (Fig. 1). Capraria frutescens has bilaterally symmetrical, tubular-campanu-late corollas pubescent along the base of the throat, and four fertile included stamens as is typical of most Scrophulariaceae. Capraria mexicana and C. peruviana have glabrous, actinomorphic, rotate corollas with five distinct petals, five separate fertile stamens, no obvious corolla tube, and exserted stamens. The flowers of C. biflora are intermediate between those of C. frutescens and C. mexicana. The flowers are pubescent, and tubular, with five distinct petals lobes (two posterior and three anterior) that are bilaterally arranged, and exserted stamens. In addition, stamen number in C. biflora varies between four and five.

Corolla color ranges from white (Capraria mexicana and C. peruviana) to white (or pale lavender) with purple splotches running along the ventral internal surface of the tube (C. biflora and C. frutescens), these probably serving as nectar guides (Henrickson and Flyr, 1985).

Gynoecium: The gynoecium in Capraria is a superior bi-loculate ovary with a solitary terminal style that varies in length from 3–5 mm (C. biflora, C. frutescens and C. mexicana) to 0.75–1.50 mm (C. peruviana). The ovary possesses numerous ovules that are arranged axially. In general, the pistil is glabrous, although the ovary is occasionally glandular in C. frutescens (Fig. 2a) and the style is occasionally pubescent in C. biflora. The styles are exserted in C. biflora and C. mexicana but included in C. frutescens and C. peruviana. Like Leucophyllum (Henrickson and Flyr, 1985), Capraria exhibits interspecific variation in stigmatic structure. Capraria biflora, C. mexicana and C. peruviana (Figs. 2b, c, d, respectively) all possess stigmas that are elongated and linear in shape, while C. frutescens has a stigma that is shortened, bi-lobed and reniform in shape (Fig. 2a). In the subtribe Maurandyinae, Elisens (1985) proposed that “divergent or lobed stigmas … suggest derived character states and evolutionary advancement” from the more primitive conical/linear-shaped stigmas.

Pollen: Niezgoda and Tomb (1975) described the pollen of Capraria as spherical, 3-colpate, and diorate with a reticulate surface (Fig. 3); this contrasts with the 4-colpate pollen found in most Scrophulariaceae. No apparent variation in pollen grain sculpturing exists among the four species of Capraria.

Fig. 1.

Flowers of Capraria frutescens (a. front view, b. side view); C. biflora (c. front view); C. mexicana. (d. front view, e. side view). Photos by the author.

Fruits: The capsules (Fig. 4d) of Capraria are woody and ellipsoid to ovoid. They dehisce both septicidally and loculicidally to the base, releasing numerous seeds. The capsules are similar in form and structure to those of Leucophyllum and Gratiola. The structure of the fruits readily separates Capraria from the fleshy, 1-10-seeded, indehiscent fruits of the Myoporaceae.

Seeds: The seeds of Capraria are brown and have outer tangential walls forming a scalariform-reticulate surface of square cells (Fig. 5). Their surface structures are very similar to those found in the Gratioleae (Thieret, 1967; Pastor and Fernández, 2000) and the Leucophylleae (Henrickson and Flyr, 1985), thereby providing no specific insight into tribal placement. Hakki (1975) described the endosperm as “ab initio cellular.”

Variation exists in seed size among the four species of Capraria. Both C. biflora and C. frutescens have seeds that = > 0.5 mm long while those of C. mexicana and C. peruviana are ± 0.35 mm.

Chromosome Numbers

The first chromosome number report for Capraria was by Borgen (1980), as a mitotic count of 2n = ca. 60 for C. biflora. Zhao (1996) reported meiotic counts for C. biflora and C. frutescens as In = ca. 28 pairs. These two approximate counts suggest a base number of x = 14 or 15. The latter numbers do not support a relationship of Capraria to the Myoporaceae (x = 27, Watson and Dallwitz, 1992 and onwards) nor to the genus Leucophyllum (x = 17; Read and Simpson 1989), as suggested by Olmstead (2004). The genus Gratiola has a reported base number of x = 14 (Kapoor et al., 1987), thus Zhao's (1996) count of 2 n = ca. 28 for Capraria supports a possible relationship to Gratiola as proposed by Thieret (1967).

Fig. 2.

Scanning electron micrographs of the pistil of Capraria (insets on the upper left are enlarged photos of the stigma). a. C. frutescens (Williams & Siedo 99, SHST). b. C. biflora (Contreras 87, TEX). c. C. mexicana (Williams, Plum & Goldman 96-3, SHST). d. C. peruviana (Leiva & Sagastegui 1991, TEX). Large photos taken at 32× and 20 kV; insets taken at 100× and 20 kV. Photos by W. Patrick Spencer.

Fig. 3.

Scanning electron micrographs of pollen of Capraria mexicana (Williams, Plum & Goldman 96–3, SHST). Photo taken at 4.02 kx and 20 kV. Photos by W. Patrick Spencer.

Systematic Treatment

Capraria L., Sp. Pl. 628. 1753. Type Species: Capraria biflora L. Xuarezia Ruíz & Pav., Prod. 24. 1794. Type Species: Xuaresia biflora Ruíz & Pav. Pogostoma Schrad., Ind. Sem. Hort. Gott. 1831. Type Species: Capraria saxifragifolia Cham. & Schltdl.

Fig. 4.

Capraria biflora (Williams & Plum 95–48, SHST). a. Habit, b. Leaf. c. Sepals, d. Capsule, e. Corolla, side view. f. Corolla front view. Figure by Maria Thompson.

Perennial suffrutescent herbs to 2 m high, erect, with one to several stems developing from the basal stem, monopodial or ramified, glabrous to densely glandular-pubescent. Leaves alternate, sessile; laminae spatulate or lanceolate; bases cuneate; margins serrate along the upper half with 8–10 teeth along each side at midstem; surface with punctate glands and numerous scattered internal secretory oil cavities, with 3 principal nerves arising from or near the base, glabrous or villous. Inflorescence consisting of 2–5 pedicellate flowers in a leaf axil, with 5–20 flowering nodes per stem. Pedicels 1–22 mm long, glabrous or glandular-pubescent; bracts absent. Flowers 4–5-merous, perfect, actinomorphic or zygomorphic. Calyx regular, lobes five, free to the base or nearly so, 3–6 mm long, linear-oblanceolate to linear-lanceolate, glabrous, villous, or glandular-pubescent. Corolla white, rotate or bilabiate, tubular- funnelform, glabrous externally. Corolla TUBE wholly white or white above with purple spots along the ventral side within, glabrous internally or with trichomes along the ventral portion of throat. Petallobes spreading, narrowly triangular or ovate andapically acute. Stamens 4 or 5, didynamous or isomerous, included or exserted, alternate or opposite the corolla lobes; filaments glabrous. Anthers creamy white, introse, dorsifixed, bithecal, 3 locular with the basal portions of the thecae divaricate, inner locules shorter (0.4–0.7 mm long) with the outer locule longer (0.8–1.4 mm) and confluent across the anther tip. Style straight, included or exserted, glabrous or pubescent. Stigma ellipsoid or reniform. Ovary superior, bilocular, ovoid, glabrous or apically glandular-pubescent; ovules numerous, axially arranged, 4–6 mm long, 34 mm wide. Fruita glabrous capsule, elliptical, glandular-punctuate, loculicidally dehiscent, the placenta and calyx persisting. Seeds numerous, minute, brown, with outer tangential walls forming a scalariform- reticulate surface of square cells.

Fig. 5.

Scanning electron micrographs of the seeds of Capraria. a. C. biflora (Contrera 87, TEX). b. C. mexicana (King 3879, TEX). c. C. frutescens (Hinton 12824, TEX). d. C. peruviana (Leiva & Sagastegui 1991, TEX). Scale equals 0.1 mm. Photos by the author.

The genus contains four species distribeas. Because of its medicinal use, Capraria uted throughout the neotropics, occuring biflora has been propagated in China and mostly along beaches and moist seepage arAfrica.

Key to Species

1. Plants hirsute, pilose or glabrous throughout; leaf blades mostly (2-)3–4(-6) times as long as wide; corollas zygmorphic, tubular-campanulate, the tubes 4–7 mm long, villous and with purple markings ventrally within; stamens 4(5), included to occasionally slightly exserted, didynamous.

2. Stems ramified; calyx lobes not glandular-pubescent, linear-lanceolate, widest well below the middle; pedicels mostly 5–25 mm long; stigmas linear 1. C. biflora

2. Stems monopodial; calyx lobes glandular-pubescent, linear-oblanceolate, widest at or above the middle; pedicels mostly 1–4 mm long; stigmas reniform 2. C. frutescens

1. Plants glabrous throughout; leaf blades mostly 4–6 times as long as wide; corollas actinomorphic, rotate, the tubes 1–2 (-3) mm long, without hairs or colored markings within; stamens 5, exserted, isomerous.

3. Mature styles 3–5 mm long; corollas 8–10 mm long; Mexico and Belize 3. C. mexicana

3. Mature styles 0.75–1.50 mm long; corollas 5–6 mm long; South America and the Galapagos Islands 4. C. peruviana

1. Capraria biflora L., Sp. Pl. 875. 1753.

  • Type: SWEDEN. Cultivated, Uppsala, (Lectotype: LINN 785.1!; photo-LINN at F!. Howard (1989) reported the type of C. biflora as LINN 912348.50. That is actually a specimen of C. frutescens collected by Houston and is without location. The correct type of C. biflora is LINN 785.1 as selected by D'Arcy (1979)).

  • Capraria lanceolata Vahl [non. L. f.], Eclog. Amer. 2: 47. 1798. Capraria semiserrata Willd., Sp. PI. 3:324. 1800. Type: COLOMBIA. St. Martha, s.d., Rohr 20 (Holotype: C n.v.; photo-C at F!, US!). (Willdenow recognized C. lanceolata Vahl as a homonym of the earlier C. lanceolata L. f. [= Freylinia lanceolata (L. f.) G. Don] and renamed it Capraria semiserrata Willd.)

  • Capraria semiserrata Willd. var. berterii A. DC. in DC., Prodr. 10: 429. 1846. Type: COLOMBIA. St. Martha, 1829, Bertero s.n. (Holotype: G-DC!; photo-G-DC at F!, NY!, US!).

  • Capraria biflora L. var. pilosa Griseb., FI. Brit. W. I. 427. 1861. Type: none designated.

  • Capraria biflora L. forma hirsuta Loes., Bull. Herb. Boissier, Ser. 2, 3: 284. 1903. Type: GUATEMALA. Dept. Chiguimula, San Juan, 7 Jan 1897, Seler 3314 (Holotype: B; Isotypes: NY!, US!).

  • Capraria biflora L. subsp. havanensis Tzvelev, Bot. Zh. (Lenningrad) 72: 1663. 1986. Type: CUBA. Valle Bacuranao, near Havana, 8 Feb 1984, Tzvelev 511 (Holotype: LE).

