LANKESTERIANA 6(2): 73-81. 2006.

THREE NEW SPECIES OF MARCGRAVIACEAE FROM COSTA RICA,

WITH REFERENCES TO RELATED SPECIES AND NOTES ON THE
GENERIC PLACEMENT OF SCHWARTZIA JIMENEZII
BARRY E. HAMMEL
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, USA
Instituto Nacional de Biodiversidad (INBio), apdo. 22-3100, Santo Domingo, Heredia, Costa Rica.
ABSTRACT. Marcgravia glandulosomarginata, Ruyschia moralesii and Schwartzia tarrazuensis, woody epiphytes known mostly from mid-elevation rainforest of Costa Ricas pacic coastal mountains just south of
San Jos, are described, compared to related species and illustrated. The new Marcgravia species is known
also from Panama and Colombia, but the other two species are endemic to Costa Rica.
RESUMEN. Se describen, ilustran y comparan con especies anes Marcgravia glanduloso-marginata, Ruyschia moralesii y Schwartzia tarrazuensis, eptas leosas principalmente de bosques lluviosos de alturas
medias en la cordillera costea pacca de Costa Rica, al sur de San Jos. La nueva especie de Marcgravia
se conoce tambin de Panam y Colombia; las otras dos especies son endmicas en Costa Rica.
KEY WORDS / PALABRAS
Colombia.

CLAVE:

Marcgraviaceae, Marcgravia, Ruyschia, Schwartzia, Costa Rica, Panama,

During the preparation of the Marcgraviaceae treatment for the Manual de Plantas de Costa Rica, three
new species for the family have come to light. All of
them are woody epiphytes, and are known from and
can even be found locally sympatric in, though not all
are restricted to, the botanically rich pacic coastal
range just south of San Jos, along the border between
San Jos and Puntarenas provinces. The elevational
range within which these species occur [(700) 9001800 m] encompasses that generally cited as highest
in diversity, especially for epiphytes (see Gentry &
Dodson 1987, Zamora et al. 2004: 195-216). Other
epiphytic or climbing species that have recently been
described from this specic region of Costa Rica include: Apocynaceae - Allomarkgraa insignis J.F.
Morales, Araceae - Philodendron dominicalense Croat
& Grayum, Asclepiadaceae - Matelea costaricensis
W.D. Stevens, Bromeliaceae - Werauhia tiquirensis
(J.F. Morales) J.F. Morales, Cactaceae - Weberocereus
frohningiorum Ralf Bauer, Gesneriaceae -Paradrymonia bullata Gmez-Laur. & M.M. Chavarra (= Nautilocalyx biserrulatus Kriebel), Orchidaceae - Kefersteinia orbicularis Pupulin, Macroclinium confertum
Pupulin, Ornithocephalus castelfrancoi Pupulin. New
country records or rediscoveries of rare epiphytic or
climbing species also from this area include: Asclepiadaceae - Marsdenia laxiora Donn. Sm., Gesneriaceae
- Drymonia peltata (Oliv.) H.E. Moore, Lomariopsidaceae - Elaphoglossum moralesii A. Rojas, Malpighiaceae - Stigmaphyllon bogotense Triana & Planch.,

Orchidaceae - Catasetum integerrimum Hook.

The very diverse morphology of the nectariferous
inorescence bracts, as well as the very strong and
unusual (fruity to rumlike, pers. obs.) oral fragrances
in many species, should make this family fertile ground
for a study of pollination and oral biology. As pointed
out by Ward & Price (2005), few such studies have been
made but known pollinators include bats and birds (see
Dressler & Tschapka 2002), and oddly, cleistogamy.
The presence of oil-bearing pollen, observed on live
material of two of the species described below, as well
as on herbarium specimens of several other species
in different genera of Marcgraviaceae, appears not to
have been mentioned previously in the literature.
Marcgravia glandulosomarginata Hammel, sp. nova
Haec species nova Marcgravia schippii similis,
sed imprimis differt sepalorum glandulis obscuris
glandulisque poriformibus submarginaliter lineatis (ad
vicem inornatis), etiam laminarum foliorum glandulis
obscuris, submarginalibus, conspicuoribus lineatis
quasi a basi ad apicem complectentibus.
[This species is similar to M. schippii, but differs
principally by its sepals with submarginal rows of dark
glands and poriform glands and by its leaf blades with
a more conspicuous submarginal row of dark glands
extending from nearer the base to the tip]

