## HINTONIA

Bullock (1935) separated *Hintonia* from *Coutarea* because of its cylindrical capsules (vs. laterally compressed in *Coutarea*) and basipetal-imbricate seeds (vs. vertical in *Coutarea*). He dedicated the genus name to George B. Hinton (1882–1943), a London-born metallurgist and avid plant collector based in southern Mexico. The taxonomic placement of *Hintonia* within the family has differed according to various authors. Aiello (1979) treated *Coutarea* and *Hintonia* as members of the *Portlandia* complex. Robbrecht (1988, 1994) removed *Hintonia* from Condamineae and listed it among the *genera incertae sedis*. Andersson and Persson (1991), according their phylogenetic study using morphological data, removed *Coutarea* and *Exostema* from Cinchoneae, and returned them to Condamineae. Bremer (1992) and Delprete (1996b) in their morphology-based phylogenies, retrieved *Coutarea* and *Hintonia* as sister genera, closely related to *Portlandia*. Bremer (1992) placed all three genera in the amended Chiococceae. Ochoterena-Booth (2000) revised *Hintonia* in an unpublished doctoral dissertation, maintained the distinction between *Hintonia* and *Coutarea*, and their inclusion in the *Portlandia* group sensu Delprete (1996b). Paudyal et al. (2018) were able to include only *Hintonia latiflora* Bullock and *H. octomera* Bullock in their molecular phylogenetic study, and confirmed that the genus is monophyletic, and was positioned as a sister taxon to the clade including *Coutarea* and the two South American species of *Exostema* (which were transferred to other genera).

*Hintonia* is a genus of three species occurring in Mexico and Central America of shrubs or trees to 10 m tall, with axillary uniflorous inflorescences, 6–8-merous flowers, fragrant from the afternoon to the evening, stigmatic surface as two lines along the style, capsules ellipsoid to subspherical, round in cross-section, and seeds acropetally or centripetally aligned (Ochoterena-Booth, 2012b).

**Hintonia** Bullock, in Hook. Ic. Pl. 33, tab. 3295. 1935. Ochoterena-Booth, Fl. Mesoamericana 4.2:118–119. 2012; Borhidi et al., Rubiáceas México 263–267, f. 54. 2006; Borhidi et al., Rubiáceas México, 2nd ed. 282–286, f. 62. 2012; Paudyal et al., Bot. J. Linn. Soc. 187(3):365–396. 2018. TYPE: *Hintonia latiflora* (DC.) Bullock

1. **Hintonia latiflora** (DC.) Bullock, in Hooker's Icon. Pl. 33(4):tab. 3295, p. 4. 1935. *Coutarea latiflora* DC., Prodr. 4:350. 1830. TYPE: [MEXICO]. [icon.]: Torner Collection Plate No. 0856, preserved in the Hunt Institute for Botanical Documentation (LECTOTYPE (McVaugh 2000:463)).

**Distribution.**—Mexico, Guatemala, and El Salvador.

**Notes.**—Augustin Pyramus de Candolle (1830:350) in the protologue of *Coutarea latiflora* DC., stated "C. latiflora (fl. mex. ic. ined.) pedicellis I-floris, ubi dicitur *Copalchi*." Therefore, the authority of this name should be attributed solely to him. There is no original specimen of *C. latiflora* in G or G-DC. In MA, there are two specimens in the Sessé & Mociño Herbarium, one with barcode MA605137, *Sessé & Mociño 5470*, which has a label with the handwritten annotation "Portlandia grandiflora. N° 135," and the other with barcode MA605138, *Sessé & Mociño 1620*, which has a label with the handwritten annotation "Portlandia hexandra. N° 474." The two specimens in MA are not original material, because they were not examined by Candolle.

The history of the Sessé and Mociño Expedition (1781–1803), itinerary, herbarium, illustrations, and impact in the botanical world, was addressed in great detail by McVaugh (1969, 1972, 1977, 1980, 1982, 1987, 1990, 1998, 2000), and is here summarized. The King Charles III of Spain, established the Royal Botanical Expedition to New Spain (*Expedición Botánica al Virreinato de Nueva España*), headed by the Spanish doctor Martín Sessé y Lacasta (1751–1808). The team of botanists that took part in the expedition included the Mexican-born Doctor José Mariano Mociño Suárez Lozano (1757–1820), who later became Sessé's most important collaborator. During the expedition, aside from the botanical collections, a team of artists made about 2000 paintings on the spot, using living material. After Sessé and Mociño return to Spain, Mociño carried all those paintings to Montpellier, to be examined by A.P. de Candolle. Candolle studied them in great detail and travelled with Mociño and those paintings to Geneva, and used them to describe hundreds of new species. Learning about the imminent departure of Mociño to return to Spain, Candolle hired several local artists to make copies of those paintings, which are now kept in G-DC. Candolle organized the copies of the original paintings, numbered them, and provided a handwritten index.
Mociño left Montpellier with the bulk of about 2000 original paintings and arrived at Barcelona, where he died shortly after, in 1820. The paintings disappeared for a long time, and were eventually found in 1979 in a private library in Barcelona. They were acquired in 1981 by the Hunt Institute for Botanical Documentation, in Pittsburgh, Pennsylvania (McVaugh, 1982), where they comprise the Torner Collection of Sessé & Mociño Biological Illustration (https://huntbot.org/torner/). When the paintings arrived at the Hunt Institute, they were bound in volumes. They were separated as single pages at the Hunt Institute, without particular systematic arrangement, and to each painting was assigned a "Torner Number." The paintings of the Torner collection represent the original material of the new taxa described by Candolle, as they were studied and annotated by him in Montpellier during 1813–1816, and in Geneva in 1816–1817.

McVaugh (2000) explained in detail the typification of the names published by Candolle, using the original illustrations made during the Sessé and Mociño expedition, which are now part of the Torner Collection. McVaugh (2000:463) designated the lectotype of *Coutarea latiflora* DC. as follows, "Lectotype [DC.]: No. 0856 of the Torner Collection, annotated by de Candolle "Coutarea latiflora" and "colpachi." DC. plate 458, as cited in Calques des Dessins (Field Mus. neg. 30687), is an original painting, nearly identical with Torner 0856, annotated "Portlandia hexandra" and "colpachi"."

Ochterena-Booth (2012:119) cited the type of "Coutarea latiflora Sessé et Moc. ex DC." as "Holotipo: México, Michoacán, Sessé y Mociño s.n. (G-DC!)." However, as explained above, the authority of this name should be attributed solely to Candolle, and there is no original specimen associated with this name in G-DC.

*Exostema canescens* Bartl. ex DC., Prodr. 4:359. 1830 (as "Exostemma"). *Cinchona canescens* (Bartl. ex DC.) Brign., Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. II, 1:63. 1862. Type: MEXICO: without locality, 1791, *Haenke s.n.* (LECTOTYPE, here designated: PR [No. 612234]; isolectotypes: G-DC [G00665709], GOET [GOET010233], HAL [HAL0114173], PR [No. 612235]).

**Notes.**—Candolle (1830:359) attributed the authority of *Exostema canescens* to Bartling, as "E. canescens (Bartl! in h. Haenk.), and described it as being villous-canescent, with axillary inflorescences, and pedicels as long as the capsules, and capsules not crowned by the calyx. He also stated that it was collected by Haenke in Mexico, and "v. sine fl. in h. Haenke." The last statement means that he saw a specimen in the Haenke Herbarium. According to Stafleu and Cowan (1979:6), Thaddeus Haenke's herbarium is in PR.

In PR there are two original specimens associated with this name. Specimen with accession number 612234 has three labels. At the bottom of the sheet there is a label with heading "Herbarium Generale Musei Nationalis Pragae" and the stamps "1791" and "Thaddeus Haenke." On this label is glued a smaller label with the handwritten annotation "Mexico Rubiacea." Above the bottom label is glued another label with the annotation "Exostemma canescens Bartl." handwritten by an unknown author. A third label has the handwritten annotation "Mexico Rubiacea." The specimen consists a branch with lateral branches with several leaves and several capsules, some of them still closed, and some of them dehiscend. This specimen, collected in Mexico by Haenke in 1791, with accession number 612234, is here designated the lectotype of *Exostema canescens*.

The PR specimen with accession number 612235 has two labels. At the bottom of the sheet there is a label with heading "Herbarium Generale Musei Nationalis Pragae" and the stamps "1791" and "Thaddeus Haenke." On this label is glued a smaller label with the handwritten annotation "Mexico Rubiacea." Above that label, is affixed another label with the annotation "Peruanae montanae Rubiacea" handwritten by Sternberg. The second label is certainly an error, because the specimen is from Mexico, and not from Peru. The specimen consists of a small branch with two leaves and a dehiscend capsule. This specimen is an isotype of *Exostema canescens*.

In G-DC there is a sheet, with barcode G00665709, which has a label with the annotation "Exost. canescens Mexico Hb. Haenke." The specimen consist of just a few fragments contained in an envelope. Those fragments were extracted by Candolle from a specimen in the Haenke herbarium, which was examined by him. This G-DC specimen is an isotype.

A specimen in GOET, barcode GOET010233, has two labels with the annotations "I. Exostemma canescens Bartl. Mexico" and "Exostemma canescens Bartl. In Mexico leg. Haenke" handwritten by unknown authors. On the sheet there is a third label with the annotation "possibly Coutarea sp." by Tim McDowell. The specimen consists of a small branch with a few leaves, and fragments of leaves and capsules included in an envelope. This specimen is an isotype.

In HAL there is a specimen, with barcode HAL0114173, which a label with the annotations "Exostemma canescens Bartl. Mexicis Haenke" handwritten by two unknown authors. The specimen consists of a small branch with a few leaves, and capsule fragments included in the attached envelope. This specimen is an isotype.

Standley (1921:126) regarding this taxon wrote, "Described from Mexico; inflorescence axillary, 1-flowered; leaves pubescent. Probably not of this genus." Because of the peculiar pubescence of the vegetative parts, and the impression on the capsule internal walls, indicating the horizontal position of the seeds, this taxon is a synonym of *Hintonia latiflora* (DC.) Bullock.

*Portlandia pterosperma* S. Watson, Proc. Amer. Acad. Arts 24:52. 1889. *Coutarea pterosperma* (S. Wats.) Standl., N. Amer. Fl. 32:127. 1921. TYPE: MEXICO. SONORA: In deep canyons near Guaymas, 1887, *E. Palmer* 298 (HOLOTYPE: BM n.v.; ISOTYPES: C [without barcode],
E [E00285404], GH, K [K000173483], MEXU [MEXU 00031099], NDG [NDG50849], NY [02684370], US [4 sheets, No. 1390863 (without barcode), No. 1390864 (barcode 01013598), No. 47978 (barcode 00137335), No. 47979 (barcode 00137334)], YU [YU.001750]).

*Hintonia latiflora* var. *leiantha* Bullock in Hooker's Icon. Pl. 33(4):1, 5, tab. 3295. 1935. **TYPE:** MEXICO. **MEXICO:** Dist. of Temascaltepec, Ixtapan, 1000 m, 5 Aug 1932, G.B. Hinton et al. 1258 (**HOLOTYPE:** K n.v.; **ISOTYPES:** A [01154801], BM n.v., G [G00436275], RSA [RSA0005799]).

*Hintonia standleyana* Bullock, in Hooker's Icon. Pl. 33(4):6, tab. 3295. 1935. *nom. inval.* **TYPE:** Based on the description of *Coutarea latiflora* in Standley, N. Amer. Fl. 32:127. 1921, and Contr. U.S. Natl. Herb. 23:167. 1926.

**2. Hintonia lumana** (Baill.) Bullock, in Hooker's Icon. Pl. 33 (4):5, tab. 3295. 1935 (as "*Lumaeana*"). *Coutarea lumana* Baill., Adansonia 12:301. 1879 (as "*Lumaeana*" but honoring M.G. Luma). **TYPE:** GUATEMALA: without locality, s.d., G. Luma (or *Luma*) s.n. (**HOLOTYPE:** P [P00559099]).

**Distribution.**—Mexico, Guatemala, El Salvador, and Costa Rica.

**Notes.**—Baillon (1879:301) discussed *Coutarea lumana* Baill. as followed “Léspece du Guatemala m’a été remise, il y a deux ans, par un jeune médecin du pays, M.G. Luma, d’où le nom de C. Lumeana que je lui ai donné.” [The species from Guatemala was given to me, two years ago, by a young doctor, M.G. Luma, hence I gave it the name C. Lumeana]. According to the *Code* (Turland et al. 2018), because the specific epithet is dedicated to Luma, it should be corrected as *lumana*. A specimen in P, barcode P00559099, has a label with the annotations “Portlandia! *Coutarea* Lumaeana H. Bl. Sect. Pacourea (capsula septicida!). Guatemala. M.G. Luma (1878)” handwritten by Baillon. The annotation was corrected by an anonymous author, who stroked the letter “m” and substituted it with “n,” with the intention to correct the last name of the collector as Luna. On the bottom center of the sheet is affixed another label with the annotation “A Monsieur Baillon. Professeur de Botanique à l'Ecole de Medecine de la part de George Luna de Guatemala” by an unknown author. Therefore, the doubt as whether the last name of the collector is Luma or Luna, remains. The specimen consists of two small branches with flowers in anthesis, and a loose, wrinkled-up corolla. This specimen is the holotype.

**3. Hintonia octomera** (Hemsl.) Bullock, in Hooker's Icon. Pl. 33(4):6, tab. 3295. 1935. *Coutarea octomera* Hemsl., Biol. Centr. Amer. 4:101. 1886. **TYPE:** MEXICO. **YUCATÁN:** Cozumel Island, 8 Apr 1885, G.F. Gaumer 148 (**HOLOTYPE:** K n.v.).

**Distribution.**—Southern Mexico and Guatemala.

*Coutarea acamptoclada* B.L. Rob. & Millsp., Bot. Jarhb. Syst. 36 (Beibl. 80):28. 1905. **TYPE:** MEXICO. **YUCATÁN:** “im Walde bei Umán, 7 Apr 1902, C. Seler & E. Seler 3944** (**LECTOTYPE** (Lorence 1999:44): F n.v. (photo in PTGB)).

## HINTONIA EXCLUDED TAXA

*Hintonia pulchra* D.R. Simpson, Phytologia 29:277. 1974.
= *Osa pulchra* (D.R. Simpson) Aiello (See below).

## ISIDOREA

Achille Richard completed a monograph of the family Rubiaceae, and gave a manuscript to A.P. de Candolle, which was supposed to be published soon, but it was instead published in December 1830. The volume of the *Prodromus* where Candolle treated the Rubiaceae was published in September 1830, preceding the publication of Richard's Rubiaceae monograph by three months. Richard (December 1830) included *Isidorea* in the tribe Cinchoneae, and explained that he dedicated the genus name to the zoologist Isidore Geoffroy Saint Hilaire (1805–1861, son of Étienne Geoffroy Saint-Hilaire). Candolle (September 1830:405) described the genus *Isidorea* A. Rich. ex DC., with the sole species *Isidorea amoena* A. Rich. ex DC., Prodr. 4:406. 1830, which is a superfluous name, because he cited *Ernodea pedunculata* Poir. and *Ernodea pungens* Lam. in synonymy. The epithet “pungens” was available for this species, which is what Candolle should have used to make a new combination. Robinson (1910:401) published the combination *Isidorea pungens* (Lam.) B.L. Rob. Urban (1923a) published the descriptions of all the species of *Isidorea* known to him. Aiello (1979) transferred three species from *Portlandia* to *Isidorea*, without any discussion about the species previously included in *Isidorea*, and esteemed that the genus has 11 species. Borhidi (1981), and Aiello and Borhidi (1986) described several
new species in this genus. Liogier (1995) estimated *Isidorea* to have approximately 20 species occurring in Cuba and Española, and produced a key to the Españolan species. In Paudyal et al.'s (2018) molecular phylog- enies, *Isidorea* was confirmed to be monophyletic, and retrieved in a clade sister to the *Portlandia* clade. The *Isidorea* clade was divided into two subclades, one with the species from Cuba, and the other with species from the Dominican Republic. *Isidorea* differs from *Portlandia* in having stiff, pungent, coriaceous leaves and stipules divided at the base into two units, looking like four, apically pungent stipules per node, and smaller flowers. In the present treatment, 13 species are recognized in this genus.

**Isidorea** A. Rich ex DC., Prodr. 4:405. 1830; A. Richard, Mém. Rubiacées 284. 1830; A. Richard, Mem. Soc. Hist. Nat. Paris 5:284, tab. 25, f. 1. 1834; Standley, N. Amer. Fl. 32(1):14–15. 1918; Liogier, Fl. Cuba 5:28–29. 1962; Borhidi, Acta Bot. Hung. 29:181–216. 1981 [“1980”; Liogier, Fl. Española 7:311, 313– 318, f. 198–27. 1995; Borhidi et al., Rubiáceas Cuba 163–168, f. 45. 2017; Paudyal et al., Bot. J. Linn. Soc. 187(3):365–396. 2018. TYPE: *Isidorea amoena* A. Rich ex DC., Prodr. 4:405. 1830, nom. superfl. [= *Isidorea pungens* (Lam.) B.L. Rob.]

**1. Isidorea brachyantha** Urb., Ark. Bot. 17(7):57. 1922. TYPE: HAITI. DEPART. DU SUD: prope Port à Piment in collibus siccis calcareis rara, [27 Jul 1917], E.L. Ekman 401 (NEOTYPE, here designated: S [No. S-07-14592]; ISONEOTYPE: S [S-R-3048]).

*Distribution*.—Dominican Republic, Haiti.

**Notes.**—Urban (1922:57), in the protologue of *Isidorea brachyantha* Urb., cited the material studied as “Départ. du Sud prope Port à Piment in collibus siccis rara, m. Jul. flor.: [Ekman] n. 401.” The original material in B studied by Urban was destroyed during WWII. In S there are two specimens of *Ekman 401*. The specimen with accession number S-07-14592 has a label with the heading “Mus. Botan. Stockholm” and the annotation “*Isidorea brachyantha* Urb. Typus. det. I. Urban” handwritten by an unknown author (not Urban). This specimen consists of three branches with numerous leaves and several capsules, and is here designated as the neotype of *I. brachyantha*.

The S specimen with accession number S-R-3048 has a label with the heading “Mus. Botan. Stockholm” and the annotation “*Isidorea brachyantha* Urb. Typus. det. I. Urban” handwritten by an unknown author (not Urban). This specimen consists of two branches with numerous leaves and several capsules, and is an isotype.

**2. Isidorea brachycarpa** (Urb.) Aiello, J. Arnold Arbor. 60:114. 1979. *Portlandia brachycarpa* Urb., Symb. Antill. 9:135. 1923. TYPE: CUBA. GUANTÁNAMO: Baracoa, ad Taco Bay, 5 Dec 1914, E.L. Ekman 3758 (LECTOTYPE (Aiello 1979:114): S [No. S-R-5213]; ISOLECTOTYPES: F [No. 635301 (frag. ex G)], G [G00436336], UPS [No. V-084588]).

*Distribution*.—Cuba (Guantánamo: Bahia de Taco, Costa Sur-Maisí).

**Notes.**—Urban (1923a:135–136) in the protologue of *Portlandia brachycarpa* Urb., cited the material studied as “Prov. Oriente prope Baracoa ad Taco Bay, in rupibus calcareis rara, m. Dec. flor. et fruct.: [Ekman] n. 3758.” The original material in B studied by Urban was destroyed during WWII. Aiello (1979:114) designated the lectotype of this name a specimen at S. The specimen with accession number S-R-5213 has a label with the annotation “*Portlandia brachycarpa* Urb. (typus)” handwritten by Urban. It consists of five branch tips with numerous leaves and dehisced capsules, and is the specimen designated as the lectotype by Aiello.

*Isidorea leonis* Alain, Contr. Ocas. Mus. Hist. Nat. La Salle 17:5. 1959, **syn. nov.** TYPE: CUBA. GUANTÁNAMO: Maisí, coastal thickets, Jan 1940, *Bro. León & P. Matos* LS No. 17945 (HOLOTYPE: HAC [Specimen A, ex LS]; ISOTYPES: HAC [Specimen B, ex LS], NY [00115159], US [00026703]).

**Notes.**—In HAC there two specimens of *Bro. León & P. Matos* LS No. 17945, which where originally part of the LS herbarium, which was later integrated in HAC. These two specimens are identified by the letters “A” and “B” handwritten on the sheets to distinguish them from each other. Specimen with the letter “A” has a label with the annotation “*Isidorea leonis* Alain” handwritten by Alain Liogier, and is the holotype. Specimen with the letter “B” has the typewritten annotation “*Isidorea leonis* Alain” and it has is no evidence that was seen by Alain Liogier.

**3. Isidorea elliptica** Alain, Contr. Ocas. Mus. Hist. Nat. Colegio De La Salle 17:4. 1959. TYPE: CUBA. ORIENTE: near Macambo, Vía Azul, dry coastal thickets, 28 Dec 1954, Hno. Alain [A.H. Liogier] & M. López Figueiras 4198 (HOLOTYPE: HAC [ex LS]; ISOTYPES: BP n.v., HAJB [HAJB G 000520], NY [00115157]).

*Distribution*.—Cuba (Guantánamo: Macambo, San Antonio del Sur, Abra Mariana, Cajobabo-Jauco).
*Isidorea rheidioides* Borhidi, Abstr. Bot. Budapest 5:37. 1977, **syn. nov.** TYPE: CUBA. ORIENTE: in fruticetis litoralibus calcareis inter Cajobabo et Jauco, 31 Dec 1959, *Bro. Alain* [*A. Liogier* & *López Figueiras* 7097 (HOLOTYPE: HAC [ex LS]; ISOTYPES: HAJB [3 sheets, HAJB G 000518, HAJB G 000989, HAJB G 000990]).

*Isidorea microphylla* Borhidi, Acta Bot. Acad. Sci. Hung. 26:268, fig. 4. 1981 ["1980"], **syn. nov.** TYPE: CUBA. GUANTÁNAMO: San Antonio del Sur, 4 km NW del pueblo (Abra Mariana), 200-400 m, 10 Feb 1976, A. *Areces*, J. *Bisse*, J. *Gutiérrez* & H. *Manitz* HAJB No. 29924 [HAC No. 28363] (HOLOTYPE: HAJB [HAJB G 000519]; ISOTYPES: B [B 10 0385011], BP n.v., HAC [No. 28363], JE [JE00004995]).

4. **Isidorea gonavensis** Aiello & Borhidi, Acta Bot. Hung. 32:221. 1986. TYPE: HAITI. ÎLE DE GONÂVE: Pte.-à-Raquettes, Marne Fort-Fiel, 24 Jul 1927, E.L. *Ekman* H-8664 (HOLOTYPE: S [No. S14-28111]; ISOTYPES: G [G00436335], GH [00058986], LL [00000280]).

**Distribution.**—Haiti (Gonâve Island).

5. **Isidorea leonardi** Urb., Repert. Spec. Nov. Regni Veg. 19:8. 1923. TYPE: HAITI: vicinity of Fond Parisien, Étang Saumâtre, upright shrub 4 to 5 ft high, scarce, flowers cream-white, very fragrant, 5–13 May 1920, *E.C. Leonard* 4041 (LECTOTYPE, here designated: US [00026702]; ISOLECTOTYPES: BM [000081661 (specimen "a")], GH [00058985], NY [00115158]).

**Distribution.**—Dominican Republic, Haiti.

**Notes.**—Urban (1923c:8), in the protologue of *Isidorea leonardi* Urb., cited the material studied as "Vicinity of Fond Parisien, Etang Saumatre, Leonard 4041" without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. The US specimen with barcode 00026702, has a label the annotation "Isidorea" handwritten by an unknown author, and "Leonardii Urb. (typus)" handwritten by Urban. The specimen consists of two branches with numerous leaves and several flowers in anthesis, and two separate portions of a stem. This specimen is here designated as the lectotype of *I. leonardi*.

6. **Isidorea leptantha** Urb., Symb. Antill. 7:391. 1912. TYPE: DOMINICAN REPUBLIC. BARAHONA: Barahona, 200 m, Aug 1910, M.D. *Fuertes* [Pater *Fuertes*] 634 (LECTOTYPE, here designated: BR [000000531699]; ISOLECTOTYPES: E [E00505337], F [Nos. 768324 (ex G-DC), 635309 (frag. ex G)], NY [00115160], P [P00559098], US [00026704]).

**Distribution.**—Dominican Republic.

**Notes.**—Urban (1912:391-392), in the protologue of *Isidorea leptantha* Urb., cited the material studied as "In Santo Domingo prope Barahona 200 m., m. aug. flor. et fruct.: Fuertes n. 634" without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. The BR specimen with barcode 000000531699 has a label with the heading "H. von Türkheim: Plantae Dominicensis—Herb. Krug et Urban." On the label there is the annotation "634. Isidorea (n. sp?) handwritten by Pater Fuertes, and "leptantha Urb. n. sp." handwritten by Urban. The specimen consists of two branches with numerous leaves and several flowers in anthesis. In the envelope affixed on the sheet is included one dehisced capsule. This specimen is here designated as the lectotype of *I. leptantha*.

7. **Isidorea oblanceolata** (Urb.) Aiello, J. Arnold Arbor. 60:115. 1979. *Portlandia oblanceolata* Urb., Symb. Antill. 9:136. 1923. TYPE: CUBA. ORIENTE: Sierra de Nipe, ad viam Bio dictam, 27 Apr 1919, E.L. *Ekman* 9588 (LECTOTYPE (Aiello 1979:114): S [No. S-R-5216]; ISOLECTOTYPES: F [No. 604819], NY [00126758 (one leaf)]).

**Distribution.**—Cuba (Holguín: Sierra de Nipe).

**Notes.**—Urban (1923a:136-137), in the protologue of *Portlandia oblanceolata* Urb., cited the material studied as "Prov. Oriente in Sierra de Nipe ad viam Bio dictam in charrascales, m. April. fruct.: [Ekman] 9588" without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. The specimen in S with accession number S-R-5216 has a label with the annotation "Portlandia oblanceolata Urb. (typus)" handwritten by Urban. It consists of three branches with numerous narrowly oblanceolate leaves and a few capsules. This specimen was designated as the lectotype of this name by Aiello (1979:114).

