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Transcript of THE STATUS AND FUTURE OF ORCHID CONSERVATION ...
THE STATUS AND FUTURE OF Gary A. Krupnick,2 Melissa K. McCormick,3
ORCHID CONSERVATION IN Thomas Mirenda,4 and Dennis F. Whigham3
NORTH AMERICA1
ABSTRACT
The status and trends of issues related to the conservation of orchids native to the United States, Canada, and Greenland areconsidered. We focus on nine of the 16 Targets of the Global Strategy for Plant Conservation (GSPC). The first two targets, whichall other targets rely upon, appear to have been adequately achieved, in addition to Target 11. Limited progress has been madeon six other GSPC targets. Three case studies of efforts to conserve the native threatened orchids, Platanthera leucophaea (Nutt.)Lindl., Isotria medeoloides (Pursh) Raf., and Tolumnia bahamensis (Nash) Braem, are presented to demonstrate the difficulties aswell as the issues associated with effective conservation. We describe our efforts to establish an international program toconserve all native orchids in the United States and Canada. The North American Orchid Conservation Center (NAOCC) is aninternationally focused effort that is based on public-private partnerships. The goal of NAOCC is to conserve the geneticdiversity of all native orchids through efforts to develop an international collection of seeds and orchid fungi. The NAOCC alsofocuses on the cultivation of all native orchids in an international network of botanic gardens, and they partner with private andpublic landowners to develop techniques to conserve and restore all native orchid species.Key words: Endangered species, Global Strategy for Plant Conservation (GSPC), North American Orchid Conservation
Center (NAOCC), Orchidaceae, restoration.
Beautiful, diverse, and often bearing large and the species around them, orchids, highly sensitive toshowy flowers, orchids are an ancient plant family habitat change, are among the first casualties fromthat has evolved an amazing array of bizarre flower environmental degradation. Most orchid generatypes, unique pollination syndromes, and complex contain threatened or endangered species (Swarts &symbiotic interactions with animals and fungi. In the Dixon, 2009). Orchids are found throughout Northplant world, orchids reign supreme as about 10% of America, and many of the approximately 210 speciesall flowering plant species are members of the found north of Mexico are threatened, endangered, orOrchidaceae. No other plant family can match the extirpated in at least part of their ranges because ofpeculiar array of evolutionary features that orchids habitat loss and alteration. Most North Americancollectively possess. Along with the Asteraceae, the orchids are terrestrial. Globally, terrestrial orchidsOrchidaceae has more species, estimated to be make up only one third of orchid species with thebetween 20,000 and 35,000 taxa (Cribb et al., other two thirds being epiphytes and lithophytes.2003), than any other family of flowering plants, However, terrestrial herbaceous perennials are dis-and individual orchid species are often rare in nature, proportionately represented in the extinct plantoccurring in restricted and specific niches and species listed by The World Conservation Unionhabitats. Collectors prize orchids for their seemingly (IUCN, 1999). Consequently, terrestrial orchids areinfinite variety of showy flowers; scientists have long likely subject to a greater extinction risk thanbeen fascinated by the relationships between the epiphytes, particularly in response to current climateplants and their pollinators and other symbionts. changes.Today, orchids have taken on even greater Much of orchids’ sensitivity to habitat change
significance. Due to their interconnectedness with likely can be traced to their dependence on two, often
1 We are thankful to the United States Botanic Garden and the Smithsonian Institution for financial support of the NorthAmerican Orchid Conservation Center (NAOCC) through a Smithsonian Grand Challenges Award grant. Frank Clements,Vickie Dibella, Tony Dove, Barbara Faust, Christine Flanagan, Jim Kaufmann, John Kress, William McLaughlin, RayMims, Jay O’Neill, Holly Shimizu, Teresa Vetick, and Kyle Wallick contributed substantially to discussions about thestructure and development of NAOCC. Intern Jean Linsky assisted in collecting data. Information related to the case studydescription of Isotria medeoloides was obtained during a research project funded by Contract Number H399207001/J370010018 from the National Park Service.
2 Department of Botany, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 166,Washington, D.C. 20013-7012, U.S.A. [email protected].
3 Smithsonian Environmental Research Center, P.O. Box 28, Edgewater, Maryland 21037, U.S.A. [email protected];[email protected].
4 Smithsonian Gardens, Smithsonian Institution, P.O. Box 37012, MRC 506, Washington, D.C. 20013-7012, [email protected].
doi: 10.3417/2011108
ANN. MISSOURI BOT. GARD. 99: 180–198. PUBLISHED ON 13 DECEMBER 2013.
