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Title: Lankesteriana
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Permanent Link: http://ufdc.ufl.edu/UF00098723/00021
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Title: Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Physical Description: v. : ill. (some col.) ; 25 cm.
Language: English
Creator: Jardi´n Bota´nico Lankester
Jardín Botánico Lankester
Publisher: Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Place of Publication: Cartago Costa Rica
Cartago Costa Rica
Publication Date: August 2008
Frequency: three times a year[2002-]
irregular[ former 2001]
three times a year
Subject: Botany -- Periodicals -- Costa Rica   ( lcsh )
Epiphytes -- Periodicals -- Costa Rica   ( lcsh )
Orchids -- Periodicals -- Costa Rica   ( lcsh )
Plantkunde   ( gtt )
Botanische tuinen   ( gtt )
Genre: periodical   ( marcgt )
Spatial Coverage: Costa Rica
Language: In English and Spanish.
Dates or Sequential Designation: No. 1 (mayo 2001)-
Numbering Peculiarities: Issues for May 2001-Oct. 2003 designated no.1-8; issues for Apr. 2004- designated vol. 4, no. 1-
General Note: Latest issue consulted: Vol. 4, no. 1 (abr. 2004).
General Note: International journal on orchidology.
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Bibliographic ID: UF00098723
Volume ID: VID00021
Source Institution: University of Florida
Holding Location: University of Florida
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Resource Identifier: oclc - 48491453
lccn - 2001240973
issn - 1409-3871


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Table of Contents
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Full Text

I W I.l 1( ', H A1 I N A 1,

ISSN 1409-3871

VOL. 8, No. 2 AUGUST 2008

Capsule development, in vitro germination
and plantlet acclimatization in Phragmipedium humboldtii,
P longifolium and P pearcei

Stanhopeinae Mesoamericanae IV:
las Coryanthes de Charles W. Powell

The Botanical Cabinet

New species and records of Orchidaceae from Costa Rica

Book reviews 75



Copyright C 2008 Lankester Botanical Garden, University of Costa Rica
Effective publication date: August 29, 2008

Layout: Jardin Botanico Lankester.
Cover: Plant of Epidendrum zunigae Hagsater, Karremans & Bogarin. Drawing by D. Bogarin.
Printer: Litografia Ediciones Sanabria S.A.
Printed copies: 500

Printed in Costa Rica / Impreso en Costa Rica

R Lankesteriana/ The International Journal on Orchidology
No. 1 (2001)-- -- San Jose, Costa Rica: Editorial
Universidad de Costa Rica, 2001--

1. Botanica Publicaciones peri6dicas, 2. Publicaciones
peri6dicas costarricenses


LANKESTERIANA8(2) 23-31 2008





CIGRAS, Universidad de Costa Rica, 2060 San Pedro, Costa Rica
Jardin Botanico Lankester, Universidad de Costa Rica, P.O. Box 1031, 7050 Cartago, Costa Rica
imelaniamunozg @yahoo.com; 2victor.jimenez @ucr.ac.cr

ABSTRACT. Capsule development from pollination to full ripeness was evaluated inPhragmipedium longifolium,
P. pearcei and P. humboldtii. Besides, seed viability, analyzed in each capsule by means of the tetrazolium
chloride staining, was determined. Considering seed viability, germination rate was corrected and expressed as
the rate of viable seeds that germinated in the presence and absence of light, on Knudson C and on half-strength
Murashige and Skoog culture media. Capsule length remained constant during the evaluation period, while the
diameter increased during the first 6-8 weeks and then stagnated. Capsule opening occurred 16 weeks after
pollination in P. longifolium, after 9.8 weeks in P. pearcei and after 32 weeks in P. humboldtii. Seed viability
averaged 44.7% in P. longifolium, 82.3% in P. pearcei and 34.3% in P. humboldtii. No significant effect of
light conditions was evident in any of the species. However, a higher proportion of seeds of P. longifolium and
P. pearcei germinated earlier on half-strength Murashige and Skoog medium than on Knudson C. Only 2.9%
of the viable seeds of P. humboldtii germinated, while approximately 40% germination occurred in the other
two species. Initial growth of the embryos was better in the dark on Knudson C medium, compared to the
other treatments studied. Further growth of the seedlings took place under light conditions. Developed plants
formed roots and were successfully acclimatized in the greenhouse.

KEY WORDS: capsule development, in vitro seed germination, pollination, Phragmipedium, terrestrial orchids,
tetrazolium chloride, tropical orchids

Slipper orchids belonging to the genus
Phragmipedium (Subfamily Cypripedioideae Lindl.)
are distributed in Meso and South America (Cox et al.
1998, Dressler 2003). They are seriously threatened
because of alteration and destruction of their habitat
and over collection from their natural environment
(Arditti 1992, Salazar 1996).
Use of in vitro protocols has been foreseen as a
successful approach for ex-situ conservation and
reintroduction of endangered orchids (Stenberg
and Kane 1998, Decruse et al. 2003, Sarasan et al.
2006). Plants regenerated from seeds have a broader
genetic background than those developed by clonal
propagation methods. Therefore, the former meet
the goals of a reintroduction program better, in the
sense of warranting sufficient genetic resources in

the reintroduced population to undergo adaptive
evolutionary change (Guerrant and Kaye 2007).
This strategy has been successfully employed for
the reintroduction of the orchid species Bletia urbana
(Rubluo et al. 1989), Ipsea malabarica (Gangaprasad
et al. 1999) and Spiranthes brevilabris (Steward et al.
2003). An additional advantage of mass-propagating
orchids for conservation purposes is that increasing
availability of plants from preferred species with
adequate phytosanitary standards and at affordable
prices would reduce illegal collection from the wild
populations (Ramsay and Dixon 2003, Salazar and
Mata 2003).
Propagation of Phragmipedium through seeds
does not seem to be extremely difficult, because
commercial formulations for asymbiotic germination
in this genus are available. However, the composition


of the culture media has remained elusive to the
scientific community for commercial reasons. Most
research on this and related genera is limited to studies
on systematics and evolution (e.g., Cox et al. 1997,
1998). Investigations on phenology in this genus,
describing characteristics of seed capsules and their
development, are scarce (Arditti and Ghani 2000).
When proper conditions for germination are to
be assessed, it is important to know the viability of
the seeds under study. This allows distinguishing
the proportion of seeds that do not germinate as a
consequence of unfavorable germination conditions
from those seeds that are not capable of germinating at
all due to lack in viability. The most common method
used to evaluate seed viability is the tetrazolium
(C19H,1C1N4) stain. Dehydrogenases, which are active
in living tissues, reduce the colorless tetrazolium
chloride to a red compound, coloring viable seeds. The
intensity of the tint could vary from pink to dark red
(Singh 1981). This technique has been successfully
used to test viability of orchid seeds (Lauzer et al.
1994, Vujanovic et al. 2000).

The aims of this study were to describe the
development of P. humboldtii, P. longifolium, and P.
pearcei capsules, to evaluate the seed viability and to
establish a method for in vitro germination of mature
seeds and for growing and acclimatizing plantlets
of these three Phragmipedium species aiming at

Materials and methods

Pollination of flowers and capsule development.
Manual pollination of flowers of Phragmipedium
humboldtii, P. longifolium, and P. pearcei, growing at
Lankester Botanical Garden, Universidad de Costa
Rica, was conducted using pollen from a different
plant. Capsule length, diameter and color were
evaluated weekly, until full maturity. The time taken
for each capsule to open was also recorded.

Seed viability. The percentage of viable seeds was
determined in each ripe capsule using the method of
tetrazolium stain (Singh 1981). For that purpose, four
subsamples of each of the two P. humboldtii, one P.
longifolium, and four P. pearcei capsules were placed
in tetrazolium chloride (1%, pH 6-7), in a water bath

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.

(300C) and in dark conditions for 24 h. Subsequently,
they were transferred to Petri dishes and the percentage
of viable (stained) seeds was determined with aid of
a stereomicroscope (model 222279, Nikon). Seeds
from the corresponding capsules were used in the
germination studies.

In vitro germination. Seeds from open capsules
were sterilized in sodium hypochlorite (NaOC1,
0.6% w/v) and Tween 20 (1 drop/100 ml) for 10
min. They were subsequently screened through
sterile Albet filter paper (quantitative quality), in
the laminar hood. They were then rinsed three times
with sterile distilled water, while adhered to the filter
paper and, after decanting the water of the last rinse,
distributed evenly, with a scalpel (blade No. 22) on
20 ml of semisolid culture medium contained in 90
mm-Petri dishes.
Knudson C (Knudson 1946) and half-strength
Murashige and Skoog (1962) mineral salts, both
supplemented with 1 mg 1-1 thiamine, nicotinic acid
and pyridoxine, together with 20 g 1-1 sucrose, were
compared for seed germination. pH was adjusted to
5.7 and media gelled with agar (0.8%). Media were
autoclaved at 1.05 kg cm2 for 25 min.
Additionally, two light regimes were evaluated
during germination: dark conditions and photoperiod
of 12 hours (10.9 imol m2 s-1, Sylvania Supersaver
Cool White, 32 W, F48%12/CW/SS). Cultures were
grown at 25+10C.
To determine the percentage of germination, three
squares, of 1 cm2 each, were drawn in each Petri dish.
Areas of the Petri dishes in which individual seeds
could be clearly observed using a stereomicroscope
were selected to draw the squares. Total number of
seeds in each section was counted at the first day
of culture. Afterwards, the number of germinating
seeds (those in which rupture of the testa occurred
by the enlarging embryo) was assessed weekly. Three
Petri dishes were evaluated for every combination
of culture medium, light condition and species.
Considering seed viability, germination rate was
corrected and expressed as the rate of viable seeds
that germinated in each treatment. For example, if
50% of the seeds of a capsule with a 50% germination
rate, according to the tetrazolium test, germinated on
culture medium, it was considered that 100% of the

MUNoz & JIMENEZ Capsule development and germination in Phragmipedium

viable seeds germinated. Besides, the percentage
of dead protocorms (those that turned brown after
germination) was calculated.
Protocorms growing in the dark were transferred
to the same light conditions of the other treatment
after six weeks of culture. Growing protocorms were
subcultured every six weeks on the corresponding
culture medium. Only actively growing and
green protocorms were selected and individually
subcultured. When the plantlets reached 1 cm in
height, they were transferred into glass flasks (150
ml), eight plants per flask. Later on, when they were
2-3 cm tall, only five plants were cultured per flask.
Plants were acclimatized approximately 15 months
after culture beginning, when they were 4-5 cm tall
and had 2-3 roots around 1 cm in length. For that
purpose, all remnants of the gelled culture medium
were carefully removed from the roots with running
water. One plant was potted per plastic container (3
x 3 x 4 cm), carefully covering the roots completely
with the potting mix (peat moss). Plants were placed
under mist irrigation (4 s watering every 15 min
during the daylight hours). After six months in the
greenhouse, the average number of roots and the size
of the longest root were annotated.

Statistical analysis. Capsule length and diameter,
as well as the effect of culture medium and light
conditions on germination for each species were
analyzed with a multivariate test of significance (for
repeated measures). Days needed for full maturity
were compared between genotypes with an analysis of
variance. Subsequently, the Post hoc Tukey's Honest-
Significant-Difference-Test (HSD) for unequal N
(Spjotvoll/Stoline) was used to determine significant
differences in the parameters tested (p<0.05). All
analyses were conducted using Statistica 6.1 (StatSoft
Inc., Tulsa, Oklahoma, U.S.A.).


Pollination of flowers and capsule development.
Blooming was observed in plants ofP. longifolium and
P. pearcei during the entire year, while P. humboldtii
plants bloomed only in May and June. All manual-
pollinated flowers (three of P. humboldtii, nine of P.
longifolium and 16 ofP. pearcei) developed capsules.


4 -
--P. humbolidtii
2 A P. longifolium
P pearcei

0 5 10 15 20 25 30 35
Time (weeks)
FIGURE 1. Diameter measured in capsules ofP. humboldtii,
P. longifolium and P. pearcei during development.

Capsule development, from pollination to opening,
took longer in P humboldtii (320.6 weeks), than in
P. longifolium and P pearcei (162.05 and 9.80.6
weeks, respectively).
Significant differences (p<0.001) were measured in
the capsule size of the three species (Fig 1). Largest
capsules were those of P humboldtii, with an average
length of 182.73.2 mm and a diameter of 7.00.2 mm
one week before opening. Capsules of P longifolium
were 60.022.0 mm in length and 5.60.1 mm in
diameter, while those of P pearcei were 42.51.4
mm in length and 4.20.1 mm in diameter at the same
moment (Fig. 1). During development, length of the
capsules did not vary in any of the species studied
(p>0.05), while diameter increased during the first 6-8
weeks after pollination and then remained constant
(Fig 1).
Capsules of P humboldtii were light green and
pubescent (Fig. 2A), while those of P longifolium
were dark purple without pubescence (Fig. 2B);
capsules of P pearcei were green-purple or totally
purple, with short hairs (Fig. 2C). The capsules did not
present notable changes in color or pubescence degree
during maturation, except for those of P humboldtii,
which turned yellow 4-6 weeks before opening.

Seed viability. A great difference in seed viability
was observed between P. pearcei and the other species
(Fig. 3). Average seed viability in the capsules of P.
humboldtii was 34.3%, in P. longifolium 44.7% and
in P. pearcei 82.3%. When the seed viability among
different capsules of a single species was compared,
values ranging from 20 to 80% were observed in P.
pearcei and from 18 to 51% in P. humboldtii.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rca, 2008.


FIGURE 2. Development of capsules of P. humboldtii (A, A'), P. longifolium (B B1) and P. pearcei (C C1) at pollination
day (left) and one week before capsule opening (right).

In vitro germination. In order to consider only the viable
seeds when calculating germination rate, percentage of
seeds that germinated in the culture medium was corrected
with the rate of viable seed measured for each capsule,
as described in Materials and Methods. The corrected

LANKESTERIANA8(2), August 2008. 0 Umversidad de Costa Rica, 2008.

percentage of germination was lower in P humboldtii
(2.9%) than in P pearcei (38.7%) and P longifolium
(41.3%), without significant differences between the last
two (Fig. 3). Developing protocorms that subsequently
died were observed only in P pearcei (Fig. 3).

MUNoz & JIMENEZ Capsule development and germination in Phragmipedium

OP. humboldtii
80 O- P. longifolium
SP. pearcei
' 60 *

4e 40


Seed viability Germination Dead protocorms

FIGURE 3. Seed viability, germination rate corrected with the
percentage of seed viability of each capsule and death
rate of germinated seeds in P humboldtii, P. longifolium
and P pearcei. Data represent the mean of at least three
replicates SD. *Four weeks after sowing. **Eight
weeks after sowing.

Viable seeds of P. longifolium and P pearcei started
geminating during the first two weeks of culture.
Afterwards, the germination rate decreased until, atweek
four, no additional germinating seeds were observed. On
the other hand, germination in P humboldtii occurred in
a very low rate, with only a slight increase during the
evaluated period of eight weeks (Fig. 4).
Culture medium had a significant effect on seed
germination (Table 1), evident in P. longifolium and
P. pearcei only during the first week by a higher
germination rate on half-strength Murashige and
Skoog medium. Thereafter, seed germination in these
two species behaved very similarly (Fig. 4), without
significant differences according to the Post-Hoc

Knudson C _
40 --
P- P. humboldtii
S20 P. longifolium
S-*- P. pearcei
50 i^ -- -- ...... .... .* ...* *.... ...
MS 50%



0 I----* -- 4 ...... .....--t" "T-. .
0 1 2 3 4 5 6 7 8 9
FIGURE 4. Germination rate in seeds of P. humboldtii, P.
longifolium and P. pearcei along time on Knudson C and
half-strength Murashige and Skoog media (MS 50%).

