TERNSTROEMIACEAE Anna L. Weitzman FREZIERA Freziera Willdenow, Sp. pl. 2: 2: 1179. Dec. 1799, nomen conservandum. Swartz, Fl. ind. occ. 2: 971. Oct. 1800. Type species: Freziera undulata ( Swartz ) Willdenow (conserved). Eroteum Swartz , Prodr. 85. 1788; no type specified; nomen rejecienda; non Eroteum Blanco, 1857 (= Trichospermum Bl., Tiliaceae). Lettsomia Ruiz & Pavón, Fl. peruv. prodr. 4: 77. 1794; no type specified; nomen rejecienda; non Letsomia Roxburgh, 1814 (= Argyreia Loureiro, Convolvulaceae). Eurya Thunberg Freziera Szyszylowicz in Engler & Prantl, Nat. Pflanzenfam. III, 6: 190. 1893. Patascoya Urban, Ber. Deutsch. Bot. Ges. 14: 283. 1896. Type: Patascoya stuebelii ( Hieronymus) Urban. Eurya Thunberg Freziera Melchior in Engler & Prantl, Nat. Pflanzenfam. ed. 2. 21: 148. 1925. Killipiodendron Kobuski, J. Arnold Arbor. 23: 231. 1942. Type: Killipiodendron colombianum Kobuski. Dioecious small trees or shrubs; mature branches terete; twigs frequently with indumentum, lenticellate; terminal bud conduplicate-involute, or merely conduplicate, usually with indumentum. Leaves alternate, distichous, simple, exstipulate, petiole, when present, winged, colleter(s) often present in petiole base, narrowly triangular, flattened, red to black; blade usually coriaceous, apex terminating in caducous, thick, conical, black seta, the margin usually sinuous, with forward pointing setae associated with each tooth, the surface often with indumentum above and/or below. Inflorescence axillary, solitary, short-racemose or rarely single-flowered, with flowers 1-14, pedicel scars absent or 1-14; bract nearly always persistent, sclerotic, the base clasping, the apex nearly always terminating in thick, conical, black seta, the outer surface often with indumentum, the inner surface nearly always glabrous; pedicel present (or absent) often with indumentum; bracteoles two, nearly always apical on pedicel, usually opposite, persistent, convex, sclerotic at least centrally, the outer surface often with indumentum, the inner surface nearly always glabrous. Flowers small,  15mm long, actinomorphic, hypogynous, unisexual, though staminate flowers frequently appear hermaphroditic and carpellate flowers have staminodia; sepals five, distinct, imbricate, broadly ovate, nearly always convex and sclerotic (at least centrally), the outer surface often with indumentum; corolla urceolate, the petals five, distinct or lightly connate basally, imbricate, membranaceous below, sclerotic above. Staminate flowers with stamens (8–)15–35(–48), uniseriate, free or adnate basally, equal or unequal (if unequal nearly always in no particular order), the anthers 2-celled, latrorse, connective usually extending into a flap-like apiculus; gynoecium conical or pyriform, (2–)3–5(–6-)carpellate, with locules containing (0–)18–60(–110) ovules, the style tapering, the stigmatic lobes erect (or flaring), the surfaces abaxial and adaxial (or only adaxial), minutely papillate. Carpellate flowers with staminodia (6–)15–35(–38), uniseriate, free or adnate basally, usually equal, gynoecium pyriform or conical, (2–)3–5(–6-)carpellate, with locules containing (6–)16–100(–150) ovules, the style tapering, the stigmatic lobes flaring (or erect), with surfaces abaxial and adaxial (or only adaxial), papillate. Fruit usually a fleshy berry (or a pyrene), when ripe, reportedly black to purple to dark blue; but usually immature, and green, when collected, tapering abruptly into persistent style, seeds reniform (rarely hippocrepiform), (0–3–)25–60(–128) per locule, dark red (or brown or black), the testa reticulate (or areolate). A neotropical genus of 56 species, ranging through from southern México south and the West Indies through Central and South America to southern Bolivia, and east through the Guayana Highland; primarily montane.montane. Generic local name. " Ajicillo Venezuela, " ( Aristeguieta, 1973). Key to Species of Freziera 1a. Twigs (at least when young) with four paired wings decurrent from base of petiole and descending through two internodes, young twigs square or flat, only older twigs and branches rounded 2. 