Volume 99, Number 2 Krupnick et al. 1812013 Status and Future of Orchid Conservation in
North America
very specific, types of symbiotic associations. for and actively participating in efforts to conserveOrchids’ relationships with specific pollinators have native orchids.been a subject of interest since before Darwin, but We begin by identifying successes and major gapsmore recently orchid dependence on mycorrhizal in conservation strategies for North American nativefungi has also received substantial research attention orchids by addressing nine of the 16 targets of the(Waterman & Bidartondo, 2008). Identification of the 2011–2020 Global Strategy for Plant Conservation
fungi on which orchids depend requires DNA (GSPC) in relation to the conservation of orchid
sequencing and analysis, but it is clear at this point species native to the United States, Canada, and
that some orchids are dependent upon fungi that are Greenland. GSPC targets 4, 5, 6, 9, 10, 12, and 13 do
free living in the soil, while others associate with not apply directly to native North American orchid
fungi that are also connected to other plants, species. We then propose a centralized NorthAmerican Orchid Conservation Center (NAOCC) thatespecially trees. As habitats change, the fungaladdresses each deficiency.community changes, and orchids may lose fungi
upon which they depend for their survival.GAs popular, and charismatic subjects for LOBAL STRATEGY OF PLANT Cdesirable, ONSERVATION TARGETS
cultivation, orchids face another serious threat. Like TARGET 1, AN ONLINE FLORA OF ALL KNOWN PLANTSprecious gems, the most unique and rare orchids aresought by the most enthusiastic collectors. As word A total of 210 orchid species are native to the
spreads about the location of a rare orchid, more and United States, Canada, and Greenland (see Table 1),including one described as recently as 2007, asmore demand is placed on already fragile popula-Platanthera yosemitensis Colwell, Sheviak & P. E.tions. Unfortunately, without efforts to cultivate theMoore (Colwell et al., 2007). The novel orchid list is asymbiotic fungi, most of these plants are doomed incompilation of species from Kartesz (1994), the Floracultivation. Due to the combination of habitat lossof North America (Romero-Gonzalez et al., 2002),and poaching, many orchid species, which were onceBrown (2009), and NatureServe Explorer (Nature-widespread, are now found in small, ecologicallyServe, 2011), and each name is accepted by Thefragile, fragmented populations. The majority ofPlant List (,http://www.thePlantList.org.) and theorchids are represented globally by tropical epi-World Checklist of Selected Plant Families (WCSP,phytes, but temperate zone terrestrials make up a2011). Additionally, 135 orchid species are native to
significant proportion (approximately 28%; Grave-Puerto Rico and the Virgin Islands (Acevedo-
ndeel et al., 2004) and conservation of terrestrialRodrıguez & Strong, 2007), but of these species only
species has proven to be especially challenging (e.g.,those that extend into the mainland are treated here.
Stewart & Hicks, 2010). No orchid has ever been de-The 210 described species (Table 1) are distrib-
listed, though one, Isotria medeoloides (Pursh) Raf.,uted among 66 genera (Table 2), with 49 genera
was upgraded from endangered to threatened in 1994 (74%) represented by only one or two species and a(U.S. Fish and Wildlife Service, 1994) after extensive few genera being represented by more than threedirected searching uncovered new populations. A species (e.g., Platanthera Rich. with 35 species,comprehensive holistic approach to species conser- Spiranthes Rich. with 23 species). While there arevation has to be fully realized in conservation and always questions of whether taxa should be split orrestoration plans. lumped and questions about the treatment of hybrids,The orchid flora of North America represents an relationships between most genera in the Orchid-
important scientific challenge for conservation biol- aceae are well established (e.g., Gorniak et al., 2010).ogists. Unlike many other plant species, where Native orchid species are found in all 50 statessignificant efforts are often employed to cultivate within the United States and all 10 provinces andand reintroduce rare plants and to store germplasm, three territories of Canada. On average, there are 40the majority of organizations that identify and protect native species per state, with Hawaii having theorchids on public and private lands rely solely on fewest native species (three) and Florida with thehabitat conservation for management. This conserva- most (106). Other species-rich areas include thetion strategy has largely been dictated by the unique states along the East Coast associated with theaspects of orchid biology that make them not Appalachian Mountains, especially North Carolinaamenable to standard plant conservation techniques. (67 species), Virginia (59 species), and New York (57We propose a model of centralized but integrated species). Of the Canadian provinces, Prince Edwardorchid conservation that can provide one-stop Island and Ontario have an especially diverse orchidshopping for agencies and organizations responsible assemblage (59 and 58 species, respectively). All
situ
ex2011).
holding
gardens
(NatureServe,
botanic
of Explorer
umber
nthe
NatureServe
and
and
protections,
(200
9),
Brown
legal
assessments,
2002
),al.,
etconservation le
z´
(Rom
ero-Gonza
includ
ing
America
Greenland
,North
ofand
Flora
Canada,
the
(199
4),
States,
Kartesz
United
from
the
ofspecies
species
oforchid
compilation
native
ais
210
list
The This
1.Table
collections.
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Anoectochilu
ssandvicensisLindl.
G3;
S3Vulnerable
1Aplectrum
hyem
ale(M
uhl.ex
Willd.)Nutt.
G5;
SH,S1
-S5,
SNR
End
angered
11ArethusabulbosaL.
G4;
SX,SH
,S1
-S4,
SNR
End
angered
0Basiphylla
eacorallicola
(Small)Ames
G2;
S1End
angered
0Beloglottiscostaricensis(Rchb.
f.)Schltr.
G4;
S1*
End
angered
0Bletia
patula
Hook.
G4;
SH5
Bletia
purpurea
(Lam
.)DC.
G5;
S3Threatened
17Brassia
caudata(L.)Lindl.
G3;
SXEnd
angered
37Bulbophyllum
pachyrachis(A.Rich.)Griseb.
G4;
SX0
Calopogon
barbatus
(Walter)Ames
G4;
S1-S3,
SNR
1Calopogon
multiflorusLindl.