Tukey's Honest-Significant-Difference-Test (HSD)
for unequal N (Spjotvoll/Stoline).
No effect of the light conditions tested could be
observed on seed germination in any of the evaluated
species (p>0.05) as presented in Table 1. However,
among the protocorms cultured on the Knudson
C medium, a higher growth rate was observed
afterwards in those germinated under dark conditions.
The protocorms germinated in the dark were placed
under a photoperiod of 12 hours of light after six
weeks of culture, and one week later they began to
change their color from white to green. By that time,

Table 1. Multivariate test of significance of seed germination corrected with the viability rate for each capsule of the three
Phragmipedium species studied in response to culture medium and light conditions.

Source of variation Test Value F Effect df Error df p

Intercept Wilks 0.031021 226.4653 4 29 0.000000
Species (A) Wilks 0.038002 29.9406 8 58 0.000000
Culture medium (B) Wilks 0.451880 8.7941 4 29 0.000089
Light conditions (C) Wilks 0.757612 2.3195 4 29 0.080649
AxB Wilks 0.469325 3.3328 8 58 0.003338
AxC Wilks 0.421866 3.9122 8 58 0.000948
BxC Wilks 0.691800 3.2299 4 29 0.026159
AxBxC Wilks 0.454910 3.4992 8 58 0.002320

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


FIGURE 5. Stages during in vitro development of P pearcei, P. longifolium and P humboldtii cultured on MS 50%.

the protocorms of P. pearcei and P. longifolium began
to develop the first leaf. On the other hand, the first
leaf of P. humboldtii appeared in the eighth week of
culture. The seedlings of P. pearcei and P. longifolium
developed two or more leaves within the first three
months, and began to develop roots in 3.5 months
(Fig. 5). The plantlets of P. humboldtii developed
their leaves in three months too, but began to develop
roots only after five months in culture (Fig 5). Higher
growth rate and better leaf development was observed
with Knudson C (data not shown).
All plants were successfully acclimatized after
ca. 15 months of culture in vitro (Fig. 5). After six
months in the greenhouse, each plant of P. pearcei
had 7.32.5 roots, with the longest one measuring
10.41.7 cm in length. At the same moment, the
plants ofP. longifolium had 3.60.9 roots, the longest
one with 11.73.5 cm.


The time it takes an orchid capsule to reach full-
maturity varies according to genus and species. Times
registered in this work for P. pearcei, P. longifolium
and P. humboldtii (10, 16 and 32 weeks, respectively)

LANKESTERIANA8(2), August 2008. O Unversidad de Costa Rica, 2008.

were considerably shorter than those measured in
other species belonging to the Cypripedioideae
subfamily. For example, Paphiopedilum callosum
capsules required between 47 and 48 weeks to open
(Arditti 1992). Moreover, it is not unusual to observe
differences in this respect between species belonging
to the same genus. For example, while capsules of
Laelia gouldiana needed 17 weeks to open, those
of L. purpurata required 31 weeks. Similarly, in
Dendrobium kingianum the time until full ripeness
was 20 weeks, while in D. nobile it was 41 weeks
(Arditti 1992).
Capsule development in the three Phragmipedium
species studied characterized by an initial growing
period of around eight weeks, followed by a
maturation period in which the fruit did not increase
in size until opening (Fig. 1). This growing pattern is
commonly observed in many fruits. It is characterized
by a sigmoid arrangement, in which cell division and
elongation initially occur, followed by growth halt
and finally the fruit reaches maturity (Agusti 2000).
Those phases could vary in duration, depending on
the species or genotype, as it was observed in the
three Phragmipedium species studied here.

MuNOZ & JIMENEZ Capsule development and germination in Phragmipedium

Knowing the time the capsule of a particular
orchid species requires until full maturity is very
useful to determine proper harvesting time for in
vitro germination. Ripe capsules usually have larger
amounts of viable seeds than unripe ones. Moreover,
collecting seeds after capsule opening usually
reduces success during in vitro establishment due to
contamination problems and damage caused by the
sterilization process, and should be avoided when
Seed viability measured in Phragmipedium (Fig. 3)
is similar to that observed in other terrestrial orchids.
For example, the seed viability of Cypripedium acaule
varied from 20% to 40% in the capsules studied by
Lauzer et al. (1994), and the viability of Calopogon
tuberosus was 35% in the study conducted by Kauth
et al. i2 *.. ,. High values, as measured inP. pearcei in
this study, have been observed in mature capsules of
Ophrys (Kitsaki et al. 2004). Given that seed viability
can vary significantly among capsules within the
same species (up to 60%, as observed in P. pearcei),
it is important to evaluate viability in each capsule.
This is necessary to assess the real effect of particular
conditions on seed germination based on the actual
amount of viable seeds per capsule.
The seeds of P. pearcei and P. longifolium
germinated faster (2-3 weeks) (Fig. 4) than those
of several terrestrial orchids. Henrich et al. (1981)
showed that 7-12 weeks were necessary for the
beginning of germination in 17 terrestrial orchid
species. In the study of Shiau et al. i2 ', the seeds
of Anoectochilusformosanus germinated eight weeks
after sowing. On the other hand, there are other
terrestrial orchids that germinate faster, as in the case
of C. tuberosus, whose seeds germinated one week
after sowing and reached the maximum germination
in 4-6 weeks (Kauth et al. 2006).
The percentage of germination in terrestrial
orchids varies among species. Henrich et al. (1981)
found 100% germination in Orchis fucssi and
Epipactis gigantea, 75% in Goodyera oblongifolia
and Spiranthes nrolton/offinima. 50% in Platanthera
stricta and Orchis macula, and 25% in other seven
species, Platanthera dilatata, Liparis laeselii and
Cypripedium reginae among them. These authors
also measured very low germination rates (1%) in

Cypripedium calceolus, C. candidum, Plantanthera
hiperboria and P. flava. However, they did not
evaluate the viability of the seeds used; therefore the
real percentage of germination of viable seeds could
not be determined.
The seeds of P. humboldtii had very low
germination rate (2.9%) in spite of their relatively
high viability (34.3%). However, the large quantity of
seeds in each capsule (data not shown) and the low rate
of dead protocorms (Fig. 3) allowed the regeneration
of enough plants using the protocol developed in
this work for conservation and reintroduction of this
It has been considered that both germination and
embryo staining (e.g., with tetrazolium chloride)
are comparable viability tests. However, although
the latter test has been successfully employed with
epiphytic tropical orchids and several European
and North American terrestrial orchids, there are
some reports on inconsistencies in several species,
probably attributed to variation in the permeability
of the seed coat (reviewed by Vujanovic et al.
2000). In one of these cases, Lauzer et al. (1994)
did not find any correlation between the percentage
of germination and the percentage of seeds stained
with tetrazolium chloride in Cypripedium acaule.
They attributed these differences to a prolonged
pretreatment with NaOC1 (40 min with 0.6% w/v
NaOC1, as compared to 10 min with the same
concentration in our study), a compound that can
promote dormancy release in terrestrial orchid
seeds (St-Arnaud et al. 1992).
Numerous orchid species grow better in the
dark, mainly during the first phases of development.
Activated carbon has also been used to darken the
culture medium and, in this way, improve germination
(Arditti and Ernst 1993). In some species, the role of
light during germination is not clear; for example,
some authors recommend germination of C.
tuberosus in the dark and others in light conditions
(reviewed by Kauth et al. 2006). The light or dark
treatments did not have any effect over germination
in the Phragmipedium species studied. However,
larger protocorms in the three species were obtained
in dark conditions. The protocorms that germinated
in the dark were white because they do not produce

LANKESTERIANA8(2), August 2008. 0 Umversidad de Costa Rica, 2008.


chlorophyll; when they were placed in the light, the
chlorophyll synthesis began, and their color changed
to green, as observed in other orchids (Arditti &
Ernst 1993).
A considerable number of culture media has been
successfully employed and even devised specifically
for orchid seed germination, some of them being only
slight modifications of others (Arditti 1967). Knudson
C (Knudson 1946) and Murashige and Skoog (1962)
mineral salts are probably the most commonly used
formulations for asymbiotic germination of orchid
seeds. Since the latter one contains high amounts of
nitrogen, especially in form of ammonium, dilutions
are frequently employed, such as in this investigation.
For many orchids, the effect of media on germination
and development appears to be genotype dependent.
When Knudson C and half-strength Murashige
and Skoog media were compared, some genotypes
developed better in the former and others in the latter
(Johnson and Kane 2007). There are also examples of
equal response inbothmedia (Roy andBanerjee 2002).
This behavior could be related to the differences in
the ammonioum:nitrate ratio of both media. Contrary
to the results obtained with Phragmipedium in this
work, half-strength Murashige and Skoog promoted
better development of the developing seedlings of
hybrid Vanda, as compared to Knudson C (Johnson
and Kane 2007).
Although, to the best of our knowledge,
acclimatization of in vitro-generated Phragmipedium
orchids has not been reported in the scientific literature,
this task should not be very difficult, since it seems to
be routinely conducted by commercial growers. Very
high survival rates during acclimatization, as reported
in the present work, indicate that plants generated in
vitro were in adequate conditions and could adapt
with few problems to the reduced relative humidity in
the greenhouse and become autotrophic. This seems
to be the case in the present work, since 100% success
was obtained in this step.
In this work an efficient protocol for seed
germination and plantlet growth of three
Phragmipedium species was described. The
combination of Knudson C medium and culture
in the dark during the first six weeks seems to be
appropriate to reach a good germination percentage

LANKESTERIANA8(2), August 2008. 0 Umversidad de Costa Rica, 2008.

and initial protocorm development. Following the
method described in this work it was possible to
grow plants of the three Phragmipedium species
studied and to achieve their acclimatization in the
greenhouse. A great quantity of plants could be
produced by in vitro germination, maintaining larger
genetic variability than with clonal propagation. This
is a useful tool for the conservation of these species.

ACKNOWLEDGMENTS. This investigation was partially
funded by the Universidad de Costa Rica (project number
VI-814-A6-107). Authors wish to thank J. Warner for
facilitating the plants used in this study and for additional
support. Helpful comments from anonymous reviewers are
gratefully acknowledged.

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LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.


LANKESTERIANA8(2) 33-42 2008




1 Botanischer Garten Miinchen-Nymphenburg, Menzinger Str. 65, D-80638 Mtinchen, Germany;
2 Orchid Herbarium of Oakes Ames, Harvard University Herbaria, 22 Divinity Avenue, Cambridge,
Massachusetts 02138-2020, U.S.A.; romero@oeb.harvard.edu

ABSTRACT. The descriptions of Coryanthes hunteriana and Coryanthes powellii described by Schlechter
are critically compared. We conclude that the same species was described twice. Because of the loss of the
holotypes, the specimens collected by Powell with the same collection number at AMES, K and MO are
discussed in detail.

RESUMEN: Se comparan criticamente las descripciones de Coryanthes hunteriana y Coryanthespowellii ambas
descritas por Schlechter. Concluimos que la misma especie se describi6 dos veces. Como los holotipos se
perdieron, se discuten en detalle las muestras existentes del collector Powell en los herbarios AMES, K y MO
con el mismo numero de colecci6n de los tipos.

ZUSAMMENFASSUNG: Die Beschreibungen von Coryanthes hunteriana und Coryanthes powellii, beide
beschrieben von Schlechter, werden kritisch verglichen. Wir kamen zu dem SchluB, daB ein und dieselbe Art
zweimal beschrieben wurde. Wegen dem Verlust der Holotypen werden die existierenden Herbarbelege des
Sammlers Powell in den Herbarien AMES, K und MO mit denselben Sammelnummern der Typen eingehend

KEY WORDS: Orchidaceae, Stanhopeinae, Coryanthes, Powell, Schlechter, Panama, Costa Rica

La morfologia de las flores del g6nero Coryanthes
Hook. es sumamente complicada, y casi impossible
de describir con palabras. Cada lector puede hacer
interpretaciones diferentes de las descripciones y por
consiguiente es de gran importancia acompafiarlas de
un dibujo y, de ser possible, de fotografias mostrando
las flores desde diferentes angulos.
De las dos species descritas por Rudolf Schlechter
de Panama (C. hunteriana y C. powellii) no existen
dibujos, pero si algunas fotografias las cuales, por
cierto, al parecer no estuvieron al alcance del autor (al
menos 1l no las mencion6 en sus descripciones). En
la 6poca de Charles W. Powell (1854 1927), quien
colect6 y cultiv6 orquideas principalmente de Panama
desde principios de siglo pasado hasta su muerte, el
process de tomar e imprimir fotografias de plants era
dificil y relativamente costoso, y por estas razones hay
pocas y su calidad no es buena.
Aqui debemos aclarar que el sistema de

numeraci6n de muestras empleado por Powell no
se ajusta al que esta en uso hoy en dia: sus numerous
agrupaban especimenes que de acuerdo a su criterio
representaban la misma especie, irrespectivamente
de done y cuando hubieran sido colectados y
preservados (mientras que ahora numeramos las
colecciones de herbario siguiendo el criterio de
McNeill et al., 2006: 8, pie de pagina 1, "... a single
gathering of a single species... made by a collector at
one time"). En este articulo precisamente tratamos
de elucidar las diferentes species de Coryanthes
que Powell distribuy6 bajo un mismo numero de
colecci6n. En algunos casos se puede descartar que
algunas de estas muestras puedan ser tipos, porque la
fecha que aparece en los r6tulos es diferente a la citada
en el prot6logo. El caso de las muestras sin fecha es
ambiguo, aunque en el pasado se han propuesto como
lectotipos (Christenson 1991). Aqui continuamos
asumiendo que los especimenes sin fecha son tipos,


0.:4 G himLe .oA an. 2 h \

00RT &a Cabrian 5&i

D oai ma Albt sa lDa l

FIGURA 1. Lectotipo de Coryanthes hunteriana Schltr., AMES (26894).
LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


GERLACH & ROMERO-GONZALEZ Las Coryanthes de Powell

Tabla 1. Comparaci6n de las descripciones originales en latin de Coryanthes hunteriana Schltr. y C. powellii Schltr.