74

LANKESTERIANA

TYPE: Costa Rica; San Jos, cantn de Tarraz, entre

Cerro Toro y Cerro Hormiguero, ca. 15 km al suroeste
de San Marcos, camino a Quepos, 1100 m, 14 jul 2005
(, fr), Hammel et al. 23714 (holotype: INB, isotypes:
COL, CR, FR, MICH, MO, P). Figs. 1, 4A.
Woody liana; fertile branches terete, drying tan to
light grayish tan, sometimes exfoliating, with lines
of darker, conspicuous lenticels to 1.5 x 0.7 mm.
Mature leaves indistinctly petiolate, the petiolar region
to only ca. 1 mm, attened above, rounded below;
blade 10.5-13 (2.5-)3-5 cm, oblong-elliptic, drying
grayish to orangish tan, usually lighter or more orangecolored below, the base often slightly asymmetrical,
abruptly obtuse to rounded, rarely acute, the apex
acute to broadly acuminate; secondary veins indistinct;
hypophyllous glands in two distinct rows on each side
of the midrib, the submarginal of dark, solid, circular
to ovate dots running from a few mm of the midrib at
base to a few mm of the tip, the laminar of 4-8(-14),
mostly narrowly elliptic, poriform glands in a diagonal
line from the base of the midrib towards the margin at
about midleaf. Inorescence erect, the peduncle 0.31 cm with 5-9 owers and 1 nectary, the rachis 2-4
mm, shortly conical; pedicels 4.5-6 cm, ascending at
mostly < 45, conspicuously lenticellate, the lenticels
to ca. 2 mm in diam. and often 4-lobed, the epidermis
drying light grayish tan, often transversely splitting,
exfoliating. Nectary 1, erect, green to dark greenish
purple, lenticellate as on the ower pedicels, the free
pedicel 5-8 mm, the cup 35-50 mm, tubular, clavate,
basally anged to spurred, the spur 2-5 mm, apically
sometimes with a small (to 6 mm) bud. Flower buds
green, erect 13-18(-20) mm, conical with an acute- to
obtusely rounded apex, the 2 bracteoles to 3.5 x 4 mm,
semicircular to broadly ovate, born ca. 2 mm below
the calyx, usually minute or lacking; sepals appressed,
equal (the outer 2 slightly wider than the inner 2), ca.
7 x 11 mm, broadly semicircular, thin-margined with
a continous submarginal row of conspicuous, dark
glandular dots, and just inward of them, at the lower
edges and sometimes nearly continuous, an additional
row of few to many poriform glands; corolla calyptrate
(see ower buds above). Stamens 34-44, prior to
anthesis the laments 4-5.5 mm, attened, the anthers
(3-)5-7.5 mm, oblong-linear, basally sagittate, apically
retuse, releasing pollen in bud. Ovary ca. 8 mm,
obturbinate, 9-loculate. Fruits 12-18 mm, slightly
depressed globose, the sepals spreading.
DISTRIBUTION. This somewhat rare and overlooked
species is known from Costa Rica to Colombia in rain

Vol. 6, N 2

forest at (700-) 900-1750 m elevation.