G2;
SH,S1
-S3
End
angered
0Calopogon
oklahomensisD.H.Goldm
anG3;
SH,S1
,S2
,SN
R0
Calopogon
pallidusChapm
.G4;
S1-S5,
SNR
2Calopogon
tuberosus(L.)Britton,
Sterns
&G5;
SX,S1
-S4,
SNR
End
angered
16Poggenb.
Calypso
bulbosa(L.)Oakes
G5;
SH,S1
-S5,
SNR
End
angered
8Cam
aridium
vestitu
m(Sw.)Lindl.
G4;
S1*
5*Cam
pylocentrum
pachyrrhizum
(Rchb.
f.)Rolfe
G4;
S1End
angered
1Cepha
lanthera
austiniae(A.Gray)
A.Heller
G4;
S2,S3
,SN
RThreatened
1Cleistesiopsisbifaria(Fernald)Pansarin&
F.Barros
G4;
S1-S4,
SNR*
3*Cleistesiopsisdivaricata
(L.)Pansarin&
F.Barros
G4;
SX,S1
,S3
,S4
,SN
R*
End
angered*
1Cleistesiopsisoricam
porum
P.M.Br.
GNR;S2
0Corallorhizabentleyi
Freudenst.
G1;
S10
Corallorhizamaculata(Raf.)Raf.
G5;
SH,S1
-S4,
SNR
End
angered
12Corallorhizamertensiana
Bong.
G4;
S2,S3
,S5
,SU
,SN
R1
Corallorhizaodontorhiza(W
illd.)Nutt.
G5;
SH,S1
-S5,
SNR
End
angered
2CorallorhizastriataLindl.
G5;
SH,S1
-S4,
SNR
End
angered
0Corallorhizatrifida
Chatel.
G5;
S1-S5,
SNR
End
angered
2CorallorhizawisterianaConrad
G5;
SX,SH
,S1
-S5,
SNR
End
angered
2Cranichismuscosa
Sw.
G4;
S1End
angered
2Cyclopogoncranichoides
(Griseb.)Schltr.
G4;
SNR*
0Cyclopogonelatus
(Sw.)Schltr.
G4;
SH*
End
angered
4*Cypripedium
acauleAiton
G5;
S1,S3
-S5,
SNR
End
angered
22
182 Annals of theMissouri Botanical Garden
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Cypripedium
arietin
umR.Br.
G3;
SH,S1
-S3
End
angered
1Cypripedium
californicum
A.Gray
G3;
S37
Cypripedium
candidum
Muhl.ex
Willd.
G4;
SX,SH
,S1
-S3,
SNR
End
angered
Endangered
10Cypripedium
fasciculatum
Kellogg
exS.
Watson
G4;
S1-S3
2Cypripedium
gutta
tum
Sw.
G5;
S2,SN
R7
Cypripedium
kentuckiense
C.F.Reed
G3;
S1-S3
End
angered
14Cypripedium
montanum
Douglas
exLindl.
G4;
S1-S4,
SNR
3Cypripedium
parviflorum
Salisb.
G5;
SH,S1
-S4,
SNR
End
angered
12Cypripedium
passerinum
Richardson
G4;
S1-S4,
SNR
2Cypripedium
reginaeWalter
G4;
SX,S1
-S4,
SU,SN
R,SN
AEnd
angered
33Cypripedium
yatabeanum
Makino
G4;
SNR*
1Cyrtopodium
punctatum
(L.)Lindl.
G5;
S1End
angered
22Dactylorhizaaristata
(Fisch.ex
Lindl.)Soo
G4;
S40
Dactylorhizaviridis(L.)R.M.Batem
an,Pridgeon
G5;
SX,SH
,S1
-S5,
SNR*
End
angered
11*
&M.W.Chase
Dendrophylaxlin
denii(Lindl.)Benth.ex
Rolfe
G3;
S2*
End
angered
7*Dendrophylaxporrectus(Rchb.
f.)Carlsward&
GU;SN
R*
0Whitten
Dichrom
anthus
cinnabarinus
(LaLlave
&Lex.)
G5;
SNR*
2Garay
Dichrom
anthus
michuacanus
(LaLlave
&Lex.)
G4;
S3,SN
R*
0Salazar&
Soto
Arenas
Eltroplectriscalcarata(Sw.)Garay
&H.R.Sw
eet
G4;
S1End
angered
1Encyclia
tampensis(Lindl.)Sm
all
G4;
SNR
23Epidend
rum
amphistomum
A.Rich.
GNR;S3
End
angered
0Epidend
rum
anceps
Jacq.
10Epidend
rum
blancheanum
Urb.
G4;
SXEnd
angered*
1*Epidend
rum
floridenseHagsater
5Epidend
rum
magnolia
eMuhl.
G4;
S2,S3
,SN
R*
4*Epidend
rum
nocturnum
Jacq.
G4;
S2End
angered
25Epidend
rum
rigidum
Jacq.
G4;
S3End
angered
19Epidend
rum
strobiliferum
Rchb.
f.G4;
S1End
angered
1Epipactisgigantea
Douglas
exHook.
G4;
S1-S3,
SU,SN
RSpecialConcern
37Eulophiaalta
(L.)Faw
c.&
Rendle
G4;
S46
Eulophiaecristata(Fernald)Ames
G2;
S1,S2
*End
angered
0
Volume 99, Number 2 Krupnick et al. 1832013 Status and Future of Orchid Conservation in
North America
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Galeand
rabeyrichiiRchb.
f.G4;
SNR
0Galeand
rabicarinata
G.A.Rom
ero&
P.M.Br.