Coryanthes hunteriana

Coryanthes powellii

"sepalo intermedio plus minusve crispato, ovato-
suborbiculari, apiculato, vix 3 cm long et lato,
lateralibus oblique dolabriformibus, a basi usque in
apicem 6,5 cm longis, supra basin margin anteriore

"petalis oblique lanceolato ligulatis, margin crispatis,
c. 3,5 cm longis;"

"labelli hipochilio ovatli-cucullato, apice subacuto,
dorso glabro, later utrinque striguloso-tomentello, 2,5
cm long, medio fere 1,8 cm alto,"

"mesochilio canaliculato, nudo, hypochilium, paululo
tantum superante,"

"epichilio generis cucullato, c. 4 cm long, lobis
lateralibus amplis semiovato-triangularis, obtusis,
antice truncatis, intermedio multo minore semiovalis,

aunque siempre podran ser cuestionados (K. Gandhi,
comunicaci6n personal, 2008).
De las Coryanthes de Powell existen cuatro
fotografias. Todas estas fotografias estan montadas en
muestras con su numero 19. De estas, una represent
C. misasii G. A. Romero & G. Gerlach, dos representan
C. panamensis G. Gerlach y la cuarta tal vez se
pueda asignar a una de las dos species descritas por
Schlechter (C. hunteriana, C. powellii).
Las muestras tipos de Schlechter, depositadas
en el Herbario de Berlin, fueron destruidas durante
la Segunda Guerra Mundial (Merrill, 1943; Ames,
1944). Aunque Schlechter elabor6 bosquejos de la
mayoria de sus species y envi6 copias a su amigo
y colega Oakes Ames (Universidad de Harvard,
E.E.U.U.; Ames, 1944), no se conocen dibujos de
las Coryanthes de Powell. De manera que solamente
nos pueden ayudar a elucidar sus dos species
panamefias de Coryanthes las muestras de Powell

"sepalo intermedio orbiculari, obtuso, c. 2,5 cm long
et lato, lateralibus dolabriformibus acuminatis, antice
basin versus sublobato-dilatatis, e basi usque apicem
c. 7 cm longis;"

"petalis oblique ligulatis, obtusiusculis, undulatis 3,5 cm

"labelli ungue 1, 7 cm long, hypochilio cucullato, obtuse
acuminato, dorso glabrato, lateribus linea strigillosa
ornato, 2,7 cm long, medio 1,8 cm alto,"

"mesochilio canaliculato, nudo, hypochilium, paulo sed
manifesto superante,"

"epichilio ovali-cucullato, c. 4 cm long, lobis
lateralibus amplis semiovatis, antice valde obtusis,
intermedio multo minore oblongo-triangulo trilobulato,"

hoy en dia presents en el Herbario de Orquideas
de los Jardines Reales de Kew (K), en el Herbario
de Orquideas de Oakes Ames en la Universidad de
Harvard (AMES) y en el Herbario de Jardin Botanico
de Missouri (MO).
En estos tres herbarios hemos localizado siete
muestras que llevan el numero 19 de Powell, las que,
de acuerdo al criterio de Powell y a la descripcion
de Schlechter, deben pertenecer todas a Coryanthes
hunteriana. Ademas, hay dos muestras de herbario con
el numero 156 de Powell que, siguiendo de nuevo a
Powell y a Schlechter, serian asignadas a C. powellii.
Christenson(1991) lectotipific6 lasmuestras destruidas
en Berlin y aclar6 que hubo un error en la cita del
numero del tipo de esta ultima especie (Schlechter, por
error propio o por un error tipografico, cit6 el numero
158 de Powell en el prot6logo aunque no hay duda que
debi6 haber citado el numero 156).
Enseguida examinamos una lista de las

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


No. 1C(. or/antii en
I;ear CiNty of 'Pnnrm nolinnll' oo a l.vel.
Sepala on backs ,)rpliaini hro n with trine
iowia center. Petali olI bIak unie vCe't -'
inner 1iide witu red p-iaplao nots..Hollm t
oruntia, buckut .ollnw, spotted on inner
sir;e *ith red brusrn paots. Colur n" le
Jrean. OwvIr tube paarilin : brown,



11 L NR
IItIlI1cn.1 iYI'iyrtini A^A

1all B..t u..tu M* C. O rnwm.i

Idl*d by Oak Amt


FIGURA 2. Lectotipo de Coryanthes powellii Schltr., K. Reproducido con gentil permiso del Board of Trustees of Royal
Botanic Gardens, Kew.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

GERLACH & ROMERO-GONZALEZ Las Coryanthes de Powell

Figura 3. Hija de Charles W. Powell (originalmente
montada en la muestra AMES 31853) con Coryanthes

muestras existentes de Coryanthes de Powell con
algunos comentarios.

Bajo Powell 19 todo este material se ha identificado
como Coryanthes hunteriana Schltr.

1) AMES 26894: Lectotypus (Fig. 1), no contradice
el prot6logo, sin fecha; labelo 5,9 cm alto;
2) AMES 25007: Isolectotypus, no contradice el
prot6logo, sin fecha; labelo 5,7 cm alto;
3) AMES 25008: la plant, por falta de flores, no es
determinable, la muestra incluye una foto de C.
misasii, tiene fecha del 5 de mayo de 1923;
4) K (sin nunero de herbario): Isolectotypus, cinco
flores de C. panamensis, dos fotografias (la del
habito no se puede determinar, la de la flor la
identificamos como C. panamensis), tiene fecha
del 15 de noviembre de 1918; labelo 6,3 cm
5) MO 955890: no contradice el prot6logo, sin
fecha; labelo 5,6 cm alto;

Figura 4. Coryanthes hunteriana Schltr., Powell 156
(conservado en liquid en AMES); la flecha muestra el
hipoquilo alargado y truncado, el character mas distintivo
de la especie.

6) AMES (sin numero de herbario) Powell 3546 y
Powell 19 (ambas anotaciones sobre r6tulo), con
fecha de abril de 1924;
7) AMES 31853: Powell 19 y Oakes Ames 3519;
una flor y una foto de la hija de Powell (Fig.
3) con una flor de C. panamensis, con fecha de
marzo de 1924.

Bajo Powell 156 todo este material se ha identificado
como Coryanthespowellii Schltr.

1) K (sin numero de herbario): Lectotypus (Fig. 2),
una una flor montada al reves, la flor mas delgada
que la de C. hunteriana, tiene fecha del 4 de mayo
de 1920, labelo 5,9 cm alto;
2)AMES 26893: se parece aAMES 26894 y todavia
mas a AMES 25007 (C. hunteriana), sin fecha;
labelo 5,8 cm alto.

En este material de herbario se encuentran dos
species de Coryanthes adicionales que fueron descritas

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


Figura 5. Coryanthes hunteriana Schltr., Panama, Prov. Panama, cerca de Arraijan, 02/2948, vista lateral; la flecha muestra
el hipoquilo alargado, el caracter mas distintivo de la especie. Foto G. Gerlach.

Figura 6. Coryanthes hunteriana Schltr., Panama, Prov. Figura 7. Coryanthes hunteriana Schltr., Costa Rica,
Panama, cercadeArraijan, 02/2948, vistadorsal; laflecha Prov. Limon, Rio Pacuare, 0-18363 formaa rosada que
muestra el hipoquilo alargado y truncado, el caracter mas representando el color de tipo de C. horichiana). Foto
distintivo de la especie. Foto G. Gerlach. G. Gerlach.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

GERLACH & ROMERO-GONZALEZ Las Coryanthes de Powell

Figura 8. Coryanthes panamensis G. Gerlach, Panama, Figura 9. Coryanthes panamensis G. Gerlach, Panama,
Prov. Panama, Rio Piedras Zonaa del canal) 0-18594 Prov. Panama, Rio Piedras Zonaa del canal) 0-18594
(fotografia del material del holotipo), vista lateral. Foto (fotografia del material del holotipo), vista dorsal. Foto
G. Gerlach. G. Gerlach.

decadas despu6s como C. misasii G.A. Romero & G.
Gerlach (Gerlach & Romero 1990) y C. panamensis
G. Gerlach (Gerlach & Schill 1993). Comparando las
flores de las llamadas C. hunteriana y C. powellii, no
encontramos diferencias que justifiquen el status de
C. powellii como especie o subespecie. Comparando
las descripciones originales (i.e., los prot6logos) de las
dos species tampoco se detectan grandes diferencias.
Ademas, aqui enfatizamos que estas descripciones se
basan en caracteres que tienen poco peso para distinguir
las species. La tabla 1 contrast las descripciones
originales en latin.
Trabajando con material extenso del g6nero
Coryanthes, tanto cultivado como de herbario, hemos
determinado que los caracteres que permitenidentificar
las species (i.e., los caracteres diagn6sticos) no
estan siempre incluidos en las descripciones. Gerlach
(1993) diferenciaba su C. panamensis por el apice

del hipoquilo profundamente trapeziforme y sinuado
("... Cum apice hypochilii profundo trapeziforme-
sinuato"). En sus descripciones Schlechter mencion6
al character del hipoquilo: "apice subacuto" en
C. hunteriana y "hypochilio cucullato, obtuse
acuminato" en C. powellii. En realidad, el hipoquilo
se ve subagudo o obtuso acuminado en todas las flores
menos en aquellas que nosotros hemos identificado
como Coryanthes panamensis y C. misasii. Estas
dos ultimas species no presentan el caricter del
apice del hipoquilo mencionado por Schlechter (Fig.
8 12) y por consiguiente no se pueden confundir
con sus concepts de C. hunteriana o C. powellii.
Comparando todo este material con los numeros
mencionados 19 y 156 de Powell, creemos que
Schlechter tuvo en sus manos la misma especie pero
en dos oportunidades con diferentes nuneros y, sin
darse cuenta, la describi6 dos veces (un error que este

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.


3,3 cm

Figura 10. Coryanthes panamensis G. Gerlach. A. habito, B. Flor en vista lateral, C. Osmoforo, parties del hipoquilo
removido, D. Hipoquilo en vista dorsal. Dibujo por H.G. Seeger.

LANKESTERIANA8(2), August 2008. O Unversidad de Costa Rica, 2008.

GERLACH & ROMERO-GONZALEZ Las Coryanthes de Powell

Figura 11. Fotografia de Coryanthes misasii G. A. Romero
& G. Gerlach (originalmente montada en la muestra
AMES 25008).
autor cometi6 envarias oportunidades). Esta hip6tesis
se apoya en la falta de diferencias significativas en las
flores prensadas por Powell con estos numerous y la
falta de diferencias importantes en las descripciones
de Schlechter de C. hunteriana y C. powellii (Tabla
1). Basandonos en estos arguments, referimos a C.
powellii a la sinonimia de C. hunteriana.
Allen (1949) fue el primer botanico en relegar las
dos species de Schlechter a la sinonimia a Coryanthes
maculata Hook., aunque tambi6n incluy6 en su lista
de sin6nimos a C. albertinae H. Karst. y C. splendens
Barb. Rodr. Pensamos que el concept de Coryanthes
de Allen era muy amplio y que incluia muchas
species que ahora son aceptadas separadamente
por muchos autores, como C. albertinae, restringida
a Venezuela, C. maculata, un element tipico de
la Flora del escudo Guayan6s, y C. splendens, un
sin6nimo de C. speciosa (Hook.) Hook., una especie
endemica de Brasil.
Debido a la falta de caracteres diagn6sticos descritos

Figura 12. C. misasii G. A. Romero & G. Gerlach. Dibujo
por K. Brown-Wing.

por Schlechter que pudieran servir para identificar
Coryanthes hunteriana, aqui complementamos su
descripci6n para que esta especie pueda ser identificada
con seguridad en el future. Se distingue facilmente
por la forma del apice del hipoquilo: en vista lateral
6ste se ve largamente elongado (Fig. 5 y 7) y en vista
dorsal truncado o obtuso (Fig. 4 y 6). En la mayoria
de las species de la secci6n Coryanthes del g6nero
Coryanthes (en Mesoamerica no hay reports de la otra
otra secci6n) el apice del hipoquilo se ve emarginado y
sinuado (por ejemplo en C. panamensis, Fig. 8, 9 y 10)
o agudo, pero en ningun caso truncado o obtuso.
La interpretation de C. hunteriana por Gerlach &
Schill (1993) en su monografia del g6nero Coryanthes
estuvo errada: la especie ilustrada con este nombre
hoy se asigna a C. picturata, la especie mas coming en
Mesoamerica, que se encuentra desde Mexico hasta
Panama. Es bastante variable y tiene formas unicolores
de color crema o amarillo palido igual que formas
punteadas o manchadas. La especie que esta ilustrada

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rca, 2008.


en la misma obra con el nombre Coryanthes picturata
(Gerlach & Schill 1993: pagina 79, figure 10 y pagina
146 figures 90 & 91), tampoco fue correctamente
interpretada: esta fue descrita diez afios mas tarde como
C. kaiseriana G. Gerlach (Gerlach & Dressier 2003).
Para concluir, hoy en dia el conocimiento y
los datos que tenemos disponibles de Coryanthes
claramente son much mejores que en el pasado,
debido a una mejor infraestructura de informaci6n
y a que se han colectado mas muestras de las
diferentes species del genero. Las species de
Coryanthes descritas por Schlechter de Panama ya
han sido colectadas en Costa Rica, aunque las flores
presentan una coloraci6n algo diferente (Fig. 7).
Supuestamente por causes de la descripci6n pobre
en caracteres y por falta de ilustraciones, Jenny
(1986) la describi6 como C. horichiana sin darse
cuenta que se trataba de la C. hunteriana.

AGRADECIMIENTOS. Agradecemos a los herbarios y los
curadores del Royal Botanical Gardens, Kew en Inglaterra
(K), y del Jardin Botanico de Missouri, San Louis, Missouri,
E.E.U.U. (MO), por permitiros estudiar sus colecciones y
publicar algunas de sus muestras. Los autores agradecen
el apoyo financiero de la Sociedad de Orquideologia de
Arizona (Orchid Society of Arizona, a G.A.R.-G.).

Allen, PH. 1949. Flora of Panama. Part II. Fascicle 4.
(Orchidaceae, third part). Ann. Miss. Bot. Gard. 36(1):
Ames, O. 1944. Destruction of the Schlechter Herbarium by
bombing. Amer. Orchid Soc. Bull. 13: 105-106.
Christenson, E.A. 1991. Mesoamerican orchid studies II:
Powell's panamanian orchidaceae. Lindleyana 6(3):
Gerlach, G. & Romero, G.A. 1990. Beitrige zur Gattung
Coryanthes Hook. VI: Coryanthes gemotii und
Coryanthes misasii zwei neue Orchideen aus den
Waldern Siidamerikas. Die Orchidee 42(2): 64-68.
Gerlach, G. & Schill, R. 1993. Die Gattung Coryanthes
Hook. (Orchidaceae); Eine monographische Bearbeitung
unter besonderer Berficksichtigung der Blftenduftstoffe.
Tropische und subtropische Pflanzenwelt 83: 1-205.
Gerlach, G. & Dressier, R. 2003. Stanhopeinae
Mesoamericanae I. Lankesteriana 8: 23-30.
Jenny, R. 1986. Einige Notizen zur Gattung Coryanthes
Hooker und eine neue Art: Coryanthes horichiana. Die
Orchidee 37(3): 124-132.
McNeill, J. [Chairman]. 2006. International Code of
Botanical Nomenclature (Vienna Code). A. R. G.
Gantner Verlag, Ruggell, Liechtenstein.
Merrill, E. D. 1943. Destruction of the Berlin Herbarium.
Science 98, No. 2553: 490-491.
Schlechter, R. 1922. Beitrige zur Orchideenkunde von
Zentralamerika. I. OrchidaceaePowellianaePanamenses.
Repert. Spec. Nov. Regni Veg. Beih. 17: 1-95.

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rca, 2008.