ETYMOLOGY. The epithet glandulosomarginata refers to
the distinctive rows of dark and poriform glands on the
margins of the sepals and the submarginal row of dark
glands along nearly the entire length of the leaf blade.
DISCUSSION. This species is distinct for its submarginal
rows of dark glands and poriform glands on the sepals,
for its very obvious submarginal row of dark glands
running from very near (1-5 mm from) the base to very
near the tip of the leaf, and for the light grayish tan,
transversely splitting and exfoliating epidermis of the
pedicels. It somewhat resembles M. brownei (Triana &
Planch.) Krug & Urb. in leaf size and shape, and the
diagonal row of numerous poriform hypophyllous
glands, but in addition to the rst stated characters,
differs from that species by having fewer owers and
nectaries, and the pedicels ascending rather than at
a right angle or reexed. However, overall, the new
species is more like M. schippii Standl., with which
it has been confused and with which it agrees in most
aspects, including that of having only one nectary per
inorescence. In addition to the rst stated characters,
it differs from M. schippii by having usually 4-8 or
more instead of only 1-4 poriform hypophyllous
glands on each side of the midrib. The submarginal
hypophyllous glands of M. schippii are relatively pale,
visible beginning no closer than 12 mm from the base
of the blade, and are sometimes accompanied by a
few somewhat poriform glands. Flowering pedicels
of most material of M. glandulosomarginata lack
bracteoles, whereas they are always present on M.
schippii. These two species are somewhat separated
elevationally, M. schippii known mostly from sea level
to below 1000 m, and the new species not known from
below 700 m, is found mostly between 1000-1750 m.
The Colombian material included here has up to 14
hypophyllous glands in the diagonal row, whereas in
Costa Rica, the most seen was eight.
ADDITIONAL SPECIMENS. COSTA RICA. Heredia:
Cantn de Sarapiqu, Parque Nacional Braulio Carrillo,
1060 m, 29 Nov 1992 (st), Boyle 1324 (MO); John &
Kathy Utley 5632 (DUKE, MO). San Jos: Cantn
de Prez Zeledn, Fila Tinamastes, 1140 m, 14 Febr
1996 (fr), Hammel et al. 20205 (INB, MO); cantn de
Tarraz, entre Npoles y Santa Marta, 1400 m, 5 ago
1994 (), O. Vargas 460 (INB, MO); 1360 m, 5 Aug
2005 (, fr) Hammel & Prez 23760 (CR, INB, MO);
13 Jan 2006 (, fr), Hammel et al. 24043 (CR, INB,
F, MO); 1100 m, 17 Febr 2006 (, fr), Hammel et al.

LANKESTERIANA 6(2):

HAMMEL New species of Marcgraviaceae from Costa Rica

24114 (FR, INB, MO). PANAMA. Chiriqu: 1000 m,

2 Aug 1984 (), Churchill 5919 (MO). Cocl: 930 m,
6 Dec 1979 (fr), Croat 49214 (MO); 700-750 m, 27
Nov 1985 (fr), McPherson 7685 (MO). COLOMBIA.
Antioquia: Urrao, Las Orqudeas, 890 m, 1 abr 1988
(fr), Cogollo et al. 2903 (JAUM, MO); 1450 m, 18 oct
1993 (), Cogollo et al. 7066 (JAUM, MO); 1450 m, 5
dic 1993 (, fr), Cogollo et al. 7819 (MO); 1750 m, 24
Sept 1987 (fr), Zarucchi et al. 5722 (MO).

75

5, basally connate to about the level of the sepals, the

lobes reexed, 4-5 x 3.5-4 mm, ovoid. Stamens 5, the
laments green, 3-4 mm, narrowly triangular-attened,
the anthers orangish yellow, 1.5-1.7 x ca. 1 mm,
triangular, basally sagittate, apically obtuse, the pollen
orangish yellow, apparently mixed with oil. Ovary ca.
4 mm (including the ca. 1 mm style), bottle-shaped,
3-locular, the stigma indistinctly lobed. Fruits greenish
pink, ca. 8 mm (including the ca. 2 mm stylar beak),
globose-ovoid.