G1;
S1End
angered
0Galearisspectabilis
(L.)Raf.
G5;
S1-S5,
SNR
End
angered
4Goodyeraoblongifolia
Raf.
G5;
S1-S5,
SNR
End
angered
9Goodyerapubescens(W
illd.)R.Br.
G5;
S1-S5,
SNR
End
angered
27Goodyerarepens
(L.)R.Br.
G5;
SH,S1
-S5,
SNR
End
angered
14GoodyeratesselataLodd.
G5;
SX,SH
,S1
-S5,
SU,SN
REnd
angered
3Govenia
floridana
P.M.Br.
End
angered
0Habenaria
distansGriseb.
G5;
S1End
angered
1Habenaria
floribundaLindl.
G4;
SNR*
2*Habenaria
quinqueseta(M
ichx.)Sw
.G4;
S1,SN
R1
Habenaria
repens
Nutt.
G5;
S1-S3,
SNR
4Ham
marbyapaludosa
(L.)Kun
tze
G4;
S1-S3,
SNR*
End
angered*
2Heterotaxissessilis(Sw.)F.Barros
G4;
S1*
18*
Hexalectrisgrandiflora
(A.Rich.
&Galeotti)L.O.
G4;
S22
Williams
Hexalectrisnitid
aL.O.Williams
G3;
S1,S3
End
angered
2Hexalectrisrevoluta
Correll
G1;
S1,SN
R0
Hexalectrisspicata(W
alter)Barnh
art
G5;
SH,S1
-S4,
SNR
End
angered
3HexalectriswarnockiiAmes
&Correll
G2;
S1,S2
0Ionopsisutricularioides(Sw.)Lindl.
G4;
S1End
angered
10Isotriamedeoloides
(Pursh)Raf.
G2;
SX,SH
,S1
,S2
Threatened
End
angered
Endangered
1Isotriaverticillata(M
uehl.ex
Willd.)Raf.
G5;
SX,S1
-S5,
SNR
End
angered
Endangered
5Lepanthopsismelanantha(Rchb.
f.)Ames
G3;
SHEnd
angered
1LiparishawaiensisH.Mann
G3;
S33
Liparisliliifolia
(L.)Rich.
exLindl.
G5;
SX,S1
-S5,
SNR
End
angered
Endangered
4Liparisloeselii(L.)Rich.
G5;
SX,SH
,S1
-S5,
SNR
End
angered
8Liparisnervosa(Thunb.)Lindl.
G4;
S2End
angered*
10MacradenialutescensR.Br.
G4;
SHEnd
angered
0MalaxisabieticolaSalazar&
Soto
Arenas
G4;
S1,S3
*0
MalaxisbayardiiFernald
G1;
SH,S1
,SU
End
angered
0Malaxisbrachystachys(Rchb.
f.)Kun
tze
G4;
S3,S4
*0
Malaxismacrostachya(Lex.)Kun
tze
G4;
SNR,SN
A3*
Malaxismonophyllos(L.)Sw
.G4;
SH,S1
-S4,
SNR*
End
angered*
1Malaxisporphyrea(Ridl.)
Kun
tze
G4;
S2,SN
R0
184 Annals of theMissouri Botanical Garden
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
MalaxisspicataSw
.G4;
S1-S3,
SNR
0Malaxisunifolia
Michx.
G5;
S1-S5,
SX,SN
REnd
angered
2MalaxiswendtiiSalazar
G2;
S10
Mesad
enus
lucayanus(Britton)
Schltr.
G4;
S1,S2
*End
angered
1*Microthelys
rubrocallosa
(B.L.Rob.&
Greenm.)
GNR;S1
0Garay
Neottiaauriculata
(Wiegand)Szlach.
G3;
S1-S3,
SNR*
End
angered
0Neottiabanksiana(Lindl.)Rchb.
f.G4;
S1-S4,
SNR*
4*Neottiabifolia
(Raf.)Baumbach
G4;
S1-S4,
SNR*
End
angered*
5*Neottiaborealis(M
orong)
Szlach.
G4;
SH,S1
-S4,
SNR*
0Neottiaconvallarioides(Sw.)Rich.
G5;
S1-S4,
SNR*
0Neottiacordata(L.)Rich.
G5;
SH,S1
-S5,
SNR*
End
angered*
1Neottiasm
allii
(Wiegand)Szlach.
G4;
S1-S4*
End
angered*
0Oncidium
ensatum
Lindl.
GNR;S1
End
angered*
9*Pelexia
adnata
(Sw.)Poit.ex
Rich.
G5;
S1End
angered
1Peristylusholochila
(Hillebr.)N.Halle
G1;
S1*
End
angered*
3*Piperia
candidaRand.
Morgan&
Ackerman
G3;
S2,S3
,SN
R0
Piperia
colemaniiRand.Morgan&
Glic.
G3;
S30
Piperia
cooperi(S.Watson)
Rydb.
G4;
S30
Piperia
dilatata
(Pursh)Szlach.&
Rutk.
G5;
SH,S1
-S5,
SU,SN
R*
5*Piperia
elegans(Lindl.)Rydb.
G4;
S3,S4
,SN
R1
Piperia
elongata
Rydb.
G4;
S3,S4
,SN
R0
Piperia
leptopetalaRydb.
G3;
S30
Piperia
transversa
Suksd.