8. **Isidorea ophiticola** (Borhidi) Borhidi & Moncada, Acta Bot. Acad. Sci. Hung. 26:267, fig. 2 (photo of holotype). 1981 ["1980"). *Portlandia ophiticola* Borhidi, Növennyrendsz. Novényföldr Tanz., Eötvös Lórand Tudományegyet. Budapest 5:34. 1977. TYPE: CUBA. ORIENTE [HOLGUÍN]: Sierra de Nipe, charrascal de la Cueva, Pinares de Mayarí, 27-31 May 1960, M. López Figueiras 1037 (HOLOTYPE: HAC [ex SV]; ISOTYPE: HAJB [HAJB G 000797, ex SV]).

**Distribution.**—Cuba (Holguín: Sierra de Nipe).
complete description of the species, handwritten by him. The specimen consists of two branches, one with an infructescence, and one with an inflorescence with flower buds. It is here designated as the lectotype of this name.

In P-Bonpl., there is another original specimen of *Exostema peruvianum*, with barcode P00671150. The specimen consists of one branch with numerous leaves and an infructescence with dehisced capsules, and two branchlets with a few leaves. This specimen is an isotype of this name.

In B-W there is a specimen, with barcode B-W 04027-01 0, which has a label with the annotation "Exostema Peruviana Pl. Eq. [...] (Humboldt)." On the bottom right corner of the sheet there is the annotation "Humboldt. W" handwritten by Willdenow. The specimen consists of a branch with several leaves, an inflorescences with a flower bud, and an infructescence with numerous dehisced capsules. Just above the branch is affixed a flower in anthesis. Because Bonpland collected this specimen and is the author of the name, this specimen is a second isotype of *E. peruvianum*.

*Exostema bicolor* Poepp., Nov. Gen. Sp. Pl. 3:32. 1841. Type: PERU. Huánuco: Cerro de San Cristóbal, near Cuchero, in rocky windswept sites, 1829, E.F. Poeppig *Diar. 1352* (*LECTOTYPE*, here designated: W [No. W 0049011]; possible isotype: GOET [GOET010234]; isotype fragment: F [No. 776569]).

*Notes*.—Poeppig (1841:32), in the protologue of *Exostema bicolor* Poepp., cited his own collection as "Crescit in scopulosis ventosis Peruviae transandinae, in monte Cerro San Cristobal juxta Cuchero. Septembre florebat" without indicating the herbarium of deposit. According to Stafleu and Cowan (1983:310), Poeppig's herbarium and types are at W, with duplicates in many herbaria. In W there is a specimen, with accession number W 0049011, that has a label with the annotation "Diar. 1352, Cuchero Peruvia 1829" handwritten by Poeppig. On the same label there is also the annotation "Hb. Endl." handwritten by an unknown author. This specimen, consisting of a branch with numerous leaves and several inflorescences with flower buds, is here designated as the lectotype of *E. bicolor*.

In GOET there is a specimen with barcode GOET010234, which has a label with the annotation "I. Exostemma peruvianum Humb. et Bonpl. in Peruvia orient. ad Huallagam superiorem leg. Poeppig Diar. n. 1352, May 1839" handwritten by an unknown author. The collection locality and date do not correspond to the locality described by Poeppig. However, the annotation "Poeppig Diar. n. 1352" is the same of the W specimen. This specimen consists of a branch with a few leaves and two inflorescences with flower buds, and it corresponds entirely with the W 0049011 specimen. Hence, it is a possible isotype of *E. bicolor*.

## NERNSTIA

Candolle (1830:350) published *Coutarea mexicana* Zucc. & Mart. ex DC., and described it as "foliis utrique glabris, pedunculis 1-floris bibracteolatis, corollis infundibulariformis, lobis obtusissimis. Mexico." [Leaves glabrous, peduncle with 1 flower, subtended by two bracteoles, corolla infundibular, lobes broadly obtuse. Mexico]. Urban (1923a:146) transferred this species to *Nernstia* Urb, proposing the new combination *N. mexicana* (Zucc. & Mart. ex DC.) Urb. Urban dedicated the genus name to the German chemist Walter Nernst (1864–1941), who won the Nobel Prize in Chemistry in 1920. Aiello (1979) apparently overlooked the genus *Nernstia* previously published by Urban, and published the superfluous generic name *Cigarilla*. Paudyal et al. (2018), in their molecular phylogenetic study, retrieved *Nernstia* and *Osa*, on a strongly supported clade, sister to the *Catesbaea-Portlandia-Isidorea* clade. *Nernstia* is similar to *Osa* by the axillary, 1-flowered inflorescences, 5-merous flowers, and linear anthers. *Nernstia* differs from *Osa* by being shrubs to 5 m tall, endemic to limestone areas (vs. shrubs or trees to 15 m tall, endemic to tall forest, on rich soil, in *Osa*), with leaves 3–7.5 cm long, coriaceous, with revolute margins, (vs. 16.5–23.5 cm long, membranaceous to chartaceous, with planar margins), corollas campanulate, 6–8 cm long (vs. trumpet-shaped, 30–35 cm long), capsules ovoid-obovoid, 1.3–1.8 cm long (vs. ellipsoid, 3.2–3.4 cm long), and by being endemic to Costa Rica and Panama (vs. endemic to central Mexico, states of Hidalgo and San Luis Potosí).

*Nernstia* differs from *Portlandia* by its colliculate seeds (vs. tuberculate) with acropetally imbricate arrangement, non-persistent funicle (vs. persistent), and large, spongy placenta (vs. linear, adnate to the septum). *Nernstia* is a monospecific genus endemic to central Mexico.

*Nernstia* Urb, Symb. Antill. 9:145. 1923; Candolle, Prodr. 4:350. 1830; Aiello, J. Arnold Arbor. 60:109–111 (as *Nernstia*). 1979; Borhidi et al., Rubiáceas México 306–307, f. 65. 2006; Borhidi et al., Rubiáceas México, 2nd ed. 352, f. 77. 2012; Paudyal & Delprete, Bot. J. Linn. Soc. 187:365–396. 2018. TYPE: *Nernstia mexicana* Urb.

*Cigarilla* Aiello, J. Arnold Arbor. 60:109. 1979. TYPE: *Cigarilla mexicana* (Zucc. & Mart. ex DC.) Aiello, J. Arnold Arbor. 60:109. 1979. (= *Nernstia mexicana*).
1. **Nernstia mexicana** (Zucc. & Mart. ex DC.) Urb., Symb. Antill. 9:146. 1923. *Coutarea mexicana* Zucc. & Mart. ex DC., Prodr. 4:350. 1830. *Cigarilla mexicana* (Zucc. & Mart. ex DC.) Aiello, J. Arnold Arbor. 60:109. 1979. TYPE: MEXICO: without locality, s.d., *Collector Unknown* s.n. [Communicated by Martius in 1829] (HOLOTYPE: G-DC [G00665779]).

*Distribution*.—Mexico (Hidalgo, San Luis Potosí).

*Notes*.—Candolle (1830:350) attributed the name *Coutarea mexicana* Zucc. & Mart. ex DC. to an unpublished manuscript of Zuccarini and Martius, and indicated the collection locality as “Mexico. (v.s.).” The expression “(v. s.)” means that he saw a herbarium specimen.

Aiello (1979:110) cited the type of *Coutarea mexicana* as “Type: Mexico, without further locality, Martius in 1829 (holotype, G-DC, Microfiche, IDC 800. 680: 11.7!).” The G-DC specimen cited by Aiello has barcode G00665779. On the bottom right corner of the sheet is affixed a label with the annotation “Coutarea mexicana Zucc.” handwritten by Candolle. The specimen consists of a small branch with a few leaves, a flower bud, and a flower in anthesis. In the attached envelope are included four leaves. At the base of the branch is pinned a label with the annotations “Coutarea mexicana Zuccar. M. Martius 1829. Mexico” handwritten by an unknown author. The date “1829” corresponds to the year when Martius sent the specimen to Candolle, and the collector and collection date of this specimen are unknown. This specimen is the holotype of *C. mexicana*.

## OSA

Simpson (1974:277) published *Hintonia pulchra* D.R. Simpson, and described it as a tree 15 m tall, with 5-merous flowers, axillary, in pairs at each node (one in each leaf axil), calyx lobes linear or filiform, corollas white, trumpet-shaped, 27 cm long, 11 cm wide at mouth, stamens included, style as long as the stamens, undivided, fruit capsular, and seeds flattened, not winged, ca. 6 mm long, 1.5–2.5 mm thick. Aiello (1979) transferred *H. pulchra* to the new genus *Osa* Aiello, dedicating the generic name to the Osa Peninsula, Costa Rica, where it is endemic. Aiello differentiated *Osa* from *Hintonia* by the large, wingless seeds (vs. small winged seeds) with tuberculate testa (vs. reticulate) and persistent funicle (vs. non-persistent), long trumpet-shaped corollas (vs. funnelform), and large leaves with attenuate apex (vs. medium-sized leaves with acute to acuminate apex). Paudyal et al. (2018), in their molecular phylogenies, found the two monospecific *Nernstia* (from Mexico) and *Osa* (from Costa Rica) on a strongly supported clade, sister to the *Catesbaea–Portlandia-Isidorea* clade, supporting the close relationship of these two genera. Hodel and Hannon (2024) dedicated an amply illustrated article on the morphology and cultural techniques of *Osa pulchra*.

*Osa* Aiello, J. Arnold Arbor. 60:115. 1979; Simpson, Phytologia 29(4):277-279, f. 1. 1974; Aiello, J. Arnold Arbor. 60:115-116. 1979; Ochoterena-Booth, in Davidse et al., Fl. Mesoamericana 4.2:163. 2012; Paudyal & Delprete, Bot. J. Linn. Soc. 187:365-396. 2018; Hodel & Hannon, Palmarbor 8:1-40. 2024. TYPE: *Osa pulchra* (D.R. Simpson) Aiello.

1. **Osa pulchra** (D.R. Simpson) Aiello, J. Arnold Arbor. 60:116. 1979. *Hintonia pulchra* D. Simpson, Phytologia 29(4):277, fig. 1. 1974. TYPE: COSTA RICA. Puntarenas: Osa Peninsula, ca. 5 km W of Rincón de Osa, areas of secondary vegetation N and W of air field, 50–60 m, 8°42'N, 83°31'W, 9–12 Jan 1970, W.C. Burger & R.L. Liesner 7320 (HOLOTYPE: F [No. 1729029]; ISOTYPES: MO [No. 2262984, barcode MO-2532201], U [U 0006042]).

*Distribution*.—Costa Rica (Peninsula de Osa), Panama.

## PHIALANTHUS

Grisebach (1861:335) published the genus *Phialanthus* Griseb., with the single species *P. myrtilloides* Griseb., without explaining the etymology of the name. From the Greek, phialé- (φιάλη), meaning vial, small glass container, and -anthos (ἀνθοσ) meaning flower, referring to the general shape of the flower. Correll and Correll (1982:1408) explained the etymology of the genus name as “Greek for “urn-flower.” Grisebach (1861:335) described *Phialanthus* as a viscid shrub, with shortly campanulate-infundibular, 4-lobed corollas, exserted stamens, style simple with blunt style lobes, 2-celled drupe, and cylindrical seeds. In addition, he stated that “the character, though not yet quite complete, shows this genus to be nearly related to *Scolosanthus*,
while from its resinous excretions *Stenostomum*, sect. *Laugeria*, is analogous.” *Phialanthus* has been included into the Chiococcae by most Rubiaceae specialists (e.g., Hooker 1873a, Bremer 1992; Robbrecht 1988, 1994; Delprete 1996). Bremer (1992) in an analysis using molecular and morphological data, excluded it from the Chiococcae due to the presence of free filaments. Rova et al. (2002) showed that it is closely related to several members of Chiococcae. Motley et al. (2005), in a phylogenetic analysis of the Chiococcae using molecular data, *Phialanthus* was supported as monophyletic. Paudyal et al. (2018) included nine *Phialanthus* species in their molecular phylogenetic study, and in both combined and separate analyses, it was retrieved as non-monophyletic. Eight species of *Phialanthus* were found on a strongly supported clade, and *P. hispaniolae* Alain & R.G. García at a basal position of the clade, next to *Eosanthe*. *Phialanthus* has never been subject of monographic revision. In the present treatment, 22 species are recognized in this genus, most of them endemic to Cuba, with some species occurring in the Bahamas, and in the Greater and Lesser Antilles.

**Phialanthus** Griseb., Fl. Brit. W. I.:335. 1861; Standley, North Amer. Flora 32(4):281-284. 1934; Liogier, Fl. Cuba 5:90-92. 1962; Correll & Correll, Fl. Bahama Arch. 1408-1410, f. 616. 1982; Borhidi, Acta Bot. Hung. 29:194-205. 1983; Liogier, Fl. Española 7:357, 359. 1995; Borhidi et al., Rubiáceas Cuba 250-260. 2017. TYPE: *Phialanthus myrtilloides* Griseb.

1. **Phialanthus acunae** Borhidi, Acta Bot. Hung. 29:196, fig. 2. 1983. TYPE: CUBA. HOLGUIN: Cananova, Camino de Centeno, Jul 1949, *Bro. Alain* [A.H. Liogier] & *Bro. Clemente* 975 [as "Clemente, Alain & Crisógone 975"] (HOLOTYPE: HAC; ISOTYPES: BP n.v., GH [00257260], HAJB n.v., NY [02200749], US [00902069]).

**Distribution.**—Cuba (Holguín, Guantánamo).

2. **Phialanthus alainii** Borhidi, Acta Bot. Hung. 50:278. 2008. TYPE: CUBA. GUANTÁNAMO: Toa, Magdalena, Peña Prieta, serpentine barrens, 700 m, 30 Jul 1952, *Bro. Alain* [Liogier] 3471 (HOLOTYPE: HAC n.v. [ex LS]; ISOTYPES: GH [00257527], HAC n.v., NY [02200761], US [00955899]).

**Distribution.**—Cuba (Guantánamo).

3. **Phialanthus bissei** (Borhidi) Borhidi, Acta Bot. Hung. 51:277. 2009. *Phialanthus rigidus* ssp. *bissei* Borhidi, Acta Bot. Hung. 38:168. 1994. TYPE: CUBA. ISLA DE PINOS: Camino entre Cayo Piedras y Punta del Este, sobre caliza, 24 Jul 1971, *J. Bisse s.n.* HFC No. 19844 (HOLOTYPE: HAJB n.v.; ISOTYPES: BP n.v., HAC, JE [JE00001228]).

**Distribution.**—Cuba (Isla de la Juventud).

**Notes.**—As a result of a detailed search in HAJB, no original material *Phialanthus rigidus* ssp. *bissei* Borhidi was found. The HAJB curator, Eldis R. Béquer (pers. comm.), is not aware of the possible location of the holotype specimen of this name. He confirmed that all the HAJB Rubiaceae specimens that where on loan to JPU, to be studied by Attila Borhidi, were returned to HAJB.

4. **Phialanthus ellipticus** Urb, Symb. Antill. 9:161. 1923. TYPE: CUBA. HOLGUIN: Río Piloto, in charrascales, 1 Sep 1914, E.L. *Ekman* 2716 (LECTOTYPE, here designated: S [No. S-07-14888]).

**Distribution.**—Cuba (Holguín: Sierra de Nipe, Río Piloto).

**Notes.**—Urban (1923a:161-162), in the protologue of *Phialanthus ellipticus* Urb., cited the gathering "Prov. Oriente prope Río Piloto in charrascales, m. Sept. flor.: [Ekman] n. 2716," without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. Borhidi (2017:252) cited the type of *Phialanthus ellipticus* as "Cuba. Oriente prope Río Piloto, [...], *Ekman* 2716 Holotipo: B†. Lectotipo: S!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017) lectotype designation is not valid. The specimen in S, with accession number S-07-14888, has a label with the annotation "Phialanthus ellipticus Urb. (typus)" handwritten by Urban. The specimen consists of several branches with numerous leaves, and several small flowers present among the leaves. This specimen is here designated as the lectotype of this name.

5. **Phialanthus glaberrimus** Borhidi, Acta Bot. Hung. 29:194. 1983. TYPE: CUBA. GUANTÁNAMO: Baracoa, Loma de Buena Vista, 500-600 m, 12 Aug 1975, A. Álvarez, *J. Bisse & K.F. Meyer* HFC No. 27393 (LECTOTYPE, here designated: HAJB [HAJB G 000749]; ISOLCTOTYPES: B [B 10 0385019], HAJB [3 sheets, HAJB G 000750, HAJB G 000751, HAJB G 000752], JE [JE00001214]).

**Distribution.**—Cuba (Guantánamo).
**Notes.**—Borhidi (1983:194), in the protologue of *Phialanthus glaberrimus* Borhidi, cited the type of this name as "Holotypus: HAJB 27393. Baracoa: Loma de Buena Vista 500-600 m.s.m., Leg. Alvarez, Bisse, Meyer 12.8.1975." The number "27393" pertains to the series *Herbario de la Flora de Cuba* (HFC), which was used by the collectors that participated in the Project *Flora de Cuba*, an international convention between the University of La Habana, Cuba, the Friedrich Schiller University in Jena, Germany, and the Humboldt University in Berlin, Germany (Regalado Gabancho et al. 2008). All the duplicates of this gathering were distributed as HFC No. 27393. In HAJB there are four specimens of this gathering, all of them with the typewritten annotation "Phialanthus glaberrimus Borhidi. sp. nov." The specimen with barcode HAJB G 000749 is here designated as the lectotype of this name.

6. **Phialanthus grandifolius** Alain, Bull. Torrey Bot. Club 92(4):302. 1965. **TYPE:** PUERTO RICO: Maricao State Forest, serpentine barrens, 800 m, 26 Jun 1963, A.H. Liogier [Bro. Alain] 9715 (LECTotype, here designated: NY [02200755]; ISOLECTOTYPES: DUKE [10000654], GH [00094964], IJ n.v., US [00138506]).

**Distribution.**—Puerto Rico.

**Notes.**—Liogier (1965:302), in the protologue of *Phialanthus grandifolius* Alain, cited the type as "PUERTO RICO. Serpentine barrens, Maricao State Forest, alt. 800 m, June 26, 1963, Alain 9717 (NY, GH, IJ, US)." He did not specify which of those specimens is the holotype. The specimen in NY, barcode 02200755, is here designated as the lectotype of this name.

7. **Phialanthus guantanamensis** Borhidi, Acta Bot. Hung. 29:195. 1983. **TYPE:** CUBA. GUANTÁNAMO: monte seco sobre diente de perro, en la subida a Monte Cristi, 300 m, Jun 1967, *J. Bisse & L. Rojas HFC No. 3517* (HOLOTYPE: HAJB [HAJB G 000753]; ISOTYPE: JE [JE00004986 (HFC No. 0035517)]).

**Distribution.**—Cuba (Guantánamo).

8. **Phialanthus hispaniolae** Alain & R.G. García, Moscosoa 8:8, fig. 2. 1994. **TYPE:** DOMINICAN REPUBLIC. INDEPENDENCIA: Sierra de Bahoruco, S de Duvergé, en el Firme La Cañita, al S de Monte Palma, 18°17'N, 71°30'W, 700-750 m, 3 Dec 1993, R. García, G. Caminero, D. Hönér & T. Montilla 5288 (HOLOTYPE: UPR n.v.; ISOTYPES: B [B 10 0413571], F [No 2226303], JBSD n.v., NY n.v. [probably not there], USD n.v.).

**Distribution.**—Dominican Republic (Sierra de Bahoruco).

9. **Phialanthus inflatus** Borhidi, Acta Bot. Hung. 29:195. 1983. **TYPE:** CUBA. GUANTÁNAMO: Baracoa, charrascales cerca de la desembocadura de Arroyo Maguana, Feb 1968, *J. Bisse & E. Köhler HFC No. 5756* (HOLOTYPE: HAJB [HAJB G 000754]; ISOTYPE: JE [JE00001205 (HFC No. 005756)]).

**Distribution.**—Cuba (Guantánamo: Baracoa).

10. **Phialanthus jamaicensis** Urb., Symb. Antill. 5:515. 1908. **TYPE:** JAMAICA: road to Wareka, 10 Oct 1905, W. Harris 9023 (LECTotype, here designated: F [No. 189197]; isolectotypes, A [00094965], C [without barcode], NY [2 sheets, 02200756, 02200757], RSA [RSA0005812], US [0013507]).

**Distribution.**—Jamaica.

**Notes.**—Urban (1908:515), in the protologue of *Phialanthus jamaicensis* Urb., cited two gatherings as follows: "Jamaica. Juxta viam ad Wareka, 260 m. alt., m. Oct. florifer: Harris n. 9023, in Long Mountain ad latus australe, 300 m. alt., m. Jun. fl.: Harris 9587," without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. There is a specimen in F, with accession number 189197, that has a label with the heading "Herbarium Krug et Urban" and the annotation "9023. Phialanthus jamaicensis Urb. (typus)" handwritten by Urban. The specimen consists of a branch with numerous leaves and a few inflorescences with some flowers in anthesis. This specimen is here designated as the lectotype of *P. jamaicensis*.

11. **Phialanthus linearis** Alain, Contr. Ocas. Mus. Hist. Nat. La Salle 17:7. 1959. **TYPE:** CUBA. HOLGUÍN: Moa, Cuabales de Playa la Vaca, 9 Nov 1945, *J. Acuña* 13369 (HOLOTYPE: HAC [ex SV] (Fig. 3); ISOTYPE: HAJB [HAJB G 000755, ex LS]).

**Distribution.**—Cuba (Holguín: Moa).

12. **Phialanthus macrocalyx** Borhidi, Acta Bot. Hung. 29:195. 1983. **TYPE:** CUBA. HOLGUÍN: Moa, Sierra de Moa, Charrascal del Cayo Coco, 200-300 m, 13 Aug 1970, *J. Bisse & H. Lippold HFC No. 17596* (HOLOTYPE: HAJB [HAJB G 000756]).

**Distribution.**—Cuba (Holguín: Moa).
<figure><img src="image_6.png" /><figcaption>FIG. 3. Holotype of *Phialanthus linearis* Alain (Acuña 13369, HAC [ex SV]). Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
17. *Phialanthus parvifolius* Urb., Symb. Antill. 9:160. 1923. TYPE: CUBA. GUANTÁNAMO: Baracoa, ad Rio Yoa, in charrascales valde communis, 28 Nov 1914, E.L. Ekman 3680 (*LECTOTYPE*, here designated: S [S07-14889]; *ISOLECTOTYPE*: G [G00436461]).

*Distribution*.—Cuba (Guantánamo: Baracoa).

*Notes*.—Urban (1923a:160–161), in the protologue of *Phialanthus parvifolius* Urb., cited the material studied as “Prov. Oriente prope Baracoa ad Rio Yoa in charrascales valde communis, m. Nov. flor.: [Ekman] n. 3680,” without citing the herbarium of deposit. The original material studied by Urban in B was destroyed during WWII. Borhidi (2017:257) cited the type of *Phialanthus parvifolius* as “Tipo: Cuba, Oriente prope Baracoa, [...], Ekman 3680, Holotipo: B†. Lectotipo: S!” According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al.’s (2017) lectotype designation is not valid.

In S there is a specimens, with accession number S07-14889, which has a label with the annotation “Phialanthus parvifolius Urb. (typus)” handwritten by Urban. The specimen consists of two branches with numerous, dense leaves, and several axillary flowers in anthesis. This specimen is here designated as the lectotype of this name.

18. *Phialanthus peduncularis* Borhidi, Acta Bot. Hung. 38:167. “1993–94” [1995]. TYPE: CUBA. HOLGUÍN: Sierra del Cristal, Río Lebisa, 26 Aug 1959, M. López Figueiras 201 (HOLOTYPE: HAC [ex LS]).

*Distribution*.—Cuba (Holguín: Sierra del Cristal).

19. *Phialanthus resinifluus* Griseb., Cat. Pl. Cub. 140. 1866. TYPE: CUBA. ORIENTE [HOLGUÍN or GUANTÁNAMO]: Monteverde, Mar 1859, C. Wright 1269 (*LECTOTYPE*, here designated: GOET [GOET010424]; *ISOLECTOTYPES*: BM [BM000839311], BR [000000531468], G [5 sheets, G00436458, G00436459, G00436460], GH [2 sheets, 00094966, 00094967], GOET [GOET010425], HAC, K [K000432640], PH [00029581], S [2 sheets, Nos. S07-14892, S07-14893], YU [2 sheets, YU.001779, YU.001780]).

*Distribution*.—Cuba (Pinar del Rio, Artemisa, Sancti Spiritus, Holguín, Guantánamo).

*Notes*.—Grisebach (1866:140), in the protologue of *Phialanthus resinifluus* Griseb., cited the material studied as “Cuba or., pr. Monteverde (Wr. [Wright] 1269); eundem fruticem e gemmis guttas resinosas stillantem in Jamaica leg. March. E.,” without citing the herbarium of deposit. For information regarding Grisebach’s original specimens, see notes under *P. myrtilloides*. Borhidi et al. (2017:259) cited the type of *P. resinifluus* as “Tipo: Cuba, Oriente, Monteverde, [...], C. Wright 1269, Holotipo: GOET, isotipos: GH, S-HAC, US.” According to the *Code*, starting from 1 January 2001, Borhidi et al.’s (2017) holotype citation cannot be treated as an inadvertent lectotypification because it is not accompanied by “here designated” or a similar expression.

In GOET there are two specimens of Wright 1269 annotated with this name. The GOET specimen with barcode GOET010425 has a label with the heading “Plantae Cubenses Wrightianae,” with the number “1269” handwritten by Wright, and “Phialanthus resinifluus m.” handwritten by Grisebach. The specimen consists of a branch with numerous leaves and a few flowers in anthesis. In the envelope affixed on the sheet are included numerous leaves and a small branch with some flowers in anthesis.

The GOET specimen with barcode GOET010424 has a label with the heading “Plantae Cubenses Wrightianae,” with the numbers “221 = 1269” handwritten by unknown authors, and “Phialanthus resinifluus m.” handwritten by Grisebach. On the right side of that label are affixed two labels with a detail description of the specimen, handwritten by Grisebach. The specimen consists of a large, ramose branch with numerous leaves and numerous inflorescences with many flowers in anthesis. This specimen is here designated as the lectotype of *Phialanthus resinifluus*.