2 a. Lamina base conspicuously unequal 3. 3a. Lamina base unequal but symmetric, not cordate, auriculate, or clasping on one side 2. F. calophylla. 3b. Lamina base unequal, asymmetric, one side cordate, auriculate, and clasping 5. F. forerorum. 2b. Lamina base equal (or nearly so) 4. . 4a. Petiole (2.6–)3–5 cm long; seeds > 100 per locule 6. F. friedrichsthaliana. 4b. Petiole 0.8–2(–3.7) cm long, seeds fewer than 30 per locule 2. F. calophylla 1b. Twigs not conspicuously winged (although sometimes ridged below petioles), young twigs square, flat, or rounded 5. 5a. Lamina base deeply cordate and asymmetric with one auricle as long as petiole, the other longer and clasping or overlapping stem 4. F. cordata. 5 b. Lamina base truncate, rounded, cuneate, or very slightly cordate, symmetric or asymmetric 6. 6a. Upper lamina surface with at least sparse indumentum (sometimes only when leaf first unfolds, or only basally, or only on lateral veins; upper midrib surface with or without indumentum) 7. Indumentum 7a. Lamina margin deeply serrate, mostly with 1 or 2 smaller serrations between major teeth, the serrations somewhat hooked; apex very long acuminate 1. F. biserrata. 7b. Lamina margin merely crenate, serrulate, sinuous, or entire; lamina apex rounded, obtuse, acute, acuminate, or apiculate 8. 8a. Lamina margins entire or only barely serrate or sinuous, often narrowly revolute9. F. guatemalensis. 9. a. Indumentum rust-brown or orange-brown 8. F. guatemalensis. 9b. Indumentum beige or cream-colored 2. F. caloneura. 8b. Lamina margins prominently sinuous, undulate, serrulate, or if revolute, serrations prominently visible on undersurface 10. 10a. Leaf margins crenate to sinuous; fruits (even immature) > 1½ times longer than broad; all sepals sericeous with narrow glabrous margin 7. F. grisebachii. 10b. Leaf margins sinuous to finely serrulate; fruits (even immature) < 1½ times longer than broad; sepals glabrous, hirsute, or sericeous, in varying amounts but without narrow glabrous margin 2. F. calophylla. 6b. Upper lamina surface glabrous, even when leaf first unfolds (upper midrib surface and revolute auricles may have indumentum or not) 11. 11a. Leaves with persistent, dense indumentum entirely obscuring lower lamina surface (although many hairs wear off with age) 12. 12a. Indumentum rust-brown or orange-brown 8. F. guatemalensis. 12b. Indumentum golden to beige 3. F. candicans. 11b. Leaves glabrous to covered with indumentum below, hairs not obscuring undersurface 13. 13a. Midrib above entirely glabrous 14. 14a. Lamina base revolute, base thus appearing attenuate and tapered; petiole absent or short, nearly always < 1 cm long 3. F. candicans. 14b. Lamina base obtuse, rounded, truncate, attenuate, or slightly cordate, not revolute; petiole nearly always > 1 cm long 2. F. calophylla 13b. Midrib above with indumentum, at least basally 15. 15a. Stamens and staminodia 10–19(–24); ovules in carpellate flowers ca. 10–36, in staminate flowers 0–26 per locule; flowers 2–3.5(–5.1) mm in diameter 2. F. calophylla. 15b. Stamens and staminodia (19–)22–35; ovules in carpellate flowers ca. 44–120, in staminate flowers (18–)30–50 per locule; flowers 3.2–5.7 mm in diameter 16. 16a. Lamina base revolute, base thus appearing attenuate and tapered; petiole 0.3–1(–1.5) cm long 3. F. candicans. 