  • Subshrubs or suffruticose perennial herbs, stiffly erect to somewhat sprawling, 40–200 cm tall, with several ramified stems emerging from the root stock. Stems mostly hirsute, occasionally nearly glabrous but always with a few hairs on the emergent part of young branches, 3–5 mm in diameter at midstem. Leaves spatulate to lanceolate, mostly 3.5–8.0 cm long, 0.5–3.0 cm wide, glabrous to moderately hirsute. Pedicels 5–22 mm long, glabrous or glandular-pubescent. Flowers 5-merous, perfect, zygomorphic to slightly regular, 10–13 mm long. Calyx lobes 4–6 (-7) mm long, lanceolate, glabrous to pubescent. Corolla white, bilabiate, tubular-funnelform. Corolla Tube white above with purple spots ventrally within, with villous trichomes along the ventral portion of throat, 4–8 mm long. Petal Lobes spreading, ovate, apically acute, the tips rolling back after anthesis; anterior lobes 3, 3–5 mm long, 1.8–2.2 mm wide; posterior lobes 2, 3–5 mm long, 1.8–2.2 mm wide. Stamens 4 or 5, didynamous, posterior pair exserted about 0.5–0.8 mm past the tube, opposite with the corolla, lobes; filaments glabrous. Style included, glabrous to sparsely pilose, 3–5 mm long. Stigma ellipsoid. Ovary glabrous. Seeds ca. 0.50 mm long, 0.35 mm wide,

  • Common Name: Claudiosa (Spanish, Balick 2304), Tan-chi (Mayan, Balick 2116), Pasmo-wa-xi-uil (Mayan, Balick 1768), sabadil (Huave, D. Zizumbo 63); Goatweed, Té del Pais, Hierba Té, Cola de Gallo (D'Arcy, 1979); and many others listed in Sprague (1921).

  • Chromosome Numbers: 2n = ca. 28 (Zhao, 1996); 2n = ca. 60 (Borgen, 1980).

  • Phenology: Flowering August through February with a few specimens flowering in March. Producing mature fruits December through August.

  • Distribution (Fig. 6): Southeastern U.S.A. (Florida), Caribbean Islands, Mexico, Central America, Galápagos Islands and South America east of the Andes; sea level to 1250 m. Sanders et al. (1996) reported C. biflora as for north as Sonora, Mexico (Van Devender 95–90, ARIZ, UCR). Sprague (1921) reported the species from Texas, but to date I have not found additional citations or specimens to verify this account.

  • Economic Uses: Label data on Ster- gios 7773 reports that the plant is used to control hypertension; Fishlock 270, used as a tea; M. Balick 2116, used for constipation (leaves), kidney stones, and blocked urine flow, when used in conjunction with other species; M. Balick 2304, antipyretic (the whole plant, heated in fire and rubbed over body); M. Balick 1768, used for bath for feverish infants.

  • Illustrations: Photo of flower (Fig. 1c); illustration of flower and habit (Fig. 4).

  • Representative Specimens: U.S.A. Florida. Collier Co.: 7 mi SE of Naples, road to Briggs Nature Center and Rookery Bay Aquatic Preserve, 23 Nov 1987, Alcorn 451 (FLAS). Dade Co.: County Sanitary fill, across canal from Coco Palm Drive, ca. 1 mi E of Florida Turnpike, 20 Dec 1980, Correll 51435 (NY). Lee Co.: Captive Island, 1 Jan 1972, Brumbach 7790 (FLAS). Monroe Co.: Park Key, S of Park Channel, mile marker 18.1 of Rte. 1, 13 Mar 1984, Hill 13415 (NY).

    MEXICO. Campeche. Mpio. Champotón, beach at Hotel Siho Playa, ca. 38 km S of Campeche on Hwy. 180 (90° 43′W, 19° 35′ N), 9 Mar 1990, Sanders 9564 (TEX); Tuxpeña, 4 Nov 1931, Lundell 898 (LL, US). Chiapas. Mpio. Chiapa de Corozo, Chorreadero de Tuxtla, 5.6 mi E of Chiapa de Corozo along Hwy. 190, 20 Feb 1965, Breedlove 9098 (LL); Mpio. Ocozocoautla de Espinosa, La Cima, 15 km WNW of Ocozocoautla, 15 Oct 1972, Breedlove 28999 (NY); Villa Flores, adelante de Monte Cristo, 71.3 km E del entronque Nac 196/Chis 195 sobre Chiapas 195 hacia Suchiapa y 0.8 km S de El Cielito, 790 m, 24 Dec 1984, Cowan 5016 (TEX); Nuevo Amatenango, 1300 m, 17 Jul 1941, Matuda 4734 (LL, NY); Mpio. Venustiano Carranza, slope above Finca Carmen, along road from Acala to Pugiltik, 3 Oct 1967, Ton 2979 (NY). Guerrero. Dist. Coyuca, Pungarabato, 14 Jan 1934, Hinton 5474 (NY, US). Mexico. Dist. Temascaltepec, Naranjo, 4 Oct 1932, Hinton 1979 (NY, US). Nayarit>. Tres Marías Islands, María Madre, 25 Oct 1925, Ferris 5731 (US). OAXACA. Tehuantepec, Rincón Moreno, 19 Jan 1971, MacDouga.il s.n. (NY); Puerto Angel, 18 Jul 1910, Orcutt 5009 (NY). Puebla, around San Gabriel Chilac near San Juan Atzingo and San Andrés, 24 Jul 1961, Smith 4038 (US). Quintana Roo. 5 km S of Playa del Carmen, 2 Mar 1985, Cowan 5078 (TEX); 3 km SSW of Tulum ruins, 7 Jan 1973, Taylor 12616 (NY, US). Sinaloa. Dept. Badiraguato, Tierra Blanca, 4 Mar 1940, Gentry 5805 (NY); near San Blas, 22 Mar 1910, Rose 13218 (NY, US); Villa Unión, Dec 1921, Ortega 4372 (US). Sonora. On riverbank, El Paso on the Río Cuchijaqui (26° 40′ 35″N, 108° 49′ 30″W), 150 m, 25 Feb 1995, Van Devender 95–90 (ARIZ). Tabasco. Mpio. Paraíso, road from Nicolás Bravo to Mecoacán, km 15.2, 21 km from road Paraíso-Comalcalco, 9 Sep 1980, Cowan 3188 (NY); Mpio. Nacajuca, Masateupa N of Nacajuca, 17 Jan 1979, Cowan 1883 (NY). Veracruz. 21 mi S of Veracruz, 29 Dec 1970, Taylor 7391 (NY); Mpio. Chacaltianguis, N side of Río Papaloapan on W side of Benito Juárez and 12 km SW of Cosamalopan (18° 16′N, 95° 54′W), 2 Feb 1984, Nee 29256 (NY); ca. 5 miles E of Naranjos on dirt road to Tamiahua, 15 Aug 1995, Williams & Plum 95–48 (SHST, TEX). Yucatán. Mpio. Celestún, entrance to Puerto de Celestún, 3 May 1983, Chan 2240 (TEX); 15 km NW of Humucma, along road Mérida-Sisal, 20 Jul 1985, Cabrera 9083 (NY); Progreso, along railroad through cleared flat, July 1938, Lundell 7976 (LL, US).

    BELIZE. Dist. Cayo, ca. 15 km SW of San Ignacio, near San Antonio (17° 05′N, 89° 00′W), 11 Nov 1987, Balick 1768 (US); Toll bridge area at New River, 3 mi S of Orange Walk, 25 Jan 1974, Dwyer 12177 (NY, US); Branch Mouth N of Cayo, 19 Feb 1931, Bartlett 11944 (NY).

    COSTA RICA. Río Higueron near Taboga (10° 20′N, 85° 12′W), 30 Jun 1977, Liesner 2764 (NY).

    EL SALVADOR. Dept. San Miguel, S side of Lake Olomega (13° 17′N, 88° 04′W), 27 Jan 1942, Tucker 840 (US); Dept. Sonsonate, Acajutla, 20 Mar 1922, Standley 21922 (NY, US); Comasagua, Dec 1922, Calderon 1377 (NY);

    GUATEMALA. Dept. El Peten, Tikal Nat. Park, near airfield, 16 Jul 1959, Lundell 16474 (LL); Tikal, on airfield, 20 Aug 1959, Contreras 87 (LL, US); Dept. Zacapa, Gualan, 28 Dec 1905, Kellerman 5734 (LL, US).

    HONDURAS. Dept. Copan, matorrales húmedos de Río Copan, cerca de Copan Ruinas, 650 m, 17 Apr 1956, Molina 6597 (LL); San Pedro Sula, 16 Apr 1984, Cristoff 190 (NY).

    NICARAGUA. Dept. Carazo, S boarder of dept, near boarder of Dept. Rivas (11° 30′N, 86° 10′W), 10 Sep 1982, Grijalva 1027 (NY); Dept. Chinandega, Ameya, 19 Jun 1923, Maxon 7118 (US); Calabazas, Rte 1, S of Dario, 25 Dec 1969, Seymour 2598 (NY); Prov. León, near León, 20 Dec 1975, D'Arcy 10431 (NY); along shore of Lake Managua, 24 Jun 1923, Maxon 7255 (US).

    PANAMA. Fort Sherman, Torro Point, 28 Feb 1976, Fosberg 56088 (NY, US); Panamá City, 12 Jun 1923, Maxon 6951 (NY, US); Bocas del Toro, 6 Feb 1921, Carleton 150 (NY, US); Isla Taboga, 23 Jul 1938, Woodson 1492 (NY); Gaeleta Island, 10 Jun 1954, Ritchie s.n. (NY); Isla Colón, 19 Nov 1941, Wedel 2977 (NY); San José Island (8° 15′N, 79° 08′W), 16 Sep 1945, Harlow 60 (US).