20. *Phialanthus revolutus* Urb., Repert. Spec. Nov. Veg. 17:407. 1921. TYPE: JAMAICA: Healthshire Hills, near Salt Island, 1 Sep 1908, W. Harris & N.L. Britton 10525 (*NEOTYPE*, here designated: NY [02200764]; *ISONEOTYPES*: F [No. 243383], US [00879022]).

*Distribution*.—Jamaica.

*Notes*.—Urban (1921c:407), in the protologue of *Phialanthus revolutus* Urb., cited the material studied as “Hab. in Jamaica in Healthshire Hills prope Salt Island m. Sep. deflor.: Harris et Britton no. 10525,” without citing the herbarium of deposit. The original material at B was destroyed during WWII. Three specimens of
*Harris & Britton 10525* are in F, NY, and US. None of these specimens has proof that they were examined by Urban. The specimen in NY, with barcode 02200764, has a label with the heading "FLORA JAMAICENSIS" and the annotation "Phialanthus jamaicensis Urban" handwritten by an unknown author. The specimen consists of two branches. The branch on the right side of the sheet has larger leaves and no flowers. The branch on the left side of the sheet has numerous, smaller leaves and numerous flowers in anthesis. This specimen is here designated as the neotype of *P. revolutus*.

21. **Phialanthus rigidus** Griseb., Cat. Pl. Cub. 140. 1866. **TYPE:** CUBA. SANTIAGO DE CUBA: San Marcos, 1860-1864, C. Wright 2725 (**LECTOTYPE**, here designated: GOET [GOET010426]; **ISOLECTOTYPES:** BM [BM000839310], G [G00436457], GH [00094968], HAC [2 sheets], K [K000174008], MPU [MPU021772], NY [02200765], S [S07-14890], US [00138509], YU [YU.001787]).

**Distribution.**—Cuba (Pinar del Rio, Artemisa, Villa Clara, Sancti Spiritus, Isla de Pinos).

*Notes*.—Grisebach (1866:140), in the protologue of *Phialanthus rigidus* Griseb., cited the material studied as "Cuba occ., pr. S. Marcos (Wr. [Wright] 2725). E.," without citing the herbarium of deposit. For information regarding Grisebach's original specimens, see notes under *P. myrtilloides*. Borhidi et al. (2017:259) cited the type of *P. rigidus* as "Tipo: Cuba; Prov. Pinar del Rio, San Marcos, [...], C. Wright 2725, Holotipo: GOET, isotipos: GH, HAC, NY, S, US." According to the *Code*, starting from 1 January 2001, Borhidi et al.'s (2017) holotype citation cannot be corrected as an inadvertent lectotypification because it is not accompanied by "here designated" or a similar expression, and is here superseded.

In GOET there is a specimen, with barcode GOET010426, which has a label with the heading "Plantae Cubenses Wrightianae" and the annotation "Phialanthus rigidus m." followed by a detail description handwritten by Grisebach. The specimen consists of two branches with numerous leaves. The branch on the upper left side of the sheet does not have any visible flowers. The branch on the right side of the sheet has numerous inflorescences with flowers in anthesis. This specimen is here designated as the lectotype of *P. rigidus*.

22. **Phialanthus stillans** Griseb., Cat. Pl. Cub. 140. 1866. **TYPE:** CUBA. PINAR DEL RIO: In vertice Guajarbon [Pan de Guajaibón], 16 Nov [1860-1864], C. Wright 2726 (**LECTOTYPE**, here designated: GOET [GOET010427]; **ISOLECTOTYPES:** BM [BM000839309], G [G00436456], GH [00094969], HAC, K [K000432634], YU [YU.001782]).

**Distribution.**—Cuba (Pinar del Rio, Artemisa).

*Notes*.—Grisebach (1866:140), in the protologue of *Phialanthus stillans* Griseb., cited the material studied as "Cuba occ., in vertice m. Guajarbon (Wr. [Wright] 2726). E.," without citing the herbarium of deposit. For information regarding Grisebach's original specimens, see notes under *Phialanthus myrtilloides*. Borhidi et al. (2017:260) cited the type of *P. stillans* as "Tipo: Cuba; Prov. Pinar del Rio, Pan de Guajabón, [...], C. Wright 2726, Holotipo: GOET, isotipos: GH, HAC, US." According to the *Code*, starting from 1 January 2001, Borhidi et al.'s (2017) holotype citation cannot be corrected as an inadvertent lectotypification because it is not accompanied by "here designated" or a similar expression.

The GOET specimen, with barcode GOET010427, has a label with the heading "Plantae Cubenses Wrightianae" and the annotation "Phialanthus stillans m." handwritten by Grisebach. On the verso of that label there is a detailed description of the species, handwritten by Grisebach. Above that label is affixed a small label with the annotation "Rubiac. Fruticose 10° f. Summit of Guajarbon. Nov. 16" handwritten in pencil by Charles Wright. This specimen consists of two small branches with a few flowers in anthesis, and is here designated as the lectotype of *P. stillans*.

## PHIALANTHUS EXCLUDED TAXA

*Phialanthus spicatus* Wright, Anales Acad. Ci. Méd. Fís. Nat. Habana 6:149. 1869.
= *Ceratopyxis verbenacea* (Griseb.) Hook. f., in Hooker's Icon. Pl. 12:24, tat. 1125. 1876 (See above).

## PORTLANDIA

Patrick Browne (1756:164-165) described the genus *Portlandia* P. Browne, without citing a Latin name for a species, and only published the English name "The large-leaf'd *Portlandia*." He dedicated the genus name to the Duchess of Portland, Margaret Cavendish Bentick (1715-1785). Linnaeus (1759:928) cited table 11 of
Browne's *History of Jamaica*, and published the name *P. grandiflora* L., which is the type species of the genus. *Portlandia* has a complex taxonomic history, because it was used by several authors to position numerous species with variable morphological characters, resulting in a genus of more than 20 species (e.g., Wernham, 1913; Britton, 1914; Urban, 1923a, 1923b). Aiello (1979) studied *Portlandia* and associated taxa, provided a taxonomic revision, with complete synonymy and typification, transferred numerous species to several new genera, mostly based on placentation and seed morphology, and restricted *Portlandia* to a genus of five species and one variety endemic to Jamaica. Delprete and Motley (2003), based on molecular and morphological data, elevated Aiello's variety to species level, adding to a total of six species endemic to Jamaica. In Paudyal et al.'s (2018) phylogenetic study, the six species of *Portlandia* formed a strongly supported monophyletic clade, sister to the *Isidorea* clade. A sister relationship of *Portlandia* and *Isidorea* was already found in previous studies (Delprete 1996; Rova et al. 2002; Delprete & Motley 2003; Motley et al. 2005; Robbrecht & Manen 2006; Manns & Bremer 2010). *Portlandia* is represented by shrubs or small trees growing in limestone areas, stipules broadly triangular, coriaceous, non-pungent, leaves coriaceous, non-pungent, inflorescences axillary, 1-6-flowered, corollas funnel-shaped, white, pink or red, stamens adnate to the base of corolla tube, ovary 2(3)-locular, placentation linear, adnate to the septum, stigmatic area as two lines along the style, capsules loculicidal, and seeds perpendicular, with tuberculate exotesta.

**Portlandia** P. Browne, Hist. Jamaic. 164, tab 11. 1756; Wernham, J. Bot. 51:320-324. 1913; Britton, Bull. Torrey Bot. Club 41 1-24. 1914; Standley, N. Amer. Fl. 32(1):8-13. 1918; Urban, Symb. Antill. 9:55-176. 1923; Urban, Symb. Rep. Sp. Regn. Veg. 19:1-9. 1923; Liogier, Fl. Cuba 5:27-28. 1962; Adams, Fl. Jamaica 704-705. 1972; Aiello, J. Arnold Arbor. 60 38-123. 1979; Delprete & Motley, Brittonia 55(3):233-239, f. 1-2. 2003; Paudyal et al., Bot. J. Linn. Soc. 187(3):365-396. 2018. **TYPE:** *Portlandia grandiflora* L.

1. **Portlandia coccinea** Sw., Prodr. 42. 1788. **TYPE:** JAMAICA: without locality, s.d., *Swartz s.n.* (FIRST-STEP LECTOTYPE (Aiello 1979:102), SECOND-STEP LECTOTYPE, here designated: S [No. S-R-5215]; ISOLECTOTYPES: BM n.v., F [No. 588420], M [M-0188947], S [No. S-R-5214], SBT [SBT13334]; POSSIBLE ISOLECTOTYPE: LINN-HS [321-3]).

**Distribution.**—Jamaica (Westmorland, Trelawny, St. Elizabeth, Manchester).

**Notes.**—Swartz (1788:42), in the protologue of *Portlandia coccinea* Sw., cited his own collection from Jamaica, without indicating any specimen or herbarium of deposit. Aiello (1979:102) cited the type of this name as "Type: Jamaica, without further locality, *Swartz s.n.* (lectotype, S!; probable isotype, BM!)." There are two specimens associated with this name in S, both collected by Swartz in Jamaica. Hence, Aiello's citation is a first-step lectotypification. The S specimen with accession number S-R-5215 is here designated as the second-step lectotype of *P. coccinea*.

*Portlandia coriacea* Spreng., Syst. Veg. 1.708. 1825, Sphalm. of *P. coccinea*.

2. **Portlandia grandiflora** L., Syst. Nat., ed. 10, 2:928. 1759. ["Habitat in Jamaica," Sp. Pl., ed. 2, 1:244. 1762]. **TYPE** [CON.]: "Portlandia foliis majoribus nitidis ovatis oppositis, floribus amplissimis" in Browne, Civ. Nat. Hist. Jamaica: 164: tab. 11. 1756 (LECTOTYPE (Aiello 1979:100) [Tab. 11 is reproduced in Jarvis, p. 764. 2007].

**Distribution.**—Jamaica (the whole island).

*Portlandia grandiflora* var. *parviflora* S. Moore, J. Bot. 68:108. 1930. **TYPE:** JAMAICA: without locality, s.d., *Broughton s.n.* (HOLOTYPE: BRIST n.v.).

3. **Portlandia harrisii** Britton, Bull. Torrey Bot. Club 39:8. 1912. **TYPE:** JAMAICA. Peckham Woods, Upper Clarendon, 6 Jul 1911, *W. Harris* 10975 (FIRST-STEP LECTOTYPE (Aiello 1979:104), SECOND-STEP LECTOTYPE, here designated: NY [00126749]; ISOLECTOTYPES: NY [00126748], UCWI n.v., US [00137329]).

**Distribution.**—Jamaica (Clarendon, St. Ann).

**Notes.**—Britton (1912:8), in the protologue of *Portlandia harrisii* Britton, stated "Type locality: Peckham Woods, Upper Clarendon, Jamaica," without citing any specimen or herbarium of deposit. Aiello (1979:104) cited the type of this name as "Type: Jamaica. Clarendon, Peckham Woods, *Harris* 10975 (holotype, NY!; isotypes, NY!, UCWI!, US!)." Because she did not specify which of the two NY specimen is the "holotype," her citation is a first-step lectotypification. Both NY specimens have the annotation "Portlandia harrisii Britton"
handwritten by Britton. The NY specimen with barcode 00126748 consists of two branches with numerous leaves and some capsules.

The NY specimen with barcode 00126749 consists of two branches with numerous leaves, one of them with numerous capsules. At the base of the sheet is affixed an envelope including portions of dissected capsules and two corollas in anthesis. This specimen is here designated as the second-step lectotype of *P. harrisii*.

**4. Portlandia microsepala** Urb., Repert. Spec. Nov. Regni Veg. 13:478. 1915. **TYPE:** JAMAICA. ST. ANN: A bion, 1850, R.C. Alexander s.n. (NEOTYPE: Aiello 1979:106): K [K000173526]).

*Distribution*.—Jamaica (St. Ann).

**Notes.**—Urban (1915:478), in the protologue of *Portlandia microsepala* Urb., cited the material studied as "Hab. in Jamaica prope Moneague in Union Hill, m. Majo flor.: Alexander (typus), in collibus distr. St. Ann's fruct.: Alexander." Hence, Urban cited as type a flowering specimen collected by Alexander at Moneague on Union Hill. The original material in B studied by Urban was destroyed during WWII. Aiello (1979:106) cited the type of this name as "Type: Jamaica. St. Ann, Albion, *Alexander* in 1850 (holotype, K!)," with a locality corresponding to that of the paratype cited by Urban. She annotated the K specimen with barcode K000173526 as "Neotype of Portlandia microsepala Urb., Annette Aiello 1977." Among the specimens examined of this species, she cited duplicates of Alexander in NY and P, but I was unable to trace them. Her citation as "holotype" of the K specimen is an inadvertent neotype designation.

**5. Portlandia platantha** Hook. f., Bot. Mag. 76:tab. 4534. 1850. **TYPE:** cultivated at Kew, origin unknown [Jamaica], flowering in July 1850, *Collector Unknown s.n.* (HOLOTYPE: K [K000173522]).

*Distribution*.—Jamaica (St. Ann, St. Mary, St. Andrew, Portland, St. Thomas).

*Portlandia grandiflora* var. *latifolia* DC., Prodr. 4:405. 1830. **TYPE:** JAMAICA: without locality, s.d., *Collector Unknown s.n.* (HOLOTYPE: G-DC [3 sheets, G00666674, G00666685, G00666422]).

**Notes.**—In G-DC there are three sheets associated with this name, which are kept together in the same folder. Only one of these sheets, with barcode G00666674, has the annotation "Portlandia grandifolia β DC." handwritten by Candolle. Hence, according to Art. 8.3 of the *Code* (Turland et al. 2018), these three sheets are treated as a single specimen with multiple preparation, which is the holotype of *Portlandia grandiflora* var. *latifolia*.

*Portlandia albiflora* Britton & Harris ex Standl., N. Amer. Fl. 32:12. 1918. **Type:** JAMAICA: St. Andrew, Cane River Valley, 12 Jul 1907, W. Harris 9637 (LECTOTYPE, here designated: NY [00126743]; isolectotypes: A [00058983], BM [000081655], CAS [0004142], K [K000173490], MO [No. 867111], NY [00126742], UCWI n.v, US [00137324]).

**Notes.**—Standley (1918:12), in the protologue of *Portlandia albiflora* Britton & Harris ex Standl., cited the type as "Type collected in Cane River Valley, Jul 12, 1907, William Harris 9637 (herb. N. Y. Bot. Gard.)." Aiello (1979:103) cited the type of this name as "Type. Jamaica. [...] Harris 9637 (holotype, NY!; isotypes, A!, BM!, NY!; UCWI!, US!)." Hence, she did not specify which of the two specimens in NY is the "holotype." Both NY specimens have the annotation "Portlandia albiflora Britton & Harris" handwritten by Britton, and the stamp "Examined for NORTH AMERICAN FLORA." The NY specimen with barcode 00126743 consists of three branches with numerous leaves, with flowers and capsules, and is here designated as the lectotype of *P. albiflora*.

*Portlandia latifolia* Britton & Harris ex S. Moore, J. Bot. 68:108. 1930. **TYPE:** JAMAICA: St. Andrew, Hall's Delight, 3 Oct 1917, W. Harris 12669 (HOLOTYPE: BM [2 sheets, 000081656 ("Sheet II"), 000081657 ("Sheet II"); ISOTYPES: DS n.v., F [2 sheets, Nos. 492870, 492872], GH [00058979], K [3 sheets, K000173523, K000173524, K000173525], MO [No. 867111], NY [00126751], P [P02273497], UCWI, US [00137331]).

**6. Portlandia proctorii** (Aiello) Delprete, Brittonia 55(3):238, fig. 2. 2003. *Portlandia coccinea* var. *proctorii* Aiello, J. Arnold Arbor. 60:102. 1979. **TYPE:** JAMAICA: St. Catherine, several mi N of Old Arbour, 12 Nov 1971, G. Proctor 32708 (HOLOTYPE: IJ [No. 51993]; ISOTYPE: TEX n.v.).

*Distribution*.—Jamaica (St. Catherine).

## PORTLANDIA EXCLUDED TAXA

*Portlandia acuminata* Willd. ex R. & S., Syst. 5:23. 1819.

= *Coutarea hexandra* (Jacq.) K. Schum. (See above).

*Portlandia acunae* Borhidi, Abstr. Bot. Univ. Budapest 5:35. 1977.

= *Isidorea acunae* (Borhidi) Borhidi (See above).
*Portlandia brachycarpa* Urb., Symb. Antill. 9:135. 1923.

= *Isidorea brachycarpa* (Urb.) Aiello (See above).

*Portlandia corymbosa* Ruiz & Pav., Fl. Peruv. 2:49. 1799.

= *Motleychothamnus corymbosum* (Ruiz & Pav.) Paudyal & Delprete (See above).

*Portlandia daphnoides* R. Graham, Edinburgh New Philos. J. 30:206. 1841.

= *Cubanola daphnoides* (R. Graham) Aiello (See above).

*Portlandia domingensis* Britton, Bull. Torrey Bot. Club 41:24. 1914.

= *Cubanola domingensis* (Britton) Aiello (See above).

*Portlandia elliptica* Britton, Bull. Torrey Bot. Club 41:22. 1914.

= *Schmidtottia elliptica* (Britton) Urb. (See below).

*Portlandia ghiesbreghtiana* Baill, Adansonia 12:300. 1879.

= *Coutaportla ghiesbreghtiana* (Baill.) Urb. (See above).

*Portlandia guatemalensis* Standl., J. Wash. Acad. Sci. 18:162. 1928.

= *Lorencea guatemalensis* (Standl.) Borhidi (See above).

*Portlandia gypsophila* Macfad. ex Griseb., Fl. Brit. W. Ind. 324. 1861.

= *Thogssennia lindeniana* (A. Rich.) Aiello (See below).

*Portlandia hexandra* Jacq., Sel. Stirp. Amer. Hist. 63. 1763.

= *Coutarea hexandra* (Jacq.) K. Schum. (See above).

*Portlandia hexandra* L., Mant. Pl., ed. 2, 45. 1767, nom. superfl.

= *Coutarea hexandra* (Jacq.) K. Schum. (See above).

*Portlandia involucrata* Wernham, J. Bot. 51:320. 1913. *Ceuthocarpus involucratus* (Wernham) Aiello, J. Arnold Arbor. 60:109. 1979.

= *Schmitottia involucrata* (Wernham) Alain (See below).

*Portlandia lindeniana* (A. Rich.) Britton, Bull. Torrey Bot. Club 41:23. 1914. *Gonianthes lindeniana* A. Rich., in R. de la Sagra, Hist. Fis. Cuba, Bot. 11:10. 1850.

= *Thogssennia lindeniana* (A. Rich.) Aiello (See below).

*Portlandia longiflora* Meisen ex Griseb., Cat. Pl. Cub. 126. 1866.

= *Cubanola daphnoides* (R. Graham) Aiello (See above).

*Portlandia lumana* (Baill.) Baill., Hist. Pl. 7:333. 1880. *Coutarea lumana* Baill., Adansonia 12:301. 1879.

= *Hintonia lumana* (Baill.) Bullock (See above).

*Portlandia mexicana* (Zucc. & Martius ex DC.) Hemsl., Diag. Pl. Nov. 31. 1879. *Coutarea mexicana* Zucc. & Martius ex DC., Prodr. 4:350. 1830. *Cigarilla mexicana* (Zucc. & Martius ex DC.) Aiello, J. Arnold Arbor. 60:117. 1979, nom. superfl.

= *Nernstia mexicana* (Zucc. & Martius ex DC.) Urb. (See above).

*Portlandia nitens* Britton, Bull. Torrey Bot. Club 39:10. 1912.

= *Schmidtottia nitens* (Britton) Urb. (See below).

*Portlandia oblanceolata* Urb., Symb. Antill. 9:136. 1923.

= *Isidorea oblanceolata* (Urb.) Aiello (See above).

*Portlandia ophiticola* Borhidi, Növenyrendsz. Novényföldr. Tansz., Eötvös Loránd Tudományegyet. Budapest 5:34. 1977.

= *Isidorea ophiticola* (Borhidi) Borhidi (See above).

*Portlandia pendula* C. Wright ex Griseb., Cat. Pl. Cub 126. 1866.

= *Siemensia pendula* (C. Wright ex Griseb.) Urb. (See below).

*Portlandia polyneura* Urb., Symb. Antill. 9:135. 1923.

= *Isidorea polyneura* (Urb.) Aiello (See above).

*Portlandia pterosperma* S. Watson, Proc. Amer. Acad. Arts 24:52. 1889.

= *Hintonia latiflora* (Sessé & Moçoïno ex DC.) Bullock (See above).

*Portlandia sessilifolia* Britton, Bull. Torrey Bot. Club 41:21. 1914.

= *Schmidtottia sessilifolia* (Britton) Urb. (See below).
*Portlandia shaferi* Standl., N. Amer. Fl. 32:9. 1918.

= *Schmidtottia shaferi* (Standl.) Urb. (See below).

*Portlandia speciosa* Baill., Hist. Pl. 7:381. 1880.

= *Coutarea hexandra* (Jacq.) K. Schum. (See above).

*Portlandia uliginosa* Wernham, J. Bot. 51. 320. 1913.

= *Schmidtottia uliginosa* (Wernham) Urb. (See below).

## RAMONADOXA

Urban (1921a:163) described *Chiococca cubensis* Urb., a species endemic to Sierra de Nipe, eastern Cuba. In Paudyal et al.'s (2018) phylogenetic analyses, *Chiococca cubensis* was represented by two different collections. It was retrieved as a sister taxon to *Scolosanthus* in the combined and nuclear phylogenies, or nested within the *Scolosanthus* clade in the analysis using only the plastid dataset. In their analyses, the *Scolosanthus-Chiococca cubensis* clade is sister to the *Salzmannia* clade. *Chiococca cubensis* is similar to *Salzmannia*, as delimited by Paudyal et al. (2018), by the tubular-subcylindrical corollas, from which it can be distinguished by the branches without resinous exudate (vs. with resinous exudate), cymose inflorescence (vs. subcapitate or small racemes), corollas deep purple-brown outside and yellow inside (vs. white, pale yellow, yellow orangish-yellow to greenish) and by being endemic to Cuba (vs. occurring in coastal cordillera of Venezuela and coastal Brazil). *Chiococca cubensis* is similar to *Scolosanthus* in having narrowly imbricate corollas, from which it can be distinguished by being a scandent or climbing shrub lacking thorns (vs. erect shrubs with bifurcate or trifurcate thorns in *Scolosanthus*). It can be distinguished from all the other *Chiococca* species by having corollas tubular-subcylindrical, purple-brown outside and yellow inside, while in the other *Chiococca* species the corollas are campanulate to funnelform, white, cream-white to pale yellow throughout, or exceptionally red-dish outside. According to the molecular phylogenies and morphological differences mentioned above, Paudyal et al. (2018) transferred *C. cubensis* to *Ramonadoxa* Paudyal & Delprete, dedicating the generic name to Ramona Oviedo-Prieto (Instituto de Ecología y Sistemática, Ministerio de Ciencia, Tecnología y Medio Ambiente, La Havana, Cuba), an extremely knowledgeable botanist specialized on the Cuban flora. *Ramonadoxa* is a monotypic genus known from thickets, charrascal vegetation and pinelands of eastern Cuba (Oriente).

**Ramonadoxa** Paudyal & Delprete, Bot. J. Linn. Soc. 187:389. 2018; Urban, Symb. Antill. 9:163. 1921; Standley, N. Amer. Fl. 32(4):287. 1934; Liogier, Fl. Cuba 5:94. 1962; Borhidi et al., Rubiáceas Cuba 70. 2017; Paudyal et al., Bot. J. Linn. Soc. 187(3):365–396. 2018; Borhidi et al., Acta Bot. Hung. 60(3–4):300–302. 2018. TYPE: *Ramonadoxa cubensis* (Urb.) Paudyal & Delprete

1. **Ramonadoxa cubensis** (Urb.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:389. 2018. *Chiococca cubensis* Urb., Symb. Antill. 9:163. 1921. TYPE: CUBA, HOLGUÍN: Sierra de Nipe, Río Piedra, in charrascales, 200 m, 21 Oct 1919, E.L. Ekman 9996 (LECTOTYPE (Paudyal et al. 2018:390): S [No. S-10-24102]).

**Distribution.**—Cuba (Holguín: Sierra de Nipe, Mayarí, Sierra de Moa; Guantánamo: Baracoa).

**Notes.**—Urban (1923:163), in the protologue of *Chiococca cubensis* Urb., cited the gathering *Ekman 9996* as the type, without indicating the herbarium of deposit. The original material at B studied by Urban was destroyed during WWII. Borhidi et al. (2017:70; 2018:301) cited the type of *C. cubensis* as “[...] Sierra de Nipe ad Río Piedra in charrascales 200 m alt., Ekman 9996; holotipo B†, lectotipo: S!). According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al.’s (2017, 2018) lectotype designations are not valid. In S there is a specimen of *Ekman 9996*, with accession number S-10-24102, which was designated as the lectotype by Paudyal et al. (2018:390).
SALZMANNIA

Candolle (1830:617) described the genus *Salzmannia* DC., dedicating the generic name to Philipp Salzmann (1781–1851), a German doctor, who made several expeditions in the tropics, and valuable botanical collections in northeastern Brazil in 1827–1830. In the genus, Candolle (1830:617) included the sole species *S. nitida* DC., citing material collected by Salzmann in the Brazilian state of Bahia. Jardim et al. (2015) distinguished *Chiococca* from *Salzmannia* from northeastern Brazil, by being plants without resinous exudates or with only very small quantities of these (vs. with copious resinous exudate in *Salzmannia*), inflorescences lax, cymose or racemiform (vs. subcapitate), corollas campanulate, funnelform, or tubular, white to yellow with 5 lobes, valvate or narrowly imbricate in bud (vs. corollas tubular, yellow to orange, commonly with 4 lobes, rarely varying and up to 6 within certain populations, narrowly imbricate in bud), style subcapitate to fusiform or bifid with linear lobes (vs. style bifid with linear lobes), mature fruits white or yellowish white (vs. pink to purple). In addition, Jardim (in Jardim et al. 2015) described an arboreal species of *Salzmannia*, *S. arborea* Jardim. Paudyal et al. (2018), in their molecular phylogenetic trees, found two species of *Chiococca*, *C. plowmanii* (from coastal dunes of Brazil; Delprete 2005) and *C. naiguatensis* (from coastal cordillera of Venezuela; Steyermark, 1972a) on a strongly supported clade, with two species of *Salzmannia* (from coastal dunes of Brazil; Jardim et al. 2015). Hence, they transferred the two *Chiococca* species to *Salzmannia*, providing the necessary new combinations.

