Table Of ContentRHODORA,
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).