RHODORA, Vol. 104, No. 918, 186-200, 2002 pp. NEOTROPICAL APOCYNACEAE STUDIES IN II A REVIEW OF THE GENUS FERNALDIA Morai Francisco fs J. B Nacional dc odi vers dad (INBio), InslitLito i i Domingo Santo de Heredia, Apto 22-3100, Costa Rica e-mail; [email protected] A ABSTRACT. synopsis of the three species Fernaldia, a genus of Neo- o'i A tropical Apocynaceac restricted to Central America, presented here. sum- is mary of descriptive morphology, specific relalionsliips, and synonymy pro- is vided. Key Words: Gentianales, Apocynaceac, Apt)cynoideae, Fenuildia, Neo- tropics Woodson Fernaldia a genus o{ vines characterized by eglan- is dular leaves that lack colleters along the midrib adaxially, sepals with racemose coroMa a single collctcr within, inflorescence, tube without annular corona or free corona lobes within, and corolla The was lobes usually villose adaxially. genus proposed by first Woodson 1932 honor of Merrit Lyndon Fernald (1873- in in 1950), of the Gray Herbarium Harvard University. occurs (cii), It from Mexico Panama. northern Despite ornamental and to their edible features, these plants are rarely collected and very few herbarium specimens Fernaldia was by Wood- exist. last treated son when two were known. new names (1936), species Several for species or varieties have been published since then, but no recent revision exists. Therefore, a revision of the genus pre- is A sented here. key to the species, along with descriptions, illus- trations, and citations of selected specimens are given below. Specimens from Petersburg (formerly Leningrad), Russia St. (le), cited here were examined the Missouri Botanical Garden (mo), at where some on material loan. is NOTEWORTHY MORPHOLOGICAL FEATURES Sepals. In Fernaldia, the sepals are further solitary, truncate, and sometimes very deeply The lacerate apically (Figure se- 1). pals are characterized by having a single colleter within adaxially. Within the subfamily Apocynoideae, the sepal colleters can be a very helpful character distinguish genera. to 186 — Morales Review of FernaUlia 187 2()02J A. B C www 2 B Figure Fenialdiu sepals. A. F. speciosissiiua {Morales 713/, inb); 1. F. pcuidiirata (Morales 3074, iniO; C. F. asperogloitis (Mexia 8751 inb). , When Woodson Corolla pubescence. Femaldia described (1932), one of the features mentioned by him to distinguish the was genus the conspicuously arachnoid-villous corolla lobes. Af- ter careful field study of the three species of the genus, it is pubescence mostly suitable to note that the restricted to the is corolla throat and just at the base of the lobes. The hairs are long mm), (2—3 always conspicuous and fresh material, invariably in white to greenish-white. Tn related genera (e.g., Ecliites, Temnci- clenia), the upper part of the corolla tube and the mouth are gla- brous and never with long to glabrate, hairs. Following Morales key and (1999), the species descriptions in in this paper, the lower pari of the corolla tube measured from is the base of the corolla up to the position where expanded is it The abruptly (stamens attachment). length of the upper part is measured from this point to the base of the lobes. Anthers, The anther shape Femaldia a very important in is feature to recognize this genus from other closely related genera such as Ecliites and Temnadeiiia, In Fenialdia, the anther auricles are usually obtuse to almost rounded basally, while the other in genera the auricles are conspicuously acute acuminate; only to F. Woodson pandurata (Alph. de Candolle) has auricles very shortly and broadly However, most acute. the striking difference the is Rhodora 188 104 [Vol. B A. mm 6 Figure 2. FemaUlia anthers. A. F. cisperoghfiti.s (Mexia (S751, inb); B. F. {Morales 3074, C. speciosis.sima (Morales 7I3I, pandiircitci inb); F. inb). presence of a hyaline border the anthers of Fernaldia (Figure in a character never present the three other related genera. in 2), TAXONOMIC TREATMENT Fernaldia Woodson, Ann. Missouri Bot. Gard. 48. 