Ruyschia moralesii Hammel, sp. nova

Haec species nova Ruyschia andina cum oribus
quinquestaminatis et ovario triloculari valde similis,
sed imprimis differt laminis foliorum notabile
angustioribus, tenerioribus, glandulis hypophyllis
minus conspicuibus sed magis numerosis, area latiore
in utroque costae lato dispersis.
[This species is most similar to R. andina for its
owers with 5 stamens and a 3-locular ovary, but
differs from it principally by its distinctly narrower and
much less coriaceous leaves with less conspicuous but
more numerous hypophyllous glands spread in a wider
area on each side of the midrib.]
TYPE: Costa Rica; San Jos, Cantn de Tarraz, ca.
19 km de San Marcos, camino a Quepos por Londres,
17 febr 2006 () Hammel et al. 24095 (holotype: INB,
isotypes: CR, FR, MICH, MO, P, QCA). Figs. 2, 4B.
Shrubby epiphyte or woody liana. Leaves with a
denite petiole 0.5-1 cm, slightly attened above;
blade 7.5-10 3-3.5 cm, oblanceolate, drying dark tan
to nearly black, the base symmetrical, cuneate, the
apex acute to obtuse, mucronate, sometimes retuse;
secondary veins obscure on both surfaces, slightly more
evident below; hypophyllous glands inconspicuous,
16-40 (on each side of the midrib), poriform, more or
less scattered the length of the blade in a broad band
between the margin to ca. halfway to the midrib.
Inorescence an erect raceme, 6-21 cm (including
the peduncle) with 25-70 owers; pedicels 5-10 mm
patent at anthesis, lacking lenticels. Nectaries green,
attached 1.5-2.5 mm below the ower, ca. 6 mm,
spoon shaped with a narrow stalk (ca. 2 mm) and an
orbicular, abaxially concave lamina (ca. 4 mm) with
2 hypophyllous, poriform glands. Mature ower buds
5-6 mm, globose-ovoid, the 2 bracteoles sepaloid, ca.
1.5 x 2 mm, broadly semicircular, born just below
and appressed to the sepals; sepals equal, ca. 2.5 x
3 mm, broadly semicircular; corolla green, the petals

DISTRIBUTION. This species is known only from Costa

Rica, along the central pacic slope in coastal mountains
towards the northern end of the Talamanca cordillera
in rain forest between 1100-1800 m elevation.
ETYMOLOGY. This species is dedicated to friend and
colleague, Juan Francisco Morales, instrumental in the
discovery of this and hundreds of other new species
and new records for Costa Rica, especially from his
favorite haunts in the Acosta region, along the Pacic
slope, just south of San Jos.
DISCUSSION. Among Costa Rican species, R. moralesii
resembles the rare R. enervia Lundell for its ve
stamens, but it differs markedly from that species by
having more numerous hypophyllous glands (to only
ca. 7 in R. enervia), nectariferous bracts having a
stalk, rather than being sessile, and arising somewhat
distant from the owers rather than virtually adjacent
to them, smaller owers, and a 3- (rather than 2-)
locular ovary. Only one other species currently placed
in Ruyschia, the recently described, South American
R. andina de Roon (de Roon 2005) has a 3-locular
ovary. This new Costa Rican species is most probably
sister to R. andina, differing from it most obviously
by its distinctly narrower and much less coriaceous
leaves with less conspicuous but more numerous
hypophyllous glands spread in a band, rather than a
distinct line. It also appears to have the nectaries placed
more distant from the owers, and somewhat smaller
inorescences. Otherwise, the distinctly petiolate
leaves, the shape and position (somewhat distant from
the ower) of the nectariferous glands, ower size and
number of stamens are the same as or similar to R.
andina. Ruyschia andina is known from 1800-2300 m
elevation in rain and cloud forest on the eastern slopes
of the Andes of Ecuador, Per and Bolivia.
The similarity of Ruyschia moralesii and R. andina
to another Marcgraviaceae endemic to Costa Rica,
Schwartzia jimenezii (Standl.) Bedell, needs further
investigation. The current placement of this species