    WEST INDIES AND THE CARIBBEAN. ANGUILLA. Road Bay, 9 Jan 1959, Proctor 18697 (NY, US). ANTIGUA. Royals, 11 Sep 1937, Box 1051 (US). ARUBA, near Noord, 17 Jan 1953, Stoffers 1572 (NY). BAHAMAS. San Salvador, Dec 1973, Catino 14 (FLAS); Farmer's Hill, Great Exuma, 14 Aug 1975, Eldridge s.n. (FLAS, US); Caicos Island, 29 Aug 1974, Correll 43274 (NY); Icacos Island, 7 Aug 1968, Woodbury 1–67 (NY); Nassau, 8 Jan 1890, Northop 27 (NY); Andros Island, Nicols Towm, 26 Mar 1890, Northop 381 (NY); Grand Bahama, Pine Ridge, NE of Freeport, 5 Nov 1973, Correll 40537 (NY); Berry Islands, Great Harbour Cay, 16 Oct 1974, Correll 43683 (NY); Watling's Island, 27 Nov 1907, Wilson 7290 (NY); Eleuthera, 21 Feb 1907, Britton 5581 (NY); Addin's Island, 21 Dec 1905, Brace 4318 (NY); Cat Island, near Port Howe, 22 Nov 1975, Correll 46206 (NY); Long Island, Clarence, 19 Jun 1974, Hill 2216 (NY); Providence, N side of Island, 3 Sep 1952, von Reis 249 (NY); Andros Island, 13 Mar 1966, Dawson 26850 (US). BARBADOS. Parish Saint James, 31 Jan 1969, Andrews 643 (NY). BERMUDA. Saint Davis Island, 10 Feb 1908, Brown 603 (NY, US). CUBA. Prov. Camaguey, Camaguey, 15 Mar 1909, Shafer 792 (NY, US); Prov. Havana, Río Almendares to Playa de Marianao, 22 Dec 1910, Wilson 9459 (NY); Prov. Ma- tanzas, near San Miguel, 2 Sep 1903, Britton 218 (NY); Prov. Oriente, Sabana to Maisi, 13 Dec 1910, Shafer 7889 (NY); Prov. Pinar del Río, Vedado, Habana, Sierra del Jíbaro, 20 May 1955, Alain 4295 (NY); Prov. Santa Clara, near Soledad, bank of Rio Caonoa, Jun 1941, Howard 5122 (NY). CURAçAO. Near Willemstad, 19 Feb 1917, Curran 78 (NY); Tanchi, 19 Apr 1952, Arnoldo 2054 (US). DOMINICAN REPUBLIC. Prov. Barahona, Río Baoruco, 6 Nov 1979, Smith 10026 (NY); Prov. Distrito Nacional, La Caleta, 20 m E of Santo Domingo, 25 Sep 1970, Liogier 17500 (NY); Prov. La Altagracia, 2 km E of Playa de Macaooon, road to Punta Cana Club (18° 45′N, 68° 30′W), 25 Jul 1981, Watson 1118 (FLAS); Prov. La Romana, Llano Costero, Isla Catalina (18° 22′N, 69° 02′W), 17 Dec 1986, Zanoni 37229 (FLAS, NY); Prov. Monte Christi, 8 km N of Villa Elisa (19° 44′N, 71° 15.5′W), 21 Aug 1985, Pimentel 532 (NY); Prov. Peravia, Palmar de Ocoa (18° 18′N, 70° 35′W), 7 Aug 1980, Zanoni 7769 (NY); Prov. San Pedro de Macpris, Guayacanes, 19 Mar 1965, Lavastre 1912 (NY); Prov. Puerto Plata, 35 km E of jct with Puerto Plata and Santigo road, 25 Apr 1970, Burch 2404 (NY). GRAND CAYMEN. Center of Island, 14 Feb 1899, Armour 1364 (NY). GRENADA. Parish Saint Patrick, Levera Beach, 20 Jan 1951, Hunnewell 19499 (NY). HAITI. Port-au Prince near Mariani, 20 Nov 1927, Ekman 9344 (LL, US); near Fond Parisien, Etang Saumatre, 5 May 1920, Leonard 4146 (NY, US); near La Vallee, Tortue Island, 28 Dec 1928, Leonard 11731 (NY, US). JAMAICA. Parish St. Thomas, road from Golden Grove to Morant Point, 56 mi E of Golden Grove, 19 Aug 1965, Hespenheide 1354 (LL); Saint Ann's Bay, 27 Mar 1908, Britton 2527 (NY); between Portlan Point and Rocky Point, 5 Mar 1908, Britton 1932 (NY); Kingston to Port Morant, 28 Feb 1909, Britton 3902 (NY); Parish Saint Catherine, Port Henderson, 19 Nov 1957, Yuncker 17475 (NY). MARTINIQUE. near Usine Caritan, 24 Jul 1939, Egler 39147 (NY). MONTSERRAT. N of Plymouth, 21 Jan 1907, Shafer 88 (NY). PUERTO RICO. Mona Island, 20 Dec 1913, Sterene 6350 (NY); Ponce, 23 Dec 1902, Heller s.n. (NY); Caguas, 7 Jan 1899, Armour 217 (NY); Punta Vacía Talega, 10 Jan 1980, Liogier 30173 (NY); Isla de Mona, Playa de Sardinera, E side, 27 May 1991, Acevedo 4287 (NY, US); N of Guayama, 10 Feb 1986, Taylor 6792 (NY); Mpio. Toa Boja, Sabana Seca, 27 Dec 1984, Montalvo s.n. (NY); Vieques Island, near Isabel Segunda, 24 Jan 1914, Shafer 2433 (NY, US). SAINT CROIX. W of Tague Bay, 18 Jul 1970, D'Arcy 4680 (US); Sandy Point, 28 Feb 1993, Acevedo 5296 (US). SAINT KITTS. Canada Estate, 8 Sep 1901, Britton 751 (NY). SAINT VINCENT. Par ish Saint George, Calli-aqua Bay, 12 Jan 1962, Cooley 8141 (NY). TABAGO. Saint Patrick Parish, Bon Accord Lagoon near Pigeon Point, 7 Jan 1989, Worthington 17695 (NY). TRINIDAD. Port of Spain, Jun–Aug 1939, Almandoz 1 (NY). TURKS ISLANDS. Grand Turk, 27 Aug 1905, Nash 3827 (NY). VIRGIN ISLANDS. Tortola, 15 Nov 1965, D'Arcy 327D (FLAS); Virgin Gorda, 14 Jan 1919, Fishlock 270 (NY); Water Island, 8 Nov 1969, Woodbury WI-26 (NY).

    ARGENTINA. Prov. Buenos Aires, Bolivar, 31 Jan 1945, Perdero 119 (NY); Prov. Cordillera, Santa Cruz, Cabeza, 22 Jan 1945, Perdero 44 (NY); Prov. Jujuy, Dept. Santa Bárbara, Jinalito, 10 Apr 1945, Meyer 8512 (NY); Prov. Salta, Dept. Orán, Río Ber- mejo, 16 May 1945, Pierotti 1383 (NY).

    BOLIVIA. Dept. Santa Cruz, Prov. Andrés Ibáñez, Puerto Pailas (17° 40′S, 62° 47′W), 11 Jul 1991, Nee 41641 (NY); Dept. Santa Cruz, Prov. Cordillera, Ovai, 6 km NW of Charagua (19° 49.5′S, 63° 14′W), Quevedo 70 (NY).

    BRAZIL. Estado Bahiá, by the Río Cariçã, SW of Monte Santo, 21 Feb 1974, Harley 16445 (NY, US); Estado Coará, Forrtaleanm 11 Dec 1955, Duke 2502 (NY); Estado Maranhão, Aleantara, 10 Apr 1954, Fróes 50720 (NY); Estado Paraíba, em regioes secas, Aug 1959, Smith 2122 (NY); Estado Portugesa, Guan- are, 31 Dec 1984, Stergios 7773 (NY); Estado Zulia, km 440 road from Machiques to La Fría, 24 Jul 1976, Stergios 574 (NY); Mpio. Itiuba, Bahiá (10° 43′S, 39° 50′W), 26 May 1983, Bautista 765 (NY); Maranhão, Dec 1959, Carvalho 2 (NY); Saint Paul, Sep 1892, Glaziou 19734 (NY);

    COLOMBIA. Dept. Antioquia, Vuelta de Acuña, Río Magdalena, 14 Jan 1918, Pennell 3789 (NY, US); Dept. Atlántico, bear Barranquilla, 18 Mar 1961, Dugand 5625 (NY); Dept. Bolívar, Sincé, 20 Apr 1963, Romero-Castaneda 9680 (NY); Dept. Bolivar, Mpio. Cartagena, 18 km SW of crossing of Canal Dique at Pasacaballos, Isla Barú (10° 08′ N, 75° 42′ W), 6 Aug 1985, Zarucchi 3963 (NY, US); Dept. Cun- dinamarca, Quebrada Cabaña, Hacienda El Cucharo, between Tocaima and Pubenza, 8 May 1944, Killip 38366 (US); Dept. Huila, Río Cabrera, 2 km below confluence of Río Ambicá, 3 km WSW of Colombia (3° 22′ N, 74° 50′ W), 15 Dec 1942, Fosberg 19316 (NY, US); Dept. La Guajira, 1 km W of Puerto Estrella, 4 Apr 1962, Saravia 431 (US); Dept. Magdalena, Isla de Salamanca, 20 km along the road from Ciénaga to Boquilla, 10 Dec 1966, Romero-Castañeda 10500 (NY); Dept. Magdalena, Isla de Santa Marta, 1 Apr 1918, Pennell 4771 (NY, US).

    ECUADOR. Prov. Galápagos, Indefatigable Island, 16 Feb 1939, Taylor 32 (NY); Prov. Galápagos, Santa Cruz Island, old trail to Bella Vista, 8 Feb 1964, Fournier 161 (US); Chatham Island, Wreck Bay, Jan 1925, Stewart 3424 (US).

    FRENCH GUIANA. Cayenne, 21 Jul 1985, de Granville 7282 (NY); Maripasoulas, 8 Apr 1990, Fleury 860 (NY).

    GUYANA. Dept. Demerara-Mahaica Region, Atlantic coastline facing leper colony at mouth of Mahaica river (6° 38′ N, 57° 55′W), 2 Dec 1986, Pipoly 9044 (NY, TEX, US); Queen's College, Georgetown, in grasses at edge of drainage trench, 10 May 1956, Irwin R-97 (TEX); Mahaica-Berbice Region, Abary River mouth and along canals leading into river, between coastal hwy and ocean (6° 35′N, 57° 47′W), 28 Mar 1987, Pipoly 11248 (NY, US); Pomeroon District, Moruka river, Jul 1927, de La Cruz 4575 (NY, US); Waini river (8° 20′N, 59° 40′W), 3 Apr 1923, De La Cruz 3810 (NY).

    PERU. Dept. Loreto, San Salvador along the Amozon River, Jul 1929, Williams 1564 (US); Dept. San Martin, Fundo San Isidro, 13 Feb 1976, Sagastegui 8312 (NY).

    SURINAM. Coronie, 22 Oct 1933, Lanjouw 1090 (NY); 10 km SW of Paramaribo, 25 May 1961, Hekking 823 (NY); Nickerie swamp, Schelpenrits Prodobong, 2 Feb 1943, Geijskes s.n. (NY); Zandery, 31 May 1916, Samuels 263 (NY).

    VENEZUELA. Estado Aragua, La Trinidad de Maracay, Jan 1913, Pittier 5823 (NY, US); Estado Bolivar, Rio Suapure, 5 Jun 1984, López-Palacios 4369 (NY); Estado Falcón, Distrito Silva, Cayo Sal, NE of Chichiriviche (10° 58′N, 68° 15′W), 28 Aug 1974, Steyermark 110325 (NY); Estado Sucre, Península de Araya, 20 km NW of Cariaco (10° 38′N, 63° 40′W), 17 May 1981, Liesner 11933 (NY); Estado Zulia, near Perijá, 1918, Tejera 114 (US); Prov., Trujillo, La Concepéion, 23 Mar 1931, Reed 1049 (US); road between Playa El Falucho and Playa El Americano, 5 km de el apostadero naval (11° 50′N, 65° 00′W), 30 May 1986, Rivero 1052 (FLAS); Maracay, 1928, Corivelius 286 (FLAS); Guarico, 14 km N of San Fernando de Apure along main hwy to Calaboz, 10 Nov 1973, Davidse 3952 (NY); road between San Felipe and Barquisimlleto km 102, 26 Nov 1952, Aristeguieta 1100 (NY); Dist. Federal, Mpio. Vargas, Parraquia Catia (10° 36′N, 67° 02′W), 12 Jun 1990, Ramírez 2715 (NY); Península Paraguana, ca. 1.5 km from Pueblo Nuevo along road to Santa Ana, 19 Dec 1964, Breteler 4353 (NY, US); Miranda, between Las Canales and El Encanto, Nov 1942, Lasser 688 (US).