LANKESTERIANA8(2) 43-52 2008



Moosweg 9, 3112 Allmendingen, B Bern, Switzerland

There are a few famous examples of seldom seen
botanical periodicals. The orchid plates published
in journals like Curtis's Botanical Magazine and
Edwards's Botanical Register have been reprinted,
but many others are known only from the originals
locked away in libraries. A good example is Loddiges'
Botanical Cabinet, published in 20 volumes with 2000
plates between 1817 and 1833. The history of this
journal and its content must be seen in relation to the
Loddiges & Sons Nursery, in Hackney.
At the beginning of the 19th century, tropical
orchids were almost unknown in Europe, though
some illustrations and descriptions were known
from expeditions in tropical countries, along with
herbarium material collected on such expeditions.
Nobody had a clear idea of the richness of forms
and species in this family, and tropical orchids were
treated more or less as mere curiosities of the plant
kingdom. This situation would change dramatically
about 1840, when the orchid fever in England started
to spread, first through the island and then also -
beginning in Belgium through continental Europe.
It was a part of the life-style of rich people to build
up an orchid collection, and the names of nurseries
like Sander, Linden, Low and Veitch, all trying to
satisfy the growing market for plants from tropical
countries, became famous.
Right at the beginning of this development, we find
the name of Joachim Conrad Loddiges (fig. 1) and his
establishment in Hackney, London. Joachim Conrad
Loddiges was born in 1738, the son of a gardener
working for a nobleman in Vristbergholzen near
Hannover in Germany. He was trained as a gardener
between 1758 and 1761 with Joseph Conrad Wefer
in Velzen, near Haarlem in the Netherlands. After
finishing his education he went to England to start
in the best tradition of his family as gardener with
J. B. Sylvester in Hackney, London. In 1771 Conrad

took over a small import-nursery from John Busch,
with its stock of customers and suppliers. Conrad
was a very keen and intelligent businessman, but his
starting capital was so low that that he couldn't pay
Busch for the nursery at one time, but by 1777 he had
paid everything and owned a small but flourishing
and profitable nursery with a small shop for seeds.
In the same year Loddiges published the first edition
of a whole series of catalogues as "A catalogue of
plants and seeds which are sold by Conrad Loddiges,
nursery and seedsman, at Hackney, near London."
We have a handwritten copy by Conrad Loddiges
(member of the fifth generation, great-great-grandson
of Joachim Conrad Loddiges, or great-grandson of
George Loddiges) of this first catalogue published in
the year 1914, copied from the original of Sir Joseph
Banks (which is now kept in the British Museum).
The editions 2 to 8 are very seldom seen, we know
only handwritten copies of those catalogues from
the Lindley library. In 1804 there were already 9
orchids mentioned, all of them as Epidendrum, in
1807 there were still 6 Epidendrum listed and in
1811 13 (some of them now known as Cymbidium or
Oncidium species). The orchid catalogues of 1839,
1841 and 1844 list a tremendous number of species,
in Stanhopea alone, Loddiges offered in 1839 23
species (some unnamed), in 1841 53 species, and
in 1844 65 species. Obviously Loddiges was both
fascinated by Stanhopea and able to cultivate them
rather well. Interestingly enough, some colored
illustrations of different Stanhopea species are known
that were not published in the Botanical Cabinet; one
of them is Stanhopea insignis var. major (Fig. 2). The
drawing has a label dated 4 October 1844, and the
handwritten remark: "drawn in Loddiges' garden."
It is quite unclear where these drawings are. Xerox
copies showed up in the collection of Herman Sweet,
but we have been unable to locate the originals.


FIGURE 1. Joachim Conrad Loddiges, portrait by John Renton, son of the headgardener at Loddiges' nursery.

LANKESTERIANA8(2), August 2008. O Unlversidad de Costa Rica, 2008.

JENNY The Botanical Cabinet

About the year 1790 Loddiges started to import
living plants from other continents, especially from
the United States of America. In 1776 his first son
William was born, and in 1786 his second son George
(Fig. 3): both later joined the company. Therefore,
Conrad changed the name of the company to Loddiges
& Sons. When Conrad Loddiges died at the age of 87
years, on 13 March 1826, he still was not an English
citizen, but he owned the best known and most
famous company for the importation of exotic plants
in Great Britain. Already in 1800 his nursery was the
largest in Great Britain and also the most profitable.
According to some authors, Loddiges & Sons was
then the largest nursery in the world (Yearsley,
2000). In 1818 Loddiges erected the first, at the time,
gigantic, steam-heated palm house, 24 years before
Kew got its famous palm house. The technology that
Loddiges used for the building and the steam heating
system was revolutionary. To the great surprise of the
visitors it was even possible to produce artificial rain
in the house. In those years, the loss of plants during
their transport from outside Europe was dramatic.
Loddiges himself mentioned that only 1 plant
out of 20 would survive transport. The invention
of Wardian cases in effect, small, transportable
greenhouses changed the situation and dramatically
reduced the loss. The Wardian case was invented by
Nathaniel Bagshaw Ward, physician and examiner in
botany to the Society of Apothecaries in London
(1791-1868). By using Wardian cases, only 1 out of
20 plants was lost during transport and this change
was one of the reasons for the "Orchid fever" in
the second half of the 19th century. Loddiges &
Sons was one of the very first companies to realize
the tremendous advantage of the Wardian cases,
and started to use them immediately after their
Loddiges started to import tropical orchids
around 1812; in 1826 his catalogue listed 84
species. Conrad and especially his son George saw
the tremendous business possibilities with orchids
quickly, and consequently they treated orchids as
one of the mainstays of their business. Like many
other nurseries, Loddiges also worked together with
a number of professional collectors who supplied
the nursery in Hackney with material, and from
time to time also with new species collected all

-. tL-..a
irn Enm -,~

FIGURE 2. Stanhopea insignis var.major, drawing from
Loddiges collection, detailed origin unknown.

FIGURE 3. George Loddiges, portrait by John Renton, son
of the headgardener at Loddiges' nursery, from the
collection of the Royal Horticultural Society in London.

LANKESTERIANA8(2), August 2008. 0 Universdad de Costa Rca, 2008.

L ii

I r


over the world. Hugh Cumming was one of those
early collectors, travelling for Loddiges. In 1839,
13 years after Conrad Loddiges died, his son George
published the first orchid catalogue, including 1024
different species in 25 different genera. In 1844 the
last catalogue was published by Loddiges, and it
included no less than 1900 species. In those years, the
competition from other nurseries became a problem
for Loddiges, other nurseries started to employ
collectors and to import huge numbers of plants from
tropical areas. To make the situation worse, George
Loddiges faced problems concerning the property on
which the nursery was built, and he was also faced
with dramatically increased air-pollution, which was
very damaging to the plant stock.
When Geoge Loddiges died, in 1846, his son
Conrad took over the nursery, together with George's
brother William. William Loddiges died only 5
years later, and so Conrad junior became owner of
the nursery. In the ensuing years the nursery had
more and more problems with its competitors. The
expanding city of London added pressure to the
nursery, and the contracts for the property were
cancelled. Another very serious problem was air
pollution which badly damaged the plants. George
Loddiges had built a new greenhouse for orchids,
but in 1853 the final decision was made to close
the establishment. Loddiges and sons had to clear
out everything, together with the large palm house
and all other greenhouses. In 1852 Joseph Paxton
erected the famous Crystal Palace in London, and
sold the complete collection of about 300 palms to
Paxton. Thirty-two horses were used to transport the
largest palm, with a weight of 15 tons, to its final
place in the Crystal Palace. In two big auctions, the
complete orchid collection was sold in 1856 and
1857. Many plants of the famous collection had
been sent to the Royal Botanic Gardens in Kew by
Conrad junior before the auctions. Conrad Loddiges
junior died in 1856, and with him disappeared a
great dynasty of nurserymen and a great nursery, the
best in its time and the first in England to import
plants in great numbers. Loddiges and Sons had
the most complete and largest collection of palms
in the world, a collection of roses that included all
known species and hybrids, a collection of tropical
ferns with more than 100 species, and, last but not

LANKESTERIANA8(2), August 2008. Unversidad de Costa Rica, 2008.

uwishlb a ablAMans orf tah.
bDIrnlale fhr Mauranetbki.4,

bn n- r n l 1h s fr I_

u. o*uWt urIJt itw A4ri.trr T

nJ.cArflit' jI.W Awd.

FIGURE 3. Frontpage of Vol.1 of the Botanical Cabinet from

least, a collection of trees and shrubs of more than
3000 species, of which it was said that it included
every single tree and shrub that could be cultivated
in England without a greenhouse. Aside from his
activities in the nursery, Geoge Loddiges found time
to do many other things, he was vice-president of the
Royal Horticultural Society in London and member
of the Linnean Society, the Zoological Society and
the Microscopy Society. He owned a collection of
preserved hummingbirds with more species than all
other collections in the world together.
The genus Loddigesia Sims (1808, Leguminosae)
was dedicated to Conrad Loddiges (father) and quite a
few orchids carry the name Loddiges, either for Conrad
Loddiges, or George Loddiges, his son:
Physosiphon loddigesii Lindley (1835)
Lepanthes lodddigesiana (Swartz) Rchb.f (1856)

JENNY The Botanical Cabinet


FIGURE 4. Goodyera pubescens, the first plate in the whole
series from 1817.

Octomeria loddigesii Lindley (1836)
Dendrobium loddigesii Rolfe (1887)
Cirrhaea loddigesii Lindley (1832)
Acropera loddigesii Lindley (1833)
Cycnoches loddigesii Lindley (1832)
Cattleya loddigesii Lindley (1821)
Already Conrad Loddiges had the idea of
presenting his plants to a broader public in an
illustrated form, and so George started in 1817
to publish his own series of publications under
the name "Botanical Cabinet." The full title is
"The Botanical Cabinet consisting of coloured
delineations of plants from all countries with a
short account of each, directions for management
&c. &c.", with the added maxim "Even Solomon
in all his glory was not arrayed like one of these."
(Fig. 3). In its essence, this periodical was simply

( rnO M.M I nrto' gdnm ij, i
rhmud r K, Ib il i

FIGE 5. Text of the plate of Godyera pubecen from
the Botanical Cabinet for 1817. t
.IF-, m- -l 1 -.-,l - ~ 1, 1-

an illustrated catalogue or an illustrated supplement

for the catalogues of the nursery. The idea was
to publish the best and most interesting plants
available in the nursery as coloured illustrations
photographs were not yet available at that time
W.-A h rl11 i l ll g [ ril ) oIiM, L- -H,

together with some information about origin and

Cabinet was, except for the Curtis's Botanical
Magazine and Edwards's Botanical Register,
one of the very first periodicals of this kind to

be published. The editor tried to make clear that
his Botanical Cabinet was not intended to be
a competitor of Curtis's Botanical Magazine.

George Loddiges contacted John Sims, from 1800
until 1826 the editor of the Botanical Magazine
(vol. 14 to 53), before he started the publication
and explained the situation to him. The very short
texts accompanying the plates inBotanical Cabinet

reflect Loddiges' effort to avoid any conflict with
a well established and scientifically accepted

journal like Curtis's Botanical Magazine.
oANlESTIANA(2), eAugst 2008 Uversdad de Costaia, 2008lat
together with some information about origin and
culture and a short description. The Botanical
Cabinet was, except for the Curtis's Botanical
aMagazine and Edwards's Botanical Register,

a competitor of Curtis's Botanical MMagazine

(vol. 14 to 53), before he started the publication

LANKESTERIANAX(Z, August 2008. 0 Umlversidad de Costa Rica, 2008.


The first part was published in May 1817, the
last part of volume 20 was distributed in December
1833. Each month one part with ten completely
hand-coloured plates was published for five
shillings, in parallel a cheaper version with only
partially coloured plates was sold for two shillings
and six dimes. The twenty volumes included a total
number of 2000 plates, 131 of them were orchids,
many of them created by George Loddiges,
himself. A number of drawings originated from
Edward William Cooke and Jane Loddiges. Jane
was the daughter of Conrad Loddiges (and sister of
George) and E. W. Cooke was her husband. Edward
William Cooke was a landscape and marine painter,
born in 1811 in Pentonville, London. The drawings
made by E. W. Cook and Jane Loddiges,together
with those from George Loddiges himself, and
other involved artists like T. Boys, Miss Rebello,
W. Miller, P Heath and J. P Heath were engraved
by George Cooke, father of E.W.Cooke, printed
and then coloured by different artists. George
Cooke was born 1781 in London and was a line-
engraver by profession. The originals from E. W.
Cooke and Jane Loddiges were offered in 1929
by their grandson to the Bombay Natural History
Society. The originals of all drawings are kept
today in the collection of the British Museum. In
a letter of October 1879, E. W. Cooke stated that
all 2000 engraved copper plates were stolen by one
of Loddiges' men from his library in the garden.
The exact publication dates of all 2000 plates
between May 1817 (Vol. 1, fasc. 1, t. 1 10) and
December 1833 (Vol. 20, fasc. 10, t. 1991 2000)
are published in Stafleu in Taxonomic Literature
and by Garay in the journal Taxon (1969).
The plates are almost perfect, and the colour reflects
reality and is not unlike many later similar publications
like Flore des Serres or Lindenia exaggerated. The
very first plate published in 1817 was an orchid:
Goodyera pubescens (Fig. 4, 5), described in 1813
by Robert Brown. Loddiges received it much earlier
from J. and W Bartram and had it since about 1785 in
cultivation. Kew received it from the Duke of Kent in
1802. As Loddiges wrote: "The plant is with difficulty
cultivated -- loves a shady situation, and rich bog earth.
Our specimen flowered in September." The original
drawing was made by George Loddiges himself.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

FIGURE 6. Ficus setacea, from Botanical Cabinet, Vol. 12.

As explained above, only 131 plates of the 2000
were orchids, and it is interesting to see that most of
them represent species that are not too difficult to
transport from their home to England and rather easy
of culture, or, more honestly, hard to kill. Cultivation
of orchids during Loddiges' time was rather different
from today's ideas about cultivation and only really
tough plants could survive. This is not the place to
write much about plants other than orchids, but I would
like to show at least a few examples, Ficus setacea (t.
1138) from vol. 12 (Fig. 6) after a drawing by T. Boys
and IP% %itiora herbertiana (Fig. 7) from vol. 14, both
are very good examples to document the high quality
of the drawings.
Plate 585 from 1822 takes a very special place in
the whole series, it is Cypripedium venustum (Fig.
8). The very first species of the genus Paphiopedilum
was described and illustrated by John Sims only
two years earlier in Curtis Botanical Magazine as
Cypripedium venustum. We don't know who the
artist of this plant was, but under the impression of
the strange, beautiful plant and flower, Loddiges
wrote: "The view of such pleasing and astonishing

JENNY The Botanical Cabinet

ra ils : hrst [r

FIGURE 7. Passiflora herbertiana, from Botanical Cabinet, Vol. 14.

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


FIGURE 8. Cypripedium venustum from Botanical Cabinet, 1822.

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.