16b. Lamina base flat, petiole (0.5–)1.5–3.5 cm long 7. F. grisebachii 1. Freziera biserrata A. Weitzman, ined. Figure 1. Treelet ca. 4 m tall; mature branches unknown; twigs terete, the bark dark red-brown, papillate when young, indumentum hirsute, the hairs straight, erect, ≲ 1.5 mm long, golden, the lenticels few, large, narrowly elliptic; terminal bud conduplicate-involute, ca. 4.8–6.7 cm long, densely hirsute. Leaves with petiole ca. (0.9–)1.7–2.2 cm long, canaliculate above, rounded below, hirsute, the wings involute, wide, deeply serrate with thick, conical or slightly curved, black setae terminating each tooth; colleters apparently absent; blade elliptic, ca. (6.5–)20.1–22.1 × (2.3–)6–6.4 cm, subcoriaceous, the base unequal, cuneate, the apex very long-acuminate, ultimately acute, the margin serrate, becoming revolute, mostly with one (or two) smaller serrations between major teeth, major teeth ca. 44–49 per side, with caducous, thick, curved, black setae terminating each tooth, the upper surface sericeous, hairs caducous, the lower surface densely hirsute, hairs persistent, the midrib above sunken, hirsute, hairs persistent, the midrib below rounded, densely hirsute, hairs persistent, the lateral veins ca. 11–12 per side, sunken above, rounded below. Inflorescence axis inconspicuous, hirsute, with flower buds one to three, pedicel scars absent or one to three and contiguous; bract triangular to narrowly triangular, ca. 2–3.5 × 1–1.5 mm, the apex acute, the margin entire, with several black setae, the outer surface densely hirsute; pedicel erect, terete, ca. 3.4–3.8 × 0.7–0.9 mm, densely hirsute; bracteoles triangular, unequal, one slightly keeled, ca. 3.5–4.5 × 3.4–4.4 mm, sclerotic centrally, the base clasping, the apex acuminate, with terminal, thick, conical, black seta, the margin chartaceous, entire, the outer surface densely hirsute. Flowers known only from bud, ca. 4.5 mm wide; sepals triangular to very broadly ovate to ovate, unequal, intergrading with petals, sclerotic centrally (innermost sometimes sclerotic in upper 2/3), the base clasping, the apex rounded (outermost sometimes acute with terminal seta), the margin chartaceous, usually minutely ciliolate, the outer surface strigose all over to centrally strigose (to sometimes glabrous on innermost); corolla apparently urceolate, the petals apparently distinct, ovate, nearly equal, membranaceous in lower 1/3, apex acute. Staminate buds with stamens ca. 20–22, free or slightly adnate basally, the filaments flat, unequal, the anthers basally cordate, the apiculus obtuse; gynoecium conical, 3-carpellate. Carpellate flowers unknown. Fruits unknown. Distribution. Freziera biserrata is known only from the type from a little-known area of Costa Rica near the north end of the border with Panamá, on the Gulf of México side. It was collected in elfin forest understory between 2400 and 2750 m. Type. Costa Rica. Limón: Cordillera de Talamanca, Atlantic slope, unnamed cordillera between Río Terbi and Río Siní, 9°00'–9°12' N, 82°58'–82°59' W, 2400–2750 m, 13 Sep 1984, G. Davidse, G. Herrera Ch., & M. Grayum 29054 ( holotype, MO; isotype, GH). Freziera biserrata differs from all other species of Freziera in its lamina margins with unusually long serrations with smaller ones between all of which are terminated by setae (rather than having them inserted in the sinuses); long, acuminate lamina tip; and acute or even acuminate triangular bracteoles. Some flower buds of F. biserrata have a terminal seta on the outer sepal. 2. Freziera calophylla Triana & Planchon, Ann. Sci. Nat. Bot. Sér. 4 , 18: 261. 1862. Figure 2. Eroteum calophyllum ( Triana & Planchon) O. Kuntze, Revis. gen. pl. 1: 62. 1891," Erotium". Eurya calophylla ( Triana & Planchon) Szyszylowicz in Engler & Prantl, Nat. Pflanzenfam. III, 6: 190. 