G4;
S3,S4
,SN
R0
Piperia
unalascensis(Spreng.)Rydb.
G5;
S1-S5,
SNR
0Piperia
yadoniiRand.
Morgan&
Ackerman
G2;
S2End
angered
0Platanthera
aquilonisSheviak
G5;
SX,S1
-S5,
SU,SN
R3
Platanthera
blephariglottis
(Willd.)Lindl.
G4;
S1-S4,
SNR
End
angered
4Platanthera
brevifolia
(Greene)
Senghas
G4;
SNR
0Platanthera
chapmanii(Small)Luer
G2;
S1,SH
,SN
R1
Platanthera
chorisiana
(Cham.)Rchb.
f.G3;
S2-S4
Threatened
2Platanthera
cilia
ris(L.)Lindl.
G5;
SX,SH
,S1
-S4,
SNR
End
angered
5Platanthera
clavellata
(Michx.)Luer
G5;
SH,S1
-S5,
SU,SN
R2
Platanthera
convallariifolia
(Fisch.ex
Lindl.)Lindl.
0
Volume 99, Number 2 Krupnick et al. 1852013 Status and Future of Orchid Conservation in
North America
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Platanthera
cristata
(Michx.)Lindl.
G5;
S1-S4,
SX,SN
REnd
angered
0Platanthera
flava
(L.)Lindl.
G4;
SX,S1
-S4,
SU,SN
REnd
angered
1Platanthera
grandiflora
(Bigelow
)Lindl.
G5;
SX,S1
-S4,
SNR
End
angered
3Platanthera
hookeri(Torr.)Lindl.
G4;
SX,S1
-S4,
SH,SN
REnd
angered
2Platanthera
huronensisLindl.
G5;
S1-S5,
SU,SN
R*
3Platanthera
hyperborea
(L.)Lindl.
G5;
S4End
angered
5Platanthera
integra(Nutt.)
A.Grayex
L.C.Beck
G3;
S1-S3,
SNR
End
angered
0Platanthera
integrila
bia(Correll)
Luer
G2;
SH,S1
-S3,
SUCandidate
End
angered
1Platanthera
lacera
(Michx.)G.Don
G5;
S1-S5,
SNR
7Platanthera
leucophaea
(Nutt.)
Lindl.
G2;
SX,SH
,S1
,S2
Threatened
End
angered
Endangered
2Platanthera
limosaLindl.
G4;
S4,SN
R0
Platanthera
nivea(Nutt.)
Luer
G5;
SH,S1
-S3,
SNR
End
angered
1Platanthera
obtusata
(Banks
exPursh)Lindl.
G5;
S1-S5,
SNR
Vulnerable
1Platanthera
orbiculata
(Pursh)Lindl.
G5;
SX,SH
,S1
-S4,
SNR
End
angered
1Platanthera
pallida
P.M.Br.
GNA;SU
,SN
A,SN
R*
0Platanthera
peramoena
A.Gray
G5;
SX,S1
-S4,
SNR
End
angered
1Platanthera
praeclaraSheviak&
M.L.Bow
les
G3;
SH,S1
-S3
End
angered
Threatened
End
angered
Endangered
1Platanthera
psycodes
(L.)Lindl.
G5;
SX,SH
,S1
-S5,
SNR
End
angered
2Platanthera
purpurascens
(Rydb.)Sh
eviak&
W.
0F.Jenn.
Platanthera
rotundifolia
(Banks
exPursh)Lindl.
G5;
SX,SH
,S1
-S5,
SNR*
End
angered*
1Platanthera
shriveriP.M.Br.
G1;
S1,SN
R0
Platanthera
sparsiflora
(S.Watson)
Schltr.
G4;
S1,SN
RSensitive
0Platanthera
strictaLindl.
G5;
S1-S5,
SNR
4*Platanthera
tescam
nisSheviak&
W.F.Jenn
.0
Platanthera
tipuloides(L.f.)
Lindl.
G4;
S20
Platanthera
yosemitensisColwell,Sh
eviak&
P.E.
G2;
S20
Moore
Platanthera
zothecina(L.C.Higgins
&S.
L.
G2;
S1,S2
0Welsh)Kartesz
&Gandhi
Platythelys
querceticola(Lindl.)Garay
G4;
S1,SN
R0
Platythelys
sagraeana(A.Rich.)Garay
0Pogonia
ophioglossoides(L.)Ker
Gaw
l.G5;
SX,SU
,S1
-S5,
SNR
End
angered
17Polystachya
concreta
(Jacq.)Garay
&H.R.Sw
eet
G4;
S3End
angered
13Ponthieva
brittoniaeAmes
G3;
S1End
angered
1
186 Annals of theMissouri Botanical Garden
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Ponthieva
racemosa(W
alter)C.Mohr
G4;
S1-S3,
SNR
End
angered
4Prescottia
oligantha(Sw.)Lindl.
G4;
S11
Prosthechea
boothiana(Lindl.)W.E.Higgins
G4;
S1*
6*Prosthechea
cochleata(L.)W.E.Higgins
G4;
S3*
65*
Prosthechea
pygm
aea(Hook.)W.E.Higgins
G4;
S1*
End
angered
9*Pseud
orchisalbida
(L.)A.Love&
D.Love
G5;
S3,SN
R*
3Sa
coila
lanceolata
(Aubl.)