JENNY The Botanical Cabinet


ea it


FIGURE 9. Cycnoches Loddigesii from 1833, the last plate in the work.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

^ ^



productions of Divine power and Goodness is an
endless source of gratitude, wonder and delight.
Under the influence of such impressions, who indeed
can avoid exclaiming with the inspired penman
of old, "0 Lord, how manifold are Thy works; in
wisdom hast Thou made them all."
Also the last published plate (no. 2000) is an
orchid: Cycnoches loddigesii (Fig. 9) described and
named by John Lindley. The plant was collected
by John Henry Lance, barrister and at that time
commissary judge in Surinam and imported by
Loddiges in 1830, it flowered in Loddiges' culture
in 1832 for the first time, the drawing was made by

Jane Loddiges and shows the male flowers of the
species. Beneath the few words about Cycnoches
loddigesii, Loddiges also explained why he would
stop the publication of the Botanical Cabinet
with Vol. 20; he wrote: "Having been enabled to
complete our twentieth volume, and thus to place
two thousand plants before the public, our labours
are closed; the precarious state of our draughtsman's
health not permitting him to go on any farther."
Though only about 131 of all the plates of the
Botanical Cabinet show orchids, the whole work is
an invaluable addition to any library, it documents the
first steps of the orchid fever in England.


Garay, L.A. 1969. Dating of the Botanical Cabinet and of Paxton's Flower Garden, Taxon 18:709 712.
Loddiges, G. 1804. Catalogue, handwriting.
Loddiges, G. 1807. Catalogue, handwriting.
Loddiges, G. 1811. Catalogue, handwriting.
Loddiges, G. 1814. A catalogue of plants and seeds Ed.9: 35 40.
Loddiges, G. 1818. A catalogue of plants Ed. 11: 3 11.
Loddiges, G. 1820. A catalogue of plants and seeds Ed.12: 3 13.
Loddiges, G. 1823. A catalogue of plants and seeds Ed.13: 3 11.
Loddiges, G. 1826. A catalogue of plants and seeds Ed.14: 3 18.
Loddiges, C. 1914. The following lists are compiled.....
Loddiges, G. 1839. Orchideae in the collection of Conrad Loddiges and Sons : 1 24.
Loddiges, G. 1841. Orchideae in the collection of Conrad Loddiges and Sons : 1 40.
Loddiges, G. 1844. Orchideae in the collection of Conrad Loddiges and Sons :1 45.
Loddiges, G. 1849. A catalogue of plants and seeds : 1 82.
Solman, D. 1995. Loddiges of Hackney, the largest hothouse in the world, Hackney Society, John Straker ed.
Stafleu, F. 1981. Taxonomic Literature 3(Lh-O): 133 137; Bohn, Scheltema & Holkema, Utrecht.
Yearsley, G. 2000. Conrad Loddiges & sons, the first orchid nursery; Orchid Review 108:116 121.

LANKESTERIANA8(2), August 2008. Unlversidad de Costa Rca, 2008.

LANKESTERLANA8(2) 53-74 2008



1 Jardin Botanico Lankester, Universidad de Costa Rica, P O. Box 1031-7050 Cartago, Costa Rica.
2 Centro de Investigaci6n en Orquideas de los Andes "Angel Andreetta", Universidad Alfredo Perez Guerrero,
Extension Gualaceo, Ecuador.
3Harvard University Herbaria, Cambridge, MA, USA.
4 Marie Selby Botanical Gardens, Sarasota, FL, USA.
SAuthor for correspondence: diego.bogarin@ucr.ac.cr

ABSTRACT. We present and illustrate 11 new records of Orchidaceae from Costa Rica, and propose a new
combination in Acianthera for Pleurothallis aberrans. Barbosella orbicularis and Myoxanthus speciosus,
previously recorded from Costa Rica on the basis of doubtful vouchers, are confirmed to occur in the country
and illustrated from Costa Rican vouchers. Warmingia margaritacea is reduced to the synonymy of Warmingia
zamorana, a species previously known only from Ecuador. A new species, Epidendrum zunigae is described.
This species is similar to E. guanacastense and E. isomerum, from which it differs by the narrow, lanceolate
leaves, the greenish flowers, the apical half of the column purple with the clinandrium white, the sepals 11 mm
long, and the creamy yellow, cordiform, acute, bicallose lip, which is somewhat convex in natural position.

RESUMEN: Presentamos e ilustramos 11 nuevos registros de Orchidaceae para Costa Rica, y proponemos
una nueva combinaci6n en Acianthera para Pleurothallis aberrans. Se confirm la presencia en el pais de
Barbosella orbicularis y Myoxanthus speciosus, registradas anteriormente con base en registros dudosos, y
las dos species se ilustran con material costarricense. Warmingia margaritacea se reduce a la sinonimia
de Warmingia zamorana, una especie previamente conocida solamente del Ecuador. Se describe una nueva
especie, Epidendrum zunigae. Esta especie es similar a E. guanacastense y E. isomerum, de los cuales difiere
por las hojas estrechas, lanceoladas, las flores verduzcas, con la mitad apical de la column morada y el
clinandrio blanco, los sepalos 11 mm de largo y el label cordiforme, agudo, bicalloso y amarillo crema, el
cual es algo convexo en posici6n natural.

KEY WORDS: Acianthera aberrans, Costa Rica, Epidendrum zunigae, floristics, new records, Orchidaceae,
Warmingia zamorana.

Despite its well established tradition in botanical
exploration, which started in 1846 with the visit of
Oersted (1846), Costa Rica is still far from having
a complete inventory of its orchidaceous flora. The
establishment of large and documented collections
of living plants at Lankester Botanical Garden,
associated with an increasing access to critical
documentation, have been the key to improve our
understanding of orchid diversity in Costa Rica.
After the publication of the most recent and
complete treatment of the family by Dressler (2003)
new species have been added on a regular basis
to the country's inventory. As part of the general
activities of botanical exploration, documentation

and orchid identification carried out at Lankester
Botanical Garden, we present the following new
records for the flora of Costa Rica and a new

1. Acianthera aberrans (Luer) Pupulin & Bogarin,
comb. nov.
Basionym: Pleurothallis aberrans Luer, Selbyana
2(4): 382.1978. Type: Panama. Veraguas: epiphytic
in tree north of the continental divide, alt. ca. 700
m., above Santa F6, 6 September 1976, C. Luer
& R.L. Dressier 1628 holotypee, SEL). Aberrantia
aberrans (Luer) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 95: 253. 2004, nom. inval. Aberrantia



1 cm

S 2 mm I


S 1 mm



FIGURE 1. Acianthera aberrans (Luer) Pupulin & Bogarin. A. Habit. B. Apex of inflorescence. C. Flower, side view. D.
Dissected perianth. E. Lip, frontal view. F. Lip, lateral views. G. Petals, lateral views. H. Pollinarium and anther cap.
Drawing by F. Pupulin based on Pupulin et al. 4857 (JBL-spirit).

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

BOGARIN et al. New species and records of Orchidaceae

aberrans (Luer) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 103: 310. 2005.

DISTRIBUTION: Costa Rica, Panama and Ecuador.

ETYMOLOGY: from the Latin aberrans, "away from the
usual", in reference to the unusual characteristics of
the species, according to its author.

HABITAT IN COSTA RICA: epiphytic in tropical
rain forest in the Caribbean lowlands at 275 m of

Horquetas, road to Rara Avis, ca. 6 km, granja La
Selva, 10020'15"N 8400'15" W, 275 m, tropical rain
forest, secondary vegetation with large remnant trees,
along the edge of pastures, 26 July 2003 F Pupulin
4857, M. Pupulin, C. Pupulin, C. Ossenbach & B.
Arias (JBL-spirit!) (Fig. 1, 15-A).

Although this species has not been yet
analyzed by molecular phylogenetic techniques,
morphological features suggest that it belongs to
the genus Acianthera Scheidw. Luer (1978) stated
that it vegetatively resembles to Pleurothallis
circumplexa Lindl., P. pacayana Schltr., and P.
pantasmi Rchb.f. (all now included in the genus
Acianthera) because the inflorescence emerges
from the blade of the leaf above the base. Also, he
noted that the green, glabrous, gaping flowers in
the short raceme resemble those of Pleurothallis
cogniauxiana Schltr., P. decipiens Ames &
C.Schweinf., and P. verecunda Schltr. (as well as
many others), all of them also transferred to the
genus Acianthera by several authors (Pridgeon &
Chase 2001, Luer 2004). Its bicallose, truncate
petals ending into a short apiculum, as well as the
long claw of the lip, are unusual features of this
species. Luer (2004, 2005) considered these floral
details, together with the pair of pointed calli laying
near the center of lip (rounded in our specimen) as
critical features to segregate Pleurothallis aberrans
into the monotypic genus Aberrantia Luer. We
consider the monotypic genus Aberrantia, only
defined by subtle floral features, congeneric with
Acianthera. According to Luer (2003a), the voucher
cited by Pupulin (2002a) is from Panama. Here, we
cite a Costa Rican voucher for this species.

2. Barbosella orbicularis Luer, Selbyana 3 (1,2): 10.
1976. Type: Panama. Panama: La Eneida, region
of Cerro Jefe, epiphytic, alt. 1000 m, 25 December
1967, R. L. Dressier 3285 holotypee, SEL).

DISTRIBUTION: Costa Rica, Panama, Colombia, Ecuador
and Venezuela.

ETYMOLOGY: from the Latin orbicular, rounded, in
allusion to the characteristic shape of the lenticular

HABITAT IN COSTA RICA: Epiphytic in tropical wet forest
along the Caribbean watershed of the Central Volcanic
range at about 500 m of elevation.

Siquirres, Alto Guayacan, ca 500 m, camino de
Siquirres hacia Turrialba, terrenos de "Costa Rican
Amphibian Research Center", en arbol viejo de pilon
(Hyeronima sp.) en potrero, bosque humedo tropical,
colectada por B. Kubicki, 30 septiembre 2005, D.
Bogarin 1949 (JBL-spirit) (Fig. 2, 15-B).

This species had been cited for Costa Rica by
Pupulin (2002a) based on a plant cultivated at
Lankester Botanical Garden. Unfortunately, the plant
never flowered, although it was clearly a specimen of
B. orbicularis. It was excluded from the flora of Costa
Rica by Luer (2003b), because of the absence of a
proper voucher. Fertile material now available leaves
no doubt of the occurrence of this species in Costa
Among Costa Rican species of Barbosella,
B. orbicularis is the most distinctive. It is easily
recognized by the creeping habit, the rounded smooth,
overlapping leaves and the purplish flowers with a
3-lobed lip.

3. Brenesia lappiformis (A.H.Heller & L.O.Williams)
Luer, Monogr. Syst. Bot. Missouri Bot. Gard.
95: 255. 2004. Pleurothallis lappiformis
A.H.Heller & L.O.Williams, Fieldiana, Bot. 31:
42. 1964. Myoxanthus lappiformis (A.H.Heller &
L.O.Williams) Luer, Monogr. Syst. Bot. Missouri
Bot. Gard. 15: 38. 1986; Echinella lappiformis
(A.H.Heller & L.O.Williams) Pridgeon &
M.WChase, Lindleyana 16: 253. 2001, nom.
illeg.; Echinosepala lappiformis (A.H.Heller &

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S2 mm

S1 mm I

2 mm


I 5 mm I

I 1 cm 1 1 2 mm

FIGURE 2. Barbosella orbicularis Luer. A. Habit. B. Flower. C. Dissected perianth. D. Column and lip, lateral view. E. Lip.
F. Column, frontal view. Drawing by D. Bogarin based onBogarin 1949 (JBL-spirit).

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3 mm 3 mm

and lip, lateral view. E. Lip. F. Column, frontal view. Drawing by F. Pupulin based on Bogarin 890 (JBL-spirit)
1 cm 1 cm 1

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1 cm

FIGURE 4. Comparison of dissected perianths of Epidendrum adnatum (A) and E. maduroi (B). ILLUSTRATION VOUCHERS: A,
Bogarin 2378 (JBL-spirit); B, Bogarin 601 (JBL-spirit). Drawings by A. Karremans.

L.O.Williams) Pridgeon & M.W.Chase, Lindleyana
17: 101. 2002. Type: Nicaragua: Chontales:
epiphytic at Pistacho Peak near Babilonia Mine,
alt. 650 m, July 1962, A. Heller 6620 holotypee, F,
not seen).
DISTRIBUTION: Nicaragua, Costa Rica Venezuela,

ETYMOLOGY: from the Latin lappiformis, "stone-
shaped" or "bur shaped", in reference to the round and
hard consistency of the flowers.

HABITAT IN COSTA RICA: epiphytic in tropical wet forest,
premontane belt transition, and premontane wet forest,
basal belt transition, along the Caribbean lowlands.

Horquetas, C. Ossenbach 265 (JBL-spirit!). Limon:
Pococi, Guapiles, carretera Braulio Carrillo, ca. 2 km
hacia abajo de la entrada del Teleferico del Bosque
Lluvioso, en lomas a orilla de la carretera, 520 m,
10011'33"N 84054'27"W, bosque muy humedo
tropical transicion a premontano, epifitas en arboles
caidos en bosque secundario, 9 julio 2004, D. Bogarin
890 & F Pupulin (JBL-spirit!) (Fig. 3, 15-C).

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This species is distinguished by the dark
purple, hairy flower, with the tip of the narrow
dorsal sepal touching the apex of the synsepal. The
sepals are rugose-verrucose in the inner surface.
A closely related species, Brenesia stone (Luer)
Luer, endemic to Costa Rica, is distinguished by
the broad dorsal sepal, free from the synsepal,
spathulate, rugose petals and narrowly uncinate
lateral lobes of the lip.

4. Epidendrum maduroi Hagsater & Garcia-Cruz,
Icon. Orchid. (Mexico) 3: pl. 352. 1999. Type:
Panama: Chiriqui: Norte de Guadalupe, Cerro
Punta, Volcan Baru, 2000-3000 m, collected
15 March 1981, pressed 22 February 1983, E.
Hcigsater & R.L. Dressier 6468 holotypee, AMO,
not seen; isotype, MO, not seen).

DISTRIBUTION: Costa Rica and Panama.

EPONYMY: dedicated to Andres Maduro, owner of Finca
Dracula, Panama, where the collectors were based
when collecting the type.

HABITAT IN COSTA RICA: epiphytic in premontane

BOGARIN et al. New species and records of Orchidaceae

and montane wet forests in the Central Cordillera at
elevations of 1400-1800 meters.

Piedades Norte-Zapotal, camino entire el Cerro Azahar
hacia Los Bajos y San Antonio de Zapotal, Finca de
Don Guillermo, 10010'07.9"N 84035'50.3" W, 1423
m, bosque pluvial premontano, epifitas en bosque
secundario, 24 marzo 2005. D. Bogarin 1465, E
Pupulin, A.C. Rodriguez & E. Salas (JBL-spirit!).
Heredia: San Rafael, camino del Monte de La Cruz
hacia el Refugio de Vida Silvestre Cerro Dantas, a
orillas del Rio Nuevo y la Quebrada Cabra, Reserva
Forestal Cordillera Volcanica Central, 1800 m,
1005'45"N 84002'02"W, bosque pluvial premontano,
epifitas en bosque secundario, 7 enero 2004, D.
Bogarin 601, D. Lobo &A. Vargas. (JBL-spirit!) (Fig.
4, Fig.15-D).

