Orchids Gone Wild
IT IS DIFFICULT TO IMAGINE THE
Hawaiian landscape before Polynesians
arrived. Much of it was forested, that we
know, but it would have been a landscape
without breadfruit, coconuts, taro, ti plants,
frangipani, bananas, pineapples, sugar cane,
kukui nuts and other plants commonly
associated with Hawaiian culture, past or
present.
As the population grew and immigrants
from many different cultures and regions of
the world arrived, most forests succumbed
to exploitation, followed by the spread of
agriculture (think pineapples and sugar
cane), urbanization, industrialization, mili-
tary bases, and later, tourist development.
Populations of native plants and animals
declined, displaced or replaced by exot-
ics. In the 1900s a horticultural sensibility
developed resulting in an influx of some
of the world’s most exotic and beautiful
plants … including and perhaps especially,
orchids.
Orchids were highly valued and in
Hawaii the common man could easily grow
them outside. By the middle of the 20th cen-
tury it seemed that nearly everyone had an
orchid growing in a pot or on a slab of tree
88 ORCHIDS FEBRUARY 2012 WWW.AOS.ORG
[1] Spathoglottis plicata from along a road-
side on the Big Island of Hawaii.
[2] Kalalau Valley on the windward side of
Kauai where alien orchids have not yet
become established, likely because there
is no source seed (winds blow in from
the Pacific and not over housing where
orchids may be cultivated).
fern. Many years ago as a boy visiting my
grandparents in Manoa Valley behind the
University of Hawaii in Honolulu, I would
have arguments with my friends: “My dad
has better orchids than your grandpa!”
proclaims a friend. “No way! My grandpa’s
vandas are so much better than your dad’s!”
And so it went. But I did learn something
about both bragging rights and orchids. It
was part of the culture.
Orchids naturally occur from below the
Antarctic Circle to above the Arctic Circle
and are remarkably diverse in the moist and
wet tropics. So, there must be many species
in Hawaii, right? No, as Larry Zettler, PhD,
describes in this issue, the islands only
have three species with the charisma that
only their mothers can see. It seems the
Discovering Naturalized Orchids in Hawaii
TEXT AND PHOTOGRAPHS BY JAMES D. ACKERMAN