Garay
G4;
S3*
Threatened*
8*Sa
coila
squamulosa(Kunth)Garay
0Schiedeella
arizonicaP.M.Br.
GNR;S3
,S4
,SN
R0
Schiedeella
confusa(Garay)Espejo&
Lopez-Ferr.
G3;
SNR*
0Spiran
thes
brevila
brisLindl.
G1;
S1,SN
REnd
angered
0Spiran
thes
caseiCatlin
g&
Cruise
G4;
S1-S4,
SNR
End
angered
0Spiran
thes
cernua
(L.)Rich.
G5;
S1-S5,
SU,SN
RVulnerable
28Spiran
thes
delitescens
Sheviak
G1;
S1End
angered
End
angered
1Spiran
thes
diluvialisSheviak
G2;
S1,S2
Threatened
End
angered
5Spiran
thes
eatoniiAmes
exP.M.Br.
G3;
SH,S1
-S3,
SNR
0Spiran
thes
infernalisSheviak
G1;
S10
Spiran
thes
lacera
(Raf.)Raf.
G5;
S1-S5,
SNR
Threatened
5Spiran
thes
laciniata(Small)Ames
G4;
S1-S4,
SU,SN
REnd
angered
0Spiran
thes
longila
brisLindl.
G3;
SH,S1
-S3,
SNR
Threatened
0Spiran
thes
lucida
(H.H.Eaton)Ames
G5;
SX,SH
,S1
-S5,
SNR
End
angered
2Spiran
thes
magnicamporum
Sheviak
G4;
SX,S1
-S4,
SNR
End
angered
2Spiran
thes
ochroleuca
(Rydb.)Rydb.
G4;
SH,S1
-S3,
S5,SN
REnd
angered
2Spiran
thes
odorata(Nutt.)
Lindl.
G5;
SH,S1
-S4,
SU,SN
REnd
angered
15Spiran
thes
ovalisLindl.
G5;
SH,S1
-S5,
SNR
End
angered
3Spiran
thes
parksiiCorrell
G3;
S3End
angered
End
angered
2Spiran
thes
porrifolia
Lindl.
G4;
S1,S2
,S4
,SN
RSensitive
3Spiran
thes
praecox(W
alter)S.
Watson
G5;
SH,S1
-S4,
SU1
Spiran
thes
romanzoffianaCham.
G5;
SH,S1
-S5,
SU,SN
REnd
angered
5Spiran
thes
sylvaticaP.M.Br.
GNR;S1
0Spiran
thes
torta(Thunb.)Garay
&H.R.Sw
eet
G4;
S1*
End
angered
0Spiran
thes
tuberosa
Raf.
G5;
SH,S1
-S5,
SNR
End
angered
4Spiran
thes
vernalisEngelm.&
A.Gray
G5;
S1-S5,
SNR
End
angered
5Stelisgelid
a(Lindl.)Pridgeon&
M.W.Chase
G5;
S1*
End
angered*
3Tipularia
discolor
(Pursh)Nutt.
G4;
S1,S3
-S5,
SNR
End
angered
15
Volume 99, Number 2 Krupnick et al. 1872013 Status and Future of Orchid Conservation in
North America
Contin
ued.
1.Table
NatureServe
U.S.
U.S.State
Canada’s
Num
berof
conservation
IUCN
2011
End
angered
conservation
SpeciesAt
botanicgardens
Species
status
rank
saRed
Listb
SpeciesAct
clistin
gdRiskAct
ewhere
presentf
Tolum
niabahamensis(Nashex
Britton&
Millsp.)
G3;
S1End
angered
5Braem
Trichocentrum
undulatum
(Sw.)Ackerman
&M.
G4;
S1*
End
angered*
1W.Chase
Triphoraam
azonicaSchltr.
End
angered
0TriphoracraigheadiiLuer
G1;
S1End
angered
0Triphoragentianoides
(Sw.)Nutt.ex
Ames
&G4;
SNR
0Schltr.
Triphoratrianthophora(Sw.)Rydb.
G3;
SH,S1
-S3,
SNR
End
angered
Endangered
0TriphorayucatanensisAmes
G1;
S10
Tropidiapolystachya(Sw.)Ames
G4;
S1End
angered
1Van
illabarbellata
Rchb.
f.G4;
S2End
angered
7Van
illadillonianaCorrell
G3;
SNR
End
angered
5Van
illainodoraSchiede
1Van
illamexicanaMill.
G3;
S1End
angered
2Van
illaphaeanthaRchb.
f.G4;
S1End
angered
8
*Listedin
original
publicationun
derasynonymousname.
aConservationassessmentsby
NatureServe
correspond
toGlobalthreats:G
1¼CriticallyIm
periled,
G2¼Im
periled,
G3¼Vulnerable,G4¼App
arently
Secure,G
5¼Secure,G
U¼
Unrankable,GNR¼NotYetRanked,
GNA¼NotApp
licable.S
ubnationalthreatsfollowsimilarvalues
(S1toS5
,etc.),
butapp
lyatthestateor
provinciallevels.N
atureServe
rank
ings
areavailablethrough
,http://www.natureserve.org/explorer.