Epidendrum maduroi belongs to the E. albertii
group (Hagsater 1999a). Plants of E. maduroi have
probably been confused as both E. adnatum and E.
lankesteri, described from Costa Rica. Epidendrum
maduroi can be distinguished from both species by
the wider, 8 mm vs. 4-5 mm, unlobed, widely ovate-
subchordate vs. slightly lobed, narrowly ovate lip
(Ames 1923, Ames & Schweinfurth 1925).

5. Epidendrum orthodontum Hagsater & L.Sanchez,
Icon. Orchid. 3: t. 361. 1999. Type: Panama:
Provincia Chiriqui: Cuesta de Las Palmas, southern
slopes of Cerro de La Horqueta, humid forest,
1700-2100 m, 17 March 1911, H. Pittier 3220
holotypee, US-677617, not seen; isotype, AMES-
22677, photo).

DISTRIBUTION: Costa Rica and Panama.

ETYMOLOGY: from the Greek ortho, "erect" and odonto,
"tooth" in reference to the erect tooth at the apex of
the column.

HABITAT: plants of this species were found growing in
montane wet forest at around 2500 m elevation in the
Cordillera de Talamanca.

COSTA RICAN MATERIAL: Alajuela: La Palma de San
Ramon, 1250 m, 11 octubre 1922, Brenes 337 (CR);
La Palma, 24 de octubre 1922, Brenes (7) 337 (AMES,

photo; CR); La Palma de San Ramon, 1250 m, 4 de
noviembre 1924, Brenes (1114) 256 (NY, not seen).
Cartago: El Guarco, San Isidro, Madreselva, Tres de
Junio, Carretera Interamericana Sur, km 67, 9040'27.4"
N 8351'66.5" W, 2530 m, bosque pluvial montano, en
bosque secundario a orillas de la carretera, 20 de enero
2008, A. Karremans 2255. (JBL-Spirit!) (Fig. 5).

Epidendrum orthodontum is similar to E.
nutantirhachis Ames & C. Schweinf., but it can be
distinguished by the trilobed lip, the larger flowers
(sepals 10-12 mm vs. 8-9 mm long; petals 9-10 mm vs.
6-8 mm long), and the erect mid tooth of the column,
forming a 90 angle with column (vs. semierect,
forming a 450 angle with the column) (Hagsater

6. Epidendrum scharfii Hagsater & Dodson, Icon.
Orchid. 2: t. 185. 1993. Type: Ecuador. Pichincha:
along river and hillside opposite town of Tandapi at
km 55 Santo Domingo to Quito road, 31 December
1986, C. Dodson & T. Dodson 16757 holotypee,
RPSC, not seen).

DISTRIBUTION: Costa Rica, Colombia and Ecuador. This
species may well occur in Panama.

EPONYMY: dedicated to Colonel Paul Scharf, bird
watcher and occasional orchid collector in Quito,

HABITAT IN COSTA RICA: plants of this species grow
on both the Atlantic and Pacific watersheds of the
Talamanca range, in premontane wet forests at
elevations between 700 to 1200 m.

between Turrialba and Jimenez, La Suiza, Pejivalle,
road to Esperanza, on hills close to Quebrada Puente,
9048'46.0"N 83039'10.0"W, 750 m, premontane wet
forest, epiphytic in secondary vegetation along the
sugar cane plantations, 5 February 2007, F Pupulin
6500, D. Bogarin & R.L. Dressier (JBL-spirit) (Fig. 6,
15-E). Cartago: Limite entire Turrialba y Jimenez, La
Suiza, Pejivalle, camino a Esperanza, en lomas cerca
de la Quebrada Puente, 9048'46.0"N 8339'10.0" W,
738 m, bosque muy humedo premontano, epifitas en
bosque secundario a la orilla de cafiaverales, 1 mayo
2008, D. Bogarin 4835, A. Karremans, Y Kisel &

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1cm 5,mm 5 cm

FIGURE 5. Epidendrum orthodontum Hagsater & L.Sanchez. A. Habit. B. Inflorescence. C. Flower. D. Dissected perianth. E.
Column and lip, lateral view. F. Column, frontal and lateral view. Drawing by A. Karremans based on Karremans 2255

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I 1 cm

FIGURE 6. Comparison of dissected perianths ofEpidendrum scharfii (A) and E. dentiferum (B). ILLUSTRATION VOUCHERS: A,
Pupulin 6500 (JBL-spirit); B, Karremans 1485 (JBL-spirit). Drawings by A. Karremans.

R. Phillips (JBL-Spirit). Puntarenas: Coto Brus, Las
Cruces (entrada a Concepcion), 1212 m, 8046'55.68"N
82057'51.36"O, 29 abril 2006, C. Ossenbach 570 & P
Casasa (JBL-spirit).

Epidendrum scharfii belongs to the Epidendrum
ditlormei group. It is closely related to Epidendrum
dentiferum Ames & C. Schweinf., but the latter has
white-green flowers, acuminate segments, linear petals
and lateral sepals forming a 900 angle with the dorsal
sepal (vs. green flowers, obtuse segments, elliptic
petals, and sepals forming a 1200 angle inE. scharfii).
It is also similar to E. gregorii which has shorter and
wider sepals and petals and a bilobed lip, with each
lobe obliquely notched on the distal margin.

7. Epidendrum stellidifforme Hagsater & Dodson,
Icon. Orchid. (Mexico) 4: t. 487. 2001. Type:
Ecuador: Carchi: Maldonado, 1500 m, 2 October
1981, L. Werling & S. Leth-Nissen 147 holotypee,
AMES, photo; isotype, QCA, not seen).

DISTRIBUTION: Costa Rica, Colombia and Ecuador. This
species may well occur in Panama.

ETYMOLOGY: from the Latin stella, "star", in reference
to the shape of the dry flowers and dittinrme' for the
group to which the species belongs.

HABITAT IN COSTA RICA: epiphytic in tropical wet forest,
premontane belt transition, on the eastern slope of the
Miravalles Volcano at 650-700 m of elevation.

Zapote, desvio a la izquierda despues del puente sobre
Rio Zapote en sentido Bijagua-Pueblo Nuevo, ladera
este del Volcan Miravalles, siguiendo la margen del
Rio Zapote, 10044'37.4"N 85005'14.9"W, 650-700 m,
bosque muy humedo tropical, transici6n a premontano,
epifitas en bordes de potreros y arboles aislados, 30 abril
2006, D. Bogarin 2814, F Pupulin, A. Rambelli & J.
Rambelli (CR, JBL-Spirit) (Fig. 7, 15-F).

Epidendrum r'ellilittnHrm belongs to Epidendrum
dlittroii' Jacq. group. It is similar to Epidendrum
hunterianum Schltr., from which it can be distinguished
by the larger (16 x 16 mm vs. 14 x 14 mm), shallowly
three lobed, mucronate lip, the acute-acuminate sepals
and petals, and the narrower (2.5-3 mm vs. 3.5-4 mm),
narrowly elliptic (vs. elliptic) petals. The type species
differs from the Costa Rican material in the acuminate,
flat lip, and the smaller plants with shorter and narrower
leaves (Hagsater 2001). The Colombian Epidendrum
killipii Hagsater & L. Sanchez, is very similar to this
species, and may prove to be the same. According to
the protologue, the ovary of E. killipii is not inflated,
but the drawing of the type shows both a cylindric,

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3 cm

2 mm

S 1 cm i L

FIGURE 7. Epidendrum stellidifomne Hagsater & Dodson. A. Habit. B. Flower. C. Dissected perianth. D. Column and lip,
lateral view. E. Column, frontal and lateral view. F. Pollinarium and anther cap. Drawing by A. Karremans based on
Bogarin 2814 (JBL-spirit).

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i i i
: 1

: :
~i r:

BOGARIN et al. New species and records of Orchidaceae

narrow ovary in lateral view, and an inflated ovary in
longitudinal section (Hagsater 1999c). If conspecific,
the name E. killipii has priority.

8. Epidendrum zunigae Hagsater, Karremans &
Bogarin, sp. nov.
TYPE: COSTA RICA. Puntarenas: Osa, Sierpe,
Mogos, alrededores de Quebrada Porvenir, ca.
3 km norte de Alto de Mogos, 8046'36.5"N
83021'23.6" W, 126 m, bosque muy humedo
tropical, epifitas bosque secundario, 30 marzo
2006, D. Bogarin 2680, J. Zu iiga & Curso de
Botinica Forestal-UCR holotypee, JBL-spirit)
(Fig. 8, 15-G).

Species Epidendro guanacastensi Ames & C.
Schweinf. similis, floribus majoribus viridulis, labello
acuto bicalloso, apice columnae purpurea, clinandrio
albo differt.

Plant epiphytic, monopodial, pendent, branching
herb. Roots basal, from the main stem, fleshy, filiform,
thin. Stems terete, somewhat flexuous, incipiently
branched near the base of the main stem. Leaves
numerous, distributed throughout the stems; sheath
tubular, minutely rugose; blades linear-lanceolate,
acuminate, short mucronate, coriaceous, slightly
carinate, those on the main stem ca. 12, similar in
size. Inflorescence apical, produced from the main
stem, and presumably form the secondary branches;
peduncle reduced. Floral bracts longer than the ovary,
amplexicaul, imbricating, ovate-oblong, rounded.
Flowers 3, distichous, greenish, the lip creamy
yellow, column green at base, the apical half purple,
clinandrium-hood white. Ovary terete, smooth, thin.
Sepals partly spreading, narrowly elliptic, acute,
5-veined, margin slightly revolute, entire. Petals
partly spreading, linear-elliptic, acute, 5-veined,
margin entire, spreading. Lip entire, cordiform, acute,
spreading, slightly convex in natural position, margin
entire, spreading; bicallose, the calli thickened at
the base and ending in low keels, with a prominent
median keel stretching from the base of the lip to the
apex. Column somewhat arching upwards above the
middle, short, internally provided with a pair of lateral
thickenings at the height of the rostellum, and forming
a narrow channel; clinandrium hood prominent, funnel-
shaped, fleshy, margin entire; rostellum at the middle
of the column, slit. Anther obovate, 4-celled.

DISTRIBUTION: Known only from the lowlands in
front of the Osa Penninsula, near the Pacific coast of
southern Costa Rica.

EPONYMY: named in honor of Jose Daniel Zufiiga
Delgado, research assistant at Jardin Botanico
Lankester, who participated in the type collection. He
photographed and preserved flowers in alcohol thus
permitting its classification.

HABITAT AND ECOLOGY: epiphytic in secondary tropical
wet forest at 100-150 m of elevation in the southern
Pacific lowlands in Peninsula de Osa. Flowering in
cultivation in November.

Epidendrum zunigae belongs to the Epidendrum
ramosum group, which is characterized by the
monopodial, branching stems, the spike-like, distichous
inflorescence, and to theEpidendrum isomerum subgroup,
which has long, pendent plants, very narrow, acute leaves,
and 1- (rarely 3-) flowered inflorescences. Epidendrum
zunigae is recognized by the narrow, lanceolate leaves,
the greenish flowers with sepals to 11 mm long, the
creamy yellow lip, the apical half of the column marked
with purple, with a white clinandrium, and the cordiform,
acute lip, somewhat convex in natural position. It
resembles E. isomerum Schltr., which has 3-11 x 0.1-
0.3 cm leaves, a single-flowered inflorescence produced
from short flowering branches, green to yellowish green
flowers, with very narrow floral segments, sepals 15-16
mm long, the lip rhombic-triangular, acute, with a "V"
shaped callus. It is also similar to E. guanacastense Ames
& C. Schweinf, which has thicker stems, and a zigzag
inflorescence with 2-3 flowers, sepals 8-10 mm long, a
cordiform, rounded lip, and the callus "Y" shaped.

9. Lockhartia chociensis Kraenzl. in H.G.A.Engler
(ed.), Pflanzenr., IV, 50(83): 19. 1923. Type:
Colombia: Choco, Triana s.n. holotypee, W).

DISTRIBUTION: Costa Rica, Colombia, Ecuador,
Venezuela and Peru.

ETYMOLOGY: named from the Choco region in
Colombia, the place from where the type specimen
was collected.

HABITAT IN COSTA RICA: epiphytic in secondary forest
in tropical wet forest premontane belt transition in the
Caribbean watershed of Tilaran range.

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I 1 cm I 3 mm I i 5 mm

FIGURE 8. Epidendrum zunigae Hagsater, Karremans & Bogarin. A. Habit. B. Flower in natural position. C. Dissected
perianth. D. Column and lip, lateral view. E. Lip. F. Column, lateral and frontal view. Drawn by D. Bogarin from the

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I 1 mm

I 2 mm I

I 5 mm I

1 5 mm I

FIGURE 9. Lockhartia chocOensis Kraenzl. A. Habit. B. Flower in natural position. C. Dissected perianth. D. Column and
lip, lateral view. E. Column, frontal view. F. Pollinarium and anther cap. Drawing by D. Bogarin based onBogarin 2352

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I 1 cm

I 2 cm

6 cm 1 cm

FIGURE 10. Maxillaria bolivarensis C.Schweinf. A. Habit. B. Flower in natural position. C. Dissected perianth. D. Column
and lip, lateral view. E. Column, lateral and frontal view. F. Pollinarium and anther cap. Drawing by D. Bogarin based
on Whitten 2030 (JBL-spirit).

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Aguas Claras de Buenos Aires, Hotel Termales
Azules, camino por la ladera del Volcan Rinc6n de
La Vieja, hasta las cataratas, 700-2000 m, 6 abril
2004, A. Karremans 271 (JBL-spirit!). Upala,
Bijagua, Zapote, desvio a la izquierda despues
del puente sobre Rio Zapote en sentido Bijagua-
Pueblo Nuevo, ladera este del Volcan Miravalles,
siguiendo la margen del Rio Zapote, 10044'37.4"N
8505'14.9"W, 650-700 m, bosque muy humedo
tropical, transici6n a premontano, epifitas en bordes
de potreros y arboles aislados, 1 febrero 2006, D.
Bogarin 2352, R.L. Dressier, R. Gomez & A. Rojas
(JBL-spirit!) (Fig. 9, 15-H).

This species is similar to Lockhartia micrantha
Rchb.f., but the lip is six-lobulate with a pair of
narrow basal lobules folding upward, two acute
lobules placed at the middle, and two apical, rounded
lobules. The callus is made up by two parallel keels
running to the middle and convergent at apex. The
illustration shown in Garay and Dunsterville (1966),
and identified as L. chocoensis, is consistent with
the specimens studied from Costa Rica. Lockhartia
lankesteri Ames, based on a collection by C.H.
Lankester from Aguas Zarcas, San Carlos, in the
Atlantic lowlands of Costa Rica, has been reduced into
the synonymy ofL. micrantha. With the discovery of
L. chocoensis populations in Upala, close to the San
Carlos region, it is now probable that L. lankesteri is
a later name for L. chocoensis. The drawing of the lip
from the holotype ofL. lankesteri at AMES (AMES-
101030) is somewhat schematic however it shows
two small lobes at the middle of the lip. A careful
rehydratation of the flowers of the type should clarify
the status ofL. lankesteri.

10. Maxillaria bolivarensis C.Schweinf., Bot. Mus.
Leafl. 20: 22. 1962. Type: Venezuela. State of
Bolivar, Region of Uriman, forest mesa of Aprada-
tepui, at 950 m, August 13, 1953, L. Bernardi 780
holotypee, MER, not seen; isotype, AMES-69561,

DISTRIBUTION: Costa Rica, Venezuela, Ecuador and
Peru. The Costa Rican records are not so far away from
Nicaraguan boundary. This species may well occur in
Nicaragua, Panama and Colombia.