1932. Type: 19: Woodson. pcuidurata (Alph. de Candolle) F, Browne, Echites Civ. Nat. Hist. Jamaica. 182. 1756, in part. P. Maudevilla Lindl., Edward's Bot, Reg. 26: 7. 1840, in part, jiofn, cons. t. Ainhlyanthera Milll. Arg., Fl. Bras. (Marlius) 6(1): 141. 1860, in part. Urechites Mull. Arg., Bot. Zeitung (Berlin) 18: 22. 1860, in part. Aiigadenia Miers, Apocyn. S. Ani. 173. 1878, in part. Stems SuffiTiticose lianas. terete to subterete, glabrous or gla- brate to very minutely and variously puberulent; nodes with few intrapetiolar conical to conical-fusiform inconspicuous colleters. — Morales Review of Fenialdia 189 20021 Leaves opposite, petiolaLe, petiole slightly fused the base; blade at glabrous or glabrate variously puberulent beneath, eglandular, to racemose, without basal colleters adaxially. Inflorescence axillary, usually many-flowered, very minutely puberulent to glabrous or glabrate, pedunculate, bracts scarious, inconspicuous. Sepals 5, essentially equal, very slightly imbricate basally, with a single colleter within, truncate, entire to variously and very minutely erose or fimbriate; corolla infundibuHform, glabrous or glabrate very minutely and variously puberulent without, the lobes var- to iously pubescent adaxially (very rarely almost glabrous in Fer- sometimes naldia pandiirata) with long white these hairs, re- stricted mostly to base of the lobe or around the corolla mouth; W^e tube without annular corona or corona lobes within, straight, limb actinomorphic, dextrorsely convolute; stamens the 5-parted, included, inserted the upper part of the corolla tube; anthers 5, in connivent and adnate to the pistil head, conformed by 2 parallel, uniformly thecae borne adaxially near the apex of an en- fertile larged, peltate connective; auricles almost inconspicuous, broadly rounded basally; carpels 2, united at the apex; pistil head fusiform or subcapitate; ovules numerous, several-seriate, borne on an ax- biseriate placenta; disk annular, usually 5-lobed. Follicles ile, 2, apocarpous, continuous, glabrous dehiscing along the to glabrate, comose numerous, ventral suture; seeds dry, truncate, apically, usually minutely rugose. The genus comprises ranging from Mexico three species, to Panama. northern KEY TO THE OF SPECIES I.RNALDIA I mm 8- Lower 2-5 part of the corolla tube long; corolla lobes 1. mm X 7—9 12 F. aspcroglottis \ . mm Lower 18-30 part of the corolla tube long; corolla lobes 1. mm 9-24 X 9-15 (2) Peduncles 2-4.5 cm; upper part of the corolla tube 10-15 2. mm mm 5-6 long; anthers long; dry forest pcujdurata F, 2. 19-24 Peduncles 15-23 cm; upper part of the corolla tube 2. mm mm wet long; anthers 1.5-12.5 long; forest 1 . . . . speciosissima F, 3. Fernaldia aspcroglottis Woodson, Ann. Missouri Bot. Card. 26: 1. N Madre 96. 