.bInternationalUnion
forConservationof
NatureandNatural
Resources
(IUCN),20
11.
cEnd
angeredSp
eciesAct
(ESA
),19
73;U.S.Governm
entPrintingOffice,20
12.
dU.S.Statelistin
gisat
thehighestlevelof
protectionam
ongallstates
listed,
asreported
bytheUSD
APLANTSDatabase(USD
A,NRCS,
2012
).eSp
eciesat
RiskAct
(SARA),20
02.
fThe
numberof
botanicgardenswhere
theorchid
speciesisconservedor
present,accordingto
Botanical
Gardens
ConservationInternational(201
2).
188 Annals of theMissouri Botanical Garden
state and provincial species numbers are according tothe NatureServe Explorer (NatureServe, 2011). Threespecies are endemic to California, as Piperiacolemanii Rand. Morgan & Glic., P. yadonii Rand.Morgan & Ackerman, and Platanthera yosemitensis,and three to Hawaii, as Anoectochilus sandvicensisLindl., Liparis hawaiensis H. Mann, and Peristylusholochila (Hillebr.) N. Halle. Two orchid species areendemic to Florida, Govenia floridana P. M. Br. andTriphora craigheadii Luer; one species is endemic,each to three states, with Spiranthes delitescensSheviak in Arizona, S. infernalis Sheviak in Nevada,and S. parksii Correll in Texas.
TARGET 2, AN ASSESSMENT OF THE CONSERVATION STATUS OF
ALL KNOWN PLANT SPECIES, AS FAR AS POSSIBLE, TO GUIDE
CONSERVATION ACTION
Only two North American orchids have beenassessed by the International Union for Conservationof Nature and Natural Resources (IUCN) and arelisted as threatened on the 2011 IUCN Red List ofThreatened Species (IUCN, 2011): Anoectochilussandvicensis as Vulnerable (or VU), and Platantherapraeclara Sheviak & M. L. Bowles, as Endangered (orEN).The U.S. Endangered Species Act (ESA; 1973;
U.S. Government Printing Office, 2012) federally liststhe four endangered species Piperia yadonii, Peri-stylus holochila [[ Platanthera holochila (Hillebr.)Kraenzl.], Spiranthes delitescens, and S. parksii, aswell as the four threatened species as Isotriamedeoloides, Platanthera leucophaea (Nutt.) Lindl.,P. praeclara, and S. diluvialis Sheviak, and the onecandidate species, Platanthera integrilabia (Correll)Luer. At the state level, 57% (119 species) areprotected as endangered, threatened, vulnerable, orsensitive in at least one state (cf. Table 1).The Canadian Species at Risk Act (SARA; 2002)
lists seven endangered orchid species, as Cypripedi-
Table 2. Species distribution among listed orchid genera inTable 1.
Percent ofGenus Species # total species
Platanthera Rich. 35 16.7%
Spiranthes Rich. 23 11.0%
Cypripedium L. 12 5.7%
Piperia Rydb. 10 4.8%
Malaxis Sol. ex Sw. 9 4.3%
Epidendrum L. 8 3.8%
Corallorhiza Gagnebin 7 3.3%
Neottia Guett. 7 3.3%
Calopogon R. Br. 5 2.4%
Hexalectris Raf. 5 2.4%
Triphora Nutt. 5 2.4%
Vanilla Mill. 5 2.4%
Goodyera R. Br. 4 1.9%
Habenaria Willd. 4 1.9%
Liparis Rich. 4 1.9%
Cleistesiopsis Pansarin & F. Barros 3 1.4%
Prosthechea Knowles & Westc. 3 1.4%
Bletia Ruiz & Pav. 2 1%
Cyclopogon C. Presl 2 1%
Dactylorhiza Neck. ex Nevski 2 1%
Dendrophylax Rchb. f. 2 1%
Dichromanthus Garay 2 1%
Eulophia R. Br. ex Lindl. 2 1%
Galeandra Lindl. 2 1%
Isotria Raf. 2 1%
Platythelys Garay 2 1%
Ponthieva R. Br. 2 1%
Sacoila Raf. 2 1%
Schiedeella Schltr. 2 1%
Anoectochilus Blume 1 0.5%
Aplectrum Blume 1 0.5%
Arethusa L. 1 0.5%
Basiphyllaea Schltr. 1 0.5%
Beloglottis Schltr. 1 0.5%
Brassia R. Br. 1 0.5%
Bulbophyllum Thouars 1 0.5%
Calypso Salisb. 1 0.5%
Camaridium Lindl. 1 0.5%
Campylocentrum Benth. 1 0.5%
Cephalanthera Rich. 1 0.5%
Cranichis Sw. 1 0.5%
Cyrtopodium R. Br. 1 0.5%
Eltroplectris Raf. 1 0.5%
Encyclia Hook. 1 0.5%
Epipactis Zinn 1 0.5%
Galearis Raf. 1 0.5%
Govenia Lindl. 1 0.5%
Hammarbya Kuntze 1 0.5%
Heterotaxis Lindl. 1 0.5%
Ionopsis Kunth 1 0.5%
Lepanthopsis Ames 1 0.5%
Macradenia R. Br. 1 0.5%
Mesadenus Schltr. 1 0.5%
Microthelys Garay 1 0.5%
Oncidium Sw. 1 0.5%
Pelexia Poit. ex Lindl. 1 0.5%
Table 2. Continued.