ETYMOLOGY: from the State of Bolivar, Venezuela
where the type specimen was collected.

HABITAT IN COSTA RICA: epiphytic in tropical wet forest
along the Caribbean lowlands.

Horquetas, above Horquetas, Terra Folia Reserve,
near Rara Avis, wet premontane forest, epiphytic,
10018' 14"N 840 01'36" W, 500 m, 21 July 2003,AM.
Whitten 2030 & M. Blanco (JBL-Spirit!); Sarapiqui,
Magsasay, ribera del Rio Peje, 180 m, 31 diciembre
2005, C. Ossenbach 507 & J.E Casasa (JBL-Spirit!)
(Fig. 10, 15-I).

This species is easily recognized by the
inconspicuous pseudobulbs, the linear-oblong,
coriaceous leaves less than 1.5 cm wide, and the
smooth clinandrium and anther cap. In Costa
Rica, Maxillaria confusa Ames & C.Schweinf. is
superficially similar to M. bolivarensis, however its
conspicuous pseudobulbs, broader subcoriaceous
leaves (up to 3 cm wide), fringed clinandrium and
ciliate anther cap are useful features to recognize this

11. Myoxanthus speciosus (Luer) Luer, Selbyana 7:
51. 1982. Pleurothallis speciosa Luer, Selbyana
3: 392. 1977. Type: Panama: Chiriqui: epiphytic
in cloud forest above Guadalupe, ca. 2000 m,
13 September 1976, C. Luer & H. Butcher 1371
holotypee, SEL).

DISTRIBUTION: Costa Rica and Panama.

ETYMOLOGY: from the Latin speciosus, "splendid", in
reference to its showy flowers.

HABITAT IN COSTA RICA: epiphytic in lower montane
rain forest at 2450 m of elevation in Talamanca range.

Dota-El Guarco, Jardin, entire La Chonta y Cafion,
km 56 carretera Interamericana, Finca Santa Maria
de La Selva, 9041'27.5"N 83055'20.3" W, 2450 m,
epifita en bosque pluvial montano bajo, en arboles
de potreros principalmente sobre Drymis winteri,
Alnus acuminata y Quercus sp, 19 febrero 2005, D.
Bogarin 1420, C. Ossenbach & E Pupulin (JBL-
spirit!) (Fig. 11).

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1 cm

FIGURE 11. Myoxanthus speciosus (Luer) Luer. A. Flower in natural position. B. Dissected perianth. C. Column and lip.
Drawing by F. Pupulin based on Bogarin 1420 (JBL-spirit).

Pupulin (2002a) and Luer (2003) cited this species
for Costa Rica based on a plant cultivated in Switzerland
and collected in "Cordillera de Talamanca, Sierra Hills
near Coraz6n de Jesus, alt. 2100 m" in Puntarenas
Province (Jenny P-25, MO). Luer (2003c) stated
that its origin is doubtful in all the collection details
including the country. However, material collected near
Cerro de La Muerte, along the Panamerican Highway
and flowered in the living collections at Lankester
Botanical Garden confirms the presence of this species
in Costa Rica.

12. Scaphyglottis robusta B.R.Adams, Phytologia 64:
253. 1988. Type: Panama. Panama: El Llano-Carti-
Tupile road, 10-12 km N of Inter-Am. Highway,
9 January 1975, Luteyn & Wilbur 4679 holotypee,
DUKE, not seen).

DISTRIBUTION: Costa Rica and Panama.

ETYMOLOGY: from the Latin robustus, "stout" in
reference to the robust and strong appearance of this
species compared with other members of the genus.

HABITAT IN COSTA RICA: epiphytic in tropical wet
premontane forest to 1150 m of elevation in the
Caribbean watershed of the Talamanca range.

La Esperanza, floreci6 en cultivo en la colecci6n de
Julio Carmona, La Suiza de Turrialba, 22 marzo
2006, D. Bogarin 2662 (CR, JBL-Spirit) (Fig. 12,

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15-J). Turrialba, Pacayitas, calle a La Suiza, ca. km 8,
9052'29.9"N 83035'03.6"W, 1150 m, bosque humedo
premontano, 4 de marzo 2007, A. Karremans 1661 &
D. Karremans (JBL-Spirit).

This species is similar in habit to M.,./il,./. *.i,
modest (Rchb.f.) Schltr.; however, it can be
distinguished by the larger habit, the yellow-greenish
to bright green flowers (vs. white with reddish-purplish
stripes in S. modest. Antillean plants have dull,
yellow-greenish to brownish-green), the spathulate lip
(vs. 3-lobed) and the prominent wings of the column.
\.,.!1I,-1. *i i anneliesae Brieger, a nomen nudum
published without a description or any indication of the
type, has been attributed to Costa Rica. Adams (1993)
studied Brieger's accompanying photographs and
suggested that it is referable to S. robusta. However
in describing S. robusta, he only cited Panamanian
material. Here, we report the presence ofS. robusta in
Costa Rican based on two collections.

13. Sobralia bouchei Ames & C.Schweinf., Schedul.
Orchid. 10: 4. 1930. Type: Panama. March-April
1930, 2200 feet, terrestrial and epiphytic, A.M.
Bouche s.n. holotypee, AMES-35595, photo).

DISTRIBUTION: Nicaragua, Costa Rica and Panama.

EPONYMY: in honor ofAdrien M. Bouche, who collected
the type specimen.

HABITAT IN COSTA RICA: epiphytic or terrestrial in

BOGARIN et al. New species and records of Orchidaceae


S5mm i /I

1 cm

1 cm

1 14cm I

FIGURE 12. Scaphyglottis robusta B.R. Adams. A. Habit. B. Flower in natural position. C. Dissected perianth. D. Column and
lip, lateral view. E. Column, lateral and frontal view. Drawing by D. Bogarin based on Bogarin 2662 (JBL-spirit).

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.




3 cm

1 cm

5 mm
I i

S1 cm
1 I

FIGURE 13. Warmingia zamorana Dodson. A-B. Habit. C. Flower. D. Dissected perianth. E. Column and lip, three-quarter
view. F. Column and lip, lateral view. G. Column, ventral view. In E-G the pollinarium is bending toward the stigma,
whereas the viscidium is still in place. Drawing by F. Pupulin based on Karremans 452 from Costa Rica (JBL-spirit).

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

BOGARIN et al. New species and records of Orchidaceae

3 mm

5 mm 1 cm
__ __ __ _

FIGURE 8. Warmingia zamorana Dodson. A. Habit. B. Flower in natural position. C. Dissected perianth. D. Column and
lip, lateral view. E. Column, lateral and ventral view. F. Pollinarium. Drawing by F. Pupulin based on Medina s.n. from
Ecuador (CIOA-spirit).

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.


roadcuts in tropical wet premontane forest to 900
m of elevation in the Caribbean watershed of the
Talamanca range.

Tayutic, carretera entire Tayutic y Jicotea, Platanillo,
laderas del Rio Platanillo, siguiendo la margen del rio,
9049'27.46"N 83033'13.29"W, 878 m, bosque muy
humedo premontano, 1 abril 2008, D. Bogarin 4201,
A. Russell & R. Samuel (JBL-Spirit).

This species is easily recognized by the pink
flowers and the strongly ruffled purple lip with 7
crisped golden-yellow keels. The leaves are bright
green and smooth. Plants from Panama and cultivated
at Lankester Botanical Garden produce fully opened
flowers. Nevertheless, the plant collected in Costa
Rica has flowers self-pollinated and developing fruits.
This condition has been also observed in Nicaraguan
material (Dressler, pers. comm. 2008). Sobralia
triandra A.H.Heller & A.D.Hawkes described from
Nicaragua could be conspecific with S. bouchei.

14. Warmingia zamorana Dodson, Icon. P1. Trop., II,
6: t. 599. 1989. Type: Ecuador. Zamora-Chinchipe:
Zamora, 1000 m, August 1968, C. H. Dodson
3842 holotypee: SEL). Warmingia margaritacea
B. Johans., Lindleyana 7: 194. 1992, syn. nov.
Type: Costa Rica. [Cartago]: Turrialba, C.A.T.I.E.,
epiphytic onHibiscus sp. forming hedge, 600 m, 19
November 1988, B. Johansen & M. Sorensen 138
holotypee: C, not seen).

DISTRIBUTION: Costa Rica and Ecuador.

ETYMOLOGY: named from the city of Zamora in southern
Ecuador, the locality of the type specimen.

HABITAT IN COSTA RICA: epiphytic in premontane wet
forest on garden trees and bushes in Turrialba region.

frente al edificio principal del Centro Agron6mico
Tropical de Investigaci6n y Ensefianza (CATIE) en
cerca viva a la par de un tubo de agua, 2 a 3 flores
pendientes, 3 noviembre 2003, A. Karremans 452.
(JBL-spirit) (Fig 13, 15-K). Turrialba, CATIE. Frente
al Edificio Principal del CATIE, sobre una cerca viva
de Hibiscus sp., al lado de un tubo de agua. 9053'22"

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rica, 2008.

N 8339'12" W, 600 m, 4 noviembre 2005, A.
Karremans 1123 (JBL-spirit). ECUADOR. Zamora-
Chinchipe a lo largo del Rio Zamora entire 600 a 800
m, H. Medina s.n. (CIOA). Morona-Santiago: cerca de
Patuca, H. Medina s.n. (CIOA-spirit) (Fig. 15-L).

This species was first recorded from Costa Rica
in 1992 by Johansen as a new species named W
margaritacea. The plant was collected at CATIE
campus in Turrialba, Costa Rica, growing in Hibiscus
sp. fences (Johansen 1992). After its description,
the species had been long known only by the type
collection (Atwood 1999, Dressler 2003). Fifteen years
later, in November 2003, botanical exploration carried
out at CATIE revealed more populations of this species
growing in Hibiscus sp. (Pupulin 2004, 2005).
With a careful analysis of the type specimen of W
zamorana as well as of Ecuadorian material cultivated
from the Zamora-Chinchipe region, we conclude that
the characters used to separate W margaritacea from W
zamorana are inconsistent, and both species should be
considered conspecific. Johansen (1992) stated that W
margaritacea can be distinguished from W zamorana
by the coriaceous, lanceolate, subfalcate leaves, the
pauciflorus inflorescence, the rhombic petals, the rigid
column appendices and the self-pollinated flowers.
However, this set of features were also observed
in living specimens of W zamorana from Ecuador.
Morevover, Johansen stated that the flowers of W
margaritacea are pure ivory and pearly appearance,
contrasting W zamorana as having a yellow lip callus
(Dodson 1989), but the callus of all the examined Costa
Rican records is bright yellow in color (a photograph
is given in Pupulin 2005). The serrate petals and the
three-lobed lip with a bilobate yellow callus were
also observed in both Ecuadorian specimens from
Zamora and Costa Rican material collected at CATIE.
Although we have still not found a wild population of
this species outside of CATIE, some plants have been
collected growing in Cupressus lusitanicus trees.
Warmingia zamorana has not been recorded
yet in the orchid floras of Panama and Colombia,
the two countries in the middle of its distribution.
Nevertheless, that disjunction may be attributed to
undercollection of these small and easily overlooked
epiphytes. It is remarkable that similar examples
of disjunction in the orchid floras of Costa Rica

BOGARIN et al. New species and records of Orchidaceae

FIGURE 15. Pictures of: A. Acianthera aberrans. B. Barbosella orbicularis. C. Brenesia lappiformis. D. Epidendrum
maduroi. E. Epidendrum .rellilittlorme. F. Epidendrum scharfii. G. Epidendrum zunigae. H. Lockhartia
chocoensis. I. Maxillaria bolivarensis. J. y. ,!1i ,.-.ii robusta. K. Warmingia zamorana (Costa Rica). L.
Wamingia zamorana (Ecuador).

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.


and Ecuador were documented in the case of
Ornithocephalus montealegrae Pupulin, originally
described from the Turrialba region in Costa Rica
(Pupulin 2002b) and recorded in Ecuador by Dodson
(2003), and in Epidendrum scharfii, E. \r'lliillttrhm'.
Lockhartia chocoensis and Maxillaria bolivarensis,
discussed in this paper.

ACKNOWLEDGEMENTS. We want to thank Robert L. Dressier
and Eric Hagsater for their invaluable support and their
useful comments on the paper. Hugo Medina and Jose
Daniel Ziuiiga who assisted part of this study. The scientific
services of Ministerio del Ambiente y Energia de Costa
Rica (MINAE) and its Sistema Nacional de Areas de
Conservaci6n (SINAC) for issuing the Scientific Passport
N. 1281 under which wild species treated in this study were
collected, and the Ministerio del Ambiente of Ecuador, for
the permits issued for the managing of the ex-situ orchid
collections at Gualaceo and El Pangui, Ecuador. To the
Vicerrectoria de Investigaci6n of the University of Costa
Rica for providing support under the project N. 814-A7-
015 "Inventario y taxonomia de la flora epifita de la region

Adams, B.R. 1993. A taxonomic revision of the
genus Scaphyglottis Poepp. & Endl. (Orchidaceae-
Epidendroideae). Ph.D. dissertation, Southern Illinois
University, Carbondale.
Ames, O. 1923. Additions to the orchid flora of Central
America. Schedul. Orch. 4 : 45.
Ames, O & C. Schweinfurth. 1925. New or noteworthy
species of orchids from the American Tropics. Schedul.
Orch. 8 : 41.
Atwood, J.T. 1999. Warmingia Rchb.f Pages 172-173
in: Burger, W. (Ed.). Flora Costaricensis. Family #39
Orchidaceae: Tribe Maxillarieae: Subtribes Maxillariinae
and Oncidiinae. Field., Bot., n.s. 40.
Dodson, C.H. 1989. Warmingia zamorana. Icon. P1. Trop.,
II, 6: t. 599.
Dodson, C.H. 2003. Native Ecuadorian Orchids. Volume
IV: Oncidium-Restrepiopsis. Dodson Trust, Sarasota,
Florida. USA.
Dressier, R.L. 2003. Orchidaceae. Pages 1-595 in: Hammel,
B.E., M.H. Grayum, C. Herrera and N. Zamora (Eds.).
Manual de Plantas de Costa Rica. Vol.3. Monogr. Syst.
Bot. Missouri Bot. Gard. 93.
Garay, L. A. & G.C.K. Dunsterville. 1966. Venezuelan