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isolation of the Hawaiian Islands is right at
the edge of natural dispersal probabilities
of orchids.
Yet orchids abound in gardens of
Hawaii and even the Big Island is known
as “The Orchid Isle.” As it turns out, like
species of other plant groups, some of the
orchids brought in by plant lovers have bro-
ken the chains of horticultural servitude and
have made a successful run for freedom.
The frequency of such escapes seems to
be accelerating (Ackerman 2007; Liu and
Pemberton 2010).
Invasive species are a huge problem in
Hawaii. There is even what may only be
called SWAT teams ready at a moment’s
notice to eradicate alien invaders. With so
many populations of native, endemic spe-
cies declining, population viability is a real
issue due to habitat destruction, inbreeding
and loss of mutualistic partners, many
of which are brought on by the incessant
arrival of alien competitors, parasites,
herbivores, predators and vermin of that
ilk. Yet to the delight of locals and tourists
alike, in some parts of the islands a drive
or hike through the countryside can result
in an encounter with some lovely orchids.
90 ORCHIDS FEBRUARY 2012 WWW.AOS.ORG
3 humans entertained by plant sex. Arundina
[3] Cymbidium dayanum fruits from a popu-
lation on the Big Island of Hawaii.
The most conspicuous are the tall bamboo
orchid (Arundina graminifolia), the pink
Philippine orchid (Spathoglottis plicata)
and an occasional nun’s orchid (Phaius
tankervilleae). At a conservation confer-
ence a few years ago, I posed the question
to participants from Hawaii, “With all the
concern over invasive species, why hasn’t
anybody done something about the invasive
orchids?” They looked at me with bewil-
dered expressions. Finally, somebody broke
the silence, “Uhhh, well, Jim, it’s because
they are orchids.” So it seems that not all
invasions are created equal for reasons
other than biological.
There are at least 14 species or hybrids
that have become naturalized in the Hawai-
ian Islands, but as recently as 1990 the only
documented fugitives from cultivation
were the big four: Spathoglottis, Arundina,
Phaius and a hybrid Epidendrum (Wagner
et al. 1990). The spread of the first three
comes as no surprise as they are naturally
weedy orchids, even being among the first
vascular plants to invade Krakatau after it
exploded in 1883 (Partomihardjo 2003).
Why have there been so many more
naturalized orchids reported in Hawaii in
recent years? There are several possibilities.
Propagule pressure is one. For some species
there is a lag from the time it is naturalized
to when it begins to spread aggressively,
and this involves some threshold abundance
of propagules. Another possibility is that
establishment and spread is constrained
by the lack of either mycorrhizal fungi or
pollinators. On the other hand, some or-
chids may simply be recent arrivals to the
islands. And of course, more people may
be paying attention to what is happening
and documenting it.
Except for localized vegetative re-
production, propagule pressure is directly
related to pollination success. Fruit produc-
tion is high in Polystachya concreta, which
will likely become one of the most common
naturalized species in Hawaii. How they
produce so many fruits remains a mystery,
but one recently reported escape, Dendro-
bium atennatum, is known to self-pollinate
and become weedy. Spathoglottis plicata,
one of the most commonly encountered nat-
uralized species is also self-pollinating and
fruit and seed set is very high. Nonselfing
forms of Spa. plicata are in cultivation and
occasionally we have seen fruits — even in
Waikiki — evidence of occasional selfing,
insect pollination, or the helping hand of
and Phaius are both bee-pollinated but the
Hawaiian native bees are small, not com-
mon and often absent where these orchid
species are successful. As in other parts of
the world, an invasion of one species can
facilitate the invasion of another. Thanks
to either the accidental introduction of
a carpenter bee from Mexico (Xylocopa
sonaria) or the intentional introduction of
honeybees (Apis mellifera), some exotic
orchids in Hawaii now enjoy pollinator
services, which is not necessarily a good
thing if you are an orchid grower. Roy
Tokunaga, a grower on Oahu, keeps certain
orchids in screened enclosures to keep the
pesky bees from pollinating flowers, which
causes sepals and petals to rapidly wilt
and demands an allocation of resources to
unwanted fruits.
Dendrobiums are perhaps the most
commonly cultivated genus in Hawaii and
have been for a long time. Yet the pigeon
orchid, Dendrobium crumenatum, was only
just recently reported naturalized in the
islands (Frohlich and Lau 2010). Consider-
ing the popularity of this orchid, its late es-
tablishment is somewhat surprising — but
there is evidence that it is spreading fast.
A population was first reported thriving on
 4

[4] Arundina graminifolia in the foreground

with a volcanic plume from erupting
Kilauea in the background.
[5] Dendrobium hybrid on ridge above Hono-
lulu, Oahu. According to Roy Tokunaga,
of H&R Orchids, this is similar to Den-
drobium Jacquelyn Thomas (gouldii ×
bigibbum [formerly phalaenopsis]), which
in Hawaii is grown mostly for cut flowers
and is pollinated by honeybees.
[6] Detail of the dwarf bamboo orchid
5 (Ar. graminifolia).