1939. Type: mhxico. Guerrero: Sierra del Sur, Rhodora 190 [Vol. 104 Nov of Rfo Balsas, 5 1937 Mcxia 8751 (iiolotypr: moi; (fl), isoTYPRs: ARIZ not seen, CAS!, ny!, photograph Field F!, c.!,riii!, 56468 negative inb ex n). Fi^jure at 3. Liana; branchlets sparsely puberulent to glabrate; nodal collc- ters inconspicuous or absent, only intrapetiolar colleters present. X L8-8 Leaves: petiole (0.3-) 0.6-2.1 cm; blade (-11) 0.8-6.8 cm, membranaceous, ovate, narrowly ovate caudate- to elliptic, acuminate at the apex, obtuse, attenuate to obscurely cordate ba- sally, glabrate above; densely puberulent to glabrate beneath, more rarely glabrous. Inflorescence lax, longer than the subtend- 2-16 ing leaves, many-flowered, puberulent; peduncle (-20) cm; 4-9 mm; X mm, pedicels bracts 1.5-2 scarious; sepals 1-1.5 1 X — mm, 1.5 ovate to narrowly ovate, acute, sparsely puberulent, 1 mm colleters ca. 0.5 long, variously erose apically to conspicu- ously lacerate; corolla white to white-yellow, sparsely puberulent 2-5 X 2-3 mm; to glabrate without; lower part upper part broadly mm 9-19 X 7-12 conical conical-camiDanulate, diameter to in at X mm, 8-12 7-9 the orifice; lobes obovate to narrowly-obovate, mm, mm, 4.5-6 spreading; anthers glabrous; ovary 1.5-2 gla- mm mm; brous; style head ca. 1.5 disk ca. 1,5 long, annular, 20—23 X irregularly lobed. Follicles 0.5-0.6 cm, smooth and gla- X brate; seeds 1—1.2 0.2 cm, rugose, very minutely and incon- coma spicuously puberulent, 3-3.5 cm, creamish. AND DISTRIBUTION PiinNOLOGY. This spccics restricted to Mexi- is Guerrero, Michoacan, and Mexico 200-1350 m. co, in states, at Fernaldia (isperoi^lollis flowers September to March. Fruits are borne from October to April. Feruahlici asperoi^lottis distinguished from other species is in genus by having conspicuously the a short corolla tube. Further- more, the corolla lobes are shorter than any of the other spe- in mm 8-12 may some cies, only long. This species also resemble species of the genus Mandevilla, more specifically the complex around M. convolvulacea (Alph. de Candolle) Hemsl. and M. an- which drieiixii (Miill. Arg.) Hemsl., with shares a similar leaf it shape and However, inflorescence structure. F. asperoglotiis is easily recognized by eglandular leaves, without colleters along its midrib the adaxially. SPEZCiMi^NS F.XAMiNi-n: Mi:Xico. GuciTcro: Tcmascaltcpcc, Guayahal, Feb I 1 — Morales Review Femaldia 2002 of 191 B. cm 4 . f . i 1 * " 4 m. ± \ \ : : A cm cm I 4 L Fenuddia {Mexia 875 Calyx Figure asperoi^lottis mo). A. Habit. B. 3. J, and corolla. C. Fruits. D. Seed. 5 Rhodora 192 104 [Vol. 1933 Hintcm 3372 sheds]): Pungarahato, Coyuca, 23 Feb 1934 (n. fV). (k [2 (IV). HifUoff 56<S'j (K [2 sheets]): Teniasealtepee, Guayabal, 16 Jan 193."^ (fl, fr), HifiUffi 7239 (k [2 siieels]); Coyuea, Quebracias, 22 Jan 1935 (fr). Hiutou Mar 726/ (k sheets]): Plaeeres, Cigarillo. 1936 Hinton 9786 [2 II fr), (11, N & of La Union, road Coahuayatla, 24 Oet 1983 Soto Nunez (K); to (fl), 6067 (Mi£XLi, MO). Mexieo: San Antonio Tlatlaya, 25 Jan 1953 Mutuda ct (11), 28003 (MHxu, Mo); between Siillepec and Aniatepee.. 31 Dec 1953 cil. ((1), Miilifda 30097 (MExif. mo); Los Bejucos. Tejupileo. 27 Aug 1954 Miituda (H). ct id. 31389 (MRXLJ. mo); Olzoloapan, Valle del Bra\o. 5 Sep 1954 (fl), Mofuda aL 31416 (mexu, mo); La Junta. Valle del Bravo, Sep 1954 McUudu ct (fl). I 1 ct 31649 (Mi^xu. mo). Michoacan; Tu/anlla-Paso Tierra Caliente. 10 Mar (il. 1938 Hinton 13308 (k.mo.nv); Ajuage, Apatzingan, Oct 1939 13 (fl, fr), (fl), Hinton 15329 Puente Las road Zitacuaro-Huetamo, Dee (c. k. ny. p); Pilas, 3 SW 1983 Lotr 2/57 (inh, mexu. mo); of La Huacana, 31 Dec 1977 (fl), (fr). NW Soto 592 (MF.xu, mo); of La Ercndina, road to Casacuaro, 6 Sep 1981 Soto 3016 (MHxu, mo); San Jeronimo, road Huetamo San Jeionimo, Oct (fl), 8 3227 Nucva 1981 Soto mo); Tuinbiscatio, road Playa Azul, (fl), (mi;xli. Italia 28 Oct 1981 Soto 3586 (mkxli. mo). (fl, fr), Woodson, Fernaldia pandurata (Alph. de Candollc) Ann. Mis- 2. souri Bot. Card. 48. 1932. Figure 19: 4. Echitcs pandurata Alph. de Candolle, Prodr. (DC.) 458. 