*Salzmannia sensu* Paudyal et al. (2018) is a genus of four species, with one growing on the costal cordillera of Venezuela, and other three growing on the coastal dunes, restinga, and forested vegetation of north-eastern Brazil (Delprete 2005 ["2004"); Jardim et al. 2015; Paudyal et al. 2018).

**Salzmannia** DC., Prodr. 4:617. 1830; Steyermark, Acta Bot. Venez. 6:135, fig. 5. 1972 ["1971"); Delprete, Rev. Biol. Neotrop. 1(1-2):4-10, f. 1. 2005 ["2004"); Jardim et al., Phytotaxa 202(10):15-25. 2015; emend. Paudyal et al., Bot. J. Linn. Soc. 187:390-391. 2018. **TYPE:** *Salzmannia nitida* DC.

1. *Salzmannia arborea* J.G. Jardim, Phytotaxa 202(1):20, fig. 1. 2015. **TYPE:** BRAZIL. BAHIA: Mun. Una, Reserva Biológica do Mico-Leão (IBAMA), entrada no km 46 da Rod. Ilhéus-Una, 15°9'S, 39°5'W, 21 Oct 1992, A.M. de Carvalho, A.M. Amorim, S.C. de Sant'Ana & J.G. Jardim 4085 (HOLOTYPE: CEPEC [CEPEC56640]; ISOTYPES: ALCB n.v., JPB [No. 62039], MO [MO-102917752], NY [00503586]).

**Distribution.**—Brazil (Bahia).

2. *Salzmannia naiguatensis* (Steyerm.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:391. 2018. *Chiococca naiguatensis* Steyerm., Acta Bot. Venez. 6:135, fig. 5. 1972 ["1971"). **TYPE:** VENEZUELA. DISTRITO FEDERAL: Cerro Naiguatá, cloud forest, N facing slopes, Loma de las Delicias, towards Quebrada Caja de Agua, W of Hacienda Coquizal, 1400 m, 8 Oct 1966, J.A. Steyermark 97475 (HOLOTYPE: VEN [No. 71606]; ISOTYPES: MO [No. 2270191, NY [00099437], U [U0005960]]).

**Distribution.**—Venezuela (Distrito Federal).

3. *Salzmannia nitida* DC., Prodr. 4:617. 1830. **TYPE:** BRAZIL. BAHIA: "ad bahiam in colibus aridis," s.d. [1827-1830], *P. Salzmann s.n.* (HOLOTYPE: G-DC [G00667887]; POSSIBLE ISOTYPES: BM [0005516631], F [No. 617617], HAL [2 sheets, HAL0113337, HAL113876], HBG [HBG-521019], K [K000015313], MO [No. 1797023], MPU [5 sheets, MPU021514, MPU021515, MPU021516, MPU021517, MPU021518]).

**Distribution.**—Brazil (Bahia).

4. *Salzmannia plowmanii* (Delprete) Paudyal & Delprete, Bot. J. Linn. Soc. 187:391. 2018. *Chiococca plowmanii* Delprete, Rev. Biol. Neotrop. 1:5, fig. 1, 2005 ["2004"). **TYPE:** BRAZIL. BAHIA: Mun. Salvador, along rd (Av. Otávio Mangabeira = BA-033) from Itapuá to Aeroporto Dois de Julho, at traffic circle (intersection with Av. Luis Viana Filho), near sea level, relict area with high dunes, 24 Feb 1985, *T. Plowman & I.C. Britto* 13948 (HOLOTYPE: CEPEC [No. CEPEC00036435]; ISOTYPES: F [2 sheets, Nos. 1993896, 1944115], G [G00389940], MO [No. 3509664], NY [01365079], SP [SP001597], U [U0119465]).

**Distribution.**—Brazil (Bahia).
SCHMIDTOTTIA

Urban (1923a) segregated *Schmidtottia* from *Portlandia* because of its terminal inflorescence (vs. lateral in *Portlandia*), sheathing, truncate stipules (vs. interpetiolar, triangular), septicidal capsules (vs. loculicidal), and ovate to obovate placenta (vs. linear). He dedicated the generic name to Otto Schmidt (1891–1956), a Russian scientist.

Wernham (1913:320), published *Portlandia involucrata* Wernham, and Alain Liogier (1959:1959), transferred it to *Schmidtottia involucrata* (Wernham) Alain. Aiello (1979) transferred *S. involucrata* to the monotypic genus *Ceuthocarpus* Aiello, because of its acropetally imbricated seeds and solitary flowers surrounded by an involucre of leafy bracts.

Paudyal et al. (2018), in their molecular phylogenetic analyses, found the seven species of *Schmidtottia* included in the study on a strongly supported clade along with *Ceuthocarpus involucratus*, hence they returned to *Ceuthocarpus* to *Schmidtottia*, in agreement with Alain (1959). Several new species and subspecies were published by Borhidi and Acuña (1971), Borhidi and Muñiz (1975), and Borhidi (1977, 1980, 2002). Liogier (1962) produced a key to the nine Cuban species of *Schmidtottia* known to him. Borhidi et al. (2018) published a synopsis of *Schmidtottia*, where they recognized 17 species. In the present treatment, *Schmidtottia* is treated as a genus of 11 species endemic to Cuba.

*Schmidtottia* Urb., Symb. Antill. 9:137. 1923; Liogier, Fl. Cuba 5:29–32. 1962; Aiello, J. Arnold Arbor. 60:108–109. 1979; Borhidi, Acta Bot. Hung. 29:181–216. 1981 [“1980”]; Borhidi, Acta Bot. Hung. 44(1–2):49–65. 2002; Borhidi et al., Rubiáceas Cuba 371–386, f. 111-16. 2017; Paudyal et al, Bot. J. Linn. Soc. 187(3):365–396. 2018; Borhidi et al, Acta Bot. Hung. 60(3–4):295–299. 2018. TYPE: *Schmidtottia monantha* Urb.

*Ceuthocarpus* Aiello, J. Arnold Arbor. 60:108. 1979. TYPE: *Ceuthocarpus involucratus* Aiello [= *Schmidtottia involucrata* (Wernham) Alain]

1. *Schmidtottia cubensis* (Standl.) Urb., Symb. Antill. 9:139. 1923.

1a. *Schmidtottia cubensis* ssp. *cubensis*, Symb. Antill. 9:139. 1923. *Isidorea cubensis* Standl., N. Amer. Fl. 32:15. 1918.
TYPE: CUBA. HOLGUIN: Sierra de Nipe, Arroyo del Medio, above the falls, 450–550 m, 20 Jan 1910, A. Shafer 3230 (LECTOTYPE, here designated: NY [00115156]; ISOLECTOTYPES: HAC, NY [2 sheets, 00115154, 00115155], US [00026700 (frag. ex NY)]).

*Distribution*.—Cuba (Holguín: Sierra de Nipe, Sierra del Cristal).

*Notes*.—Standley (1918:15), in the protologue of *Isidorea cubensis* Standl., cited the material studied as “Type collected among rocks, near water, Arroyo del Medio, [...], J.A. Shafer 3230 (herb. N. Y. Bot. Gard.).” In NY there are three specimens of Shafer 3230.

Borhidi (2002:55) cited the type of *I. cubensis* as “Holotipo: Cuba, Sierra de Nipe, [...] leg.: A. Shafer, 3230, 20.01.1910, NY!; isotypo: HAC.” Borhidi et al. (2017:373; 2018:298) cited the type of *I. cubensis* as “Tipo: Cuba, Sierra de Nipe, Arroyo del Medio [...], A. Shafer 3230 Holotipo: NY!, isotypo HAC!” According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi’s (2002) and Borhidi et al.’s (2017, 2018) “holotype” citations cannot be corrected as inadvertent lectotype designations.

Each NY specimen, with barcodes 00115154, 00115155, and 00115156, has a label with the annotation “Isidorea?” handwritten by an unknown author, and “cubensis Standley” handwritten by Standley. The specimen with barcode 00115156, consists of two branches, one with numerous leaves and a few capsules, and the other leafless, with a dehisced capsule. This specimen is here designated as the lectotype of *Isidorea cubensis*.

1b. *Schmidtottia cubensis* ssp. *cristalensis* (Borhidi & Muñiz) Borhidi, Acta Bot. Hung. 44(1-2):56. 2002.
*Schmidtottia cubensis* var. *cristalensis* Borhidi & Muñiz, Acta Bot. Acad. Sci. Hung. 21:230. 1975. TYPE: CUBA. HOLGUIN: Sierra del Cristal, Loma El Serrucho, 2-7 Apr 1956. Hno. Alain [A.H. Liogier], J.B. Acuña, & M. López Figueiras 5402 (LECTOTYPE, here designated: HAC [annotated as "holotypus" by Borhidi]; ISOLECTOTYPES: HAC, NY [00126819]).

*Distribution*.—Cuba (Sierra del Cristal).

*Notes*.—Borhidi & Muñiz (1975:230), in the protologue of *Schmidtottia cubensis* var. *cristalensis* Borhidi & Muñiz, did not specify which of the HAC specimens is the holotype. The HAC specimen with the
annotation "var. cristalensis Borhidi HOLOTYPUS!" handwritten by Borhidi, is here designated as the lectotype of this varietal name.

**2. Schmidtottia cucullata** Borhidi & Bisse, in Borhidi, Acta Bot. Acad. Sci. Hung. 26:271, fig. 8. 1980. **TYPE:** CUBA. HOLGUÍN: Sierra de Moa, Cayo Coco, charrascales, 200-300 m, 13 Aug 1970 (II). *J. Bisse & H. Lippold HFC No. 17618* (HOLOTYPE: HAJB [HAJB G 000839]; **ISOTYPES:** HAJB [HAJB G 000840], JE [JE00004996]).

*Distribution*.—Cuba (Holguín: Sierra de Moa).

**3. Schmidtottia elliptica** (Britton) Urb, Symb. Antill. 9:139. 1923. *Portlandia elliptica* Britton, Bull. Torrey Bot. Club 41:22. 1914. **TYPE:** CUBA. GUANTÁNAMO: Between Baracoa and Florida, on serpentine rocks, 15 Mar 1910. *J.A. Shafer 4332* (**LECTOTYPE**, here designated: NY [00126746]; **ISOLECTOTYPES:** A [2 sheets, 00058981, 00105431], HAC, NY [00126747], US [00137326]).

*Distribution*.—Cuba (Guantánamo; Holguín: Sierra de Moa).

**Notes.**—Britton (1914:22), in the protologue of *Portlandia elliptica* Britton, cited the material examined as "Thickets on serpentine rocks, between Baracoa and Florida, Oriente, Cuba, March 15, 1910 (*Shafer* 4332)," without citing the herbarium of deposit. Borhidi (2002:57) and Borhidi et al. (2017:376; 2018:298) cited as the holotype of *Portlandia elliptica* a specimen in NY. According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi's (2002) and Borhidi et al.'s (2017, 2018) "holotype" citations cannot be treated as inadvertent lectotypifications.

There are two specimens of *Shafer 4332* in NY, and one in US. The NY specimen with barcode 00126746 has a label with "Portlandia" handwritten by an unknown author, and "elliptica Britton, Type" handwritten by Britton. This specimen consists two branches with several leaves, numerous flower buds, and two flowers in anthesis, and is here designated as the lectotype of *P. elliptica*.

The NY specimen with barcode 00126747 has a label with "Portlandia" handwritten by unknown author, and "elliptica Britton" handwritten by Britton. This specimen consists two branches with several leaves and numerous dehisced capsules, and is an isolectotype.

*Schmidtottia elliptica* ssp. *oblongata* Borhidi, Acta Bot. Acad. Sci. Hung. 26:273. 1980. **syn. nov.** **TYPE:** CUBA. HOLGUÍN: Moa, Cayoguán, camino hacia la Mina Delta, 15-16 Jul 1949, *Bro. Clemente, Bro. Alain & Bro. Chysogone 6829* (HOLOTYPE: HAC [*Bro. Clemente et al.* 6829]; **ISOTYPES:** HAC [*Bro. Alain* 929], NY [00126820 (*Bro. Alain & Bro. Clemente* 929)]).

**4. Schmidtottia involucrata** (Wernham) Alain, Contr. Ocas. Mus. Hist. Nat. Colegio De la Salle 17:10. 1959. *Portlandia involucrata* Wernham, J. Bot. 51:320. 1913. *Ceuthocarpus involucratus* (Wernham) Aiello, J. Arnold Arbor. 60:109. 1979. **TYPE:** CUBA. HOLGUÍN: Camp La Gloria to Moa Bay, 31 Dec 1910-1 Jan 1911, *J.A. Shafer 8282* (HOLOTYPE: NY [00126750]; **ISOTYPES:** A [00219729], F [No. 450905], GH [00058980], MO [No. 805437], US [00137330]).

*Distribution*.—Cuba (Holguín: Moa).

*Ceuthocarpus involucratus* var. *moaensis* Borhidi, Acta Bot. Hung. 44(1-2):51. 2002. *Schmidtottia involucrata* var. *moaensis* (Borhidi) Borhidi, Acta Bot. Hung. 60:298. 2018. **syn. nov.** **TYPE:** CUBA. HOLGUÍN: Moa, orillas del Río Cayoguán, cerca de la mina, 2 Jul 1945, *Bro. Clemente, Bro. Alain & Bro. Chysogone 4485* (HOLOTYPE: HAC n.v.; **ISOTYPE:** NY n.v.).

*Ceuthocarpus involucratus* var. *elatior* Borhidi, Acta Bot. Hung. 44(1-2):51. 2002. *Schmidtottia involucrata* var. *elatior* (Borhidi) Borhidi, Acta Bot. Hung. 60:298. 2018. **syn. nov.** **TYPE:** CUBA. HOLGUÍN: Baracoa, charrascos en el valle del Río Maraví, *J. Bisse HFC No. 18240* (HOLOTYPE: HAJB n.v.).

**Notes.**—As a result of a detailed search in HAJB, no original material *Ceuthocarpus involucratus* var. *elatior* Borhidi was found. The HAJB curator, Eldis R. Bécquer (pers. comm.), is not aware of the possible location of the type specimens of this name. According to him, all the Rubiaceae specimens that were on loan to JPU, for study by Attila Borhidi, were returned to HAJB.

**5. Schmidtottia marmorata** Urb., Symb. Antill. 9:140. 1923. **TYPE:** CUBA. GUANTÁNAMO: Prope Baracoa, ad Rio Yoa, in charrascales, 28 Nov 1914, *E.L. Ekman 3675* (**LECTOTYPE**, here designated: S [No. S-R-7819]; **ISOLECTOTYPES:** HAC [ex LS], NY [00126821 (frag)]).

*Distribution*.—Cuba (Guantánamo: Toa, Baracoa).

**Notes.**—Urban (1923a:140-141), in the protologue of *Schmidtottia marmorata* Urb., cited the gathering "Prov. Oriente prope Baracoa ad Rio Yoa in charrascales, m. Nov. fruct.: [Ekman] n. 3675," without indicating the herbarium of deposit. The original material studied by Urban in B was destroyed during WWII. Borhidi (2002:58) cited the type of this name as "*Holotipo*: Cuba, Prov. Oriente, [...] leg: E L. Ekman, 3675, 28.11. 1914, S; *isotipo*: HAC." Borhidi et al. (2017:378; 2018:298) cited the type of this name as "Tipo: Cuba, Prov.
Oriente, [...] 28.11.1914, E.L. Ekman 3675; Holotipo: B†. Lectotipo: S!, isolectotipo: HAC! According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi (2002) "holotype" citation, and Borhidi et al. (2017, 2018) "lectotype" citations are not valid lectotypifications. In S there is a specimen of *Ekman* 3675, with accession number S-R-7819, which has a label with the annotation "Schmidtottia marmorata Urb. (typus)" handwritten by Urban. The specimen consists of a branch with numerous leaves, several infructescences with dehisced capsules, and is here designated as the lectotype of *S. marmorata*.

**5. Schmidtottia monantha** Urb, Symb. Antill. 9:138. 1923. **TYPE:** CUBA. GUANTÁNAMO: near Maraví, in a pine grove, 25 Dec 1914, E.L. Ekman 4028 (*LECTOTYPE*, **here designated:** S [No. S-R-7818]; *ISOLECTOTYPES:* HAC [ex LS], NY [00126823 (frag.)]).

*Distribution*.—Cuba (Holguín: Peña Prieta, Toa; Guantánamo: Maraví).

**Notes.**—Urban (1923a:138–139), in the protologue of *Schmidtottia monantha* Urb., cited the material studied as "Prov. Oriente prope Maraví in pinetis, m. Dec. fl. et fr.: [Ekman] n. 4028 (typus), inter Tabo et Nibujón in charrascales-pinales locis humilibus, m. Dec. fl. et fr.: [Ekman] n. 3741," without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. Borhidi (2002:58) cited the type of this name as "*Holotipo*: Cuba, Prov. Oriente, pinares del Rio Mayarí, leg.: E L. Ekman, 4028, 25.12.1914, S; *isotipo*: HAC." Borhidi et al. (2017:378; 2018:298) cited the type of this name as "[...] Pinares del Rio Maraví, 25.12.1914, leg. E.L. Ekman 4028, Holotipo: B†; lectotipo: S!, isolectotipo: HAC!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi's (2002) "holotype" citation cannot be treated as an inadvertent lectotype citation, and Borhidi et al. (2017, 2018) "lectotype" designations are not valid.

In S there is a specimen of *Ekman* 4028, with accession number S-R-7818, which has a label with the annotation "Schmidtottia monantha Urb. (typus)" handwritten by Urban. The specimen consists of three branches, two of them with numerous leaves and without flowers, and a small branch with a flower in anthesis. Specimen with accession number S-R-7818 is here designated as the lectotype of *S. monantha*.

*Schmidtottia parvifolia* Alain, Contr. Ocas. Mus. Hist. Nat. La Salle 17:10. 1959, **syn. nov.** **TYPE:** CUBA. HOLGUÍN: Toa, Peña Prieta, 600 m, 30 Dec 1953 (fl-fr). *Bro. Alain* [A. Liogier] LS No. 3592 (*HOLOTYPE:* HAC [ex LS]; *ISOTYPE:* NY [00126825]).

**Notes.**—Alain (1959: 10) in the protologue of *Schmidtottia parvifolia* Alain, cited his own gathering *Alain* 3592 in LS, which was later integrated in HAC. That specimen is the holotype of this name.

**6. Schmidtottia multiflora** Urb., Symb. Antill. 9:141. 1923. **TYPE:** CUBA. ORIENTE: Sierra Azul, in charrascales, 500–700 m, 23 Jan 1915, E.L. Ekman 4422 (*LECTOTYPE*, **here designated:** S [No. S-R-7816]).

*Distribution*.—Cuba (Holguín, Guantánamo).

**Notes.**—Urban (1923a:141–142), in the protologue of *Schmidtottia multiflora* Urb., cited the material studied as "Prov. Oriente in Sierra Azul in charrascales, 500–700 m. alt., m. Jan. flor.: [Ekman] n. 4422," without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. Borhidi (2002:59) cited the type of this name as "*Holotipo*: Cuba, Prov. Oriente, [...], leg.: E L. Ekman, 4422, S!" Borhidi et al. (2017:379; 2018:298) cited the type of this name as "Cuba, Prov. Oriente, [...] leg.: E.L. Ekman 4422. Holotipo: B†; lectotipo: S!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi's (2002) "holotype" citation cannot be treated as an inadvertent lectotype citation, and Borhidi et al.'s (2017, 2018) "lectotype" designations are not valid.

In S there is a specimen of *Ekman* 4422, accession number S-R-7816, which has a label with the annotation "Schmidtottia multiflora Urb. (typus)" handwritten by Urban. It consists of a branch with a terminal inflorescence and numerous flower buds. This specimen is here designated as the lectotype of *S. multiflora*.

**7. Schmidtottia nitens** (Britton) Urb., Symb. Antill. 9:141. 1923. *Portlandia nitens* Britton, Bull. Torrey Bot. Club 39:10. 1912. **TYPE:** CUBA. GUANTÁNAMO: Baracoa, Upper Valley of Rio Navas, 22 Mar 1910, J.A. Shafer 4450 (*LECTOTYPE*, **here designated:** NY [00126756]; *ISOLECTOTYPES:* HAC, NY [00126757], US [00137332 (one corolla and a leaf fragment (ex NY))]).

*Distribution*.—Cuba (Guantánamo: Baracoa).

**Notes.**—Britton (1912:10), in the protologue of *Portlandia nitens* Britton, cited the material studied as
<figure><img src="image_7.png" /><figcaption>FIG. 4. Lectotype of *Schmidtottia corymbosa* Borhidi (Bro. León & Bro. Clemente HFC No. 23224, HAC [ex LS]). = *Schmidtottia uliginosa* (Wernham) Urb. Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
as a genus of 28 species occurring in the Bahamas, and Greater and Lesser Antilles, with a high number of them endemic to Cuba.

**Scolosanthus** Vahl, Eclog. 1:11. 1796; Standley, N. Amer. Fl. 32(4):293-297. 1934; Liogier, Fl. Cuba 5:95-98. 1962; Correll & Correll, Fl. Bahama Arch. 1418, f. 620. 1982; Borhidi, Nord. J. Bot. 3:351-354. 1983; Borhidi, Acta Bot. Hung. 29:197-204, f. 2-7. 1983; Liogier, Fl. Española 7:414-419, f. 198-51. 1995; Borhidi et al., Rubiáceas Cuba 388-404, f. 118-124. 2017. TYPE: *Scolosanthus versicolor* Vahl [based on *Catesbaea parviflora* Lam. (1792), non Swartz (1788)].

**1. Scolosanthus acanthodes** (Spreng.) Urb., Symb. Antill. 1:481. 1899. *Eranthemum acanthodes* Spreng., Syst. Veg. 1:88. 1825. (See Standley, N. Amer. Fl. 32:297. 1934). TYPE: DOMINICAN REPUBLIC. PUERTO PLATA: Mun. Puerto Plata, comunidad El Toro, paraje Palo Indio, small mountain with serpentine soil, Finca del Señor Nicolas Domínguez, 19°51'N, 70°49'W, 50-150 m, 1 Aug 2001, P.G. Delprete & T. Clase 7645 (*NEOTYPE*, here designated: NY [05154375]; *ISOTYPES*: CAY [CAY253111], FTG [without barcode], JBSD [without barcode], MO [without barcode], US [without barcode]).

**Distribution.**—Hispaniola.

**Notes.**—Sprengel (1825:88), in the protologue of *Eranthemum acanthodes* Spreng, indicated the collection locality as “Hispaniola,” without citing any specimen or herbarium of deposit. According to Stafleu and Cowan (1985:806), Kurt Sprengel’s herbarium was “dismembered and sold in small portions,” and numerous specimens can be found in B-W, BP, FI, G, GOET, LE, PH, and PR. Urban (1899:481) along with the new combination *Scolosanthus acanthodes* (Spreng.) Urb., cited the material examined as “Hab. in Santo Domingo: Bertero n. 580,” without indicating the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. After a search in Jstor Global Plants, Jacq, and in the TO herbarium, I did not find any original specimen associated with this name. Hence it is necessary to designate a neotype. The gathering *Delprete & Clase 7645* has duplicates distributed in several herbaria, and has flowers in anthesis and mature fruits. The specimen in NY is here designated the neotype of *E. acanthodes* Urb.

**2. Scolosanthus acunae** Borhidi & Muñiz, Acta Bot. Hung. 29:197, fig. 2. 1983. TYPE: CUBA. PINAR DEL RÍO: Loma Cajálbana, in declivibus abruptis orienti-septentrionalibus, fruticetis sempervirens, solo latosol serpentino, 400 m, 21 Nov 1974, A. Borhidi, E. Del-Risco & R. Capote HFC No. 27694 (HOLOTYPE: HAC [ex SV No. 27815]; ISOTYPE: BP n.v.).

**Distribution.**—Cuba (Pinar del Río: Sierra de Cajálbana).

**Notes.**—Borhidi and Muñiz (in Borhidi 1983:197-198), in the protologue of *Scolosanthus acunae* Borhidi & Muñiz, cited the type as “Holotypus: Cuba, Prov. Pinar del Río, Loma cajálbana; [...] A. Borhidi et R. Capote, 21 Nov. 1974, flor. et fruct. No. 27694 SV! Isotypi: SV! et BP!” The SV specimens are now integrated in HAC. There is only one specimen in HAC, ex SV No. 27815, which has a label with the annotation “Scolosanthus acunae Borhidi et Muñiz, Falda NE de Cajálbana, Pinar del Río, Borhidi, Del-Risco, Capote, 21/Nov. 1974” handwritten by Borhidi. This specimen is the holotype of this name.

**3. Scolosanthus bahamensis** Britton, Bull. New York Bot. Gard. 3:452. 1905. TYPE: BAHAMAS: New Providence, from Coppice along Village Road, 30 Aug 1904, N.L. Britton & L.J.K. Brace 367 (HOLOTYPE: NY [00115373]; ISOTYPES: F [No. 171801], K [K000432632], US [00138531]).

**Distribution.**—Bahamas Archipelago.

**4. Scolosanthus crucifer** C. Wright, Anales Acad. Ci. Méd. Fís. Nat. Habana 6:126. 1869.

**4a. Scolosanthus crucifer** ssp. *crucifer* C. Wright, Anales Acad. Ci. Méd. Fís. Nat. Habana 6:126. 1869. TYPE: CUBA: Lagunas de Guama ... chio (?), 3 Aug 1865, C. Wright 377 (LECTOTYPE (Borhidi 1983:198): GOET [GOET010511]; ISOTYPES: HAC n.v. [lost?], K [K000432631], NY [00115374], S n.v. [not there?], US [00433459]).