  • Capraria biflora is the most widespread species of Capraria and the only species distributed on both sides of the equator (Fig. 6). The species is readily distinguished by its ramified stems, spatulate leaves, pubescent hairs, lanceolate sepals, bilaterally symmetrical corollas with 4–5 stamens, and linear stigma.

    One specimen from Oaxaca, Mexico (Orcutt 5009) and several in Panama (Hayes 322, 745, and 876) are glabrous and have linear-lanceolate leaves that are only 4–6 mm wide, superficially resembling Capraria mexicana and C. peruviana, while the flowers are zygomorphic as in C. biflora. In addition, occasional specimens from Florida, the Caribbean islands and the Atlantic coast of both hemispheres have lanceolate leaves; these appear to be largely confined to areas that are at or near sea level. However, sporadic plants of a similar nature occur throughout the range of C. biflora, and consequently such plants do not appear to merit varietal status.

    I follow Wiggins and Porter (1971) who noted that because plants that are subglabrate and densely pubescent intergrade, “it seems futile to recognize var. pilosa Griseb.”

  • Fig. 6.

    Distribution of Capraria biflora (circles) and C. peruviana (diamonds). Note: C. biflora and C. peruviana are sympatric in the Galapagos Islands.

    2. Capraria frutescens (Mill.) Britton, J. Bot. 45: 315. 1907.

  • Erinus frutescens Mill., Gard. Dict. ed. 8. 1768. Capraria cuneata R. Br. in W. T. Aiton, Hortus Kew., ed. 2, 4: 45. 1812. Type: MEXICO. Veracruz: 1730, Houston s.n. (Holotype: BM!; photo-BM at NY!, US!)

  • Capraria hirsuta Kunth in H.B.K., Nov. Gen. Sp. 2: 355. 1817. Type: MEXICO. Guerrero: “Crescit locis exustis inter La Venta del Exido et portum Acapulci,” 200 m, 1802–1804, Humbolt & Bonpland 3892. (Holotype: P!; ISOTYPES: B; photo-B at F!, US!) Sprague (1921) treated Capraria hirsuta as a synonym of C. biflora. Examination of the type, however, indicates it to be a specimen of C. frutescens.)

  • Capraria saxifragifolia Cham. & Schltdl., Linnaea 5: 105. 1830. Pogostoma saxifragifolia (Cham. & Schltdl.) Schrad., Ind. Sem. Hort. Gotting. 1831. Type: MEXICO. Veracruz: Jul 1828, Schiede s.n. (Holotype: B; photo-B at F!, NY!, US!)

  • Perennial herbs, stiffly erect to somewhat sprawling, 40–200 cm tall, with several monopodial stems emerging from the root stock. Stems pilose, 2–4 mm in diameter at midstem, the vestiture 1–2 mm long. Leaves oblanceolate, mostly 4–8 mm long, 1.5–4.0 cm wide, moderately hirsute. Pedicels 1–4 mm long, glandular-pubescent. Flowers zygomorphic, 6.5–8.5 mm long. Calyx lobes 3.0–4.5 mm long, linear-oblong, variously pubescent with glandular or eglandular hairs (these often intermixed), the apices acute. Corolla white to pale lavender, bilabiate, tubular-funnelform. Corolla tube white with purple spots ventrally within, with villous trichomes along the ventral portion of the throat, 47 mm long. Petal Lobes spreading, ovate, apically acute, tips rolling back after anthesis; anterior lobes 3, 3.5–5.0 mm long, 1.8–2.2 mm wide; posterior lobes 2, 2.5–4.0 mm long, 1.8–2.2 mm wide. Stamens 4, didynamous, included, opposite the corolla lobes; filaments 2.5–3.0 mm long, glabrous. Style included, glabrous to sparsely pilose, 3–5 mm long. Stigma reniform. Ovary pubescent apically, the hairs extending up the style shaft near its base but not much beyond. Seeds ca. 0.5 mm long, 0.35 mm wide,

  • Common Name: Claudiosa (Yucatan; Sprague, 1921).

  • Chromosome Numbers: 2n = ca. 28 (Zhao, 1996).

  • Phenology: Flowering August through December with a few specimens flowering in January. Producing mature fruits December through September.

  • Distribution (Fig. 7): Central Mexico southwards to Honduras, 0–1000 m.

  • Illustration: Photo of flower (Fig. 1a & b); illustration of flower and habit (Fig. 8).

  • Representative Specimens: MEXICO. Campeche. near the caves of Xtacumbilxunam, 3 km W of Bolonchen de Rejón, 29 Sep 1985; Cabrera 9525 (MO). Chiapas. Mpio. Chiapa de Corzo, steeped walled canyon with tropical deciduous forest, above El Chorreadero, 800 m, 9 Jul, 1972, Breedlove 26004 (LL); Ocozocuautla, 16 km W of Ocozocuautla and 3.2 km from El Aguacero, 24 Dec 1984, Cowan 5020 (TEX); Mpio. Tuxtla Gutierrez, 5 of Tuxtla Gutiérrez on Hwy. 195 (to Villaflores), 4.7 mi from jet. with bypass, on top of escarpment in roadside brush in rocky limestone soil, 860 m, 1 Nov 1980, Fryxell & Lott 3265 (F, NY, TEX). Colima. Manzanillo, 1 Dec 1890, Palmer 917 (NY, US); Colima, 20 Oct 1910, Orcutt 4522 (F). Guerrero. near Acapulco, Oct 1894, Palmer 567 (US). Jalisco. Tuxpan, 4 Nov 1926, Mexia 1041 (US); Chamela Biological Station, along main road about 50 m from the entrance to preserve headquarters (19° 30′N, 104° 50′W), 31 Dec 1996, Williams, Siedo & Wood 99 (SHST). México. Dist. Temascaltepec, Bejucos, 13 Feb 1933, Hinton 3383 (F, NY). Michoacán. Dist. Coalcomán, Coalcoman, 1000 m, Dec 28 1938, Hinton 12824 (LL, NY). Oaxaca. Tehuantepec, Potrero Villalobos, 20 Nov 1970, Dougall s.n. (NY). Querétaro . Mpio. Jalpan, 7 km SE of Tancoyol, 800 m, 8 Nov 1986, Fernández 3685 (NY, TEX). San Luis Potosé. Mpio. San Antonio, San Antonio, disturbed ground near abandoned house, 5 Sep 1978, Alcorn 1559 (TEX); Tamazunchale, 28 Nov 1937, Kenoyer 765 (F); Tamagundarb, 19 Jan 1947, Aguirre & Reko 309 (NY). Sinaloa. near Mazatlan, 31 Mar 1910, Rose 13772 (US); Pueblo de las Trancas San Ignacio, 30 Oct 1917, Montes 22 (US). Tamaulipas. near Tampico, 15 m, 1–31 Jan 1910, Palmer 43 (F, NY). Veracruz. Mpio. Axocuapán, road Coetzalán-Cueva del Abono (19° 18′N, 96° 42′W), 11 Jun 1983, Robles 362 (NY); Mpio. Coatepec, 2 km antes de Jalcomulco, 30 Jul 1980, Castillo & Tapia 1085 (F); ladera NE of Cerro Monte de Oro, 22 Jun 1972, Dorantes et al. 920 (F); Mpio. Dos Rios, Cerro Gordo, 550 m, 14 Oct, 1970, Ventura 2613 (F, NY); Mpio. Tantoyuca, Silosuchil, dooryard, 22 Dec 1978, Alcorn 2296 (TEX); Mpio. Santiago Tuxtla, Acahual, 30 m, 29 Jul 1967, Martínez-Calderón 1472 (F, LL, NY); Lagos Tamiahua, 25 mi S of Tampico, 3 Mar 1939, LeSueur 378 (F, TEX, US); near Palma, 3 Sep 1935, MacDaniels 456 (F); ca. 5 miles E of Naranjos on dirt road to Tamiahua, 15 Aug 1995, Williams & Plum 95–49 (SHST; TEX). Yucatán. Chichen Itzá, 27 Feb 1899, Mills paugh 1625 (F); in wet places near Sayil, 35 km SE of Oxkutzcab, 19 Jul 1985, Cabrera 9006 (TEX); Mpio. Santa Elena, 0.55 km SW of entrance road to Kabah archeological site on Hwy. 261, between Holpelchen and Mérida (89° 40′W; 20° 15′N), 100 m, 11 Mar 1990, Sanders 9628 (TEX); Chankon, 22 Jun 1929, Bequaert 78 (F); near the caves of Balancan-ché, 32 km W of Valladolid, 29 Jun 1985, Cabrera 8778 (MO).

    HONDURUS. Dept. Comayagua, near Comayagua, 600 m, 12–23 Mar 11947, Standley 5312 (F); Dept. El Paraíso, vicinity of Danlí, 700–800 m, 11–23 Feb 1949, Standley 17042 (F); Dept. Olancho, near Juticalpa, 380–480 m, 5–16 Mar 1949, Standley 17953 (F).

  • Capraria frutescens is a very distinct species. It is readily distinguished by its monopodial stems, glandular-pilose vestiture, oblong sepals, bilateral corolla and reniform stigma (Fig. 2a). Due to its distinctiveness, the species has been recognized as the monotypic genus Pogostoma. However, because of the suite of characters it shares with C. biflora (Table 2) and because it has alternate leaves with punctate glands (like the other species of Capraria) I retain it in Capraria.

  • Fig. 7.

    Distribution of Capraria frutescens (open circles) and C. mexicana (diamonds).

    Fig.

    8. Capraria frutescens (Williams & Siedo 99, SHST). a. Habit. b. Leaf. c. Corolla, side view. d. Corolla, front view. e. Corolla, top view. f. Sepals, note glandular hairs. Figure by Maria Thompson.

    3. Capraria mexicana Moric. ex. Benth. in DC., Prodr 10: 429. 1846. Type: MEXICO. Tamaulipas: Tampico, s.d., Berlandier s.n (Lectotype: K!, designated by Sprague (1921)).