76

LANKESTERIANA

in Schwartzia apparently balances precariously on the

position of its nectaries, at or below the middle of the
pedicel rather than near the base of the ower, and on
its somewhat elongate pedicels. It too has ve stamens
and a 3-locular ovary, a combination that is unusual, if
not unique (to these three species) in the family. It also
has similarly shaped, distinctly petiolate leaves with
numerous hypophyllous glands and stalked, rather than
sessile nectariferous bracts. In all respects Ruyschia
moralesii must certainly be more closely related to
Schwartzia jimenezii than to any Costa Rican species
now placed in Ruyschia.
Over the years, Schwartzia jimenezii, originally
described as a Ruyschia, has been shuttled, somewhat
uncomfortably, to other genera (also Norantea),
its unusual combination of characters having been
contentious, and pointing toward afnities with
species of Ruyschia (see Bedell 1985: 168). In a
recent molecular phylogenetic work (Ward & Price
2002) that sampled each of the currently recognized
genera of Marcgraviaceae, the only robust nding
within subfamily Noranteoideae was a sister group
relationship between Ruyschia phylladenia Sandwith
and Schwartzia costaricensis (Gilg) Bedell, even
though Ruyschia has commonly been considered
closely related to Soroubea (see de Roon 2005).
Neither the currently accepted subdivision of
Noranteoideae into six genera nor the older concept
with only three, was supported by the molecular study
of Ward & Price (2002). The placement of species
in this group has often been problematic, numerous
of them having been transferred among two or more
of the currently accepted genera. As pointed out by
de Roon (2005), recent studies of Marcgraviaceae at
the generic level (Ward & Price 2002, Van Evelghem
upublished thesis) have sampled very few species,
and decisions concerning taxon realignments should
await more indepth studies. With these considerations
in mind, and given that Ruyschia is the oldest generic
name in subfamily Noranteoideae, one can speculate
that further molecular and morphological phylogenetic
work in the family is likely to expand the current
concept of Ruyschia.
ADDITIONAL SPECIMENS. COSTA RICA. Puntarenas:
Cantn de Parrita, Cabeceras del Ro Negro, 11001300 m, 27 ene 2002 (bud), J.F. Morales 8295 (CR,
INB, MO); Fila Chonta, 1100-1400 m, 27 dic 1997
(bud), J.F. Morales & R.J. Abarca 6297 (INB, MO).
San Jos: Cantn de Dota, Fila Mona, 1800-1850 m,
25 febr 2001 (bud), J.F. Morales 7575 (CR, INB, MO,
NYBG, U); cantn de Tarraz, ca. 17 km suroeste de

Vol. 6, N 2

San Marcos de Tarraz, por el camino a Cerro Cura,

1500 m, 20 dic 2005 (bud, fr), Hammel et al. 23979
(CR, INB); (bud), Hammel et al. 23981 (CR, INB,
F, MO); 12 ene 2006 (bud), Hammel et al. 24030
(INB, MO, MICH, USJ); ca. 19 km de San Marcos
de Tarraz, camino a Quepos, por Londres, 1700 m,
13 ene 2006 (bud), Hammel et al. 24035 (CR, INB,
MO, F); del puente de San Marcos 18 km camino hacia
Quepos, propiedad de la familia Garro, 1370 m, 13 ene
2006 (bud), Hammel et al. 24045 (CR, INB, MO, P).
Schwartzia tarrazuensis Hammel, sp. nova
Haec species nova Schwartzia costaricensi similis,
sed nectariis galeiformibus (ad vicem cochleariformibus) glandulisque hypophyllis paucioribus differt;
etiam Schwartzia brenesii ob formam nectariorum
similis, sed nectario pedicelli basi proximo (ad vicem pedicelli medio) nato, pedicellis longioribus, 68(-10) (ad vicem 4) glandulis hypophyllis in utroque
costae lato distincta; a duabus praecedentibus speciebus polline diluto-luteo vel sordido-albo (ad vicem
purpureo vel magenteo) dignocenda.
[This species is similar to Schwartzia costaricensis,
but differs by its helmet-shaped (versus spoon-shaped)
nectaries and by fewer hypophyllous glands; it is also
similar to S. brenesii for the shape of its nectaries,
but differs by the nectary born near the base of the
pedicel (versus the middle), by its longer pedicels,
and by 6-8(-10) (versus 4) hypophyllous glands on
each side of the midrib; the new species differs from
both these species by its pollen pale yellow or sordid
white (versus purple or magenta)]
TYPE: Costa Rica; San Jos, cantn de Tarraz, ca.
13 km suroeste de San Marcos camino a Quepos, ca.
0.8 km sur del cruce con el camino a Fila San Isidro,
sobre camino a Cerro Toro, 1300 m, 14 jul 2005 (,
fr), Hammel et al. 23715 (holotype: INB, isotypes:
COL, CR, F, FR, MICH, MO, P). Figs. 3, 4C.
Shrubby epiphyte or woody liana; stems terete,
drying reddish brown. Leaves with the petiolar
region 0.1-1 cm, distinctly channeled above, thinlymargined and merging with the abruptly expanded
base of the blade; blade 10-14 4-5.5 cm, elliptic
to oblong or oblanceolate, drying reddish tan to dark
reddish or olive brown, the base symmetrical,
abruptly acute to rounded, the apex acute to rounded,
mucronate; secondary veins indistinct; hypophyllous
glands 6-8(-10) (on each side of the midrib), circular