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HANS OPPENHEMER
7 8
frangipani (Plumeria rubra). Because the
locality was near the University, my students
and I went up the valley and found the plants.
A quick check in the neighborhood revealed
other plants growing on coconut palms next
to garbage cans. This was followed by a visit
to my grandfather’s old place farther up the
valley. And there upon the trees next to the
road were many plants in flower, which was
lucky because local populations of Den.
crumenatum flower simultaneously and
the flowers last but one day. And what was
even more exciting, we watched carpenter
bees rob nectar from the spurs, honeybees
using the same holes to thieve a little for
themselves, and a few more entering the
flowers legitimately and pollinating them.
What a marvelous sight.
The naturalization of various hybrid
dendrobiums suggests that honeybees have
indeed been busy bees. Fruit production is
not high, but is certainly typical for species
that offer no pollinator rewards, which is
the case for the majority of dendrobiums
cultivated in the islands (not the pigeon
orchid, though). Honeybees are common
everywhere, from the city streets to the
forested mountains, providing pollinator
Naturalized Orchids of Hawaii
Arundina graminifolia
Cymbidium dayanum
Dendrobium crumenatum
Dendrobium antennatum
Dendrobium cf. mirbelianum
Dendrobium nobile type
Dendrobium antelope type
Dendrobium bigibbum type
(formerly phalaenopsis)
92 ORCHIDS FEBRUARY 2012 WWW.AOS.ORG
WILFREDO FALCÓN
[7] Vanda tricolor draped over a native tree
on Maui.
[8] The nun’s orchid (Phaius tankervilleae)
flowering in a naturalized population
on the grounds of the Lyon Arboretum,
Honolulu, Oahu.
service for many different kinds of plants,
likely including the hybrid Dendrobium
populations we have seen. As for the hybrid
Dendrobium populations we have found
on the ridges above Honolulu, the original
source may not be from the city since the
tradewinds would take the seeds produced
in those gardens out toward the sea. It may
be more likely that seeds produced from
plants cultivated on the windward side of
the island were carried over the mountains
and deposited on the ridges leading to the
city.
Naturalized populations of Cymbidium
dayanum on the Big Island may also be
the beneficiaries of honeybee activity.
Although I am unaware of any observa-
tions of pollinator visits, native honeybees
pollinate other Cymbidium species in their
natural habitats (see Sugahara 2006). Fruit
Epidendrum ×obrienanum
(jamiesonis × radicans)
Habenaria rodeiensis
Phaius tankervilleae
Polystachya concreta
Spathoglottis plicata
Vanda tricolor
Zeuxine strateumatica
set was more than 30 percent, and seed set
was more than 90 percent, both of which
are quite good. We certainly can expect this
species to spread (Caccia 2005; Ackerman
et al. 2011).
Orchids are appearing everywhere
where there is sufficient water. Do you find
yourself on the leeward, dry side of the
islands? No problem! Check the irrigation
ditches. My students and I were walking
through dusty red dirt alongside an active
ditch on Kauai to take a photograph of the
island of Niihau and there in the wet soil
of the ditch was a thriving population of
Spathoglottis. Although we were surprised,
we did not change our plans to drive to
more hospitable habitats in our quest to
find naturalized orchids.
Can naturalized and invasive orchids
be a problem in Hawaii or anywhere else?
The answer is an unqualified “maybe.”
The evidence for negative consequences of
orchid invasions is scant and mostly weak.
Some are concerned that the invasive Disa
bracteata in Australia may compete for my-
corrhizal fungi with native orchids, though
this is just speculation at this point (Bon-
nardeaux et al. 2007). In Puerto Rico, the
local distribution of invasive Oeceoclades
maculata is negatively associated with two
native terrestrial orchids, which may either
reflect differences in niche preferences or
competition through mycorrhizal interac-
tions (Cohen and Ackerman 2009). On
the other hand, the evidence is strong that
populations of Spa. plicata in Puerto Rico
have increased the abundance of a native,
orchid-specialist weevil to the detriment of
co-occurring native Bletia patula. Hawaii
also has orchid weevils on Spa. plicata (and
Ar. graminifolia), but they are an invasive
species, which likely came in shipments
of plants from Southeast Asia, Malaysia,
Indonesia, Philippines or Melanesia. These
beetles are uncommon in populations of
naturalized orchids that I have seen in Ha-
waii and would not likely have any effect
on rare, native orchid species, though there
may be issues with orchids in cultivation.
One orchid that has the potential to
overwhelm native vegetation is the majes-
tic Vanda tricolor, a species only recently
reported for the islands, but now several
populations are known on the islands of
Hawaii, Maui and Oahu. When first re-
ported on Maui by Hank Oppenheimer,
the vandas were “smothering” Antidesma
platyphylla, an endemic tree (Oppenheimer
2006). Could this Vanda be an orchidaceous
kudzu? Only time will tell.
When it comes to orchids in Hawaii, we
are reasonably sure of one thing: our list of
orchids gone wild will only grow longer.
 References
Ackerman, J.D. 2007. Invasive Orchids: Weeds We Hate
to Love? Lankesteriana 7:19–21.
Ackerman, J.D., W. Falcón and W. Recart. 2011. New
Records of Naturalized Orchids in the Hawaiian Islands.
Bishop Museum Occasional Papers 110:1-4.
Bonnardeaux, Y., M. Brundrett, A. Batty, K. Dixon, J. Koch
and K. Sivasithamparam. 2007. Diversity of Mycorrhizal
Fungi of Terrestrial Orchids: Compatibility Webs, Brief
Encounters, Lasting Relationships and Alien Invasions.
Mycological Research 111:51–61.
Caccia, D. 2005. Naturalized Cymbidium in Hawaii.
Orchids 74:888.
Cohen, I.M., and J.D. Ackerman. 2009. Oeceoclades macu-
lata, an Alien Tropical Orchid in a Caribbean Rainforest.
Annals of Botany 104:557–563.
Frohlich, D., and A. Lau. 2010. New Plant Records from
O’ahu for 2008. Bishop Museum Occasional Papers
107:3–18.
Liu, H., and R. Pemberton. 2010. Pollination of an Invasive
Orchid, Cyrtopodium polyphyllum (Orchidaceae), by an
Invasive Oil-Collecting Bee, Centris nitida, in Southern
Florida. Botany 88:290-295.
Oppenheimer, H.L. 2006. New Hawaii Plant Records for
2004. Bishop Museum Occasional Papers 88:10–15.
Partomihardjo, T. 2003. Colonisation of Orchids on the
Krakatau Islands. Telopea 10:299–310.
Sugahara, M. 2006. Cymbidium devonianum and Cym-
9 10
bidium suavissimum as well as Cymbidium floribundum
Attracts Japanese Honeybees (Apis cerana japonica).
Zoological Science 23:1225.
Wagner, W.L., D.R. Herbst and S.H. Sohmer. 1990. Manual
of the Flowering Plants of Hawaii. Volume 2. University
of Hawaii Press and Bishop Museum Press, Honolulu.