**Distribution.**—Cuba (Pinar del Río, Holguín, Villa Clara, Matanzas).

**Notes.**—Wright (1869:126), in the protologue of *Scolosanthus crucifer* C. Wright, cited two of his own gatherings from Cuba, Wright 377 and Wright 3583, and cited “Randia Sagreana Gris.” Borhidi (1983:198) cited the type of this name as “Holotypus: Cuba; Ch. Wright 377; Lagunas of Guama ... Chio (?) 3. aug. 1865. GOET—isotypi S! NY! HAC! US!” This citation is an inadvertent lectotypification of a specimen at GOET. In GOET there is a single specimen of Wright 377, with barcode GOET010511. It has a label with a short species description, handwritten by Wright. Another label has the heading “Plantae Cubenses Wrightianae” and the
number "377" handwritten by an unknown author. The specimen consists of three small branches with numerous minute leaves, and is the lectotype of *S. crucifer*.

*Scolosanthus crucifer* var. *acutus* Borhidi, Acta Bot. Hung. 29:198. 1983. **syn. nov.** TYPE: CUBA. VILLA CLARA: LasVillas, Sabanas de Motembo, 9-10 Aug 1930, *Bro. León* & A. *Loustalot* 9340 (HOLOTYPE: HAC!; ISOTYPES: GH [00094505], NY [01185462]).

*Scolosanthus crucifer* var. *subtomentosus* Borhidi, Acta Bot. Hung. 29:198. 1983. **syn. nov.** TYPE: CUBA. MATANZAS: San Adrián, Cuabal de Espinal, 11 Apr 1927, *J.T. Roig* & *Bro. León* 4245 (HOLOTYPE: HAC [ex LS]; ISOTYPE: NY n.v.).

**Notes.**—In GH there is the specimen *Bro. León* LS No. 12971, barcode 00094506, collected on 28 February 1972, annotated as isotype of *Scolosanthus crucifer* var. *subtomentosus* Borhidi by Borhidi, but the type of this name is *Roig & León* 4245, as cited by Borhidi (1983:198).

Borhidi (1983:198) recognized four varieties under *Scolosanthus crucifer*: 1) var. *crucifer*, with leaves oblong-elliptic, 7-15 mm long, obtuse or round at apex, glabrous throughout; 2) var. *acutus* Borhidi, with leaves lanceolate, 7-15 × 5-8 mm, shortly acuminate and acute-mucronate at apex, glabrous throughout; 3) var. *microphyllus* Borhidi, with leaves minute, elliptic or suborbicular, 2-6 × 1.5-4 mm, glabrous throughout, margins strongly revolute; 4) var. *subtomentosus* Borhidi, with leaves elliptic, 7-12 mm long, shortly tomentulose below. The typical variety and the varieties *acutus* and *subtomentosus* have overlapping characters, in terms of leaf dimensions and vestiture. Whereas, variety *microphyllus* is distinct from the other varieties in having minute leaves with revolute margins, and is here elevated to subspecies rank (See below).

**4b. Scolosanthus crucifer** ssp. *microphyllus* (Borhidi) Delprete, **comb. et stat. nov.** *Scolosanthus crucifer* var. *microphyllus* Borhidi, Acta Bot. Hung. 29:198. 1983. TYPE: CUBA. Matanzas: Canasi, Cuabal de Espinal, 16-18 Aug 1927, *Bro. León* & *J.B. Acuña* LS No. 13066 (HOLOTYPE: HAC [ex LS]; ISOTYPES: GH [00094504], NY [01185463]; POSSIBLE ISOTYPE: HAC [ex SV No. 24693]).

**Distribution.**—Cuba (Matanzas, Villa Clara).

**Notes.**—Borhidi (1983:198) cited the type of *Scolosanthus crucifer* var. *microphyllus* Borhidi as "Holotypus: León 13066 HAC; Prov. Matanzas. Cuabal del Espinal, Canasi; Leg.: León et Acuña 16-18 Aug 1927.— Isotypus: Acuña 24693, HAC!" The holotype specimen, *León & Acuña* 13066, was originally in LS, and is now integrated in HAC.

In HAC there is a second specimen, which was originally in SV, which has a label with herbarium number "24693" and the typewritten annotations "Scolosanthus crucifer Wr. ex Sauv., Cuabal del Espinal, Canasi, Prov. Matanzas, Acuná y León," without collection date. Just above that label is affixed another label with the annotation "Scolosanthus crucifer var. microphyllus Borhidi, A. Borhidi, 1979" handwritten by Borhidi. Its label reports the same locality and the same collectors as the holotype specimen, but reports the gathering as "Acuña 24693," hence it is as a possible isotype.

**5. Scolosanthus densiflorus** Urb., Symb. Antill. 3:381. 1903.

**5a. Scolosanthus densiflorus** Urb. ssp. *densiflorus*, Symb. Antill. 3:381. 1903. TYPE: HAITI: Gonaïves, Morne Bonpère, 600 m, Jun 1901, *W. Buch* 728 (LECTOTYPE, **here designated**: K [K000432630]).

**Distribution.**—Haiti (Gonaïves).

**Notes.**—Urban (1903:381), in the protologue of *Scolosanthus densiflorus* Urb., cited the material studied as "Haiti prope Gonaïves, Morne Bonpère, 600 m. alt., m. Junio flor.: Buch n. 728," without citing the herbarium of deposit. The original material in B studied by Urban was destroyed during WWII. Borhidi et al. (2017:392) cited the type of this name as "Tipo: Haiti, prope Gonaïves in Morne monpère, 600 m, *Buch* 728, holotipo: B†, isotipos: NY, US." In K there is a specimen, with barcode K000432630, with the heading "HERBARIUM KRUG ET URBAN" and the annotation "Scolosanthus densiflorus Urb." handwritten by Urban. This specimen consists of two small branches, one of them with flower buds. In the envelope affixed on the sheet are included several leaves, several flower buds, and a flower in anthesis. This specimen is here designated as the lectotype of this name.

**5b. Scolosanthus densiflorus** subsp. *maestrensis* (Alain) Borhidi, Bot. Közlem. 58:177. 1971. *Scolosanthus maestrensis* Alain, Contr. Ocas. Mus. Hist. Nat. La Salle 17:11. 1959. TYPE: CUBA. GRANMA: Sierra Maestra, Alto de la Valenzuela, mérgenes del Arroyo Peladero, 5-8 Apr 1955, *M. López Figueiras* 2181 (HOLOTYPE: HAC [ex SV]; ISOTYPES: HAJB [HAJB G 000843], NY [00115394], US [0013535]).

**Distribution.**—Cuba (Granma: Sierra Maestra; Guantánamo: Mesa de Prado; Santiago de Cuba).
6. **Scolosanthus ekmanii** Borhidi, Nord. J. Bot. 3:351, fig. 1. 1983. **TYPE:** CUBA. SANTIAGO DE CUBA: Daiquiri, "in collibus calcareis and Papayo [Loma del Papayo]," 18 Nov 1916, E.L. *Ekman* 8391 (*HOLOTYPE:* S [No. S05-1021]; **ISOTYPES:** F n.v., UPS n.v., US [00433461]).

*Distribution*.—Cuba (Santiago de Cuba).

7. **Scolosanthus grandifolius** Krug & Urb., in Urban, Symb. Antill. 1:442. 1899. **TYPE:** PUERTO RICO: Prope Maricao, Monte Alegrillo, 7 Dec 1884, P.E.E. *Sintenis* 249 (*LECTOTYPE*, **here designated:** K [K000432629NY]; **ISOLECTOTYPES:** GH [00094508], HGB [HGB-521009], NY [00115376], US [00138532]).

*Distribution*.—Puerto Rico.

*Notes*.—Krug and Urban (in Urban, 1899:442–443), in the protologue of *Scolosanthus grandifolius* Krug & Urb., cited the material studied as “in Portorico prope Maricao in sylvis montis Alegrillo, m. Dec. flor.: *Sintenis* n. 249” without citing the herbarium of deposit. The material in B studied by Krug and Urban was destroyed during WWII. Duplicates of *Sintenis* 249 are present in several herbaria. All of them have labels with the printed annotation “det. I. Urban.” The K specimen with barcode K000432629, consist of two branches with medium-sized leaves, and long, curved thorns, and is here designated as the lectotype of this name.

8. **Scolosanthus granulatus** Urb., Symb. Antill. 9:164. 1923. **TYPE:** CUBA. GUANTÁNAMO: Monte Libanon, San Fernández, 21 Dec 1919, E.L. *Ekman* 10283 (*LECTOTYPE*, **here designated:** S [No. S05-1025]; **ISOLECTOTYPE:** HAC [No. 28096, ex S]).

*Distribution*.—Cuba (Guantánamo: Monte Libano).

*Notes*.—Urban (1923a:164), in the protologue of *Scolosanthus granulatus* Urb., cited the material studied as “[Cuba] Prov. Oriente prope Guantánamo in Monte Libanon ad San Fernández in solo calcareo, m. Dec. flor.: [Ekman] N. 10283,” without citing the herbarium of deposit. The material in B studied by Urban was destroyed during WWII. Borhidi et al. (2017:393) cited the type of this name as “Tipo: Cuba, Oriente prope Guantánamo, [...] *Ekman* 10283. Holotipo: B†, lectotipo: S!” According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al.’s (2017) lectotipo designation is not valid. In S there is a specimen, with accession number S05-1025, which has a label with the annotation “Scolosanthus granulatus Urb. (typus)” handwritten by Urban. It consists of two unarmed branches with numerous minute leaves, and without flowers. This specimen is here designated as the lectotype of this name.

9. **Scolosanthus hirsutus** Borhidi, Acta Bot. Hung. 29:198. 1983. **TYPE:** CUBA. GUANTÁNAMO: Costa Sur de Baracoa, San Antonio del Sur, 4 km W del pueblo, 200–400 m, 10 Feb 1976, A. *Areces*, J. *Bisse*, J. *Gutiérrez & H. Manitz* HAJB No. 29922 (*HOLOTYPE:* HAJB [HAJB G 000842]; **ISOTYPES:** BP n.v., HAC [ex SV], JE [JE00006001]).

*Distribution*.—Cuba (Guantánamo: Baracoa).

10. **Scolosanthus hispidus** Borhidi, Acta Bot. Hung. 29:199, fig. 3. 1983. **TYPE:** CUBA. GUANTÁNAMO: Maisí, coastal thickets, Jul 1938, Bro. *León & W. Seifriz* HAC No. 18110 (*HOLOTYPE:* HAC [ex SV]; **ISOTYPE:** NY [00115377]).

*Distribution*.—Cuba (Guantánamo: Mesa de Maisí).

11. **Scolosanthus howardii** Borhidi, Acta Bot. Hung. 29:199, fig. 4. 1983. **TYPE:** JAMAICA. TRELAWNY: Cockpit Country, dry rocky hillsides, 500 m, 4 Jul 1956, R.A. *Howard & G.R. Proctor* 14386 (*HOLOTYPE:* A [00094509]).

*Distribution*.—Jamaica.

*Notes*.—Borhidi (1983:199–200), in the protologue of *Scolosanthus howardii* Borhidi, cited the type as “Holotypus: Jamaica, Prov. Trelawny Ramgoat Cave district, [...] Leg. R.A. Howard and G.R. Proctor 14386, 4 Jul. 1956,” without citing the herbarium of deposit. In A there is a sole specimen of *Howard & Proctor 14386*, which was annotated as “Holotypus! Scolosanthus howardii Borhidi” by Borhidi in 1980. That specimen is the holotype of this name.

12. **Scolosanthus leonardii** Alain, Brittonia 20:160. 1968. **TYPE:** HAITI: vicinity of Mole St. Nicolas, road up banks of Mole gorge to Bombardopolis, 13–19 Feb 1929, E.C. *Leonard & G.M. Leonard* 13200 (*HOLOTYPE:* US [00138533] (annotated as “isotype” by Borhidi]); **ISOTYPES:** GH [00094510], NY [00115390] (annotated as “holotype” by Borhidi)).

*Distribution*.—Haiti.
**Notes.**—Alain Liogier (1968:160–161), in the protologue of *Scolosanthus leonardii* Alain, clearly stated that the type specimen of this name is in US, which is the holotype. Borhidi erroneously annotated the US specimen as the isotype, and the NY specimen as the holotype.

13. **Scolosanthus liogieri** Borhidi, Acta Bot. Hung. 29:200. 1983. **TYPE:** DOMINICAN REPUBLIC. SAN JUAN: Loma El Campanario, Ciénaga de la Culata, in cloud forest, 1650-1850 m, 24 Sep 1969, *Bro. Alain H. Liogier 16051* (HOLOTYPE: NY [00115392 (annotated as "isotype" by Borhidi]); **ISOTYPE:** US [00433462 (annotated as "holotype" by Borhidi)]).

**Distribution.**—Dominican Republic.

**Notes.**—Borhidi (1983:200–201) cited the type of *Scolosanthus liogieri* Borhidi, as "Holotypus: Alain 16051. Hispaniola, Rep. Dominicana; Loma El Campanario, Ciénaga de la Culata, [...] NY! Isotypus: US!" However, he annotated the NY specimen as isotype, and the US specimen as the holotype. According to the *Code*, the information presented in the publication regarding type designation is the valid one, and annotations on specimens have no nomenclatural validity.

14. **Scolosanthus lucidus** Britton, Mem. Torrey Bot. Club 16:112. 1920. **TYPE:** CUBA. HOLGUÍN: Moa Bay, E of Río Moa, 2–3 Jan 1911, *J.A. Shafer 8355* (LECTOTYPE, here designated: NY [00115393]; ISOLECTOTYPES: A [00094511], NY [00115392], US [00138534]).

**Distribution.**—Cuba (Holguín, Guantánamo).

**Notes.**—Britton (1920:112), in the protologue of *Scolosanthus lucidus* Britton, cited the type as "Moa Bay, east of Río Moa, Oriente (*Shafer 8355, type*)" without indicating the herbarium of deposit. Borhidi et al. (2017:396) cited the type of this name as "Tipo: Cuba; prov. Oriente, Moa Bay, [...], J.A. Shafer 8355. Holotipo: NY. Isotipo: US." According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017) lectotype designation is not valid.

In NY, where Britton worked, there are two specimens of *Shafer 8355*. The specimen with barcode 00115393, has a label with the annotation "Type. Scolosanthus lucidus Britton" handwritten by Britton. This specimen consists of one branch with numerous minute, lucid leaves and several flowers, and is here designated as the lectotype of *S. lucidus*.

The other NY specimen, with barcode 00115392, has a label with the annotation "Scolosanthus lucidus Britton" handwritten by Britton. This specimen consists of two branches with numerous minute, lucid leaves, and several flowers, and is an isolectotype.

15. **Scolosanthus moanus** Borhidi & Muñiz, Acta Bot. Hung. 29:201. 1983. **TYPE:** CUBA. HOLGUÍN: Moa, Breñales de Playa Vaca, 9 Nov 1945. *J. Acuña 13381* (HOLOTYPE: HAC [ex SV]; **ISOTYPE:** HAJB [HAJB G 000844]).

**Distribution.**—Cuba (Holguín: Moa).

16. **Scolosanthus multiflorus** (Sw.) Krug & Urb.

16a. **Scolosanthus multiflorus** ssp. **multiflorus**, in Urban, Symb. Antill. 1:443. 1899. *Ixora multiflora* Sw., Prodr. 30. 1788. **TYPE:** JAMAICA: without locality, s.d., *W.T. Marsh 1679* (NEOTYPE (Borhidi 1983: 201): GOET [GOET067223]; **ISONEOTYPE:** GH n.v.; POSSIBLE **ISONEOTYPE:** NY [00115395]).

**Distribution.**—Jamaica.

**Notes.**—Swartz (1788:30), in the protologue of *Ixora multiflora* Sw., indicated the collection locality as "Jamaica," without citing any specimen. Krug and Urban (1899:443) transferred this name to *Scolosanthus*, with the new combination *S. multiflorus* (Sw.) Krug & Urb. A general search in numerous herbaria did not retrieve any original specimen collected by Swartz in Jamaica.

Borhidi (1983:201) cited the type of *Scolosanthus multiflorus* ssp. *multiflorus* [Basionym: *Ixora multiflora* Sw.], as "Lectotypus: March, [Isic, W.T. Marsh] 1679: Jamaica GOET; Isolectotypus: GH." In absence of Swartz's original material, Borhidi's (1983) citation is here treated as an inadvertent neotype designation. According to Végter (1976:505), William Thomas Marsh collected about 2000 specimens in Jamaica in 1854–1862. In GOET there is a specimen, with barcode GOET067223, which has a label with the annotation "1679 Scolosanthus versicolor var. fol. u. Jamaica, Mr. Marsh." It has a second label with the annotation "Scolosanthus
multiflorus (Sw.) Krug et Urb." handwritten by Borhidi. It consists of six small branches with numerous leaves and numerous inflorescences with flowers in anthesis. This specimen is the neotype of *I. multiflora*.

In NY there is a specimen with barcode 00115395, which has a label with the annotations "Com. ex Hb. Hooker Kew 1866, Griseb!" handwritten by an unknown author, and "Scolosanthus versicolor Jamaica, Marsh" handwritten by Marsh. This specimen is a possible isotype of *Ixora multiflora*.

*Scolosanthus versicolor* Griseb., Fl. Brit. W. Ind. 335. 1861. *nom. illeg. superfl.* [non *S. versicolor* Vahl, Eclog. Amer. 1:11. 1797 ("1796")]
**Notes.**—Grisebach (1861:335), in the protologue of *Scolosanthus versicolor* Griseb., cited *S. versicolor* Vahl, and *Ixora multiflora* Sw., hence Grisebach's name is superflous and illegitimate.

**16b. Scolosanthus multiflorus** ssp. *hirticalyx* Borhidi, Acta Bot. Hung. 29:201. 1983. **TYPE:** JAMAICA: without locality, s.d., W.T. Marsh 1716 (HOLOTYPE: GOET [GOET067222]; ISOTYPE: GH [00094513]).

*Distribution.*—Jamaica.

*Notes.*—Borhidi (1983:201–202), in the protologue of *Scolosanthus multiflorus* ssp. *hirticalyx* Borhidi, cited a specimen as "Holotypus: March, [Isic, W.T. Marsh], 1716; Jamaica, GOET." In GOET there is a specimen, with barcode GOET067222, which has a label with the annotation "Scolosanthus versicolor V. Jamaica. Marsh 1716." It has a second label with the annotation "Scolosanthus multiflorus (Sw.) Krug et Urb. ssp. hirticalyx Borhidi" handwritten by Borhidi. This specimen consists of a branch with numerous leaves and numerous inflorescences with flowers in anthesis, and is the holotype of this subspecific name.

In GH there is a specimen, with barcode 00094513, which has a label with the annotation "Scolosanthus versicolor Vahl, Jamaica, Marsh" handwritten by Marsh. Above that label is affixed another label with the annotation "HOLOTYPUS! Scolosanthus multiflorus (Sw.) Kr. et Urb. ssp. hirticalyx Borhidi" handwritten by Borhidi. This specimen is an isotype of this name.

**17. Scolosanthus nannophyllus** Borhidi, Acta Bot. Hung. 29:202. 1983. **TYPE:** CUBA. GUANTÁNAMO: Manigua de la 2da terraza de Maisí, 19 Aug 1939, *Bro. León & Bro. Marie-Victorin* LS No. 17111 (HOLOTYPE: HAC [ex LS]; ISOTYPE: NY [00115396]).

*Distribution.*—Cuba (Guantánamo: Terraza de Maisí).

**18. Scolosanthus nipensis** Borhidi ex Delprete, sp. nov. (Fig. 5). **TYPE:** CUBA. HOLGUÍN: Sierra de Nipe, Charrascal de La Cueva, 27–31 May 1960, M. López Figueiras 1033 (HOLOTYPE: HAC [specimen "A"); ISOTYPE: HAC [specimen "B"]).

*Distribution.*—Cuba (Holguín: Sierra de Nipe).

*Diagnosis.*—*Scolosanthus nipensis* Borhidi ex Delprete is similar to *S. acunae* because of the minute leaves and the narrow corollas with tubes 2.5–3 mm long. The former differs from the latter by being unarmed (vs. with trichotomous thorns), calyx completely glabrous (vs. ciliate), corolla 4-merous (vs. commonly 3-merous, rarely 4-merous), and style 3.7 mm long (vs. 4.5–5 mm long).

*Description.*—*Shrub*, unarmed; height unknown. *Stipules* deltoid, 0.5–0.6 mm long, sometimes splitting into two narrowly triangular units. *Leaves* subsessile (petioles to 0.5 mm long); blades obovate, 8.5–9.5 x 3.5–4.5 mm, acute at base, round at apex, lucid and glabrous above, scabrellous below, the secondary veins not visible (imbedded in the lamina), the margins minutely revolute. *Flowers* axillary, 2–4 per axil, glabrous; pedicels 0.5–0.6 mm long; hypanthium oblong-obovoid, 1–1.2 mm long; calyx denticulate, teeth deltoid, 0.2 mm long. *Corolla* tube narrowly cylindrical at basal 2/3 and narrowly flaring at distal 1/3, glabrous, 3–3.2 mm long; lobes ovate-deltoid, 1.2 mm long, reflexed at anthesis. *Stamens* inserted about 1 mm below the corolla mouth; filaments 0.5 mm long; anthers linear, 1.3 mm long, apically mucronate. *Style* exserted, linear, 3.7 mm long. *Fruit* unknown.

*Notes.*—In HAC there are two specimens of López Figueiras 1033. The specimen annotated "A" (Fig. 5) by Delprete is the holotype, and the specimen annotated "B" by Delprete is the isotype of this name.

**19. Scolosanthus portoricensis** Borhidi, Acta Bot. Hung. 29:202. 1983. **TYPE:** PUERTO RICO: Sosua State Forest, ser-pentine barrens, 300–400 m, 26 Jun 1963, *Bro. Alain Liogier* [A.H. Liogier] 9742 (HOLOTYPE: GH [00094514]; ISOTYPE: NY [00115397]).

*Distribution.*—Puerto Rico.
<figure><img src="image_8.png" /><figcaption>FIG. 5. Holotype of *Scolosanthus nipensis* Borhidi ex Delprete, sp. nov. (López Figueiras 1033, HAC specimen "A"). Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
**20. Scolosanthus pycnophyllus** Borhidi, Acta Bot. Hung. 29:203. 1983. **TYPE:** CUBA. HOLGUÍN: Pico Sur del Cerro Galano, entre La Palma y Limones, 15 Aug 1975. *J. Urbino s.n.* [HAC No. 27983] (*HOLOTYPE:* HAC [ex EEAB No. 28115]).

*Distribution*.—Cuba (Holguín: Cerro Galano).

**Notes.**—Borhidi (1983:203) cited the type of *Scolosanthus pycnophyllus* Borhidi as "Holotypus: 27983 HAC. Cuba: Prov. Holguín (Oriente) Pico sur del Cerro Galeano, entre La Palma y Limones.—Leg.: J. Urbino, 15 Aug 1975." In HAC there is a sole specimen annotated as "*Scolosanthus pycnophyllus* Borhidi, HOLOTYPUS! A. Borhidi, 1977." The label of this specimen has the heading "ESTACION EXPERIMENTAL AGRONOMICA— HERBARIO CH. F. BARKER," which means that was originally in the EEAB Herbarium (Regalado Gabancho et al. 2008). It also has the handwritten annotations: "28115, [...] Pico del Cerro, Holguín, Oriente, Jacobo Urbino, 15 Aug 1975." This specimen was originally in EEAB, and is now integrated in HAC. It consists of two branches, with sturdy cruciform thorns, and numerous minute leaves, and is the holotype of *S. pycnophyllus*.

**21. Scolosanthus reticulatus** Borhidi, Acta Bot. Hung. 29:203. 1983. **TYPE:** CUBA. HOLGUÍN: Sierra de Nipe, Salto del Río Guayabo, Pinares de Mayari, 27–31 May 1960 (II), *M. López Figueiras* 973 (*HOLOTYPE:* HAC [ex LS]; *ISOTYPES:* BP n.v., HAJB [HAJB G 000845]).

*Distribution*.—Cuba (Holguín: Sierra de Nipe).

**22. Scolosanthus roulstonii** Proctor, Fl. Cayman Island, 2nd ed.:620, figs. 1–2. 2012. **TYPE:** CAYMAN ISLANDS: Grand Cayman, bluff at Little Salt Creek, 19 Nov 2005, *P.A. B. Stafford s.n.* (*HOLOTYPE:* IJ [IJ 000038672]).

*Distribution*.—Cayman Islands.

**23. Scolosanthus selleanus** Urb. & Ekman, in Urban, Ark. Bot. 20A:60. 1926. **TYPE:** HAITI: Massif de la Selle, Morne La Visite, on the slope of the siliceous conglomerate, 2150–2200 m, s.d., *E.L. Ekman H-1460* (*LECTOTYPE*, **here designated:** S [S05-1031]; *ISOLECTOTYPES:* G [G00436774], GH [00094515], K [K000432628], US [00827366]).

*Distribution*.—Haiti (Massif de la Selle).

**Notes.**—Urban and Ekman (in Urban, 1926:60–61), in the protologue of *Scolosanthus selleanus* Urb. & Ekman, cited the material studied as "Massif de la Selle in Morne La Visite in conglomeratis siliceis 2,150–2,200 m alt, m. Aug fl. et fr.: [Ekman] H 1460," without citing the herbarium of deposit. The material in B studied by Urban and Ekman was destroyed during WWII. In S there is a specimen of *Ekman H-1460*, with accession number S05-1031, which has a label with the annotation "Scolosanthus Selleanus Urb. et Ekm." handwritten by Urban. The specimen consists of three branches with numerous minute leaves, some flower buds, and a few fruits. This specimen is here designated as the lectotype of this name.

**24. Scolosanthus strictus** Urb, Symb. Antill. 9:533. 1928. **TYPE:** CUBA. GUANTÁNAMO: Prope Guantánamo, ad Caimanera, in limestone terraces, 24 Nov. 1922, *E.L. Ekman 15768* (*LECTOTYPE*, **here designated:** S [S05-1034]; *ISOLECTOTYPES:* F [No. 605137], G [G00436773], NY [00115399]).

*Distribution*.—Cuba (Guantánamo).