1844. Am- 8: blyunthera pcuu/urata de Candolle) Mull. Arg., Linnaea 30: {A\\)\\. 448. 1860. Aui^adcnia pandurata (Alph. de Candolle) Miers, Apo- Am. Aug cyn. S. 182. 1878. Tvpr:: Mtixico. Oaxaca: San Dionicio, 1832 Andricux 245 (lectoiyi'I-: selected here: o-dci; isollc- (fl), TOTYPES: photograph Field negali\e 7559 mol nv. rs K!, at inb:. r-!. ex o-DC')- Urec/ntes /<ar\vins/Kii Mull. Aru., Linnaea 30: 440. 1860. Typh: Mexico. Tamaulipas'^: ^^Huefulta/' 1S41-1842 Karwinsky 474 (eexto- (fl), T^Pi- selected here: le'). & Moc, Ec/iitcs barbata Sessc Naturale/.a (Mexico City), Sen 2 2, (App.): 45. 1893, non Desvaux Hamillen 1825, nee. D. Dietrich <:a- mexko: 1839. Type: Data Uickinc August), Scssc Lacasta in \ (fl 567 ma Mo{-iuo not seen: c^ 1 (iiOLOTYPf-:: isor^pr:: :). i & MandcvHla velutiua K. Schuin. Engl. Prantl. Nat. Pflanzenfani. /// 4(2): 171. 1895. Type: Costarica. Data lacking Hoffmann 7/0 (fl), OioEorY}'!:: u-destroyed, photograph Field negative 4533 at inhi, 1. 1 MO!. i;si): COSTA RiCA. Guanacaste: Nandayure. I^aciTico Norte. Be- Aug & juco, Cerro La Picuda, 24 1994 /Estrada Rodrfi^ucz /93 (fl), (NE:o'rYPE selected here: isoneot^pes: inhi; cri, mo-). /Vlandcvilla potosina Brandegee, Univ. Calif. Publ. Bot. 276. 1912. 4: Aug Type: mexico. San Luis Potosi: Rascon, 1911 Purpus (IL fr), 5408 (holot\pe: photograph uci; ISOT^•PEs: bm'. moi. ny', Field f:. 56477 negative inb ex at :). i Columbian Echites Mus., piui^uifolia Standi.. Publ. Field Bot. Ser. 8(1): Gaunwr 35. 1930. Typi:: MEXICO. Yucatan: 1895 8/ Izanial. (fl. fr). (iiolot^pe: isotyph: MO'; photograph Field ncgalive 56462 inb F'; at ex v\). — Review 2002 Morales of Fenuthlia 193 cm 4 cm 4 Figure FernaUlia pcuidunitu {Morales 3074. inb). A. Habil; B. Fruits; 4. C. Seed. » Rhodora 194 [Vol. 104 Fenialdict hrachyphnrynx Woodson, now, Ann. Missouri Gard. Bol. syfi. 19: 380. 1932. Type: guathmaia. Along the road from Escuintla lo Aug the port of San Jose de Guatemala, 23 1860 Hayes s.n. (11), (holotype: gii')- FeniaUlia pcuidiinitci var. glabra Ant. Molina, Ceiba 3: 95. 1952. Fcr- juilclia glabra (Ant. Molina) Lundell, Wrighlia 5: 256. 1976. T^Pi-: HONDURAS. Cortes: faldas de Montana Santa Ana, Rfo Santa Ana, la 6 Dec 1950 Molina 3640 (HOLorYPF: pho- (11), i-api; isorvi'i-s: f!, 56469 tograph Field negative inb ex at \-\). Liana; branchlets sparsely pubei*ulcnt to glabrate; nodal colle- ters inconspicuous or absent, only intrapetiolar colleters present. X Leaves: 0.9-4 cm; blade 5-14 4-11 cm, membra- petiole (17) naceous, ovate, ovate-elliptic, narrowly elliptic to variously pan- din^ate, acuminate to caudate-acuminate at the apex, rounded, ob- tuse to obscurely cordate basally, glabrous to glabrate above, densely puberulent glabrate or more glabrous beneath. to rarely Tnllorescence agglomerate, usually shortei" than the subtending leaves, rarely longer, few- to many-flowered, glabrate to very 4- minutely and densely puberulent; peduncle 2-4.5 cm; pedicels mm; mm, X X 12 bracts 1.5-3 1-1.5 sccirious; sepals 1.5-3 1.5 mm, ovate narrowly very minutely and to ovate, acute, sparsely mm puberulent, colleters ca. 0.5 long, apex variously lacerate; corolla white lo greenish-white or creamish, glabrous to glabrate or very sparsely and minutely puberulent glabrate without; to X mm; lower 18—22 2—3 upper more part part conical, rarely mm X broadly conical, 10-15 8-11 in diameter at the orifice; 9—14 X mm, 9—1 lobes obovate narrowly-obovate, spreading to 1 mm, 5-6 and distally reflexed; anthers glabi-ous to glabrate, rarely mm, minutely puberulent; ovary 1.5-2 glabrous; head style ca. mm mm; 2.5 disk 1-1.5 long, 5-lobed variously to loloed. Folli- 21-36 X cles 0.5-0.6 cm, rusose, glabrous to glabrate; seeds 1.4- X coma 1.6 0.2 cm, minutely rugose, 4-4.5 cm, creamish. DISTRIBUTION, PHENOLOGY, AND LOCAL NAMES. Mexico north- to mosdy western Costa Rica, in dry forest or in open and second- 0-1200 growth m. Flowering June December. forest, ^ ^v^...