  • Subshrubs or suffruticose perennial herbs, stiffly erect to somewhat sprawling, 50–200 cm tall, with several ramified stems emerging from the root stock. Stems glabrous, 2–4 mm in diameter at midstem. Leaves lanceolate, mostly (3-) 4–10 (-11) cm long, 0.6–2.2 cm wide, glabrous. Pedicels 5–12 mm long, glabrous. Flowers 5-merous, perfect, actinomorphic, 8–10 mm long, 8–10 mm broad when open. Calyx lobes 3–5 mm long, 0.8–1.2 mm wide, lanceolate, glabrous. Corolla white, rotate. Corolla tube light green to white, glabrous internally, 2–3 mm long. Petal Lobes spreading, lanceolate, 4–7 mm long. Stamens 5, isomerous, exserted about 0.81.0 mm past the corolla tube, alternate with the corolla lobes; filaments glabrous. Style exserted, glabrous, 3–5 mm long. Stigma ellipsoid. Ovary glabrous. Seeds ca. 0.35 mm long, 0.2 mm wide.

  • Common Name: Tamaulipan Tea (Ideker, 1996).

  • Chromosome Numbers: None reported.

  • Phenology: Flowering November through April. Producing mature fruits February through August.

  • Distribution (Fig. 7): Northern Mexico with disjunct populations south into Belize, mainly eastern Mexico with one population in south Texas (Ideker, 1996). Sea level to 650 m elevation.

  • Illustration: Photo of flower (Fig. 1 d & e); illustration of flower and habit (Fig. 9).

  • Representative Specimens: MEXICO. Guanajuato. Mpio. San Luis de la Paz, paso Macuala del Realito, 8 Nov 1990, Ventura 9003 (F). Jalisco, volcanic soil on mountain side near Lake Chapala, 12 Aug 1947, Webster 7652 (F, TEX, US). Querétaro. Mpio. Arroyo Seco, Puente Coná, 4 km SE of Concá, 18 Mar 1985, Fernández 2777 (NY). San Luis Potosí. Mpio. Tanquian, Between Tamuin Pyramids and Sagrada Familia, 29 MAY 1979, Alcorn 3107 (TEX); Mpio. San Antonio, Tanjasnec, 8 Mar 1979, Alcorn 2515 (TEX); along Road Between Río Verde and San Ciro, 1000 m, 12 Aug 1954, Rzedowski 4509 (F, TEX). Tamaulipas. Barra del Tordo, ca. 45 km east of Al- dama, marshy area 1.5 mi inland from beach, 18 Mar 1996, Williams, Plum & Goldman 96–03 (SHST); Jua- mave, 1931, Rozynski 333 (F, NY, US); Rancho Cal- abazas, on the Río Sabinas, across from the village Azteca, 3 mi W of Rte. 85, at a point 20 m N of Ciudad Mante, 30 Mar 1975, Harriman et al. 10831 (F); 30 mi S of Ciudad Victoria, 4 Apr 1947, Smith &Barkley 17M191 (F, LL, MO); 14 mi E of of Victoria jet on road to Casas, 4 Feb 1960, Johnston 5020 (TEX); low shrub Oak forest 2 mi NE of Altamira, 3 Mar 1961, King 4039 (F, NY, TEX, US); Las Adjuntas, 24 Jan 1970, Gómez-Pompa 4721 (LL, MO); Mpio. Gomez Farias, NW of Gomez Farias in Sierra de Guatemala, on cretaceous limestone (23° 06′ N, 99° 12 ′ W), 17 Mar 1987, Woodruff 157 (TEX); near Tampico, 27 Apr 1910, Palmer 320 (F, NY); shores of Laguna Anda la Piedra, 2 Nov 1939, LeSueur 585 (F, US). Veracruz. 6 km from Panuco toward Tampico (22° 07′N, 98° 09′W), 20 Mar 1971, Chiang 418 (F, NY).

    BELIZE. Dist. Corozal, between Sarteneja and Chunox (18° 17′N, 88° 15′W), 18 Mar 1987, Davidse 32632 (MO); Stann Creek, 10 Jun 1932, Schipp 945 (F, NY); El Cayo, Mar 1933, Chanek 138 (F).

  • Capraria mexicana is readily distinguished by its glabrous condition, branching stems, and regular corollas with five isomerous stamens (Fig. 1d.).

    One population of Capraria mexicana has been reported in Texas and the species has been listed as endangered by the Texas Organization for Endangered Species (Ideker, 1996). Although Ideker reported the location of the population (Falcon Reservoir, Zapata Co., Texas), he did not cite a specimen.

  • Fig. 9.

    Capraria mexicana (Williams, Plum & Goldman 96–3, SHST). a. Habit, b. Leaf. c. Corolla, front view. d. Corolla, side view. Figure by Maria Thompson.

    4. Capraria peruviana Benth. in DC., Prodr. 10: 430. 1846. Type: ECUADOR. Guayaquil, s.d., Hinds s.n. (Lectotype: K!, designated here. Of the paratypes listed by Bentham I could only locate Hinds s.n., hence its selection as the lectotype.).

  • Xuaresia biflora Ruiz & Pav., Fl. Peruv. 2: 13. 1794. pl. 123. Type: ECUADOR. Prov. of Bolívar, Guaranda, 72 mi NE of Guayaquil, s.d., Née s.n. (Lectotype: MA! spec. # 488144, designated here. Although the name Xuaresia biflora Ruiz & Pav. pre-dates Capraria peruviana by 52 years, its placement within Capraria would create an illegitimate homonym of the earliest name, C. biflora L.)

  • Witheringia salicifolia Hook, [non Salisb.], Bot. Misc. 2: 231. 1831. Type: PERU. Prov. Lurin, near Lima, s.d., Cruckshanks s.n. (Holotype: K! Although the name Witheringia salicifolia Hook, pre-dates C. peruviana by fifteen years, its placement within Capraria would be illegitimate as it would be a later hononym of Capraria salicifolia Salisb. [= Freylinia lanceolata G. Don].)

  • Subshrubs or suffruticose perennial herbs, stiffly erect to somewhat sprawling, 50–200 cm tall, with several ramified stems emerging from the root stock. STEMS glabrous, 2–4 mm in diameter at midstem. Leaves lanceolate, mostly (3-) 4–10 (-11) cm long, 0.6–2.2 cm wide, glabrous. Pedicels 5–22 mm long, glabrous. Flowers 5-merous, perfect, actinomorphic, 5.0–6.0 mm long, 6–8 mm broad when open. Calyx lobes 3.0–4.0 mm long, 0.8–1.2 mm wide, lanceolate, glabrous. Corolla white, rotate. Corolla Tube light green to white, glabrous internally, 0.8–1.5 mm long. Petal Lobes spreading, lanceolate, 3–4.5 mm long. Stamens 5, isomerous, exserted about 0.8–1.0 mm past the corolla tube, alternate with the corolla lobes; filaments glabrous. Style included, glabrous, 0.75–1.50 mm long. Stigma ellipsoid. Ovary glabrous. Seeds ca. 0.35 mm long, 0.2 mm wide.

  • Common Name: Té del Perú, Te de Lima (Sprague, 1921). Smooth capraria (McMullen, 1999).

  • Chromosome Numbers: None reported.

  • Phenology: Flowering August through January. Producing mature fruits lanuary through August.

  • Distribution (Fig. 6): South America on the western side of the Andes (Peru and Ecuador) and the Galápagos Islands. Sprague (1921) and D'Arcy (1979) identified specimen Hayes 375 as C. peruviana and consequently reported the taxon as growing in Panamá. Unfortunately, I was unable to locate Hayes 375. I have examined other Capraria collections by Hayes (322, 745, 876) from the same region of Panama and I note that these specimens are rather narrowed leafed, glabrous individuals of C. biflora, which superficially resemble C. peruviana.

  • Economic Uses: Sprague (1921) reported that the leaves of this species are brewed as a tea in Peru.

  • Illustrations: A detailed line drawing is provided in Wiggins and Porter (1971) and McMullen (1999) has a beautiful color photo of the species.

  • Representative Specimens: ECUADOR. Prov. Chimborazo, Canyo of the Río Chanchan, from Naranjapata to below Huigra, 19 Jun 1945, Camp E-3854 (NY, US); Prov. El Oro, Santa Rosa, 11 Mar 1955, Asplund 15665 (LL, NY); Prov. Galápagos, Albemarle Island, Villamil, 27 Apr 1932, Howell 8950 (US); Prov. Galapagos, Charles Island, Black Beach, 3 Apr 1930, Svenson 177 (F); Prov. Galapagos, Santa Cruz Island, Academy Bay, 4 Mar 1960, Léveque 118 (US); Prov. Guayas, Guayaquil, Jan 1938, Mille 1064 (F); Prov. Esmeraldas, in and near Esmeraldas, 12 Sep 1974, Hudson 738 (NY); Prov. Guayas, 4 km S of Punta Blanca (N of Santa Elena), gallery vegetation in desert (80° 49′W, 02° 10′S), 300 m, 22 May 1973, Holm-Nielsen et al. 2459 (F, MO, NY); Prov. Manabí, 1.5–3 km W of Leonidas Plaza (00° 36′ S, 80 27′ W), 25–75 m, 16 Sep 1993, Webster 30652 (TEX).

  • PERU. Dept. Cajamarca, 30 km E of bridge over Río Maichil (79° 10′W, 06° 30′S), 1450 m, 9 Feb 1988, Gentry 61402 (F, MO); Prov. Casma, ca. 48 km N of Pativilca on Pan Am Hwy., coastal desert dunes, 10 m, 13 Oct 1984, Dillon 4011 (NY, TEX); Prov. Lambayeque, 12 km (via Pan Am Hwy) N of Olmos, 29 May 1978, Barbour 2117 (MO, NY); Prov. Santa Cruz, Lurin, 4 Jan 1925, Pennell 12209 (NY); Prov. Trujillo, Ladera, 26 Jun 1986, Mostacero 1105 (F, NY); Prov. Tumbes, Dept. Tumbes, La Cruz, 28 May 1992, Sagástegui 14611 (F, NY).

    Except for the consistently smaller styles (1–2 mm long; Fig. 2d) and somewhat smaller greenish white corollas, this species is almost identical to C. mexicana (Table 2).

  • Excluded Names

    Below is a list of names in Capraria that are presently referable to other genera. The six Capraria names highlighted in bold are those not accounted for in Sprague's (1921) study.