Acknowledgments
Wilfredo Falcón and Wilnelia Recart were
my students collaborating with me on my recent
Hawaiian adventures. Curt Daehler hosted us at
the Botany Department, University of Hawaii,
Manoa. Phil Cribb, David Caccia, Alex Lau, Hank
Oppenheimer, Christian Torres and Roy Tokunaga
shared their knowledge of Hawaii’s naturalized
orchids for which I am grateful. Some of the
observations mentioned above were the result of
grant and fellowship support from USDA-FaST,
NSF-REU (DBI-0243750), and NSF-CREST
(HRD-0734826) programs.

James Ackerman, PhD, has been a profes-

sor at the University of Puerto Rico since
1981. His research has been primarily on
orchids, covering topics such as taxonomy,
floristics, ecology, evolution, pollination
biology, biogeography, population genet-
ics, and most recently, invasive species
biology. Although he spent time pulling
weeds from his grandfather’s orchid beds
in Hawaii, he did not develop an interest
in orchid biology until he was at Humboldt
State University in California. The expe-
rience inspired him to work with Norris
Williams at Florida State University,
leading to Panama, Smithsonian Tropi-
cal Research Institute, Bob Dressler and 11
orchid bees. He has written books on the
orchids of the Greater Antilles and has [9] Honeybee thieving nectar from Dendro- [11] Orchidless, bleak landscape at high
published more than 120 scientific ar- bium crumenatum through slits cut by elevations of Mauna Kea, Hawaii. There
ticles. Department of Biology, University carpenter bees, Xylocopa. are no orchids there because of the
of Puerto Rico, PO Box 23360, San Juan, [10] Dendrobium crumenatum fruits, the inhospitable habitat — too cold and too
Puerto Rico 00931 (email ackerman.upr@ result of successful pollination by hon- dry.
gmail.com). eybees, Oahu.

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