Orchids Illustrated. Volume IV. Andre Deutsch Limited,
London. England.
Hagsater, E. 1999a. Epidendrum maduroi. Icon. Orchid.
(Mexico). 3: 352.
Hagsater, E. 1999b. Epidendrum orthodontum. Icon. Orchid.
(Mexico). 3: 361.
Hagsater, E. 1999c. Epidendrum killipii. Icon. Orchid.
(Mexico). 3: 342.
Hagsater, E. 2001. Epidendrum stellidifforme. Icon. Orchid.
(Mexico). 4: 487
Johansen. B. 1992. Warmingia margaritacea sp. nov.
(Orchidaceae) from Costa Rica and a review of the
genus. Lindleyana 7(4):194-198.
Luer, C.A. 1978. Icones Pleurothallidinarum (Orchidaceae).
Miscellaneous new species in the Pleurothallidinae.
Selbyana 2(4): 367-390.
Luer, C.A. 2003a. Pleurothallis. Pages 386-452 in: Hammel,
B.E., M.H. Grayum, C. Herrera and N. Zamora (Eds.).
Manual de Plantas de Costa Rica. Vol.3. Monogr. Syst.
Bot. Missouri Bot. Gard. 93.
Luer, C.A. 2003b. Barbosella. Pages 22-24 in: Hammel,
B.E., M.H. Grayum, C. Herrera and N. Zamora (Eds.).
Manual de Plantas de Costa Rica. Vol.3. Monogr. Syst.
Bot. Missouri Bot. Gard. 93.
Luer, C.A. 2' '3c. Iivoxanthus. Pages 339-342 in: Hammel,
B.E., M.H. Grayum, C. Herrera and N. Zamora (Eds.).
Manual de Plantas de Costa Rica. Vol.3. Monogr. Syst.
Bot. Missouri Bot. Gard. 93.
Luer, C.A. 2004. Icones Pleurothallidinarum XXVI:
Pleurothallis subgenus Acianthera and three allied
subgenera; A Second Century of New Species of Stelis
of Ecuador; Epibator, Ophidion, Zootrophion. Monogr.
Syst. Bot. Missouri Bot. Gard. 95: 1-265.
Luer, C.A. 2005. Icones Pleurothallidinarum XXVII:
Dryadella and Acronia sect. Macrophyllae-Fasciculatae
(Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard.
103: 1-311.
Oersted, A.S. 1863. L'Am6rique Centrale: recherches sur sa
flore et sa geographic physique. Imprimerie de Bianco
Luno Par F.S. Muhle. Copenhague.
Pridgeon, A.M. & M.W. Chase. 2001. A phylogenetic
reclassification of Pleurothallidinae (Orchidaceae).
Lindleyana 16: 235-271.
Pupulin, F. 2002a. Catalogo revisado y anotado de las
Orchidaceae de Costa Rica. Lankesteriana 4:1-88.
Pupulin, F. 2002b. Exploring for orchids: Orithocephalus
montealegrae is described from Costa Rica. Orchids,
Mag. Amer. Orchid Soc. 71(11): 1016-1019.
Pupulin, F. 2004. El regreso de la Warmingia. Epidendrum
Pupulin, F. 2005. The Pearl Warmingia: a rare Oncidioid
enters cultivation. Orchids, Mag. Amer. Orchid Soc.
74(4): 282-285.

LANKESTERIANA8(2), August 2008. 0 Universidad de Costa Rca, 2008.

LANKESTERIANA8(2) 75-78 2008


Hagsater, E. & M. Soto (eds.). 2008. Icones Orchidacearum. Fascicle 10. Orchids of Mexico. Part 4. Herbario

AMO, Mexico D.F., Mexico. Pp. i-xxxvi, plates 1001-

Under the experienced editorship by Eric Hagsater
and Miguel Soto, the Herbario AMO delivers the
fourth part of "Orchids of Mexico", fascicle 10 of the
series Icones Orchidacearum. The volume is dedicated
to the late Federico Halbinger Mosig (1925-2007), a
short biography of whom is provided by Hagsater and
Soto, together with checklists of the taxa named after
him and those described or transferred by Halbinger, as
well as a complete bibliography of the great Mexican
In line with the last issues of the Icones
Orchidacearum, particular attention is given to the
typification and stabilization of nomenclature, with a
much needed effort to neotypify the species originally
proposed by La Llave and Lexarza, and by Richard
and Galeotti (the last case made more problematic
by the absence of type specimens for several taxa).
Unlike the previous releases of the series, the new
fascicle eliminates the short English descriptions to
provide more information in the Spanish version and to
augment the space devoted to the very ample chapters
on specimen citation and bibliographic references.
Even so, in some cases, the length of the texts obliged
the editors to include part of the specimen citations and
the references in 21 additional pages of appendices to
list all the vouchers known to the authors
The emphasis of the volume is onthe generaBarkeria
(5 spp.), Isochilus (4 spp.), Prosthechea (11 spp.), and
Rhynchostele (9 spp.), but the treatment also includes
interesting taxa as the Mexican endemic Lockhartia
galeottiana A. Rich ex Soto Arenas [typified by Ophrys
imbricata Sess6 & Moc. (1890)], Cuitlauzina pendula
La Llave & Lex., and Homalopetalumpumilio (Rchb.f.)
Schltr. (both neotypified in the volume), as well as
some well-known species important in horticulture
(i.e., Guarianthe skinneri, Rhynchostele spp.).
Six new taxa (i.e., Bletia village Soto Arenas,
Encyclia halbingeriana Hagsater & Soto Arenas,
Isochilus oaxanus Salazar & Soto Areas, Maxillaria
chimalpana Soto Arenas & Salazar, Rhynchostele

-1100. Published on June 11, 2008.

cervantesii subsp. halbingeriana Soto Arenas &
Hagsater, and R. maculata subsp. oestlundiana fo.
perotensis Soto Arenas & R. Jimenez) and eight
new names, combinations and/or status changes
[Camaridium punctostriatum (Rchb.f.) Soto Arenas,
Lockhartia galeottiana, Myoxanthus congestus (A.
Rich. & Galeotti) Soto Arenas, Oestlundia ligulata
(La llave & Lex.) Soto Arenas, Prosthechea squalida
(La llave & Lex.) Soto Arenas, Rhyncholaelia
digbyana subsp. fimbripetala (Ames) Soto Arenas,
Rhynchostele maculata subsp. oestlundiana (L.O.
Williams) La llave & Lex.) Soto Arenas & R. Jimenez
fo. oestlundiana, and Stelis platystylis (Schltr.) R.
Solano & Soto Arenas] are proposed in the work,
making it a necessary reference for any serious library
on Neotropical botany.
Consistently with the other volumes of the series,
the fourth part 4 of "Orchids of Mexico" adopts the
new classification system based on phylogenetic
analysis of molecular data, even when the authors
express their doubts about some of the actual generic
circumscriptions. The framework of the taxonomic
system is basically that suggested by the ongoing series
of Genera Orchidacearum, and the editors' decision
to strictly follow this system is an appreciable attempt
to maintain a common (and hopefully standing)
"language" in orchid nomenclature.
With a very few exceptions, of standard quality,
most of the botanical illustrations included in Orchids
of Mexico, part 4, are outstanding for the information
they convey and the artistic interpretation of the
depicted subjects. In particular, the work of Rolando
Jimenez Machorro (who produced 51 of the 100
illustrations) has seemingly reached its full maturity;
worthy of note are also the composite plates by
Marco Antonio L6pez-Rosas, whose inked drawings
are among the most informative in contemporary
botanical production.
Franco Pupulin
Lankester Botanical Garden, University of Costa Rica


Oakeley, H.F. 2008. Lycaste, Ida and Anguloa. The essential guide. Published by the author, Bekenham, United
Kingdom, and printed by Cambrian Printers, Aberystwyth, U.K. Large 30.5 x 30.5 cm volume, pp. (v) 445, 1400
color photographs. 1st edition published on May, 2008.

A complete, gorgeously illustrated, superbly printed
and perfectly bound large book represents the long-
waited and ultimate work onLycaste, Ida, and Anguloa
by Henry Francis Oakeley, the recognized specialist
in these orchid groups of increasing horticultural
As in his previous, small-sized "essential guide"
to Lycaste (Oakeley 1993), the author claims for
completeness: "If is not in here it has not been
described". In fact, the total numbers of recognized
species, subspecies (varieties in the actual treatment)
and natural hybrid in Lycaste and Ida (both treated
under the former genus in the old guide) are
significantly different in the new, amply augmented
monograph. When split apart into two genera, the
previous work by Oakeley recognized 24 species, 7
subspecies and 8 natural hybrids of Lycaste, and 22
species of Ida. In the present work Lycaste includes
31 species (5 of which are described as new and 4
elevated to specific rank by the author), 33 varieties
(23 new) and one (new) subvariety, and 14 natural
hybrids (7 of which described in this treatment). Ida
has 39 species, 7 of which described in the text, 11
varieties (5 new) and 3 natural hybrids, all newly
proposed in the book. A natural intergeneric hybrid
between Lycaste and Ida is also included: x Lycida
mathiasae. The treatment of Anguloa does not differ
significantly from the recent monographs published
by the author (Oakeley 1999a, 1999b, 1999c, 1999d,
1999e) and, according to Oakeley's guide, the genus
comprises today 9 species, 5 varieties, and 4 natural
hybrids, one of them (Anguloa x speciosa) described
in this volume.
For each of the nearly 150 species, natural hybrids
and varieties included in the volume, the text provides a
full description, etymology, historical notes, taxonomic
discussion, distribution and voucher citations, and
bibliography. Of these, the chapters on taxonomic
history and bibliographic references are particularly
worthy, amply documented and with frequent, direct
references to the original sources, while the citation
of herbarium specimens is generally of limited use
due the relative paucity of the consulted herbaria.

LANKESTERIANA8(2), August 2008. 0 Unversidad de Costa Rica, 2008.

Species and hybrids are listed alphabetically within
each genus, following a criterion that makes easier to
find any specific taxon, but on the other hand obscures
the phylogenetic relationships among closely related
(albeit alphabetically distant) species. The taxonomic
treatments are preceded by chapters on the history of
the discovery and the introduction of the three genera,
their general and distinguishing features, and artificial
keys to the species of Anguloa, Ida, and Lycaste. An
ample, final chapter is devoted to the cultivation of
the species of these three genera, including accounts
on their pollination and the preparation of herbarium
specimens. Dr. Oakeley is the holder of the National
Plant Collection for Anguloa, Ida and Lycaste in the
United Kingdom, and his more than 50 years long
acquaintance with the growing secrets of these plants
is well evident in this chapter. Two large Appendices
complete the book. The first is devoted to "Synonyms
and errors", and the author enlists over 300 invalid,
misapplied, and synonymous names that plagued the
complex taxonomic history of the involved genera and
have been used at some time but have to be considered
incorrect or redundant. The second Appendix is an
"Annotated bibliography" with almost one thousand
bibliographical citations (including indications of
the presence of plates/pictures/photographs), and
references to relevant websites and travel accounts
involving Anguloa, Ida, and Lycaste in their habitats.
The taxonomic novelties include the following
new species: Lycaste angelae Oakeley (with var.
alba Oakeley and var. rubra Oakeley), L. crystallina
Wubben ex Oakeley, L. fuscina Oakeley, L. occulta
Oakeley and its var. alba Oakeley, L. panamensis
Fowlie ex Oakeley, Ida angustipetala Oakeley, I.
castanea H. James ex Oakeley, I. ejirii Oakeley, I.
jimenezii Oakeley, I. munaensis Oakeley, I. priscilae
I. Portilla ex Oakeley, and I. shigerui Oakeley. Lycaste
plana 'Measuresiana' B.S. Williams, L. macrophylla
subsp. puntarenasensis Fowlie, L. macrophylla var.
viridescens Oakeley, and L. macrophylla subsp.
xanthocheila Fowlie are elevated to specific rank,
while L. plana Lindl. is sunk under a variety of L.

Book reviews

The whole book is splendidly illustrated. Large,
crisp photographs of outstanding quality by the author
himself show one or (more commonly) several views
of the flower (and variations within species), with a
special attention to taxonomic relevant features, and
the plant habit, in many cases accompanied by shots of
the plants in-situ. For most of the older taxa, original
type illustrations and/or critical historic illustrations
are perfectly reproduced to help the reader understand
original species concepts. Whenever possible, the
book even includes portraits of the dignitaries honored
with the species names. For each of the species, the
author also presents an impressive series of macro-
and microphotographs showing anatomical and
morphological features that are relevant to identify the
species. These normally include views of the dissected
lip adaxiall and lateral), the callus, the column, and
the pollinarium, but other features (like the shape and
ornament of the pseudoulbs) are also presented when
they are considered taxonomically relevant.
As the author admits in his preliminary notes, the
use of types throughout the work is "more loose than
as defined by the International Code of Botanical
Nomenclature", and this unfortunately happens in
several opportunities. Lycaste panamensis Fowlie
ex Oakeley is indicated in the text as a stat. nov.,
but no reference is made to any basionym (in fact,
Lycaste macrophylla subsp. panamensis Fowlie is a
nomen nudum). In several cases (i.e. L. dowiana, L.
lasioglossa), the author "chooses" a holotype, while in
other cases he designates a lectotype when a holotype
or isotype ostensibly is in existence (i.e., L. candida,
L. tricolor). Doubts may be expressed about the
applicability of the name Lycaste candida Lindl. This
name was just cited by Lindley as a provisional name,
but in the same text the author himself considered it
(erroneously) a synonym of L. leucantha. Thus, by
a strict application of the Rules of Nomenclature,
the name L. candida was not accepted by its own
author, and was thus invalid. The next available name
should be, in this case, L. brevispatha Klotsch 1871,
regarded by Oaekeley as a synonym of L. candida.
Other, objection cases are typifications that ignore
the available materials. Thus, for example, Lycaste
bradeorum Schltr. is neotypified with a cultivated
plant reportedly from San Isidro del General, in
southern Costa Rica, ostensibly outside the natural

distribution of this species, which is limited to the
northern, dry areas of the Guanacaste province and its
Nicaraguan neighbors. At the Ames Orchid Herbarium
of the Harvard University is kept a copy of the original
Schlechter's drawings of L. bradeorum, which could
have been selected as a more appropriate lectotype.
Another taxonomic point is the apparent frequence
of natural hybrids in some groups of species, which
can be regarded with some suspicion. Costa Rican L.
candida (probably an invalid name) and L. brevispatha
(if distinct) have been traditionally difficult to tell
apart on the basis of their amply variable flower
morphology. Now, Oakeley maintains L. candida for
populations with markedly three-lobed lip, and creates
L. angelae for plants with broadly diamond-shaped,
obscurely lobed lip (his photographs ofL. angelae var.
rubra, however, show flowers with distinctly 3-lobed
lip), resolving all the intermediate forms through the
description of a supposed new natural hybrid, L. x
daniloi, which is stated to be "much commoner than
either species".
All in all, Oakeley's book is an extraordinary
work, based on a long, deep and intimate knowledge
of the groups under study, and a fundamental piece
to understand the diversity and complexity of the
involved species. There are no doubts that this splendid
monographs will stand for a long time as the essential
reference of future students of these three genera.
Franco Pupulin
Lankester Botanical Garden, University of Costa Rica

Oakeley, H.F. 1993. Lycaste species, the essential guide.
Vigo Press, London.
Oakeley, H.F. 1999a. A revision of the genus Anguloa (in
Japanese, with type descriptions in Latin and picture
captions in English). All Japan Orch. Soc. Orch. 38:
Oakeley, H.F. 1999b. Revision der Gattung Anguloa.
Orchidee (Hamburg) 50(1): 21-28; 50(3): 3442;
50(4): 51-56.
Oakeley, H.F. 1999c. Revision del g6neroAnguloa. Revision
of the genus Anguloa. Orquideologia 159-213.
Oakeley, H.F. 1999d. Ein herziening van het Geslacht
Anguloa. Orchideedn 4: 63-80.
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