1893. Eurya nitida Hieronymus, Bot. Jahrb. Syst. 20, Beiblatt 49: 50. 1895, nomen illegit. non Eurya nitida Korthals, 1842. [Type,.] Colombia Freziera subdimidiata Planchon ex Rusby, Bull. New York Bot. Gard. 4: 310. 1907. nomen. Freziera hieronymi Kobuski, Ann. Missouri Bot. Gard. 25: 355. 1937, "hieronyma", replacement name for Eurya nitida Hieronymus. Freziera cuatrecasasii Kobuski, J. Arnold Arbor. 34: 136. 1953. [Type, Colombia .] Freziera esmeraldana Little, Phytologia 18: 466. Fig. 14. 1969. [Type, Ecuador .] Tree – 10(–35) m tall; mature branches terete; twigs dorsiventrally flattened, with ridges or very narrow paired wings decurrent from base of petiole and descending through two internodes, the bark dark red-brown, papillate when young, striate and splitting with age, glabrous or indumentum sericeous, the hairs caducous (rarely persistent), golden, the lenticels few to many, very narrowly to widely elliptic, small to large; terminal bud conduplicate-involute, 1.5–5(–9) cm long, strigose to sericeous. Leaves with petiole (thin to) stout, 0.8–2.1(–3.7) cm long, canaliculate above, rounded below, glabrous to sericeous, the wings erect, slightly revolute or only slightly involute or slightly revolute, narrow; colleters one to several, but often obscured by indumentum; blade narrowly ovate to broadly ovate, (10–)12–20(–33) × (3–)5–10(–19) cm, coriaceous, the base subequal to very unequal (with one side of blade extending ≶ 2 cm below the other), cuneate to obtuse to rounded (to slightly cordate) and often attenuate at very base, sometimes slightly revolute, the apex acuminate to apiculate, ultimately acute or rounded, the margin sinuous, teeth 60–112 per side extending to base of blade, with caducous, conical or slightly curved, black setae often surrounded by tuft of hairs inserted in the sinuses, the upper surface glabrous or sericeous, hairs caducous, papillae scattered throughout (or absent or only near veins), the lower surface strigose to sericeous to hirsute, hairs persistent or caducous, papillae small, scattered throughout (associated with hair bases or rarely absent), the midrib above canaliculate with narrow central ridge (often flat or rounded near apex), glabrous to sericeous (often sericeous even when lamina glabrous), hairs caducous or persistent, papillae often in horizontal rows, the midrib below prominently rounded, strigose to sericeous, hairs persistent or caducous, papillae large, scattered, the lateral veins 15–27(–30) per side, sunken (or raised or flat) above, prominently rounded below. Inflorescence axis ≶ 7 mm long, with flowers one to seven, pedicel scars absent or one to ten and contiguous; bract triangular to narrowly triangular, 1.7–3.8 × 1.1–1.3 mm, keeled, the apex acute, the margin entire, sometimes with several black setae, the outer surface sericeous (rarely glabrous); pedicel erect, terete, 1.7–4.3 × ca. 1 mm, ridged, sericeous; bracteoles apical, opposite or subopposite, broadly to very broadly ovate, equal or subequal, 1.5–2.8 × 2–2.8 and 2.1–2.8 × 2.5–2.9 mm, sclerotic centrally (one sometimes keeled), the base clasping, the apex rounded (rarely apiculate), the margin chartaceous, often ciliolate (rarely with basal conical, dark setae), the outer surface sparsely strigose or sericeous (at least centrally). Flowers 5.4–6.8 × 3–3.5(–5.1) mm; sepals broadly ovate, nearly equal, 2.3–4 × 2.4–3.1 mm, sclerotic centrally, the base cordate, the apex rounded, the margin membranaceous, minutely ciliolate, the outer surface glabrous (especially inner sepals) to sericeous (outer sepals most often centrally sericeous); petals distinct, ovate, nearly equal, 4.1–5.3 × 2.1–3.4 mm, membranaceous in lower ¼, apex acute. Staminate