**Notes.**—Urban (1928:533), in the protologue of *Scolosanthus strictus* Urb., cited the material studied as "Prov. Oriente prope Guantánamo ad Caimanera in calcareis, m. Nov. in alab.: [Ekman] n. 15768," without citing the herbarium of deposit. The material in B studied by Urban was destroyed during WWII. Borhidi et al. (2017:401) cited the type of this name as "Tipo: Cuba: Oriente, [...], Ekman 15768. Holotipo: B†, lectotipo: S!" According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017) lectotype designation is not valid.

In S there is a specimen of *Ekman 15768*, with accession number S05-1034, which has a label with the annotation "Scolosanthus strictus Urb. (typus)" handwritten by Urban. This specimen, consisting of three branches with numerous minute leaves, and some flower buds, is here designated as the lectotype of this name.

**25. Scolosanthus subsessilis** Alain, Phytologia 25:278. 1973. **TYPE:** DOMINICAN REPUBLIC. SANTIAGO: Jaiquí Picado, 20 mi W of Santiago, 300–400 m, 26 May 1969, *Bro. Alain* [A.H. Liogier] 15368 (*HOLOTYPE:* NY [00115398]; *ISOTYPES:* GH [00094507], US [00433460]).

*Distribution*.—Dominican Republic.
**Notes.**—Alain Liogier (1973:278–270), discussed the similarities of *Scolosanthus subsessilis* Alain with other species as, “Among the small-leaved *Scolosanthus*, *S. subsessilis* has one of the smallest corollas. The nearest species are *S. wrightianus* (Griseb.) C. Wright, with leaves 7–14 mm long, the flowers short-pedicellate, the calyx lobes deltoids, the anthers 2 mm long, according to Standley; *S. versicolor* Vahl has apiculate leaves, the calyx 2–2.5 mm long, the corolla violet or yellow, 6–7 mm long. From *S. densiflorous* Urb., it is readily distinguished by its flowers on lateral branches, instead of being on the spines; *S. densiflorous* has usually larger leaves (up to 3 cm long), the calyx lobes semiorbicular, the corolla lobes almost as along as the tube.” The diagnostic features presented by Alain Liogier are here followed, and *S. subsessilis* is here recognized as a distinct species endemic to calcareous soils of Dominican Republic.

26. **Scolosanthus triacanthus** (Spreng.) DC., Prodr. 4:484. 1830. *Catesbaea triacantha* Spreng., Neue Entdeck. Pflanzenk. 3:47. 1822. TYPE: DOMINICAN REPUBLIC: without locality, s.d. [1820], C. *Bertero s.n.* (NEOTYPE, here designated: M [M-0189390]; ISONEOTYPES: MO [No. 2091990], MPU [MPU021756], S [No. S07-14401], TO).

**Distribution.**—Dominican Republic.

**Notes.**—Sprengel (1822:47), in the protologue of *Catesbaea triacantha* Spreng., cited the collection locality as “Hispaniola,” without citing the collector, collection number or herbarium of deposit. Sprengel personal herbarium was dismantled and sold to numerous specialists. A considerable portion of Sprengel’s specimens was in B, and was destroyed during WWII. As a result of a general search in numerous herbaria, five specimens associated with this name were retrieved in M, MO, MPU, S, and TO, although none of them have evidence that they were examined by Sprengel. The M specimen, barcode M-0189390, has a label with the annotation “Catesbaea triacantha Spr. S.D.” handwritten by Balbis, and “In Sto. Domingo legt. Bertero, comct. Balbis” handwritten by an unknown author. The specimen consists of a small, ramified branch with numerous small leaves and ternate spines, and is here designated as the neotype of *C. triacantha*.

In TO there is a specimen with the annotation “Catesbaea triacantha Spr. in litt. ex S. Domingo D. Bertero 1820” handwritten by Balbis. The specimen consists of a densely ramified branch with numerous small leaves and numerous flowers in anthesis. The TO specimens do not have barcode or accession numbers. This specimen is an isotype of *Catesbaea triacantha* Spreng.

27. **Scolosanthus versicolor** Vahl, Eclog. Amer. 1:11. 1797 [“1796”]. TYPE: ST. CROIX: without locality, s.d., *J. Ryan s.n.* (LECTOTYPE, here designated: C [C10018377]; ISOLECTOTYPES: BR [000000561859], C [C10018376]; POSSIBLE ISOLECTOTYPE: B-W [B-W 02797-01 0]).

**Distribution.**—Dominican Republic, Puerto Rico, St. Croix.

**Notes.**—Vahl (1797 [“1796”:11–12), in the protologue of *Scolosanthus versicolor* Vahl, cited “*Catesbaea parviflora*. La Marck Tableau Encycl. et Meth. Botan. tab. 67.—In insula St. Crucis legit *Ryan Iz*.” In Lamarck’s (1791) *Tableau Encyclopédique et Méthodique. Requeil de Planches de Botanique*, Plate 67, figure 2, is represented “*Catesbaea parviflora*” with a branch with small leaves, subtended by pairs of acicular thorns, and the details of a dissected corolla, style, anthers, and a fruit in side view and in cross section. This plate represents original material of *S. versicolor*. According to the *Code*, a specimen should be preferred over an illustration for the typification and application of a name.

According to Stafleu and Cowan (1986:628), the original material studied by Martin Vahl is in C. In C there are two specimens annotated by Vahl with this name. The specimen with barcode C10018377, on the back of the sheet has the annotation “Scolosanthus versicolor Dr Ryan” handwritten by Vahl, and the stamp “HB. VAHLII.” On the front of the sheet is affixed a branch with numerous minute leaves, bifurcate thorns in many nodes, and a few flowers in anthesis. This specimen is here designated as the lectotype of *S. versicolor*.

The C specimen with barcode C10018376, on the back of the sheet has the annotations “Scolosanthus versicolor I” and “West ex Ins. St. Crucis” handwritten by Vahl, and the stamp “HB. VAHLII.” On the front of the sheet is affixed a branch with numerous minute leaves, bifurcate thorns in many nodes, and numerous minute fruits. This specimen is an isotype.

In BR there is a specimen with barcode 000000561859, which has a label with the annotation “Scolosanthus versicolor mihi ex Ins. St. Crucis” handwritten by Vahl. The specimen consists of a branch
with numerous minute leaves, bifurcate thorns in many nodes, and several flower buds and flowers in anthesis. This specimen is an isotype.

In B-W there is a specimen with barcode B-W 02797-01 0, bearing the annotation "Vahl. W" handwritten by Willdenow. This specimen consists of a branch with numerous minute leaves, bifurcate thorns in many nodes, and a few minute fruits. This specimen is a possible isotype of *S. versicolor*.

Although the type of this species is from St. Croix, there is no mention of *Scolosanthus* in Howard's (1989) *Flora of the Lesser Antilles*.

**28. Scolosanthus wrightianus** (Griseb.) Wright, in Sauv., Anal. Acad. Ci. Habana 6:126. 1869. *Randia wrightiana* Griseb., Cat. Pl. Cub. 122. 1866. **TYPE:** CUBA. GUANTÁNAMO: "Cuba Or.," s.d. [1861], C. Wright 2715 (LECTotype, here designated: GOET [GOET010513]; ISOLECTOTYPES: GH [00094516], HAC [3 sheets, one sheet ex SV], MO [No. 2091759], S [No. S05-1195], YU [YU.001806]).

**Distribution.**—Cuba (Guantánamo).

*Notes*.—Grisebach (1866:122–123), in the protologue of *Randia wrightiana* Griseb., cited two Cuban gatherings collected by Wright as “Cuba or. (Wr. 2660, 2715)” without indicating the herbarium of deposit. For information regarding Grisebach’s original specimens, see notes under *Phialanthus myrtilloides*. Borhidi et al. (2017:401) cited the type of *Randia wrightiana* as “Tipo: Cuba Oriental, Sta Ana, Mt. Toro, C. Wright 2660 p.p. Holotipo: GOET, isotipos: GH, HAC.” According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by “here designated” or a similar expression. Hence, Borhidi et al. (2017) “holotype” citation cannot be treated as an inadvertent lectotypification.

In GOET there are two specimens annotated as *Randia wrightiana* corresponding with the two gatherings cited by Grisebach. The specimen Wright 2715 with barcode GOET010513 has a label with the heading "Plantae Cubenses Wrightianae" and the annotation "2715. Randia wrightiana m." handwritten by Grisebach. This specimen consists of a branch with numerous leaves, bifurcate thorns with one or two nodes, and a few flower buds. This specimen is here designated as the lectotype of *Randia wrightiana*.

The specimen Wright 2660 in GOET, with barcode GOET010512, has a label with the heading "Plantae Cubenses Wrightianae" and the annotation "2660. Randia wrightiana m." handwritten by Grisebach. This specimen consists of a branch with numerous leaves, bifurcate thorns with one or two nodes, and no flowers. In the envelope affixed on the sheet are included numerous leaves and several minute fruits. In the image of this specimen available in Jstor Global Plants is included an insert of an image showing a large fruit, which does not belong to any species of *Scolosanthus*.

## SCOLOSANTHUS EXCLUDED TAXA

*Scolosanthus parviflorus* (Sw.) Wright, in Sauv., Anal. Acad. Ci. Habana 6:126. 1869.
= *Catesbaea parviflora* Sw. (See above).

*Scolosanthus sagraenus* (A. Rich.) Millsp., Publ. Field Mus., Bot. 2(1):102. 1900 (as "Sagraeanus"). *Gardenia sagraeana* A. Rich. in R. de La Sagra, Hist. Fís. Cuba, Bot. 11:10. 1850. **TYPE:** CUBA: Havana, s.d., R. de La Sagra s.n. (P not found).

**Notes.**—Achille Richard (1850:10) in the protologue of *Gardenia sagraeana* A. Rich., cited the material studied as "[Cuba] crescit in ruderatis circa Havanam, mense maio florens (Ramon de la Sagra), et prope Santiago de Cuba (Linden)," without citing the herbarium of deposit. A general search in Jstor Global Plants and in the P virtual herbarium no original specimen was found.
= *Randia spinifex* (Roem. & Schult.) Standl., Contr. U.S. Natl. Herb. 20(6):201. 1919.

## SHAFEROCHARIS

Urban (1912:412) described the genus *Shaferocharis* Urb., dedicating the name to the American botanist John A. Shafer (1863–1918), who made numerous important collections in Cuba between 1903 and 1912. This genus has been included into the Chiococceae by several Rubiaceae specialists (e.g., Robbrecht 1988, 1994). Delprete (1996), according to his phylogenetic analyses using morphological characters, maintained it in the Chiococceae. It has never been included in any molecular phylogenetic study, mostly due to lack of recent collections. According to Borhidi (1972 ["1971"], 1982 ["1981"], 1983, 2017), this genus is composed of three species endemic to Cuba, which are here recognized.
**Shafercharis** Urb., Symb. Antill. 7:412. 1912. Standley, N. Amer. Fl. 32(4):284-285. 1934; Liogier, Fl. Cuba 5:92-93. 1962; Borhidi & Muñiz, Acta Bot. Acad. Sci. Hung. 17:33-34, f. 15. 1972 ["1971"); Borhidi, Acta Bot. Acad. Sc. Hung. 27(1-2):29-36. 1982 ["1981"); Borhidi, Acta Bot. Hung. 29:181-215. 1983; Borhidi et al., Rubiáceas Cuba 404-409, f. 126-127. 2017. TYPE: *Shafercharis cubensis* Urb.

**1. Shafercharis cubensis** Urb, Symb. Antill. 7:413. 1912. TYPE: CUBA. HOLGUIN: Sierra Moa ad austrum versus, in Campo La Gloria, 30 Dec 1910. J.A. Shafer 8245 (NEOTYPE, here designated: A [00094520]; ISONEOTYPES: HAC [ex LS], NY [00115372], US [00138505] (frag. ex NY)).

**Distribution.**—Cuba (Holguín: Sierra de Moa).

**Notes.**—Urban (1912:413), in the protologue of *Shafercharis cubensis* Urb., cited the material studied as "in Cuba prov. Oriente in campo la Gloria prope Sierra Moa ad austrum versum, m. Dec. fruct.: Shafer n. 8245 (typus), ibidem: Shafer n. 8185" without citing the herbarium of deposit. The original material in B was destroyed during WWII. Four specimens of *Shafer 8245* are in A, HAC, NY, and US. None of them is annotated by Urban.

Borhidi (2017:404) cited the type of *S. cubensis* as "Tipo: Cuba; prov. Oriente in camp la Gloria prope Sierra de Moa [...] J.A. Shafer 8245. Holotipo: NY! Isotipo: (LS)"! According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:404) "holotype" citation cannot be treated as a valid lectotypification.

In A there is a specimen with barcode 00094520, which has a label with the typewritten annotations "Shafercharis cubensis Urban, Camp La Gloria, south of Sierra Moa, Oriente, Symb. Antill. 7: 413. 1912. J.A. Shafer 8245 30 Dec. 1910." This specimen consists of several branches with numerous leaves and several axillary infructescences. Because this specimen is not annotated by Urban, it is here designated as the neotype of *S. cubensis*.

**2. Shafercharis multiflora** Borhidi & Muñiz, Acta Bot. Acad. Sci. Hung. 17:33. 1972 ["1971"). TYPE: CUBA. HOLGUIN: Región de Moa, Río Yamanigüey, suelo de serpentina, 25 m, 24 Mar 1970, A. Borhidi, O. Muñiz & S. Vazquez SV No. 27122 (HOLOTYPE: HAC [ex SV No. 27122]); ISOTYPE: BP [No. 50311]).

**Distribution.**—Cuba (Holguín: Sierra de Moa).

**3. Shafercharis villosa** Borhidi & Bisse, Acta Bot. Sci. Hung. 27:33. 1982 ["1981"). TYPE: CUBA. GUANTÁNAMO-BARACOA: En charrascales húmedos del Río Maraví, Mar 1970, J. Bisse & E. Köhler HFC No. 16189 (HOLOTYPE: HAJB [HAJB G 000860 (HFC No. 5808)]; ISOTYPES: JE [JE00004987 (HFC No. 5808)], SOF n.v.).

**Distribution.**—Cuba (Guantánamo: Baracoa, Río Maraví).

## SIEMENSIA

Urban (1923a:145) separated *Siemensia* Urb. from *Portlandia sensu lato* (as delimited in those days) based on the linear placenta, loculicidal capsules, large seeds with a cupuliform funiculum, and axillary inflorescences. He dedicated the genus name to Ernst Werner von Siemens (1816-1892), inventor and industrialist. Aiello (1979) maintained *Siemensia* as a distinct monospecific genus, and described its placentae and seeds as "placentae kidney-shaped in cross section. The tiny (1 mm or less) seeds, which are perpendicular to the placenta, cover it except to the side opposite the septum. After the seeds fall, a small hole is left in the placenta where each was attached" and further described the seeds as "broadly ovate to broadly elliptic, 0.9 mm long; testa with closely packed, deep pits, interspaces densely granulate." (Aiello, 1979:69, 77, figs. 65, 83, 84).

Motley et al. (2005) obtained contrasting results regarding the position and relationships of *Siemensia* in separate molecular analyses using *trnL-F* and ITS sequences. *Siemensia* was found as sister to a New Caledonian clade in the ITS analyses, or on a trichotomy with *Bikkia sensu stricto* clade (excluding the species endemic to New Caledonia, later transferred to *Thiollierea*) and a large *Scolosanthus-Erithalis-Chiococca* clade in the *trnL-F* analyses. Following these contrasting results, Motley et al. (2005) suggested that one or two dispersal events are required to account for the two western Pacific clades, one event is required to give rise to the New Caledonia clade (*Thiollierea*), and the other the *Badusa-Bikkia sensu stricto* clade.

Paudyal et al.'s (2018) molecular phylogenies were also ambiguous in establishing the phylogenetic position of *Siemensia*. In the analysis using nuclear and plastid data combined, *Siemensia* was resolved sister to the
*Badusa-Bikkia* clade, two genera occurring in the South Pacific. In the analyses using nuclear datasets, *Siemensia* was strongly supported as sister to the *Thiolliera-Morierina* clade, a group endemic to New Caledonia. In their phylogenies using plastid datasets, *Siemensia* was resolved in a trichotomy with the *Badusa-Bikkia* clade and *Chiococca-Scolosanthus* clade as sister clades.

*Siemensia* is a monotypic genus endemic to the limestone Haystack Mountains of the Province of Pinar del Río, western Cuba (Liogier 1962; Borhidi et al. 2017).

*Siemensia* Urb., Symb. Antill. 9:145. 1923; Liogier, Fl. Cuba 5:29. 1962; Aiello, J. Arnold Arbor. 60:119. 1979; Borhidi et al., Rubiáceas Cuba 409–410, f. 128. 2017; Paudyal et al., Bot. J. Linn. Soc. 187(3):365–396. 2018. *TYPE: Siemensia pendula* (C. Wright ex Griseb.) Urb.

1. **Siemensia pendula** (C. Wright ex Griseb.) Urb., Symb. Antill. 9: 145. 1923. *Portlandia pendula* C. Wright ex Griseb., Cat. Pl. Cub. 126. 1866. **TYPE:** CUBA. PINAR DEL RÍO: "Cuba Occ.," without locality, s.d., C. Wright 2677 (HOLOTYPE: GOET [GOET010517]; ISOTYPES: AC [00320379], BM [000081666], F [No. 635156 (frag. ex G)], G [2 sheets, G00436770, G00436771], GH [00058977], NY [00126759], P [P02273498], S [2 sheets, No. S-R-7826, No. S07-14585], US [00137333], YU [YU.001824]).

**Distribution.**—Cuba (Pinar del Río, Artemisa).

**Notes.**—Grisebach (1866:126–127), in the protologue of *Portlandia pendula* C. Wright ex Griseb., cited the material studied as “Cuba occ. (Wr. 2677)” without citing the herbarium of deposit. According to Stafleu and Cowan (1976:1007), the GOET herbarium contains most of Grisebach’s types. In GOET there is a single specimens annotated with this name. The specimen with barcode GOET010517, has a label with the annotation “Portlandia pendula” and a detailed description handwritten by Grisebach. This specimen is the holotype of this name.

## SOLENANDRA

Joseph Dalton Hooker published the genus *Solenandra* in April 1873, in both Bentham & Hooker's *Genera Plantarum* and in *Hooker's Icones Plantarum*. He did not explain the etymology of the name. From the Greek, Solen- (σωλήν), tube, and -andros (άυδρος), male, anther, probably referring to the stamens with filaments basally connate, forming a basal tube. Stafleu and Cowan (1970:294–296), for the entry of *Hooker's Icones plantarum*, on page 294, supplied the date of publication for volume 12 as 1876, which is the date reported on the title page of the volume. However, on page 296, they cited different dates for the four parts of volume 12. They reported that page 43 and plate 1150 of *Solenandra ixoroides* Hook.f. are in the second part of volume 12, which was published in April 1873 (Hooker, 1873c). This publication date was determined from notes in the copies at BM and K. Article 31.1 of the *Code* (Turland et al. 2018) makes the exception that proof establishing the date of publication supersedes the date on the title page. As the publication of *Genera Plantarum* vol. 2, part 1, has the date of 7–9 April 1873 (Hooker, 1873a), it is here treated as the first publication date of *Solenandra*.

*Solenandra* was treated as a synonym of *Exostema* in the classic Rubiaceae classifications of Schumann (1891), Standley (1921:117), and Robbrecht (1988). McDowell (1996) maintained *Solenandra* as a synonym of *Exostema*, and included *Solenandra ixoroides* Hook.f. (as "*E. ixoroides*" (Hook.f.) T. McDowell *ined.*"), the type species of *Solenandra*, in *Exostema* section *Brachyantha* DC. sensu McDowell (1996). The new combination *E. ixoroides* (Hook.f.) T. McDowell ex Greuter was later published by Greuter (in Berazain et al. 2009:82), which is here treated as a synonym of *Solenandra parviflora* (Bonpl.) Delprete, comb. nov. (See below).

Rova (1999), in a molecular phylogenetic study of the Condamineae-Rondeletieae-Sipaneeae complex using *rps16* sequence data, retrieved *Exostema*, as circumscribed by McDowell (1996), as paraphyletic. Borhidi (2002), stimulated by the results of Rova (1999), resurrected *Solenandra* as a genus of 12 species endemic to Cuba, Hispaniola and Mexico, into which he transferred all the species of *Exostema* section *Brachyantha sensu* Borhidi & Fernández-Zequeira (1989) except *E. corymbosum* (which was transferred to *Motleyothamnus* by Paudyal et al (2018)). Borhidi (2002) differentiated *Exostema* from *Solenandra* by the branches with black lenticels (vs. white in *Solenandra*), flowers 4–21 cm long, blooming at night, fragrant (vs. 1–4 cm long, blooming during the day, not fragrant), with corollas pink or violet after anthesis (vs. pale yellow), corolla lobes linear (vs. ovate or lanceolate), style longer than stamens (vs. shorter than stamens), cylindrical capsules (vs. laterally compressed), containing 10–400 seeds (vs. 1–10 seeds).
C. *coriacea* Poiret rami laeves striati, tecti cortice cinereo; folia opposita petiolata, obtusa, nervis lateralibus alternis prominentibus apice bifurcatis. Paniculae rami subdichotomi; pedunculi rigidi glabri floribus subsessilis glabris. Cal. oblongus margine dentibus 5 rectis acutis. Cor. 2 poll., tubo rectotereti, limbi laciniis angustis obtusis, longitudine tubi, glabris, reflexis. Antherae rectae, filiformes. Fructus cylindricus, nigrescens, pollicaris. *In S. Domingo. Herb. Juss. et Lam.*" [Poiret undoubtedly associated this species with C. *nitida* Ruiz and Pav., although it is not of the same genus, as it has a larger panicle, corolla tube half the lobe length, fruit elongated, slightly tapered at tip. *Cinchona coriacea* has smooth, striated branches, with gray bark; leaves opposite, petiolate, obtuse, with secondary veins alternate, prominent, distally bifurcate. Panicles with subdichotomous ramification; peduncles rigid, glabrous, and flowers glabrous, subsessile. Calyx oblong, with 5 acute teeth. Corolla 2 inches [ca. 5 cm] long, tube straight, terete, with lobes narrow-linear, obtuse, as long as the tube, glabrous, reflexes. The anthers are straight, filiform. Fruits cylindrical, blackish, 1 inch [2.5 cm] long. *In Santo Domingo* [Hispaniola]. *Jussieu et Lamarck Herbaria.*]

In P-JU, there is a specimen, barcode P00680235, which has a label with the annotation "St. Domingue—herb. d. Desportes sans nom" handwritten by an unknown author. The annotation means that this specimen was collected by Jean-Baptiste-René Pouppé Desportes (1704–1748) in Haiti ("Saint Domingue"). On that label is glued a smaller label with the annotation "Cinchona coriacea Poir. Dict." handwritten by Poiret. On the sheet are affixed five branchlets, each of them with several leaves and a terminal inflorescence without flowers. On the bottom of the sheet is affixed an envelope containing numerous broken corollas, showing exserted stamens and styles. This specimen is here designated as the lectotype of *Cinchona coriacea*. https://mediaphoto.mnhn.fr/media/14418033301640yz3fs17wY8pIJeh

In P-LA there is a specimen, barcode P00308387, which has a label with the annotation "de St. Domingue ex D. Juss. Cinchona coriacea Dict." On the sheet are affixed two separate leaves, and a branchlet with several leaves, and a terminal inflorescence without flowers. On the sheet is affixed a small envelope containing a flower in anthesis, corolla lobes broken off and several portions of floral parts. The flower has exserted stamens with linear anthers, and exserted style. The material on this sheet was probably extracted from the specimen in P-JU, and is an isolectotype of *Cinchona coriacea*.

Andersson (1992:96) cited the type of *Cinchona coriacea* Poir. as "Type: Collector not cited; Hispaniola (P-Juss., P-Lam.)" and treated it as synonym of *Exostema coriaceum* (Poir.) Schult. Andersson (1998:63) maintained *C. coriacea* as a synonym of *E. coriaceum*, without citing the type.

Liogier (1995:259) recognized *Exostema coriaceum* as a distinct species. He provided a detailed description, and added "H: Herbario de *Jussieu y Larmark* (P); endémica. Una especie poco conocida, no coletada por botánicos modernos." (H [Haiti]: Herbaria of *Jussieu and Larmark* (P); endemic. A little-known species, not collected by modern botanists"). In the key to the species of *Exostema sensu lato* from Hispaniola, he distinguished *E. coriaceum* from *E. ellipticum* by the subsessile flowers (vs. with pedicels to 2 cm long in *E. ellipticum*), corolla lobes as long as corolla tube (vs. corolla lobes shorted than tube), and leaves with prominent venation (vs. not prominent). In the type specimens of *E. coriaceum* the flowers are pedicellate to short-pedicellate, while in *E. ellipticum* they are long-pedicellate. After examining the type specimens in P-JU and P-LA, and the descriptions provided by Poiret (1804:38), Schultes (in Roemer & Schultes, Syst. Veg. ed. 15 [bis], 5:20. 1819), and Liogier (1995:259), it became obvious that this species should be transferred to *Solenandra*. Paudyal et al. (2018), following their molecular phylogenies, characterized *Solenandra* by the terminal inflorescences, white corollas, basipetally septicidal capsules, among other characters, and divided it into two sections: Sect. *Solenandra*, with corollas 0.6–2.2(–3.0) cm long, turning pale yellow after anthesis, and Sect. *Pitonia* (DC.) Paudyal & Delprete, with corollas 4–21 cm long, turning pink to maroon after anthesis. Hence, this species belongs to Sect. *Pitonia*, and the new combination *Solenandra coriacea* (Poir.) Delprete is here provided.

4. **Solenandra elliptica** (Griseb.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:388. 2018. *Exostema ellipticum* Griseb., Pl. Wright. 2:504. 1862. TYPE: CUBA. HOLGUIN: near villa Monte Verde, Jan.–Jul 1859. C. Wright 1257 (LECTOTYPE) (Paudyal et al. 2018: 388): GOET [GOET003383]; ISOLECTOTYPES: A [00041000], BR [000000549057], G [G00436071, G00436072], GH [00046001], K [K000173632], MO [Nos. 2091694, 2091695], NY [00077371, 00077383], PH [00013986], S [No. S11-19749, S-R-8288], US [00130602], YU [YU.001735]).
**Distribution.**—Cuba (Pinar del Rio, Artemisa, Isla de la Joventud, Camagüey, Holguín, Santiago de Cuba), Haiti, Dominican Republic, and Puerto Rico.