    • Capraria aegyptiaca Steud. & Hochst. ex Endl., Nov. Strip. Dec. 23. 1839. = Anticharis Arabica (Steud. & Hochst. ex Endl.) Endl. (fide Sprague, 1921)

    • Capraria annua (Cham, et Schltdl.) Kuntze, Rev. Gen. Pl. 2: 459. 1891. = Scoparia annua Cham, et Schltdl. (fide Sprague, 1921)

    • Capraria arabica Steud. ' Hochst. ex Endl., Iconogr. Gen. Pl. pl. 7. 1837. = Anticharis arabica (Steud. & Hochst. ex Endl.) Endl. (fide Sprague, 1921)

    • Capraria calycina A. Gray, Proc. Am. Acad. Arts. 6: 49. 1862. = Myoporum debile R. Br. (fide Sprague, 1921)

    • Capraria crustacea L., Mant. Pl. 87. 1767. = Lindernia crustacea (L.) F. Muell. (fide Howard, 1989)

    • Capraria diffusa Roxb., Hort. Bengal. 47. 1814. = Ebermaiera thysoidea Wall. (fide Sprague, 1921)

    • Capraria dissecta Delile, Descr. Egypte, Hist. Nat., 95. t. 32. fig. 3. 1812. = Sutera glandulosa Roth, (fide Sprague, 1921)

    • Capraria dulcis (L.) Kuntze, Rev. Gen. Pl. 2: 459. 1891. = Scoparia dulcis L. (fide Sprague, 1921)

    • Capraria durantifolia L., Syst. Nat., ed. 10. 1116. 1759. = Stemodia durantafolia (L.) Swartz (fide Turner & Cowan, 1993)

    • Capraria elliptica (Cham.) Kuntze, Rev. Gen. Pl. 2: 459. 1891. = Scoparia elliptica Cham, (fide Sprague, 1921)

    • Capraria ericacea (Cham. & Schltdl.) Kuntze, Rev. Gen. Pl 2: 459. 1891. = Scoparia ericacea Cham. & Schltdl. (fide Sprague, 1921)

    • Capraria gratioloides L., Syst. Nat., ed. 10. 1117. 1759. = Ilysanthes gratioloides (L.) Benth. (fide Sprague, 1921)

    • Capraria gratissima Roxb., Hort. Bengal. 47. 1814. = Limnophila roxburghii G. Don. (fide Sprague, 1921)

    • Capraria humifusa Buch.-Ham. ex Wall., Cat. n. 3883. 1831. = Centranthera humifusa Wall, (fide Sprague, 1921)

    • Capraria humilis Solander in W. T. Aiton, Hortus Kew., ed. I. 2: 354. 1789. = Stemodia verticillata (Miller) Hassler (fide Turner & Cowan, 1993)

    • Capraria integerrima Miq., Linnaea, 22: 476. 1849. = Sesamum indicum L. (fide Sprague, 1921)

    • Capraria integrifolia Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles, 12: 20. 1845. Type: MEXICO. Oaxaca. Oaxaca, 1840, Galeotti 653 (Holotype: BR!). = Nama jamaicense L. (In his treatment of Capraria, Sprague (1921) recognized this as a valid species of Capraria. Examination of the type material, however, reveals that this is, in fact, a species of Nama.)

    • Capraria lanceolata L. f., Suppl. Pl. 284. 1781. = Freylinia lanceolata (L. f.) G. Don. (fide Sprague, 1921)

    • Capraria longiflora Thunb., Mus. Nat. Acad. Upsal. 17: 150. 1794. = Freylinia undulata (L. f.) Benth. (fide Sprague, 1921)

    • Capraria lucida W. T. Aiton, Hortus Kew., ed. I. 2: 353. 1789. = Teedia lucida (W. T. Aiton) Rudolphi (fide Sprague, 1921)

    • Capraria monnieria Roxb., Hort. Bengal. 47. 1814. = Herpestis monnieria (Roxb.) H.B.K. (fide Sprague, 1921)

    • Capraria montevidensis Kuntze, Rev. Gen. Pl. 2: 459. 1891. = Scoparia montevidensis (Kuntze) R. E. Fries; S. flava Cham. & Schltdl. (fide Sprague, 1921)

    • Capraria multifida Michx., FL Bor.-Amer. 2: 22. t. 35. 1803. = Conobea multifida (Michx.) Benth. (fide Sprague, 1921)

    • Capraria multiflora Steud., Nom. ed I. 149. 1820. = Conobea multifida (Michx.) Benth. (fide Sprague, 1921)

    • Capraria oppositifolia L., Fl. Jamaic. 380. 1759. = Stemodia durantifolia (L.) Swartz (fide Turner and Cowan, 1993).

    • Capraria pinnatifida Kuntze, Rev. Gen. Pl. 2: 459. 1891. = Scoparia flava Cham. & Schltdl. (fide Index Kewensis)

    • Capraria pusilla Torr., Ann. Lyceum Nat. Hist. New York, 1: 36. 1824. = Mimulus floribundus Dougl. (fide Sprague, 1921)

    • Capraria rigida Buch.-Ham. ex Hook, f., Fl. Brit. India 4: 301. 1884. = Centranthera hispida R. Br. (fide Sprague, 1921)

    • Capraria rigida Thunb., Prodr. Pl. Cap. 103. 1800. = Ehretia rigida (Thunb.) Druce (fide Sprague, 1921)

    • Capraria salicifolia Link & Otto, Icon. Pl. Rar. Hort. Berol. 11. t. 4. = Freylinia cestroides Colla (fide Index Kewensis)

    • Capraria salicifolia Salisb., Prodr. Strip. Chap. Allerton 94. 1796. = Freylinia lanceolata (L. f.) G. Don. (fide Sprague, 1921)

    • Capraria ternifolia Sessé & Mo¸., Fl. Mexic., ed. 2, 145. 1894. = Stemodia tenuifolia Minod (fide McVaugh, 2000)

    • Capraria undulata L. f., Suppl. Pl. 284. 1781. = Freylinia undulata (L.f.) Benth. (fide Sprague, 1921)

    • Capraria uniflora Burm. f., Fl. Indica 133. fig. 3. 1768. = Lysimachia sp. (fide Sprague, 1921)

    • Capraria verticillata Sessé & Moç., Pl. N. Hispan., ed. 1, 98. 1887–90. = Russelia sp. (fide McVaugh, 2000)

    Acknowledgements

    Appreciation is given to Billie Turner fpr suggesting this project. W. Patrick Spencer provided valuable assistance and superb technique during the preparation and photography of the SEM samples. I am equally indebted to Kerry Barringer, James Henrickson, Sarah Jackson, Heidi Meudt, and Beryl Simpson for patiently reviewing and improving the manuscript. I also thank Maria Thompson (“La Mulata”) for the illustrations, and the curators of the following herbaria for loan of their material: F, FLAS, K, LL, MO, NY, TEX, US.

    Literature Cited

    1.

    Argue, C. L. 1980. Pollen morphology in the genus Mimulus (Scrophulariaceae) and its taxonomic significance. Amer. J. Bot. 67: 68–87. Google Scholar

    2.

    Arekal, D., S. Rajeshwari and S. N. R. Swamy. 1971. Contribution to the embryology of Scoparia dulcis L. Bot. Not. 124: 237–248. Google Scholar

    3.

    Barringer, K. 1993. Five new tribes in the Scrophulariaceae. Novon 3: 15–17. Google Scholar

    4.

    Bentham, G. 1846. Scrophulariaceae, in A. De Candolle, Prodromus Systematis Naturalis Regni Vegetabilis 10: 186–598. Google Scholar

    5.

    Bentham, G. 1876. Scrophulariaceae, in G. Bentham and J. D. Hooker, Genera Plantarum 2: 913–980. Google Scholar

    6.

    Borgen, L. 1980. Chromosome numbers of Macaronesian flowering plants III. Bot. Macaronés 7: 67–76. Google Scholar

    7.

    D'Arcy, W. G. 1979. Capraria. in Flora of Panamá. Ann. Missouri Bot. Gard. 66: 208–211. Google Scholar

    8.

    Elisens, W. J. 1985. Monograph of the Maurandyinae (Scrophulariaceae-Anthirrhineae). Syst. Bot. Monogr. 5: 1–97. Google Scholar

    9.

    Hakki, M. I. 1975. Embryologische und morpholo- gische Untersuchungen aus Pflanzen aus Westin- dien: 1. Zur Embryologie von Capraria biflora L. (Scrophulariaceae). Bot. Jahrb. Syst. 96: 125–153. Google Scholar

    10.

    Henrickson, J. and L. D. Flyr. 1985. Systematics of Leucophyllum and Eremogeton (Scrophulariaceae). Sida 11: 107–172. Google Scholar

    11.

    Howard, R. A. 1989. Flora of the Lesser Antilles: Leeward and Windward Islands. Jamaica Plain, MA: Arnold Arboretum. Google Scholar

    12.

    Ideker, J. 1996. Capraria mexicana (Scrophulariaceae), an endangered addition to the United States flora. Sida 17: 523–526. Google Scholar

    13.

    Kapoor, B.M., S. Ramcharitar and C. Gervais. 1987. Liste annotée de nombres chromosomiques de la flore vasculaire du nord-est de l'Amérique. Le Naturaliste Canadien 114: 105–116. Google Scholar

    14.

    Lersten, N. R. and Curtis, J. D. 2001. Idioblasts and other unusual internal foliar secretory structures in Scrophulariaceae. PL Syst Evol. 227: 63–73. Google Scholar

    15.

    Linnaeus, C. 1753. Species Plantarum. 2: 875. Fascimile edition. 1957. London: The Ray Society. Google Scholar

    16.

    Mabberley, D. J. 1987., The Plant Book, a Portable Dictionary of the Higher Plants. Cambridge: Cambridge University Press. Google Scholar

    17.

    McMullen, C. K. 1999. Flowering Plants of the Galápagos. Ithaca: Comstock Publishing Associates, Cornell University Press. Google Scholar

    18.

    McVaugh, R. 2000. Botanical Results of the Sessé & Moçiño Expedition (1787–1803): VII, A Guide to Relevant Scientific Names of Plants. Pittsburgh: Hunt Institute for Botanical Documentation, Carnegie Mellon University. Google Scholar

    19.

    Méndez Larios, I. and Villaseñor Ríos, J. L 2001. La familia Scrophulariaceae en Mexico: diversidad y distribution. Bol. Soc. Bot. México. 69: 101–121. Google Scholar

    20.

    Niezgoda, C. and S. A. Tomb. 1975. Systematic pal- ynology of the tribe Leucophylleae (Scrophulariaceae) and selected Myoporaceae. Pollen & Spores 17: 497–516. Google Scholar

    21.

    Olmstead, R. 2004. A synoptical classification of the Lamiales (in progress). January 2004.  http://depts.Washington.edu/phylo/classifications/Lamiales.html  Google Scholar

    22.

    Pastor, R. J. J. and I. Fernández. 2000. SEM and light microscope observations on fruit and seeds in Scrophulariaceae from Southwest Spain and their systematic significance. Ann. Bot. (London) 86: 323–338. Google Scholar

    23.

    Pennell, F. W. 1920. Scrophulariaceae of Colombia. I. Proc. Acad. Nat. Sci. Philadelphia 72: 136–188. Google Scholar

    24.

    Pennell, F. W. 1935. The Scrophulariaceae of eastern temperate North America. Acad. Nat. Sci. Philadelphia Monogr. 1: 1–650. Google Scholar

    25.

    Raman, S. 1991. The trichomes on the corolla of the Scrophulariaceae. X: Taxa of uncertain relationships to, or within, the family. Beitr. Biol. Pflanzen 66: 127–143. Google Scholar

    26.

    Read, J. C. and B. J. Simpson. 1989. Documented plant chromosome numbers 1989: 2. Chromosome counts of three species in the genus Leucophyllum. Sida 13: 369. Google Scholar

    27.