LANKESTERIANA 6(2):

HAMMEL New species of Marcgraviaceae from Costa Rica

to narrowly elliptic, poriform, in a distinct diagonal

row on the lamina from the base of the midrib toward
the margin at ca. 3/4 the length. Inorescence an erect
raceme, the rachis (including peduncle) 5-9 cm with
6-12 owers; pedicels 3-6.5 cm, ascending at mostly
< 45, lacking lenticels. Nectaries green, attached 310 mm from the base of the pedicel, 12-20 x 5-12
mm (at the rim), elongate helmet-shaped, anged and
sometimes with a spur on the rim. Flower buds green,
erect, (9)13-16 mm, ovoid, the 2 bracteoles sepaloid,
3.5-6 x 5-7 mm, semicircular, directly subtending
and appressed to the sepals; sepals equal, 5.5-8 x
7-10 mm, broadly semicircular; petals 5, adaxially
greenish red, strongly reexed at anthesis 13-20 x 810 mm, narrowly ovate with an obtuse apex, connate
in the basal 1/3. Stamens 12-14, the laments bright
red, 9-10 mm at anthesis, attened, linear-ligulate,
adnate to the corolla tube, the anthers pale yellow, 910 mm, narrowly triangular, basally sagittate, apically
obtuse, introrsly dehiscent, the pollen pale yellow to
sordid white, apparently mixed with oil. Ovary 8-10
mm (including the 2-3 mm style), obturbinate, 4- or
(rarely and only apically?) 5-loculate, the stigma 4- or
(rarely) 5-lobed. Fruits green to reddish green, 18-25
mm, globose-apiculate.
DISTRIBUTION. Schwartzia tarrazuensis is known only
from rain forest at 900-1400 m elevation, on the Pacic
slope of Costa Ricas Cordillera de Talamanca.
ETYMOLOGY. This species is named for the cantn
Tarraz, locality of the type and most other
collections.
DISCUSSION. Vegetatively, this species is very similar
to Schwartzia costaricensis, with which it is locally
sympatric, but differs by its helmet-, rather than
spoon-shaped nectaries. It usually also has fewer
hypophyllous glands, 6-8 being most common in the
new species while S. costaricensis usually has more
than 10 and as many as 20, on each half of the blade.
In the shape of its nectaries the new species resembles
S. brenesii (Standl.) Bedell, but that species has
shorter pedicels (3-4 cm), and the nectary is born near
the middle, rather than near the base of each pedicel.
Schwartzia brenesii also has only four hypophyllous
glands per lamina side. The pollen of both S.
costaricensis and S. brenesii is purple or magenta
while that of the new species is pale yellow to sordid
white. The South American S. weddelliana (Baill.)
Bedell, thought to be closely related to S. brenesii
(Bedell 1985), has yellow pollen and helmet-shaped