**Notes.**—Grisebach (1862:503) in the protologue of *Exostema ellipticum* Griseb., cited the gathering *Wright* 1257, without citing the herbarium of deposit. According to Stafleu and Cowan (1979:1007) in GOET are kept most of Grisebach's types. Borhidi et al. (2017:108) cited the type of *E. ellipticum* as "Tipo: [...] Jan-Jun 1859, Wright 1257; Holotipo: GOET; isotipos: GH, US." Borhidi et al. (2018:308) cited the type of *E. ellipticum* as "Tipo: [...] Jan-Jun 1859, Wright 1257; Lectotipo: GOET; isolectotipos: BR, K, MO, NY, PH, S, US, YU."

According to the *Code* (Turland et al. 2018), starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:108; 2018:308) lectotype designations are not valid.

In GOET there is a specimen of Wright 1257, with barcode GOET003383, which has a label with the annotation "Exostemma ellipticum m." handwritten by Grisebach. The specimen consists of numerous small branches with infructescences and dehisced capsules. This specimen was designated as the lectotype of *E. ellipticum* by Paudyal et al. (2018:388).

*Exostema monticola* Borhidi & M. Fernández, Acta Bot. Hung. 35:303. 1989. *Solenandra elliptica* ssp. *monticola* (Borhidi & M. Fernández) Borhidi & M. Fernández, Acta Bot. Hung. 60:309. 2018. TYPE: CUBA. GUANTÁNAMO: Sierra Maestra, Loma Pino del Agua, Alto de Valenzuela, 11 Aug 1955, M. López Figueiras 2314 (*LECTOTYPE*, here designated: HAC [ex SV] (Fig. 6); *ISOLECTOTYPES*: HAC [ex LS], HAJB [HAJB G 000460], US [00589158]).

**Notes.**—Borhidi & Fernández Zequeira (1989:303) cited the type of *Exostema monticola* Borhidi & M. Fernández as "Holotypus: López Figueira L.F. 2314 HAC; Prov. Oriente; Sierra Maestra, Loma Pino del Agua, [...], LOPEZ FIGUEIRAS, 11.08.1955. Isotypus: HAC, HAJB." Borhidi et al. (2017:108; 2018:309) cited the type of this name as "Tipo: Cuba, Prov. Oriente, Sierra Maestra, [...] López Figueiras L.F. 2314. Holotipo: HAC; isotipos: HAC, HAJB." They did not specify which of the two specimen in HAC is the holotype, hence a lectotype needs to be designated for this name. In HAC are present two specimens of López Figueiras 2314 that were originally preserved in two different herbaria. One specimen has a label with the heading "Republica de Cuba—ESTACION EXPERIMENTAL AGRONOMICA." The label heading means that this specimen was originally in SV, a herbarium that was later integrated in HAC. This specimen has another label with the annotation "HOLOTYPUS. Exostema monticola Borhidi et Fernández" handwritten by Borhidi. This specimen consists of two branches with numerous leaves, numerous flower buds, and several flowers in anthesis (Fig. 6), and is here designated as the lectotype of *E. monticola*.

The other specimen in HAC has label with the heading "HERBARIO DE LA SALLE." The heading means that this specimen was originally in LS, a herbarium that was later integrated in HAC. This specimen has another label with the annotation "ISOTYPUS. Exostema monticola Borhidi et Fernández" handwritten by Borhidi. This specimen consists of two branches with numerous leaves, numerous flower buds, and several flowers in anthesis, and is an isolectotype of *E. monticola*.

**5. Solenandra lineata** (Vahl) Paudyal & Delprete, Bot. J. Linn. Soc. 187:388. 2018. *Exostema lineatum* (Vahl) Schult., in Roemer & Schultes, Syst. Veg. ed. 15[bis], 5:18. 1819. *Cinchona lineata* Vahl., Skr. Naturhist.-Selsk. 1:22. 1790. TYPE: HISPANIOLA [HAITI]: without locality, s.d., *A. Thouin s.n.* (FIRST-STEP LECTotype (Andersson 1992: 96); SECOND-STEP LECTotype (Paudyal et al. 2018:389): C [No C10018139]).

**Distribution.**—Haiti and Dominical Republic.

*Exostema subcordatum* Krug & Urb., in Urban, Symb. Antill. 1:421. 1899. TYPE: HAITI: mountains near La Coupe, above Rivière, s.d., B.P. Jaeger 210 (*LECTOTYPE*, here designated: C [C10018140]; *ISOLECTOTYPES*: BM [2 sheets, 000028115, 000028116], BR [000000530624], F [No. 189174], GH [00046015], GOET [GOET010236], JE [JE00004869], K [2 sheets, K000173619], M [M-0187319], US [00130610]).

*Exostema subcordatum* var. *buchii* Urb., Symb. Antill. 8:666. 1921. TYPE: HAITI: Plateau St. Roc, 24 Mar 1920, *W. Buch 1834* (*NEOTYPE*, here designated: US [00589160]).

*Exostema picardae* Krug & Urb. in Urban, Symb. Antill. 1:422. 1899. TYPE: HAITI: Plateau St. Roc, 24 Mar 1920, *W. Buch 1834* (*NEOTYPE*, here designated: US [00589160]).

**Notes.**—Krug and Urban (in Urban 1899:422), in the protologue of *Exostema picardae* Krug & Urb., cited the gathering *Picarda* 720, without citing the herbarium of deposit. The original material at B was destroyed, and no extant specimens of *Picarda* 720 was found in the herbaria consulted. Hence a neotype needs to be designated. The specimen *Buch 1834* in US, with barcode 00589160, is here designated as the neotype of *E. picardae*.

*Exostema rupicola* Urb., Symb. Antill. 7:399. 1912 (as "*rupicolum*"). TYPE: DOMINICAN REPUBLIC. BARAHONA: near Baoruco, Jun 1911, *P. Fuertes 1169* (*NEOTYPE*, here designated: A [00046009]).

**Notes.**—Urban (1912:399), in the protologue of *Exostema rupicola* Urb., cited the gathering *Fuertes 1169*. The original material in B was destroyed during WWII. There is a specimen of *Fuertes 1169* in A, without any evidence that it was seen by Urban; hence it is here designated as the neotype of this name.
<figure><img src="image_9.png" /><figcaption>Fig. 6. Lectotype of *Exostema monticola* Borhidi & M. Fernández (López Figueiras 2314, HAC [ex SV]). = *Solenandra elliptica* (Griseb.) Paudyal & Delprete.
Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
**6. Solenandra longiflora** (Lamb.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:388. 2018. *Exostema longiflorum* (Lamb.) Schult., in Roemer & Schultes, Syst. Veg., ed. 15 [bis], 5:18. 1819. *Cinchona longiflora* Lamb., Descr. Cinchona 38, fig. 12. 1797. TYPE: HAITI: without locality, s.d. [1764-1765], J.B.C. Aublet s.n. (LECTOTYPE (Paudyal et al. 2018: 388): BM [BM000028085]).

**Distribution.**—Cuba (the whole island), Haiti, and Dominican Republic.

Oxyanthus versicolor Lindl., in Edwards's Bot. Reg. 26:tab. 69. 1840. TYPE: Not found.

**7. Solenandra mexicana** (A. Gray) Borhidi, Acta Bot. Hung. 44:227. 2002. *Exostema mexicanum* A. Gray, Proc. Amer. Acad. 5:180. 1861. TYPE: MEXICO: Tantoyuca, Aug 1858, L.C. Ervendberg 125 (HOLOTYPE: GH [00046002]; ISOTYPES: GOET [GOET010238], K [K000173631], P [P01900538]).

**Distribution.**—Mexico, Belize, El Salvador, Honduras, Guatemala, and Costa Rica.

**Notes.**—Gray (1861:180), in the protologue of *Exostema mexicanum* A. Gray, cited the gathering Ervendberg 125, without indicating the herbarium of deposit. According to Stafleu and Cowan (1976:983), Asa Gray's herbarium and types are at GH. In GH there is a specimen that has a label with the annotation "Exostema Mexicanum n. sp." handwritten by Gray, which is the holotype.

*Exostema indutum* Standl., N. Amer. Fl. 32(2):126. 1921. TYPE: MEXICO. OAXACA: Villa Alta, Aug 184? [last digit missing], H. Galeotti 2664 (HOLOTYPE: US [00130603]; ISOTYPES: BR [2 sheets, 000000530656, 000000530657], F [No. 775706], US [00589161]).

**8. Solenandra myrtifolia** (Griseb.) Borhidi, Acta Bot. Hung. 44:227. 2002. *Exostema myrtifolium* Griseb., Cat. Pl. Cub. 125. 1866. TYPE: CUBA. GUANTÁNAMO: Baracoa, 14 May 1861, C. Wright 2673 (HOLOTYPE: GOET [GOET010239]; ISOTYPES: BM [000028151], G [G00436070], GH [000460003], HAC n.v., K [K000173629], MO [No. 2091701], YU [YU.001736]).

**Distribution.**—Cuba (Holguín, Guantánamo).

*Exostema shaferi* Standl., N. Amer. Fl. 32(2):124. 1921. TYPE: CUBA. GUANTÁNAMO: Near El Yunque, Loma Santa Teresa, 2 Dec 1910, J.A. Shafer 7746 (HOLOTYPE: NY [00077356]; ISOTYPES: A [00046014], BM [000028138], F [No. 493027], US [00589163]).

*Exostema crassifolium* Standl., N. Amer. Fl. 32:124. 1921. TYPE: CUBA. GUANTÁNAMO: Between Sabanilla and Yamuri Arriba, 30 Jan–1 Feb 1911, J.A. Shafer 8416 (HOLOTYPE: NY [00077382]; ISOTYPES: A n.v., F n.v., US [00130599]).

*Exostema nipense* Urb., Symb. Antill. 9:521. 1928. TYPE: CUBA. GUANTÁNAMO: Sierra de Nipe, in charrascales, at Río Piloto, 30 Jul 1914, E.L. Ekman 2303 (B†; LECTOTYPE, here designated: S [No. S07-14970]; ISOLECTOTYPES: A [00061486], BM [000028094], MO [No. 1006605]).

*Exostema dumosum* Alain, Contr. Ocas. Mus. La Salle 17:3. 1959. TYPE: CUBA. GUANTÁNAMO: 32 km to the S of Baracoa, Via Azúl, charrasco, 14 Feb 1956, Bro. Alain [A.H. Liogier] & C.V. Morton 5156 (HOLOTYPE: HAC; ISOTYPES: NY [2 sheets, 00380583, 00077355]).

*Exostema barbatum* Standl., N. Amer. Fl. 32:125. 1921. *Exostema myrtifolium* var. *barbatum* (Standl.) Borhidi & M. Fernández, Acta Bot. Hung. 35:298. 1989. *Solenandra myrtifolia* var. *barbata* (Standl.) Borhidi, Acta Bot. Hung. 44:230. 2002. TYPE: CUBA. GUANTÁNAMO: Baracoa, on pine hills, March 1903, L.M. Underwood & F.S. Earle 1360 (HOLOTYPE: NY [00077373]; ISOTYPE: US [00147046 (frag. ex NY)]).

**9. Solenandra parviflora** (Bonpl.) Delprete, **comb. nov.** *Exostema parviflorum* Bonpl., in Humboldt & Bonpland, Pl. Aequinoct. 1:132. [Apr.] 1807 (as "*parviflora*"). "*Solenandra parviflora* (L.C.M. Rich.) Borhidi," Acta Bot. Hung. 44:230. 2002, **comb. inval.** (Without bibliographic reference of the basionym). TYPE: ANTILLES [HAITI]; without locality, s.d., L.C.M. Richard s.n. (HOLOTYPE: P [P03947017]; ISOTYPE: C n.v.).

**Distribution.**—Cuba (Pinar del Río, Villa Clara, Cienfuegos, Sancti Spiritus, Las Tunas, Santiago de Cuba, Guantánamo), Haiti, Dominican Republic.

**Notes.**—Bonpland (1807:132), for the genus *Exostema* stated "Ex accurata permultarum Rubiacearum analysi, jam dudum, etiam per Antillas itinerans, detexi ipse duorum generum discrepantiam et nomine diverso in schedis meis designavi. Spontaneas in supra dictis insulis septem novi species Exostemae; quarum una, ni fallor, adhuncum nova: [*Exostema parviflorum* Bonpl.]." [From the accurate analysis of many Rubiaceae, travelling through the Antilles, I detected the discrepancy of two genera and I designated them with different names in my notes. Spontaneous in the seven aforementioned islands, a new species of *Exostema*; one of which, if I am not mistaken, is new: [*Exostema parviflorum* Bonpl.]" On the same page, in the protologue of *Exostema parviflorum* Bonpl. (as "*parviflora*"), he cited "mirror, quod neque in territorio Paraensi Brasiliae, neque in Guyanna Rubiaceis ditissima, speciem hujusce generi invenerim ullam." [I am surprised that neither in the territory of Pará, Brazil, nor in Guyana, where Rubiaceae are most abundant, I did not find any specimen of this genus].
In P, there are three specimens annotated as "Exostema parviflorum Rich." with barcodes P0394715, P03947016, and P03947018, collected by Ramón de la Sagra in Cuba. These specimens have labels with the heading "Herbarium Richard" handwritten in red ink, and they are not original material of *Exostema parviflorum*.

The P specimen with barcode P03947017, has a label with the annotations "Exostema parviflora Pl. Eq. t. 1. p.,” (“Bonpland scripsit”), and “Antilles. Rich. D.D.” The latter annotations means that the specimen was collected in the Antilles by L.C.M. Richard, most likely in Haiti [“St. Domingue”]. On the sheet are affixed two small branches, one with a few leaves and a terminal inflorescence with numerous flower buds, and the other without leaves and a terminal inflorescence with several flowers in anthesis. This specimen is the holotype of *Exostema parviflorum*.

The P specimen with barcode P03947019, has a label affixed on the bottom left corner, with the annotation "St. Domingue." Just above that label, there is another label with the annotation "Echantillon appartenent à l'Herbier générale. Classé parmi les Hamelia ausquels il ne parait pas appartenir" handwritten by an unknown author. A third label bears Liogier's identification "Exostema elegans Krug & Urb." [= *Solenandra parviflora* (Bonpl.) Delprete]. This specimen is not original material.

Borhidi (2002:230) published the new combination "*Solenandra parviflora*" but he incorrectly cited the author of the basionym as "*Exostema parviflorum* L.C.M. Rich.,” and did not cite the bibliographic reference of the basionym, and without citing the type. According to Art. 41.5 of the *Code* (Turland et al. 2018), "On or after 1 January 1953, a new combination, name at new rank, or replacement name is not validly published unless its basionym or replaced synonym is clearly indicated and a full and direct reference given to its author and place of valid publication, with page or plate reference and date (but see Art. 41.6 and 41.8)." Hence, Borhidi's new combination is not valid, and the new combination *Solenandra parviflora* (Bonpl.) Delprete is here published.

*Exostema valenzuelae* A. Rich., in Sagra Hist. Nat. Cuba 11:6, pl. 48. 1850. *nom. nud. illeg. pro syn.* of *Exostema parviflorum* Bonpl.

**Notes.**—Achille Richard (1850:6) under "Exostema parviflorum L.C. Rich." cited "Tab. 48. sub falso et delendo nomine Exost. Valenzuelae. Nob.," and in the *Observaciones* he stated "Esta especie ha sido representada en la lamina 48 de esta obra, bajo el nombre de *Exostemma* Valenzuelae: nombre que debe ser rempazado por el *E. parviflorum* que realmente le pertenece. Hemos comparado las muestras de Cuba con las que nuestro padre había recogido en las Antillas, y conocimos la identidad de todas ellas." [This species is depicted in plate 48 of this work, under the name *Exostemma valenzuelae*: a name that should be replaced by *E. parviflorum*, which really belongs to it. We compared the specimens from Cuba with those collected by my father in the Antilles, and we know the identity of all of them].

*Solenandra ixoroides* Hook.f., Hooker's Icon. Pl. 12:45, tab. 1150. Apr 1873. *Steudelago ixoroides* (Hook. f.) Kuntze, Rev. Gen. Pl. 1:298. 1891. *Exostema ixoroides* (Hook.f.) McDowell ex Greuter, Bissea 3 (num. esp.):82. 2009. *TYPE:* CUBA: without locality, s.d., *R. de La Sagra* 329 (*HOLOTYPE:* K [K000173630]).

**Distribution.**—Cuba (Las Villas, Santa Clara, Pinar del Río).

**Notes.**—Joseph Dalton Hooker published *Solenandra ixoroides* Hook.f. in Hooker's *Icones Plantarum* 12:45, pl. 1150, on April 1873. Stafleu and Cowan (1970:294-296), for the entry of *Hooker's Icones plantarum*, on page 294, supplied the date of publication for volume 12 as 1876, which is the date reported on the title page of the volume. However, on page 296, they reported that page 43 and plate 1150 of *S. ixoroides* are in the second part of volume 12, which was published in April 1873 (Hooker, 1873c). This publication date was determined from notes in the copies at BM and K. Article 31.1 of the *Code* (Turland et al. 2018) makes the exception that proof establishing the date of publication supersedes the date on the title page.

Borhidi et al. (2018:309) cited the type of *Solenandra ixoroides* Hook. f., as "dibujo de W.H. Fitch in Hooker J.D., Icones Plantarum 1876." However, as explained above, the plate was published in April 1873. Plate 1150 is original material of *S. ixoroides*. On the plate is depicted a branch with numerous ovate leaves, with a terminal inflorescence bearing numerous flower buds and a few flowers in anthesis. On the plate are also depicted the details of ovary and capitate style, a flower in anthesis with exserted stamens, stamens basally connected forming a basal tube, and a fruit longitudinally dissected. According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2018) inadvertent lectotype designation is not valid.

In K there is a specimen with barcode K000173630, which has a label with the annotations "329. Cuba. De La Sagra. Com. Franqueville. V/65. Ovarium biloculare." On the sheet are drawn, in pencil, the details of an ovary and capitate style, a flower in anthesis with exserted stamens, stamens basally connected forming a basal tube, and a fruit longitudinally dissected, all quite similar to those illustrated on plate 1150 of *Hooker's Icones plantarum*. The sheet also carries the stamp "Herbarium Hookerianum 1867." On the sheet is affixed a ramified branch with numerous ovate leaves, and several inflorescences with flower buds and flowers in anthesis. The specimen citation in Hooker's protologue of *Solenandra ixoroides*, "HAB. Island of Cuba, Don Ramon de la Sagra
(communicated by M. Franqueville)," is sufficient to identify the K specimen with barcode K000173630 as the one he cited, and is the holotype.

*Exostema elegans* Krug & Urb., in Urban, Symb. Antill. 1(3):423. 1899. Type: HAITI: Prope Port-au-Prince, flanc du Morne de l'Hôpital, 12 Nov. 1892. L. Picarda 1055 (LECTOTYPE, here designated: F [No. 189178]).

**Notes.**—Krug and Urban (in Urban 1899:423), in the protologue of *Exostema elegans* Krug & Urb., cited two gatherings from Haiti, *Picarda* 593 and *Picarda* 1055, without citing the herbarium of deposit. The original material at B studied by them was destroyed during WWII. Borhidi and Fernández Zequeira (1989:305–306) cited the type of *E. elegans sensu* Alain as "*Exostema elegans* Alain Fl. Cuba 5. 25. 1962, p.p. non Kr. et Urb. Holotypus: Acuña 10218 LS (HAC); Prov. Oriente; bordes del Rio Portillo, Sierra Maestra, Pico Turquino, 26.10.1936. Isotypus: Herb. ROIG 7742 (HAC)" [underline by the authors]. Their citation of the type of *E. elegans* sensu Alain is completely different than that of the gatherings cited by Krug and Urban (in Urban 1899:423) for *E. elegans* Krug & Urb.

A specimen in F, accession No. 189178, has a label with the heading "Ex Herbarium Krug et Urban" and the annotations "1055. Exostema elegans Kr. et Urb. det. Urb. 1897. Haiti. 26.V.1893. 12.Nov.1892. Leg. Picarda" handwritten by Urban. On the sheet are affixed a sterile branch and a branch with a terminal infructescence with dehisced capsules. This specimen is here designated as the lectotype of this name.

*Exostema wrightii* Krug & Urb., in Urban, Symb. Antill. 1:424. 1899. *Exostema parviflorum* ssp. wrightii (Krug & Urb.) Borhidi, Bot. Közl. 71:157. 1973. *Exostema valenzuelae* ssp. wrightii (Krug & Urb.) Borhidi, Acta Bot. Hung. 35:306. 1899. *Solenandra ixoroides* ssp. wrightii (Krug & Urb.) Borhidi, Acta Bot. Hung. 44:227. 2002. *Exostema ixoroides* ssp. wrightii (Krug & Urb.) Greuter, Bissea 3 (num. esp.):82. 2009. TYPE: CUBA. HOLGUIN: Monte Verde, 1860–1864, C. Wright 2672 (HOLOTYPE: GOET [GOET010235]; ISOTYPES: GH [00046016], MO [No. 2091687], S.n.v., YU [YU.065587]).

*Exostema velutinum* Standl., N. Amer. Fl. 32(2):125. 1921. *Solenandra velutina* (Standl.) Borhidi, Acta Bot. Hung. 44:230. 2002. TYPE: CUBA. SANTA CLARA: Rio San Juan, 24–25 March 1910, N.L. Britton, F.S. Earle & P. Wilson 5837 (HOLOTYPE: NY [00077357]; ISOTYPES: F [No. 492608], US [00131313]).

*Exostema eggersii* Urb., Symb. Antill. 9:521. 1928. *Exostema parviflorum* ssp. eggersii (Urb.) Borhidi, Bot. Közl. 71:157. 1973. *Exostema valenzuelae* ssp. eggersii (Urb.) Borhidi, in Borhidi & Fernández, Acta Bot. Hung. 35:306. 1989. *Solenandra ixoroides* ssp. eggersii (Urb.) Borhidi, Acta Bot. Hung. 44:227. 2002. *Exostema ixoroides* ssp. eggersii (Urb.) Greuter, Bissea 3 (num. esp.):82. 2009. Type: CUBA. Guantanamo: El Palenquito, March 1889, H.F.A. Eggers 4809 (NEOTYPE, here designated: US [00130601]; isotype: P [P03947012]).

**Notes.**—Urban (1928:521), in the protologue of *Exostema eggersii* Urb., cited *Eggers 4809* as the sole gathering for this name. The original material at B was destroyed during WWII. Borhidi et al. (2018:309) cited the type of *E. elegans* as "Tipo: Prov. Oriente, [...], Eggers (4809). Holotipo: B†; lectotipo: Cuba, Prov. Oriente, Mesa de Prada, [...] León (11958), 17 Jul–4 Aug 1924, HAC (LS); isolectotipos: NY, US." According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2018) lectotype designation is not valid. In US there is a specimen of *Eggers 4809*, which does not have any evidence that it was studied by Urban; hence, this specimen is here designated as the neotype of this name.

*Exostema valenzuelae* ssp. maestrense Borhidi & M. Fernández, Acta Bot. Hung. 35:305. 1989. *Solenandra ixoroides* ssp. maestrensis (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:227. 2002. *Exostema ixoroides* ssp. maestrensis (Borhidi & M. Fernández) Greuter, Bissea 3 (num. esp.):82. 2009. TYPE: CUBA. GUANTANAMO: Sierra Maestra, Pico Turquino, banks of Rio Portillo, 10–26 Jun 1936, J.B. Acuña Galé 10228 (HOLOTYPE: HAC; ISOTYPES: F [No. 867762], S [No. S07-15120], US [00589156]).

*Exostema valenzuelae* ssp. parvifolium Borhidi & M. Fernández, Acta Bot. Hung. 35:306. 1989. *Solenandra ixoroides* ssp. parvifolia (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:227. 2002 (as "parvifolium"). *Exostema ixoroides* ssp. parvifolia (Borhidi & M. Fernández) Greuter, Bissea 3 (num. esp.):82. 2009. TYPE: CUBA. GUANTANAMO: Palenque, Cuchillas de Toa, Cayo Fortuna, en el trillo de Riito a Piloto Arriba, Apr 1972, J. Bisse & R. Berazain 22258 (HOLOTYPE: HAC; ISOTYPE: JE [JE00001229]).

*Exostema curbeloi* Borhidi & M. Fernández, Acta Bot. Hung. 35:304. 1989. *Solenandra curbeloi* (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:227. 2002. Type: CUBA. Las Tunas: Puerto Padre, banks of streams, 8 Dec 1931, M. Curbelo X 105 (LECTOTYPE, here designated: HAC [Roig Herbarium - Plantas de Cuba No. 5857] (Fig. 7); ISOLECTOTYPES: HAC [ex LS No. 15840], NY [00077384, upper portion of sheet]).

**Notes.**—Borhidi and Fernández (1989:304), in the protologue of *Exostema curbeloi* Borhidi & M. Fernández, cited the type as "Oriente: Puerto Padre, banks of streams, *Herb. Roig* (= *Curbelo* 105) (holotype HAC; isotype, HAC)" Borhidi et al. (2017:107) cited the type of *Exostema curbeloi* as "Tipo: Cuba. Prov. Oriente. Puerto Padre, orillas y arroyos, 8 Dec 1931, Curbelo 105. Holotipo: Herb. Roig 5857 HAC. Isotipo: HAC." Because Borhidi and Fernández (1989:304) and Borhidi et al. (2017:107) did not specify which of the two specimens in HAC is the holotype, a lectotype needs to be designated.