    Sanders, A. C. et al. 1996. Noteworthy collections. Madroño 43: 523–538. Google Scholar

    28.

    Sparague, T. A. 1921. A revision of the genus Capraria. Kew Bull. 1921: 205–212. Google Scholar

    29.

    Subramanian, D. and R. Pondmudi. 1987. Cytotax- onomical studies of south Indian Scrophulariaceae. Cytologia. 52: 529–541. Google Scholar

    30.

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

    31.

    Thieret, J. W. 1967. Supraspecific classification in the Scrophulariaceae: A review. Sida 3: 87–106. Google Scholar

    32.

    Turner, B. L. and C. C. Cowan. 1993. Taxonomic overview of Stemodia (Scrophulariaceae) for North America and the West Indies. Phytologia 74: 61–103. Google Scholar

    33.

    Watson, L. and M. J. Dallwitz. 1992 onwards. The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval Version: 14th. December 2000.  http://biodiversity.uno.edu/delta/Google Scholar

    34.

    Wettstein, R. von. 1891. Scrophulariaceae. in A. Engler and E. K. Prantl, Die naturlichen Pflanzenfamilien 4: 30–107. Google Scholar

    35.

    Wiggins, I. L. and D. M. Porter. 1971. Flora of the Galápagos Islands. Stanford: Stanford University Press. Google Scholar

    36.

    Zhao, Z. 1996. Documented plant chromosome numbers 1996: 2. Miscellaneous U.S.A. and Mexican species, mostly Asteraceae. Sida 17: 259–263. Google Scholar

    Appendices

    NUMERICAL LIST OF TAXA

    INDEX TO THE NUMBERED COLLECTIONS EXAMINED

    • The exsiccate listed below is for all specimens examined during this study and includes those listed and those not listed in the representative specimens above. The numbers in parentheses refer to the corresponding taxa in the Numerical List of Taxa presented above.

    • Abbott, W. L. 1098, 1762 (1) .

    • Acevedo-Rodríguez, P. 633, 1936, 4287, 5296 (1) Acosta-Solís, M. 6977 (4)

    • Aguilar, I. 124 (1)

    • Aguirre, B. G. 309 (2)

    • Alain, Bro. 4295 (1)

    • Alcorn, J. B. 451 (1) 1559, 2075, 2296 (2) 2515, 3107 (3)

    • Allard, H. A. 13380, 14770 (1)

    • Allart, A. 275 (1)

    • Almandoz 1 (1)

    • Almeda, F. 3153 (1)

    • Alvarez M., D. 580 (1)

    • Aymard, G. 2169 (1)

    • Andersson, L. 1853 (1)

    • André 31 (1)

    • Andrews, L. M. 643 (1)

    • Anthony, J. Ill (4)

    • Antonio, T. 830 (1)

    • Araquistain, M. 1840 (1)

    • Archer, W. A. 2251 (1)

    • Aristeguieta, L. 1100 (1)

    • Armour 217, 1364 (1)

    • Arnason, T. 17509 (1)

    • Arnoldo, F. M. 1736, 2054 (1)

    • Artemio 54 (1)

    • Asplund, E. 15665, 15960 (4)

    • Atwater 590 (1)

    • Atwood, J. T. & J. C. Seymour 3538 (1)

    • Augusto, Bro. B. 452, 1447 (1)

    • Ayala, M. G. 447 (2)

    • Balick, M. 1768 (1)

    • Barbour, P. J. 2117 (4)

    • Barclay, G. W. 282 (4)

    • Barkley, F. A. 7652, 17M191 (3)

    • Barringer, K. 738, 795, 773 (1)

    • Bartlett, H. H. 11944, 12463, 17493 (1)

    • Bautista, H. P. 765 (1)

    • Beaman, J. 529 (1)

    • Bentley 63, 313 (1)

    • Bequaert 78 (2)

    • Bono, J. 5004 (1)

    • Bordenave, B, 128 (1)

    • Box, H. E. 1051 (1)

    • Brace, L. K. J. 1752, 4318, 4660, 5228 (1)

    • Bradburn, A. S. 1240 (2)

    • Brass, L. J. 21139 (1)

    • Breedlove, D. 9098, 28999 (1); 13544, 26004 (2) Breteler, F. J. 4353 (1)

    • Britton, N. L. 14, 143, 218, 321, 518, 751, 757, 1932, 2527, 2708, 2932, 3902, 4092, 5581, 5847, 6560 (1)

    • Broadway, W. E. 36, 108, 2468, 2688, 3580, 5139 (1)

    • Brooke, W. M. A. 5562 (1)

    • Brown, S. 603 (1)

    • Brumbach 5666, 7790, 9151 (1)

    • Burch, D. 2404, 4238 (1)

    • Cabrera, A. L. 4142, 33677 (1)

    • Cabrera, E. 9093, 9209, 10478, 10497 (1); 8778, 9006, 9525 (2)

    • Calderón, G. M. 1138 (1)

    • Calderón, S. 1377, 1460, 2618 (1)

    • Camp, W. H. E-3854 (4)

    • Cárdenas, M. 2523 (1)

    • Cardona, F. 556 (1)

    • Carleton 150 (1)

    • Carlson, M. C. 530, 1497 (1)

    • Carvalho, A. 2 (1)

    • Castañeda, R. R. 203 (1)

    • Castillo, G. & Tapia, L. 1085 (2)

    • Catino 14 (1)

    • Chan, C. 2240 (1)

    • Chanek, M. 138 (3)

    • Chardon, C. E. 1066 (1)

    • Chiang, F. 418 (3)

    • Christman 30 (1)

    • Clemente, Bro. 5991 (1)

    • Clough 44 (1)

    • Collins, F. S. 333 (1)

    • Combs, R. 482 (1)

    • Contreras, E. 87, 128, 1946, 3631 (1)

    • Conzatti, C. 1658 (1)

    • Cooley, G. R. 2433, 2450, 2512, 8141 (1)

    • Cooper, W. 49 (1)

    • Correll, D. S. 40537 43274, 43683, 45110, 45510

    • 46206, 46333, 47434, 50988, 51435 (1)

    • Corivelius 286 (1)

    • Cowan, C. 1102, 1792, 1883, 3188, 5016, 5078 (1); 5020 (2)

    • Crane, C.J. 333 (3)

    • Cristoff 190 (1)

    • Croat, T. B. 24613 (1)

    • Cuadros, H. 1085 (1)

    • Cuatrecasas, J. 24304 (1)

    • Curran, H. M. 6, 637, 1163 (1)

    • Curtiss, A. H. 541, 1901, 5648 (1)

    • D'Arcy, W. G. 327D, 4680, 10431, 14237 (1)

    • Daniel, H. 5075, 5675 (1)

    • Dash, J. S. 205 (1)

    • Davidse, G. 3952, 32632 (3)

    • Dawson 26850 (1)

    • Degener, O. 19068 (1); 26470 (2)

    • de Granville, J. J. 7282 (1)

    • de La Cruz, J. S. 3810, 4575 (1)

    • Delascio, J. F. 6946 (1)

    • Díaz, A. 197 (1)

    • Dillon, M. 4011 (4)

    • Dorantes, J. 1151 (1); 920 (2)

    • Drake, E. 889 (1)

    • Duckett, F. 92 (1)

    • Dugand, A. 202, 2720, 3256, 3300, 4902, 5625 (1)

    • Duke, J. A. 2502 (1)

    • Dunn & Harmon 17628 (3)

    • Durante, A. P. 1334 (1)

    • Dwyer, J. D. 2819, 12177 (1)

    • Earle, F. S. 39 (1)

    • Eggers, H. F. A. 14871 (4)

    • Egler, F. E. 39–147, 42–99, 42–211 (1)

    • Eiten, G. 4874, 10327 (1)

    • Ekman, E. L. 9344 (1)

    • Elias, B. 227, 249, 385, 593, 951 (1)

    • Ernst, W. R. 2119 (1)

    • Espinal, F. 139 (1)

    • Esposto 14796 (4)

    • Eyerdam, W. J. 430 (1)

    • Fendler, A. 543, 837 (1)

    • Fernández N., R. 1248 (1); 3685 (2); 2111 (3)

    • Ferreyra, R. 7611, 10715 (4)

    • Ferris, R. S. 5731 (1)

    • Fisher, G. L. 35309 (1)

    • Fisher-Meerow 788 (1)

    • Fishlock, W. C. 211, 270 (1)

    • Fleury, M. 794, 860 (1)

    • Fosberg, F. R. 19316, 44712, 53877, 56088 (1)

    • Fournier, L. A. 161, 263 (4)

    • Fróes, R. 50720 (1)

    • Fryxell, P. 3265 (2)

    • Fuertes. J. 588 (1)

    • Garber, A. P. 8076 (1)

    • Gardner, G. 3372 (1)

    • Garganta, M. 1107 (1)

    • Gaumer, G. F. 348, 1621, 1675, 23496 (1); 409, 520, 1468, 1554, 1622, 24099 (2)

    • Gentry, A. 10004, 61402 (4)

    • Gentry, H. S. 5032, 5805 (1)

    • Gillis, W. T. 7211 (1)

    • Gilly, C. L. 80 (2)

    • Glaziou, A. F. M. 19734 (1)

    • Goldman, E. A. 441 (1)

    • Goll, G. P. 552, 563 (1)

    • Gómez P., L. D. 3312 (1)

    • Gómez-Pompa, A. F. 4721 (3)

    • Greenman, J. M. 282, 378 (2)

    • Grijalva, A. 823, 1027 (1)

    • Gutiérrez B., C. 4752, 5047 (1); 4508 (2)

    • Haenke, T. 2270 (4)

    • Hall 280 (1)

    • Harley, R. M. 16196, 16445 (1)

    • Harling & Andersson 19347 (4)

    • Harlow 60 (1)

    • Harriman, N. A. 10831 (3)

    • Harris, W. 5933 (1)

    • Hartling 47316 (1)

    • Hatschbach, G. 45120 (1)

    • Haught, O. 3970 (1); F-109; 192 (4)

    • Havard, V. 118 (1)

    • Hawkins 58 (1)

    • Hayes 322, 745, 876 (1)

    • Hekking, A. 823 (1)

    • Heller, A. A. 320 (1)

    • Heriberto, Bro. 186, 299 (1)

    • Hespenheide, H. A. 1354 (1)

    • Hess, L. 1259 (1)

    • Heyde 623, 2931 (1)

    • Hill 2216, 13415, 24726 (1)

    • Hinton, G. B. 26492 (1); 1979, 3383, 5474 12824, 12876, 16112 (2)

    • Hitchcock, A. S. 258, 16645 (1)

    • Hodge, W. H. 3706 (1)

    • Hoffman, B. 758 (1)

    • Hohenacker, R. F. 628 (1)

    • Holm-Nielsen 2459, 7197 (4)

    • Holton, I. F. 582 (1)

    • Hood 22983 (1)

    • Howard, R. E. 4592, 5122, 5394, 5798, 6581 (1)