flowers with stamens 16–24, free or slightly adnate basally, unequal, the filaments flattened basally, terete above, slightly bent or straight, unequal, ca. 0.4–0.6 and 1.2–1.5 mm long, the anthers equal or unequal, 0.8–1.8 mm long, basally cordate, the apiculus ovate, ca. 0.1 mm long, obtuse apically; gynoecium conical (to broadly conical), 2.5–3.3 × 1.3–2.3 mm, 3- or 5-carpellate, with locules ca. 1 mm long, each containing ca. 0–6(–35) ovules, the stigmatic lobes erect, 0.3–0.5 mm long. Carpellate flowers with staminodia 13–19, free or adnate basally, nearly equal, 0.7–1.8 mm long, acute apically; gynoecium pyriform, 3–3.8 × 1.7–2.1 mm; 3- or 5-carpellate, with locules 1–1.7 mm long, each containing 20–30 ovules, the stigmatic lobes flaring, 0.5–0.7 mm long. Fruits (probably immature) ovoid, ca. 6–7 × 4.5–5.6 mm; seeds (4–)13–28 per locule, 0.6–0.8(–1.2) mm long, dark red or brown. Distribution. Freziera calophylla is a widespread species, known from eastern Panamá, western Colombia, southeastern Venezuela, northern Ecuador, and southern Peru. It is known from wet forests at a wide variety of elevations between 5 and 2000 m, mostly between 1200 and 2000 m. Type. Colombia. Antioquia: San Julián, 1800 m, 1851–1857, J. Triana 3248 (holotype, P (photos at F, MO, US; negative at F no. 35314); isotypes, BM (6 Feb 1852, 1827 m), COL; probable isotype, K (Triana s.n. n.v.)). Additional specimens examined. PANAMÁ. Cocle: La Mesa, 4 km N of El Valle, 850–875 m, Nee & Dwyer 9205 (MO, TENN, US). Darién: Pacific Coast, 5 km NW of Cocalito, on "tongue" of land extending into ocean, caused by 1976 earthquake, Garwood 745 (MO), Garwood 766 (F); Tongue-like landslide protruding into Pacific Ocean and adjacent valleys, 23 km SE of Jaqué, Garwood 929 (MO); Cerro Mali base camp, Colombian border, Cerro Tacarcuna, Serranía del Darién, 1400 m, Gentry & Mori 13756 (COL, MO); Cerro Tacaruna, W. ridge, trail toward Río Pucuru just below summit camp, 1500–1600 m, Gentry & Mori 14141 (COL, MEXU, MO (2 sheets), US); summit of Pico Tacarcuna, highest point of the Serranía del Darién, 1850 m, Gentry et al. 16907 (COL, MEXU, MO (2 sheets)). Panamá: Cerro Jefe, Valdespino 52 (F, MO, NY). Local Name. Colombia: "Caimito" ( Uribe & Uribe U., 1940). Freziera calophylla has been collected in bloom and fruit in July through March. It is morphologically very *heterogeneous and rather hard to characterize. All specimens share a conspicuously canaliculate, wide midrib. In addition most specimens have erect (or revolute or only slightly involute) petiole wings, small flowers, and large coriaceous laminas with regular, nearly straight lateral veins. The most conspicuous variation in F. calophylla is in petiole length, lamina size, and indumentum. Corresponding with this large range of variation, are large geographical and altitudinal ranges, although the morphological variation does not appear to correlate with either geography or altitude.This variation has resulted in four taxa recognized by previous TaxonAuthors. Given the number of new taxa described in this treatment, this may seem unusual, but there is no discrete way that I have found to define those four taxa. Although Cuatrecasas 23985 ( F. cuatrecasasii ) has some very large leaves (≶ ca. 35 cm long), the several sheets are fairly variable in lamina size, and it has the other characters listed above. Similar circumstances can be ascribed to the other taxa ( F. hieronymi and F. esmeraldana ), and to several other specimens which are somewhat different from the "norm" in this species (e.g., Nee & Dwyer 9205, Juncosa 1775, and Silverstone-Sopkin et al. 1677 and see lamina outlines in Fig. 76). Although stamen number is usually rather low (13–19) as is ovule