^ to ^^S.^AW^X.1. FX*r^u,i.tXiLu^LWi^ V^. Known March January. Loroco Guatemala; to locally as (Jutiapa, Cortez, Honduras; Ahuachapan, El Salvador). The available specimens of Fernahlici pamhirahi, the most common and widespread species, show that leaf shape and pu- bescence morphological are features that are highly variable with- Although in this taxon. there are slight differences scattered — Morales Review of Fenuildia 195 2002] throughout the plant's geographic range, they are mostly sporadic ocurrence and do not merit taxonomic recognition. Within the in Apocynaceae subfamily Apocynoideae, the acceptance of species defined only by pubescence unwarranted (Morales 1997, 1999). is Therefore, and subsequent combi- F, pcujclurata var. glabrxi the name nation based on this are not recognized here. Fenuildia bracliypharynx here considered sync^nym a F. is c^f showing Ad- pandunita, only slight differences in corolla length. specimens examined Woodson's monograph ditional since reveal that this feature variable within the geographical range. is Mesoamerica, have been In northern the flowers of this species (Woodson used as a flavoring for rice 1936). For further refer- work Morton ences, see the of (1990). et al. No MandevUla vehitina K. Schum. neotyhed here. other du- is Hoffmann plicates were found bm, or where specimens in c, K, are located. Specimrns f^xavtinf.d: cosiARiCA. Guanacastc: Canas, La Pacffica, 10 Nov 1969 Daiibemmre 256 Mar 1970 Daubenniirc 592 2 (fl, IV), (r usj). (fl, (V). Finca La Pacflica, 3 Sep 1972 Hcithaiis 350 (mo); Parque Nacional (t-;u8J); (fl), NW Palo Verde, 30 July 1994 Mojxiles 3074 (inb); La Pacifica. of (11. fr). Nov Canas, 20 1972 Oplcr 1568 (cr.rmo Palo Verde National sheets]); (fl), [2 Park, Dec 1996 Rodriguez cT 1829 (inb. mo); Palo Verde, 7 Sep (fl, fr), a!. 1 I 1973 Solomon 611 (fl, fr), (cr. r usj). RL SALVADOR. Ahuachapan: San Benito, E of San Alfonso, El Imposiblc, 12 & Jun !993 Scuidova\ Sandoval 1309 ragu,mo); El Imposilole National (fl), (b. Park, San Alfonso, 10 Jul 1990 Seniieno 232 lacui, mo). (fl), (b. Cumbre guatrmala between Ramfre/ and Je Chiquimula, Cliiquiniula: 15 Oct 1940 Standley 74496 mo). Jutiapa: vicinity of Juliapa, 1940 (k (fl), ffl), Aug 75297 Standley mo). Peten: exact locality lacking, 12 1967 Coii- (r (fl), treras 7008 Santa Rosa: vicinity of Chiquiinula, 1940 Sfandlcy (k. rr). (fr), 79683 MO). {[ HONDURAS. Cortes: Santa Ana mountain, Rfo Santa Ana, 6 Dec 1950 (fl), Molina 3640 (bnlrarmo). San Roque, Sep 1948 Standley JVlorazan: Villa (fl), 26257 (bm. r). Nov 47493 MRXico. Chiapas: near Chiapilla. 14 1980 Breedlove (mo); (fl), alons the road from Acala to Venusliano Carran/a, 25 Oct 1966 Lcm^idi}] (fl). & 2669 (mo); Oco/.ocoaulla, 19 Sep 1988 Reyes Urquijo 1015 (bm.mrxu). (fl), Nov 597 Guerrero: Sierra Madrc, 8 1898 (fl, fr), Langlasse (o [3 sheetsl, K. Acapuico, Oct 1894 - Mar 1895 Palmer 259 Tonala, La Jalisco: (k). p); (fl). Cruz, Barranca de la Cruz, 9 Jan 1975 (fr), Diaz. 5484 (rncb, inb). Nayarit: SE of Ahuacatlan, 2 Jan 1986 Teltez 937 mrxu). Oaxaca: exact (fr), J (inb. W 246 locality lacking, 1834 Andrien.x {G-dc, k); of Tuxtepcc, along road (fl), to Ixcallan. 7 Aug 1971 Slevens 1391 (Mt)). San Luis Potosf: Tamazun- (fl). chale, 7 Oct 1937 (d), Taylor 479 (mo. Tamaiilipas: Sierra de Tamauli- ri:x).