    • Howell, J. T. 8950 (4); 9105 (1)

    • Hudson, J. 738 (4)

    • Hunnewell, F. W. 11491, 19499 (1)

    • Irwin, H. S. R-97, 348 (1)

    • Játiva, C. & Epling, C. 451 (4)

    • Jahn, A. 1214 (1)

    • Jenman, G. S. 4499, 5460, 5857 (1)

    • Jimenez, G. 9 (3)

    • Johnston, M. 265, 1075 (1) 4926, 5020 (3)

    • Jones, M. E. 23206 (1)

    • Karwinsky, B. W. F. 516 (3)

    • Kenoyer 765 (2)

    • Kellerman, W. A. 5734, 7661 (1)

    • Killip, E. P. 14029, 14057, 14451, 14636, 31414, 31605, 38366, 41703 (1)

    • King, R. M. 518, 1684 (1) 3801, 3879, 4039 (3)

    • Kirkbride, J. H. 2629 (1)

    • Klawe 1516 (1)

    • Kuntze, O. 193, 279 (1)

    • Lakela 28014, 30471 (1)

    • Lanjouw, J. 1090, 1524 (1)

    • Lasser, T. 688 (1)

    • Laughlin 1320, 1664 (1)

    • Lavastre, Bro. B. A. 1912 (1)

    • Leiva, S. 1991 (4)

    • Leonard, E. L. 4146, 5305, 11020, 11022, 11207, 12206, 15458, 11731 (1)

    • LeSueur, D. H. 378 (2) 585 (3)

    • Léveque 118 (4)

    • Lewis, W. H. 4680 7210, 7444 (1)

    • Liebmann, F. M. 9459 (1)

    • Liesner, R. 2764, 4277, 4462 11933 (1)

    • Liogier, A. H. 17500, 18207, 30173 (1)

    • Lira, E. M. 419 (1)

    • Llatas Quiroz, S. 853 (4)

    • Lopéz- Figueiras, M. 199, 480 (1)

    • Lopéz-Palacios 4369 (1)

    • Lot, A. 1344 (1)

    • Lozano-C., G. 2718 (1)

    • Lundell, C. L. 898, 1904, 2212, 7854, 7976, 16474 (1); 7877 (2)

    • Maas, P. J. 547 (1)

    • MacDaniels, L. H. 456 (2)

    • Madrid, E. 497 (1)

    • Madsen, J. E. 64024 (4)

    • Maltby, T. S. 53, 135 (1)

    • Martínez, E. 27788, 28286, 28643, 28830, 28921, 28968, 29642, 31372 (1)

    • Martínez-Calderon, G. 1472 (2)

    • Matuda, E. 692, 3271, 4734, 16338, 16758 (1) Maxon 6951, 7118, 7255 (1)

    • McGregor, R. L. 16299 (3)

    • Mejía, M. M. 8951 13119 (1)

    • Mell, C. D. 249, 2154 (1)

    • Mexia, Y. E. J. 1041 (2)

    • Meyer, F. G. 8512 (1)

    • Mille, L. 1064 (4)

    • Miller, O. O. 40, 41, 58 (1)

    • Millspaugh, C. F. 348, 120, 217, 421, 710, 1076, 1364, 1513, 1527, 1732, 2284, 8989 (1); 520, 1625 (2)

    • Moldenke, H. 471, 3731 (1)

    • Molina R„ A. 3238, 3245, 6597, 15432, 24634, 34273 (1)

    • Montes, M. N. 22 (2)

    • Mori, S. 16588, 18800, 21213 (1)

    • Moreno, P. 607, 1724, 3577, 5626, 6778, 7346 (1)

    • Morrow 111 (1)

    • Morton, C. V. 3022, 5155, 10697 (1)

    • Mostacero, J. 739, 1105 (4)

    • Muenscher, W. C. 18268 (1)

    • Nash, G. V. 1082, 3827 (1)

    • Nee, M. 29256, 41641 (1)

    • Nelson, C. 7631 (1)

    • Nelson, G. 38, 419, 3097 (1); 3098, 6229 (2); 6043 (3)

    • Newman 36 (3)

    • Nicolson 2007 (1)

    • Northrop, A. R. 27, 381 (1)

    • Orcutt, C. R. 5009 (1); 4522 (2)

    • Orozco, J. M. 215 (1)

    • Ortega, J. S. 4372, 5146 (1)

    • Ortíz, R. T. 399, 718, 726, 737, 2036 (1)

    • Padilla, I. 121 (1)

    • Palmer, E. 43 (US), 361, 488 (1); 43 (F, NY), 567, 917, 5280 (2); 2, 188, 320, 1125 (3)

    • Paul, B. 814 (1)

    • Pennell, F. W. 3, 20, 3704, 3789, 4039, 4711, 4761, 4771, 9559, 9598, 9610, 9633, 12043 (1); 12209 (4)

    • Perdero 44, 119 (1)

    • Pflanz 4043 (1)

    • Pickel, B. J. 441 (1)

    • Pierottii, S. A. 1383 (1)

    • Pimentel, J. D. 532 (1)

    • Pipoly, J. 9044, 11248 (1)

    • Pippen, R. W. 905 (1)

    • Pires, J. M. 3429 (1)

    • Pittier, H. F. 1611, 3533, 5822, 5823, 9083, 10236 (1)

    • Plowman, T. 13539 (1)

    • Pollard, C. L. 13 (1)

    • Proctor, G. R. 3297, 16828, 18026, 18697, 32608, 38438, 42625 (1)

    • Purpus, C. A. 363 (1); 2247, 5696, 14224, 15025, 16793 (2)

    • Questel 214 (1)

    • Quevedo, R. 70 (1)

    • Ramírez, N. 2488, 2715 (1)

    • Reed, D. 1049 (1)

    • Renson 189 (1)

    • Ricksecker, A. E. 143, 301 (1)

    • Rivero 1052 (1)

    • Robles, J. 363 (2)

    • Robleto, W. 1661 (1)

    • Rodriguez, J. V. 2323, 2324, 2458, 3637 (1); 2329 (2)

    • Romero 988 (1)

    • Romero-Castañeda, R. 9680, 10500 (1)

    • Rose, J. N. 4002, 4280, 13218 (1); 13772 (2)

    • Rothrock, J. T. 381 (1)

    • Rowlee 164 (1)

    • Rozynski, H. W. 333 (3)

    • Runyon, R. 963 (3)

    • Rusby, H. H. 37 (1)

    • Rzedowski, J. 109, 4509 (3)

    • Saer d'Heguert, J. 340 (1)

    • Sagástegui, A. 8312, 10158, 11453, 12416, 14611 (4)

    • Sagot, P. A. 422 (1)

    • Samuels 263 (1)

    • Sanders, A. C. 9564 (1); 9628 (2)

    • Saravia, C. 431 (1)

    • Sargent 150 (1)

    • Schimpff, H. J. F. 2 (1) 7, 99 (4)

    • Schipp, W. A. 945 (3)

    • Schott, A. 2 (1)

    • Schulz, J. P. 5370 (1)

    • Seler, E. 1754, 3314, 3679 (1)

    • Seymore 2598, 5428 (1)

    • Shafer, J. A. 14, 88, 135, 792, 2433, 7889, 13431 (1)

    • Shillingford, C. A. 322 (1)

    • Sieber, F. W. 1620 (1)

    • Small, J. K. 726, 1314, 3088, 3593, 3732, 4940, 5694, 6014, 7374, 7659 (1)

    • Smith 2122, 10026 (1)

    • Smith, H. 551, 1331, 2823, 4038 (1)

    • Standley, P. C. 11136, 12717, 15374, 15682, 17164, 21146, 25323, 30798, 31209, 32113, 55077, 57641, 63971, 74392, 71992 (1); 5312, 17042, 17953 (2)

    • Steere, W. C. 1053, 1313 (1), 1083, 11006 (2)

    • Sterene 6350 (1)

    • Stern, P. 219A, 1457, 3340 (1)

    • Stevens, W. D. 3593, 3778, 11224, 16387 (1)

    • Stevenson, P. 1419, 2709 (1)

    • Stewart 3424, 3430 (4)

    • Steyermerk, J. A. 31772, 38303, 46284, 51349, 75227,110325, 129423 (1).

    • Staffers 1572, 3371 (1)

    • Stergios, B. G. 574, 7773 (1)

    • Struthers 62 (3)

    • Svenson, H. K. 177 (4); 243 (1)

    • Tamayo, F. 997, 2573 (1)

    • Taylor 32, 6792, 7391, 12616 (1); 7219 (3)

    • Tejera 114 (1)

    • Tellez, O. 1111, 1359 (1)

    • Tharp, B. C. 4614 (3)

    • Ton, A. M. 2979 (1); 9380 (2)

    • Toro 11 (1)

    • Tracy 7606 (1)

    • Trochez, L. 141 (1)

    • Tucker 840 (1)

    • Türckheim, H. 2695 (1)

    • Underwood, L. M. 160, 378, 1425 (1)

    • Urban, I. 785, 3399 (1)

    • Utrera, A. 135 (1)

    • Valdez, P. 26 (1)

    • Valverde, M. 50, 328 (4)

    • Van Devender 95–90 (1)

    • Van Hermann, H. A. 157, 497, 522, 607 (1)

    • Varela 5404 (1)

    • Ventura, A. 2259 (1); 2613 (2); 9003 (3)

    • Venturi, S. 5061, 7772 (1)

    • Vogel, C. 286 (1)

    • Von Reis 249 (1)

    • Watson 1118 (1)

    • Webster, G. 13451 (1); 7652 (3); 30652 (4)

    • Wedel, H. 509 1910, 2977 (1)

    • Whiteford 4076 (1)

    • Wiggins, I. 18320 (1)

    • Wight, A. E. 43, 234 (1)

    • Wilbur, R. L. 2557 (1)

    • Williams, J. K. 95–48 (1); 95–49, 99 (2); 96–3 (3)

    • Williams, L. 3070 (1)

    • Williams, L. O. 1564, 8971, 9808, 10227, 11450, 12450, 41844 (1)

    • Wilson 7290, 7640 (1); 12274 (3)

    • Woodbury, R. 1–66, 1–67, M–18, MB-68, WI-26 (1)

    • Woodruff, L. 157 (3)

    • Woodson, R. E. 1492 (1)

    • Worthington, R. D. 17695 (1)

    • Wright, C. 375 (1)

    • Wunderlin, R. P. 8329, 8488 (1)

    • Yanez, C. V. 844 (1)

    • Young 132 (1)

    • Yuncker, T. G. 17475, 18179, 18186 (1)

    • Zanoni, T. A. 3963, 7769, 9373, 13036, 37229 (1)

    • Zelaya, L. 81 (1)

    Justin K. Williams "A Revision of Capraria (Scrophulariaceae)," Lundellia 2004(7), 53-78, (1 December 2004). https://doi.org/10.25224/1097-993X-7.1.53
    Published: 1 December 2004
    JOURNAL ARTICLE
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