number, especially in staminate flowers (ca. zero to six), Asplund 8543 is unusual in having a relatively high number of stamens (24) and a relatively high number of ovules (ca. 35 per locule). The outlying collections from the Venezuelan Guayana and Peru match particularly well with the type of F. calophylla, so despite their geographic distance from the center of distribution of this species, morphologically they fit well in the species. Freziera calophylla sometimes has narrow wings on the young twigs. Most other species which have this character are listed and compared in Table VIII. Those species generally have wider wings than F. calophylla, and except for F. angulosa (and unlike F. calophylla) they are believed to have unilacunar nodes. Several of those species, F. uniauriculata, F. forerorum, and F. angulosa, have unequal and lamina bases, but unlike the specimens of F. calophylla which have unequal lamina bases, those three species have very asymmetrical lamina bases which clasp the stem on one side. The other species have symmetric lamina bases, but those differ from symmetric F. calophylla in conspicuously revolute lamina bases. Freziera verrucosa and F. nervosa are often confused with F. calophylla, apparently due to the lack of obvious distinguishing features in all three. This problem is especially apparent in Colombia, where the three species overlap. F. verrucosa, which has the largest flowers of the three species, usually has revolute lamina margins, and nearly always has long, thin, flat, and twisted hairs. In contrast F. nervosa and F. calophylla have small flowers, conspicuously sinuous lamina margins, and shorter, more robust, terete hairs. Freziera nervosa has membranaceous to subcoriaceous laminas, while F. verrucosa and F. calophylla have coriaceous laminas. Freziera calophylla has midribs that are wide and canaliculate and erect petiole wings, while the other two species have narrow midribs which are sunken or rounded, and involute petiole wings. The variation within and between these species is not sufficiently well known. It appears that field work is necessary to help sort out this variation. 3. Taxon Not Tagged 4. Taxon Not Tagged 5. Taxon Not Tagged 6. F. friedrichsthaliana. (Taxon Not Tagged> 7. Taxon Not Tagged 8. Taxon Not Tagged 9. Taxon Not Tagged Excluded taxa cited in Biologia Centrali-Americana Freziera undulata (Swartz) Willdenow, Sp. Pl. 2(2): 1179. 1799; Swartz, Fl. ind. occ. 2: 974. 1800. Eroteum undulatum Swartz, Prodr. 85. 1788; [Lectotype. St. Kitts]. Freziera hirsuta J. E. Smith in Rees, Cycl. 15: unpaginated. 1810; [Type: St. Vincent]. Ternstroemia salicifolia A. P. de Candolle, Mém. Soc. Phys. Genève 1: 411. 1822; [Type: Guadeloupe]. [Ternstroemia dentata K. Sprengel ex A. P. de Candolle, Mém. Soc. Phys. Genève 1: 411. 1822. nom. in syn., non Ternstroemia dentata (Aublet) Swartz, Prodr. 81. 1788. ] Freziera perrottetiana Tulasne, Ann. Sci. Nat. Bot. Sér. 3, 8: 332. 1847; [Type: Guadeloupe]. Freziera elegans Tulasne, Ann. Sci. Nat. Bot. Sér. 3, 8: 336. 1847; [Syntypes: Guadeloupe]. Cleyera elegans (Tulasne) Choisy, Mém. Soc. Phys. Genève 14: 110. 1855. Freziera salicifolia (A. P. de Candolle) Choisy, Mém. Soc. Phys. Genève 14: 122. 1855. Eurya undulata (Swartz) Blume, Ann. Mus. Bot. Lugduno-Batavum 2: 105. 1856. Freziera salicifolia (A. P. de Candolle) Choisy var. undulata (Swartz) Wawra, in Martius, Fl. bras. 12(1): 283. 1886. Eroteum hirsutum (J. E. Smith) Gómez de la Maza, Anal. Soc. Esp. Hist. Nat. 19: 222. 1890, excl. spec. cit. Freziera undulata (Swartz) Willdenow var. hirsuta (J. E. Smith) Krug & Urban, Bot. Jahrb. Syst. 21: 543. 1896. Freziera undulata (Swartz) Willdenow var. elegans (Tulasne) Krug & Urban, Bot. Jahrb. Syst. 21: 543. 1896.