77

nectaries, but much longer (2.5-4 cm) nectaries, and

more stamens (22-25) than the new species.
ADDITIONAL SPECIMENS. COSTA RICA. Puntarenas:
Cantn de Buenos Aires, Altamira de Biolley, 11501350 m, 20 mar 1997 (), J. Quesada 1900 (CR, INB,
MO); cantn de Parrita, ca. 22 km suroeste de San
Marcos de Tarraz, rumbo a Fila Chonta, cabeceras
del Ro Negro y Ro Palo Seco, 1200 m, 20 dic 2005
(.), Hammel et al. 23996 (INB, USJ). San Jos:
Cantn de Tarraz, camino a Esquipulas, 1000-1100
m, 5 sept 1996 (, fr), Hammel et al. 20413 (CR, INB,
MO); camino a Quepos, 900 m, 24 febr 1996 (bud),
J.F. Morales & V. Urea 5270 (CR, INB, MO).
ACKNOWLEDGMENTS. I thank Silvia Troyo for her, as
ever, suitable-for-framing and precise illustrations,
and Francisco Morales for leading me, once again,
to his favorite haunts. This manuscript was nished
during a 6 week tenureship as a visiting research
scientist at the Musum National dHistoire Naturelle,
Departement Systmatique et Evolution, Herbier
National (P). I am very grateful to that institution for
providing nancial assistantship, as well as to all the
curators, librarians and technicians who so graciously
facilitated our visit.
LITERATURE CITED
Bedell, H.G. 1985. A generic revision of Marcgraviaceae, I. The Norantea complex. Unpublished
Ph.D. thesis, Univ. Maryland, College Park.
de Roon, A.C. 2005. A new species of Ruyschia
(Marcgraviaceae) from the South American Andes.
Novon 15: 414-415.
--------- & S. Dressler. 1997. New taxa of Norantea
Aubl. s.l. (Marcgraviaceae) from Central America
and adjacent South America. Bot. Jahrb. Syst. 119:
327-335.
Dressler, S. & M. Tschapka. 2002. Bird versus
bat pollination in the genus Marcgravia and the
description of a new species. Bot. Mag. 19: 104114.
Gentry, A.H. & C.H. Dodson. 1987. Diversity and
biogeography of vascular epiphytes. Ann. Missouri
Bot. Gard. 74: 205-233.
Van Evelghem, L. 2003. Houtanatomie en Pollenmorfologie van de Marcgraviaceae. Unpublished
undergraduate thesis, Institute of Botany and Microbiology, K.U. Leuven.
Ward, N.M. & R.A. Price. 2002. Phylogenetic
relationships of Marcgraviaceae: insights from
three chloroplast genes. Syst. Bot. 27: 149-160.

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Zamora, N., B.E. Hammel & M.H. Grayum. 2004.

Novelties. In: Hammel, B.E., M.H. Grayum, C.
Herrera & N. Zamora (eds.). Manual de Plantas de

Vol. 6, N 2

C o s t a R i ca. Vo l . I . I n t r o d u cci n . M o n o g r.
S y s t . B o t . M i s s o u r i B o t . G ar d . 9 7 : 2 1 7 240.

Fig. 1. Marcgravia glandulosomarginata Hammel -A. Branch with mature inorescence just prior to anthesis.
-B. Close-up of calyx showing submarginal rows of glands. -C. Cross section of ovary. -D. Nectary. -E. Fruiting
pedicel with detail of lenticels. -F. Lower surface of leaf with detail showing submarginal glands. E from Hammel
et al. 20205 (INB), all others from Hammel et al. 23714 (INB).

LANKESTERIANA 6(2):

HAMMEL New species of Marcgraviaceae from Costa Rica

79

Fig. 2. Ruyschia moralesii Hammel -A. Branch with inorescence prior to anthesis with detail of leaf lower
surface showing hypophyllous glands. -B. Open ower with pedicel and nectary. -C. Ovary with calyx (corolla
removed) and partial corolla showing adnate stamens. -D. Mature ower bud. -E. Cross section of ovary. -F. Open
owers at rachis. -G. Fruit. A from Morales 6297 (INB), B-F from Hammel et al. 24095 (INB), G from Hammel
et al. 23979 (INB).

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Fig. 3. Schwartzia tarrazuensis Hammel -A. Branch with mature inorescence just prior to anthesis. -B. Lateral,
front and longitudinal section views of nectary. -C. Fruit at top of pedicel. -D. Ovary and top view of stigma.
-E. Stamens. -F. Corolla lobe, showing adnate stamens. -G. Mature bud with calyx and corolla lobes removed.
B from Quesada 1900 (INB), all others from Hammel et al. 20413 (INB).

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Fig. 4. Three new species of Marcgraviaceae. -A. Marcgravia glandulosomarginata (Hammel et al. 23714).
-B. Ruyschia moralesii (Hammel et al. 24095). -C. Schwartzia tarrazuensis (Hammel et al. 23996).