In HAC there are two original specimens that were originally in two different herbaria. One specimen has a label with the heading "PLANTAS DE CUBA—HERBARIO DE JUAN T ROIG" and the handwritten annotations "No. 5857 [...] Puerto Padre, Diciembre 8 de 1931 [...] M. Curbelo X 105." On the sheet is affixed another label with the annotation "HOLOYPUS. Exostema curbeloi Borhidi et Fernández" handwritten by Borhidi in 1988. This specimen (Fig. 7) consists of seven branchlets with numerous leaves. Several branchlets have terminal infructescences with immature fruits. One branchlet has terminal infructescences
<figure><img src="image_10.png" /><figcaption>FIG. 7. Lectotype of *Exostema curbeloi* Borhidi & M. Fernández (*Curbelo X 105*, HAC [Roig Herbarium - Plantas de Cuba No. 5857]). = *Solenandra parviflora* (Bonpl.) Delprete. Reproduced with permission by the Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana, Cuba.</figcaption></figure>
with dehisced capsules, and the other branchlet, without leaves, has an inflorescence with flower buds. This specimen is here designated as the lectotype of *Exostema curbeloi*.

The other specimen in HAC has a label with the heading "HERBARIO DEL COLEGIO DE LA SALLE" and the annotations "Núm. 15840 [...] Pto. Padre (Oriente), M. Curbelo X 105, Fecha Dic. 8. 1931." The label heading means that this specimen was originally part of the LS herbarium, which was later integrated in HAC. On the sheet is affixed another label with the annotation "Exostema curbeloi Borhidi et Fernández" handwritten by Borhidi in 1988. This specimen consists of four branchlets with numerous leaves and terminal infructescences with immature fruits, and is an isolectotype of *Exostema curbeloi*.

*Exostema microcarpum* Borhidi & M. Fernández, Acta Bot. Hung. 35:304. 1989. *Solenandra microcarpa* (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:227. 2002. Type: CUBA. Santiago de Cuba: Cercanías de la desembocadura del Río San Juan, Playa de Aguadores, 26 Oct 1952. M. López-Figueiras 718 (LECTOTYPE, here designated: HAC [ex SV]; isolectotypes: HAC [ex LS], NY [00077354], US [00589157]).

**Notes.**—Borhidi and Fernández Zequeira (1989:304), in the protologue of *Exostema microcarpum* Borhidi & M. Fernández, cited the type as "Holotypus: L. F. 718. HAC; Cuba; prov. Oriente; Santiago de Cuba, cerca de la desembocadura del Río San Juan, Playa de Aguadores. Leg.: M. LOPEZ FIGUEIRAS 26. 10. 1952. Isotypus: HAC, HAJB, NY." Borhidi et al. (2017:111; 2018:310) cited the type of *E. microcarpum* as "Tipo: L.F. 718. Cuba; prov. Oriente; Santiago de Cuba, [...], 26. Oct 1952. Holotipo: HAC. Isotipos: HAC, HAJB, NY." Because Borhidi and Fernández Zequeira (1989:304), and Borhidi et al. (2017:111; 2018:310) did not cite which of the two specimens in HAC is the holotype, a lectotype needs to be designated for this name.

In HAC there are two specimens of *Lopez-Figueiras 718* that originated from two different herbaria. One specimen has a label with the heading "REPUBLICA DE CUBA—ESTACION EXPERIMENTAL AGRONOMICA" and the handwritten annotations "L.F. 718 [...] M.Lopez Figueiras, Octubre 26/952." The label heading means that this specimen was originally in SV, which is now integrated in HAC. On the sheet is also affixed a label with the annotation "HOLOTYPUS: Exostema microcarpum Borhidi et Fernández" handwritten by Borhidi in 1988. The specimen consists of a ramified branch, with numerous leaves, and numerous inflorescences; some them with flowers in anthesis, and some others with flower buds. This specimen is here designated as the lectotype of *Exostema microcarpum*.

The other specimen of *López-Figueiras 718* in HAC has a label with the heading "HERBARIO DE LA SALLE" and the typewritten annotations "718. Fecha: Oct 26, 1952. Col.: M. López Figueiras." The label heading means that this specimen was originally in LS, which is now integrated in HAC. On the sheet is also affixed a label with the annotation "ISOTYPUS: Exostema microcarpum Borhidi et Fernández" handwritten by Borhidi in 1988. The specimen consists of a ramified branch, with numerous leaves, and numerous inflorescences; some of them with flowers in anthesis, and some others with flower buds. This specimen is an isolectotype of *Exostema microcarpum*.

*Exostema pervestitum* Borhidi & M. Fernández, Acta Bot. Hung. 35:307. 1989. *Solenandra pervestita* (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:230. 2002. TYPE: CUBA. GUANTANAMO: Baracoa, Maísi, vertientes, Apr 1939, *Bro. León 18964* (HOLOTYPE: HAC [ex LS]; ISOTYPES: GH [00061491], MO [No. 2285941], NY [00077360], US [00811016]).

*Exostema myrtoides* Alain, Phytologia 70:151. 1991. TYPE: DOMINICAN REPUBLIC. BARAHONA: Polo—Los Arroyos, 28 Feb 1969, *Bro. A. Liogier 14342* (HOLOTYPE: NY [00077381]; ISOTYPE: GH [00061488]).

*Exostema pulverulentum* Borhidi, Acta Bot. Hung. 37:79. 1992. *Solenandra pulverulenta* (Borhidi) Borhidi, Acta Bot. Hung. 44:230. 2002. TYPE: CUBA. SANTA CLARA: Trinidad Mountains, San Joss, on low cliffs, 29 Jul 1936, L.B. Smith, A.R. Hodgdon & F. González 3268 (HOLOTYPE: NY n.v. [probably lost]); ISOTYPES: F [No. 858347], GH n.v., JPU n.v., S [No. S07-15068], US [00589155]).

**10. Solenandra polyphylla** (Urb. & Ekm.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:388. 2018. *Exostema polyphyllum* Urb. & Ekman, Ark. Bot. 22A(10):86. 1919. HAITI: Massif de la Hotte, central group, St-Louis du Sud, 1 Nov 1937, E.L. Ekman 9212a (LECTOTYPE (Paudyal et al. 2018:388): S [No. 07-14969]; ISOLECTOTYPES: S [No. 05-557, No. 07-14959], US [No. 1413356, barcode 00589159]).

**Distribution.**—Haiti (Massif de la Hotte).

**11. Solenandra rotundata** (Griseb.) Paudyal & Delprete, Bot. J. Linn. Soc. 187:388. 2018. *Exostema rotundatum* Griseb., Pl. Wright. 2:504. 1862. *Exostema ellipticum* var. *rotundatum* (Griseb.) Maza, Anales Soc. Esp. Hist. Nat. 23:286. 1894. TYPE: CUBA. HOLGUÍN: Monte Verde, "prope villam Monte Verde," Jan–Jul 1859, C. Wright 1258 (FIRST-STEP LECTOTYPE (Borhidi & M. Fernández 1989: 295); SECOND-STEP LECTOTYPE (Paudyal et al. 2018: 388): GOET [GOET003385]; ISOLECTOTYPES: BR [00000530580], K [K000173633], MO [No. 2091691], PH [00013987], YU [YU001738]).

**Distribution.**—Cuba (Holguín: Moa, Monte Verde).

**Notes.**—Grisebach (1862:504), in the protologue of *Exostema rotundatum* Griseb., cited the two gatherings Wright 1258 and Wright 1259. Borhidi and Fernández-Zequeira (1989:295) cited the type of this taxon as "Tipo: Wr. [Wright] 1258, Monteverde," which is a first-step lectotypification, because they did not cite the herbarium of deposit. Borhidi et al. (2017:115) cited the type of *E. rotundatum* as "Tipo: Cuba Oriental, Monteverde, C. Wright 1258. Holotipo: GOET, isotipos: GH, HAC." Borhidi et al. (2018:311) cited the type of this name as "Tipo: Cuba, Oriente: Guantánamo Prov. [sic! Holguín Prov.], [...], C. Wright (1258), Jan–Jul
1859 (fl, fr). Lectotipo: GOET; isolectotipos: BR, HAC, K, MO, PH, MO, PH, YU." According to the *Code*, starting from 1 January 2001, the designation of a lectotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al.'s (2017:115) "holotype" citation, and Borhidi et al.'s (2018:311) "lectotype" citation cannot be treated as valid lectotypifications. Paudyal et al. (2018:388) designated the GOET specimen with barcode GOET 003385 as the second-step lectotype for this name.

*Exostema obovatum* Alain, Contr. Ocas. Hist. Nat. Colegio De la Salle 17:2. 1959. **TYPE:** CUBA. **HOLGUIN:** Moa, Mina Cayoguan, 2 Jul 1945, *Bro. Clemente* 4483 (**HOLOTYPE:** HAC [ex LS]; **ISOTYPES:** GH [00046004], NY [00077380], US [00130604]).

**12. Solenandra sanctae-luciae** (Kentish) Paudyal & Delprete, Bot. J. Linn. Soc. 187:389. 2018. *Exostema sanctae-luciae* (Kentish) Britten, J. Bot. 53:138. 1915. *Cinchona sanctae-luciae* Kentish, Exp. Obs. New Sp. Bark 51. 1784. *Cinchona caribaea santae* luciae Davidson in Morgan, Trans. Amer. Phil. Soc. 2:292. 1786. *Cinchona santae-luciae* Davids. ex Roem. & Schult., Syst. Veg. 5:19. 1819. *nom. illeg. superfl.* **TYPE:** ST. VINCENT: Leeward side of the island, s.d., A. *Anderson s.n.* (**LECTOTYPE** (McDowell 1996: 293): BM [BM000028083]).

**Distribution.**—Lesser Antilles (St. Vincent, St. Lucia, Guadeloupe, Dominica, Martinique).

*Cinchona floribunda* Sw., Prodr. 41. 1788. *Exostema floribundum* (Sw.) Schult., in Roemer & Schultes (as "Exostemma"), Syst. Veg. 5:19. 1819. **TYPE:** ANTILLES [St. Lucia, Martinique, Hispaniola], s.d. [1784–1786], *O. Swartz s.n.* (**LECTOTYPE**, **here designated:** M [M-0187320]).

**Notes.**—Swartz (1788:41), in the protologue of *Cinchona floribunda* Sw., cited the localities "St. Lucia, Martinica, Hipaniola." According to Stafleu and Cowan (1986:117) "The Swartz herbarium constitutes, with the Alstrolmer herbarium, the basis of [...] S. [...] The West-Indian collections (1784–1786) at S, however, are not complete." Stafleu and Cowan (1986:117) added "Swartz was very liberal with his specimens and Swartz types will be found for instance in BM and LD." Hence, Swartz's specimens are present in many herbaria.

There is a specimen in M, barcode M-0187320, which has a label with the annotations "Cinchona floribunda Sw., Prodr. p. 41, Fl. Ind. Occ. 1 p. 375. Exostema floribunda R. & Sch., Syst. Veg. V p. 19. India occident. Ol. Swartz. Herbarium Schreber." The specimen consists of a small branch with several leaf pairs and a terminal inflorescence with flower buds. This specimen is here designated as the lectotype of *Cinchona floribunda*.

*Cinchona luciana* Vitman, Summa PL, Suppl. 1:264. 1802. **TYPE:** Not found.

**13. Solenandra selleana** (Urb. & Ekm.) Borhidi, Acta Bot. Hung. 44:230. 2002. *Exostema selleanum* Urb. & Ekm., Ark. Bot. 22A(10):85. 1929. **TYPE:** HAITI: Massif de la Selle, Ganthier, gorge of Rivière Blanche, 12 Dec 1926, E.L. *Ekman H-7358* (**LECTOTYPE**, **here designated:** S [No S07-14966]; **ISOLECTOTYPES:** A [00046012], G [G00436064], GH [00046011], K [000173628], NY [00077398], S [Nos. S05-561, S05-563, S07-14965], U [U0044609], US [00130607, 00589162]).

**Distribution.**—Haiti (Massif de la Selle, Montagnes Noires) and Cuba (Guantánamo: San Antonio del Sur, Abra de Mariana; Santiago de Cuba).

**Notes.**—Urban and Ekman (in Urban, 1929:85) in the protologue of *Exostema selleanum* Urb. & Ekm., cited the material studied as "Haiti, Plaine Cul-de-Sac prope Ganthier in faucibus ad Riv. Blanche 250 m, alt., m. Dec. flor. et fruct.: [Ekman] n. H 7358," without citing the herbarium of deposit. The original material at B that they studied was destroyed during WWII. In S there is specimen, accession number S07-14966, that has a label with the heading "Mus. Botan. Stockholm" and the collection data handwritten by Ekman. On the sheet are mounted several branches with infructescences and inflorescences with flowers in anthesis. Because Ekman is the collector and the author of the name, this specimen is here designated as the lectotype of *E. selleanum*.

*Exostema scabrum* Borhidi & M. Fernández, Acta Bot. Hung. 35:304. 1989. **TYPE:** CUBA. **GUANTÁNAMO:** San Antonio del Sur, Abra de Mariana, Monte seco, 21 May 1982, *J. Bisse*, *M. Bassen*, *M. Díaz*, *H. Dietrich*, *L. Glez* & K. *Günther* 48177 (**HOLOTYPE:** HAJB [HAJB G 000464]).

*Exostema cordatum* Borhidi & M. Fernández, Acta Bot. Hung. 35:303. 1989. *Solenandra cordata* (Borhidi & M. Fernández) Borhidi, Acta Bot. Hung. 44:227. 2002. **TYPE:** CUBA. **SANTIAGO DE CUBA:** Sardinero, Dec 1948, *Hermelia Casas* LS No. 6331a (**HOLOTYPE:** HAC [ex LS]; **ISOTYPES:** loc. cit., Dec 1948, *Bro. Clemente* LS No. 6331b (GH [00061492, 00061493], HAC [ex LS], US [00589153, 00589154]).

**Notes.**—Borhidi and Fernández Zequeira (1989:303–304) cited the type of *Exostema cordatum* Borhidi & M. Fernández as "Holotypus: 6331ª Sra HERMELIA CASAS (HAC); Cuba, prov. Oriente, Sardinero, en paredón. Dec. 1948.—Isotypus: 6331ᵇ Hno CLEMENTE (HAC)" [underline by the authors]. The gatherings *Hermelia Casas* LS No. 6331a and *Bro. Clemente* LS No. 6331b where made on December 1948 from the same plants.
14. Solenandra stenophylla (Britton) Paudyal & Delprete, Bot. J. Linn. Soc. 187:389. 2018. *Exostema stenophyllum* Britton, Bull. Torrey Bot. Club 42:517. 1915. TYPE: CUBA. ORIENTE: Río Guayabo, 21 Jan 1920, J.A. Shafer 3623 (HOLOTYPE: NY [0007358]; ISOTYPES: F [No 450871, MO [No. 805456, barcode 716665], US [No. 792667, barcode 00130609]]).

*Distribution*.—Cuba (Holguín: Sierra de Nipe, Sierra de Moa).

15. Solenandra triflora (W. Wright) Paudyal & Delprete, Bot. J. Linn. Soc. 187:389. 2018. *Exostema triflorum* (W. Wright) G. Don, Gen. Hist. 3:481. 1834. *Cinchona triflora* W. Wright, London Med. J. 8:240. 1787. TYPE: JAMAICA: Manchioneel Parish, s.d., *T. Dancer & O.P. Swartz s.n.* (FIRST-STEP LECTotype (Andersson 1992: 97), SECOND-STEP LECTotype (Paudyal et al. 2018: 389): BM [BM000028092]).

*Distribution*.—Jamaica (St. Ann's and Manchioneel Parishes).

THOGSENNIA

Aiello (1979) transferred *Gonianthes lindeniana* to the monospecific genus *Thogsennia*, because the name *Gonianthes* was previously used by Blume (1823) for a genus of Burmanniaceae. The name *Thogsennia*, as explained by Aiello, is an anagram of *Gonianthes*. Robbrecht and Bridson (1994 ["1993"]) discussed the nomenclatural problems of *Gonianthes*, *Thogsennia*, and *Cubanola*. Aiello (1979:117) stated that *Thogsennia* is similar to *Portlandia* in having large flowers and horizontal seeds, and that the former differs from the latter by having thinner leaves, placenta forking in cross-section (vs. placenta linear and adnate for the whole length to the midline of the septum), indehiscent fruits (vs. loculicidal capsules), and seed testa collapsed-colliculate to reticulate, with cells elongated and wrinkled (vs. testa tuberculate with the tuberculae collapsed, funicle persistent and with cup-shaped swelling at apex). *Thogsennia* is a monotypic genus occurring in Cuba and Dominican Republic.

*Thogsennia* Aiello, J. Arnold Arbor. 60:116. 1979; Standley, N. Amer. Fl. 32(1):11-12. 1918 (as *Portlandia lindeniana*); Liogier, Fl. Cuba 5:27. 1962 (as *Portlandia lindeniana*); Liogier, Fl. Española 7:438-439. 1995; Aiello, J. Arnold Arbor. 60:116-118. 1979; Borhidi et al., Rubiáceas Cuba 445. 2017. TYPE: *Thogsennia lindeniana* (A. Rich.) Aiello

1. **Thogsennia lindeniana** (A. Rich.) Aiello, J. Arnold Arbor. 60:117. 1979. *Gonianthes lindeniana* A. Rich., in R. de la Sagra, Hist. Fis. Cuba, Bot. 11:10, tab. 49bis. 1850. TYPE: CUBA. SANTIAGO DE CUBA: ["San Yago"], forêt entre Jagua et Tanamo, May 1844, *J. Linden* 1799 (HOLOTYPE: P [P02273499]; ISOTYPES: F [No. 635306 (frag. ex G)], G [G00436806], P [P00582089], MPU [MPU021510]).

*Distribution*.—Cuba (Santiago de Cuba, Guantánamo, Holguín), Dominican Republic (Santo Domingo, Santiago).

*Notes*.—Achille Richard (in Sagra 1850:10), in the protologue of *Gonianthes lindeniana* A. Rich., cited the material studied as "Crescit in insula Cuba, ubi detexit et benigne communicavit clar. Linden," without citing the herbarium of deposit.

Borhidi et al. (2017:445) cited the type of *Gonianthes lindeniana* as "Tipo: Linden, tab. 49." It is unknown to me to which Linden table he made this reference. Also, according to the *Code*, starting from 1 January 2001, the designation of a lectotype or a neotype must be accompanied by "here designated" or a similar expression. Hence, Borhidi et al. (2017:445) type citation cannot be treated as an inadvertent neotype designation.

In P, where Achille Richard worked, there are two specimens of *Linden* 1799 associated with this name. The specimen with barcode P02273499, has a label with the heading "Herbarium Richard" handwritten in red ink, and the annotation "Gonianthes Lindeniana nob. Cuba. M. Linden (no. 1799)" handwritten in black ink by Achille Richard. This specimen, consisting of five leaves and two corollas, is the holotype of *G. lindeniana*.

The other P specimen, with barcode P00582089, has a label the printed heading "HERB. MUS. PARIS" and the annotations "Portlandia gypsophyla Macf.? San Yago_Cuba_M. Linden Cat. N°_1799." This specimen, not annotated by Richard, consists of a branch with several leaves, two loose leaves, and a separate corolla, and is an isotype.
I started working on members of the tribe Chiococceae while I was a Ph.D. Student at University of Texas at Austin, during 1990–1996, under the guidance of Billie L. Turner (1925–2020). During the following decades, I continued to study this group, with projects focusing on morphology, floristics, palynology, taxonomy, phylogeny, and systematics. Due to my moves to different institutions and numerous concurrent long-term projects, the study of this tribe was stopped and restarted several times. This gave me the opportunity to visit numerous herbaria and undertake important field work in many Neotropical countries.

My first expedition dedicated to the Chiococceae was made in Jamaica, in July 2001, accompanied by Tim Motley (1965–2013; then at the New York Botanical Garden), George Proctor (1920–2015; Institute of Jamaica, Natural History Division, Kingston), Andreas Oberli (Hope Gardens, Kingston), Luke Lee, and Kevin Boswell (Forestry Department of Jamaica). During that expedition, I observed and collected natural populations of all the species of *Portlandia*, and several species of *Chiococca*, *Exostema*, *Erithalis*, and *Phialanthus*. My second expedition dedicated to the Chiococceae was made in Dominican Republic, also in July 2001, accompanied by Teodoro Clase (Jardín Botánico Nacional, Santo Domingo). During the expedition in Dominican Republic, I had the occasion to observe and collect natural populations of numerous species of *Catesbaea*, *Chiococca*, *Cubanola*, *Erithalis*, *Exostema*, *Isidorea*, *Scolosanthus*, and *Solenandra*. Funds for field-work in Jamaica and Dominican Republic were provided by the Lewis B. and Dorothy Cullman Foundation, the Plant Genomics Consortium, and The New York Botanical Garden.

My third expedition dedicated to the Chiococceae took place in Cuba, from June to August 2002. Numerous local botanists helped me with logistics, obtaining permits, collecting and processing specimens, assistance with herbarium work, and accompanying me during field trips: Ramona Oviedo-Prieto, Lázara Sotolongo Molina, Daisy Albert, and Mayra Fernández, all of them of Instituto de Ecología y Sistemática, Cuban Academy of Sciences, La Habana. Their contribution was crucial for the realization of field work, during which I observed and collected the natural populations of numerous species, most of them endemic to Cuba, of *Catesbaea*, *Ceratopyxis*, *Chiococca*, *Cubanola*, *Exostema*, *Isidorea*, *Phialanthus*, *Ramonadoxa*, *Schmidtottia* (incl. *Ceuthocarpus*), *Scolosanthus*, and *Siemensia*. I flew from Cuba to the USA with eight boxes of herbarium specimens, half of the boxes containing my collections, and the other half containing loans of Chiococceae specimens from HAC and HAJB. At my arrival at the New York airport on 31 August 2002, all the boxes were confiscated by the US Department of Agriculture for custom control, and were released to me three months after. Funds for field work in Cuba were provided by the American Council of Learned Societies/Social Science Research Council Working Group on Cuba.

Field observations of *Chiococca alba* and *Coutarea hexandra* in Central-Western Brazil, in the states of Goiás, Tocantins and Mato Grosso, were made during 2004–2008, while I was working as a Visiting Scientist at the Federal University of Goiás (UFG), with a fellowship from the Conselho Nacional de Pesquisa (CNPq) of the Brazilian Government (grant 309885/2003–5). During that period I also had the occasion to study several populations of *Salzmannia* on the costal dunes of the state of Bahia. Special help during that period was provided by Juliana Costa da Silva (now my wife), who accompanied me during numerous expeditions in those regions.

Field work in French Guiana was made from 2009 to present, while working at the IRD. During this period, Juliana Costa da Silva accompanied me during field excursions, and helped me processing herbarium specimens of *Chiococca alba* and *Coutarea hexandra*. Funds for field work in French Guiana were provided by the IRD (Institut de Recherche pour le Développement).

The following members of the *Muséum National d'Histoire Naturelle* of Paris are here acknowledged: Odile Poncy and Cecile Aupic, for help with historical herbaria (P-JJR, P-JU, P-AD, P-LA), Sovanmoloy Hul, for her supervision during the study of Rubiaceae specimens of the general herbarium (P), and Gerard G. Aymonin (1934–2014), for conversations about Aublet specimens, help with taxonomic matters, facilitating access to historical collection, supplying historical literature, and identifying the authors of historical handwritings on herbarium specimens.
Nicolas Fumeaux and Laurent Gautier helped me during the consultation of Rubiaceae specimens in the general herbarium (G) and in the Candolle herbarium (G-DC) of the *Conservatoire et Jardin Botanique de la Ville de Genève*. Fernand Jacquemoud, also of that institution, produced important digital images of specimens in the Candolle Herbarium (G-DC). Funds for that project were provided by the IRD.

I am extremely grateful to the directors and curators of the following herbaria for loans of material and/or for providing working space during my visits: B, B-W, BBS, BM, BR, BRB, C, CAS, CAY, CEN, COL, E, F, FI, FT, G, G-DC, HEPH, HTO, IAN, IBGE, INPA, K, L, LINN-SM, MBM, MG, MO, MPU, NY, NX, P, P-AD, P-BONPL, P-JJR, P-JU, P-LA, R, RB, U, UB, UFG, UFMT, UMP, US, and VEN. A special thank is due to the staff of the CAY Herbarium—Herbier IRD de Guyane, https://dx.doi.org/10.23708/herbier-guyane-ird—for their help in handling the loans of herbarium specimens from numerous institutions.

I also wish to thank the staff members that contributed to the following virtual herbaria: Jstor Global Plants (https://plants.jstor.org/), Reflora—Brazilian Plants: Historic Rescue and Virtual Herbarium for Knowledge and Conservation of the Brazilian Flora (http://reflora.jbrj.gov.br/reflora/PrincipalUC/PrincipalUC.do), and INCT—Herbario Virtual da Flora e Fungos (http://inct.splink.org.br/) for making millions of images of herbarium specimens available to the general public.

A special thank is due to Rosie Jones, Special Collection Librarian of the Natural History Museum, London, who, in 2024, carefully searched for a possible drawing of *Cinchona caribaea* in the volume "Von Jacquin botanical notes. 18th-century botanical drawings and notes made by French scientist and botanist Nicolaus Joseph von Jacquin (1727–1817)." I am also grateful to Ranee Prakash, herbarium curator of the Natural History Museum of London, for searching for type specimens of *Cinchona caribaea*.

In March 2025, I studied additional Chiococcaceae specimens, including types, in three Cuban herbaria at La Habana. During that study, I was assisted by Ramona Oviedo-Prieto, Daisy Albert, and Ilsa Fuentes (HAC, Instituto de Ecología y Sistemática, Cuban Academy of Sciences), Heldis R. Bécquer (HAJB, Jardin Botánico Nacional, Universidad de La Habana), and Amarillys Leyva Hernández (MNHN, Museo Nacional de Historia Natural de Cuba). I am very grateful to Bruno Enrique Tellez Baños (Universidad Autónoma Metropolitana de México, México City, México) for taking high resolution images of numerous Rubiaceae type specimens of the HAC Herbarium.

Important help regarding nomenclatural issues was provided by Joseph H. Kirkbride Jr., John H. Wiersema (both at the Smithsonian Institution, Washington, D.C., USA), Kanchi Gandhi (Harvard University, Cambridge, Massachusetts, U.S.A.), and John McNeill (Royal Botanic Garden, Edinburgh, Scotland). Last, but not less important, I wish to express my gratitude to the Joseph H. Kirkbride and Michael Nee for the detailed revision of the manuscript, and for their significant comments and corrections.

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