Percent ofGenus Species # total species
Peristylus Blume 1 0.5%
Pogonia Juss. 1 0.5%
Polystachya Hook. 1 0.5%
Prescottia Lindl. 1 0.5%
Pseudorchis Seg. 1 0.5%
Stelis Sw. 1 0.5%
Tipularia Nutt. 1 0.5%
Tolumnia Raf. 1 0.5%
Trichocentrum Poepp. & Endl. 1 0.5%
Tropidia Lindl. 1 0.5%
Volume 99, Number 2 Krupnick et al. 1892013 Status and Future of Orchid Conservation in
North America
190 Annals of theMissouri Botanical Garden
um candidum Muhl. ex Willd., Isotria medeoloides, I. specialized equipment (Liu et al., 2010; Seaton et al.,verticillata (Muhl. ex Willd.) Raf., Liparis liliifolia 2010; Stewart & Hicks, 2010). However problematic,(L.) Rich. ex Lindl., Platanthera leucophaea, P. this is essential for propagation and establishment ofpraeclara, and Triphora trianthophora (Sw.) Rydb. self-sustaining populations. Few organizations haveFurther, one threatened species, Cephalanthera the capacity to handle these unique aspects of orchidaustiniae (A. Gray) A. Heller, is included, as well biology, placing cultivation and reintroduction beyondas one species that is noted of special concern for the abilities of nearly all conservation agencies.conservation, Epipactis gigantea Douglas ex Hook. Scientific research has made substantial progress in(cf. Table 1). overcoming these difficult aspects of orchid ecology,All but 10 species of the 210 orchid names but additional efforts are needed on all key elements of
included in Table 1 have been assessed by Nature- orchid life histories that must be understood if we areServe (2011), with 24% (50 species) listed as globally to successfully support conservation, reintroduction,threatened. Of these 50 threatened species, 11 were and propagation efforts for native orchids. Theassessed at the global scale as critically imperiled techniques being developed by scientists are still,(G1), 13 are imperiled (G2), and 26 are vulnerable and likely will remain, beyond the capacity of most(G3; cf. Table 1). For subnational assessments by conservation agencies, and there exists no currentNatureServe, 14% (30 species) are presumed to be network for scientific researchers either to supportextirpated from at least one state or province, and an conservation program managers or to communicateoverlapping 23% (48 species) are possibly extirpated with commercial or private growers and gardenfrom at least one state or province. Also at the enthusiasts who would benefit from a more completesubnational level, 84% (176 species) are variably understanding of all aspects of orchid growth,threatened (ranging from S1 or critically imperiled, cultivation, and conservation.S2 or imperiled, or S3 or vulnerable) in at least onestate or province. In all cases, where an orchid had
TARGET 7, AT LEAST 75% OF KNOWN THREATENED PLANTbeen extirpated in one or more states, it was also
SPECIES CONSERVED IN SITUthreatened or endangered in at least one other state.
When considering the proportion of threatenedTARGET 3, INFORMATION, RESEARCH AND ASSOCIATED orchids that are conserved in situ, it is important toOUTPUTS, AND METHODS NECESSARY TO IMPLEMENT THE distinguish between species that are consideredSTRATEGY DEVELOPED AND SHARED globally threatened and the majority of orchids that
are threatened within a portion of their ranges.As indicated by Stewart (2008), much of theConserved in situ means ‘‘that biologically viableinformation on conservation and reintroduction ofpopulations of these species occur in at least oneorchids has been published in the ‘‘gray’’ literatureprotected area or the species is effectively managedand obtained from unreplicated efforts that are rarelyoutside the protected area network, e.g., as part of adesigned as scientific studies to obtain statisticallymanagement plan’’ according to a recent GSPC Plantsignificant data. Academic research occasionallyConservation Report (Convention on Biologicaladdresses aspects of orchid biology, but rarelyDiversity [CBD], 2009: 23–24). As previouslyincludes the whole process from basic biology tooutlined (cf. Table 1), of the 50 globally threatenedapplication for conservation or reintroduction (e.g.,(i.e., ranked G1, G2, or G3 by NatureServe) orchidKindlmann et al., 2002; Dixon et al., 2003).species, only eight are protected under the ESA andReplicated assessment of conservation is sorelyone additional species, Triphora trianthophora, underlacking. There are three major areas of orchid biology
that urgently need additional research: (1) identifi- Canada’s SARA. An additional 22 species are
cation of fungi associated with nearly all orchids, (2) protected at the state or province level. Taken
understanding of how those fungi contribute to seed together, a total of 62% (31 of 50 species, cf. Table
germination in situ and in vitro, and (3) how to 1) of native orchid species assessed as threatened are
maximize survival of cultured seedlings or plants. thus conserved in situ (i.e., have legal protection),
Seed banking alone cannot successfully preserve which is far below the goal of Target 7. It isorchids, because using the seeds to eventually noteworthy that six of the 10 globally imperiled orcultivate and restore plants in nature requires that critically imperiled species (G1 or G2) that currentlyappropriate mycorrhizal fungi are present, especially at have no protection at the federal or state level havethe orchid’s protocorm stage. Identifying, maintaining, only recently been described, suggesting that accu-and establishing the symbiotic fungi needed for orchid rate identification and species delimitation haveseed germination is technically difficult and requires hampered attempts at species protection.
Volume 99, Number 2 Krupnick et al. 1912013 Status and Future of Orchid Conservation in
North America
Beyond recognition of the need to protect orchid preserve. The reserve’s 17,424 acres are owned andtaxa is the degree to which protection is actually managed by The Nature Conservancy, and itsaccomplished. The level of protection provided by longleaf pine savannas must be periodically subject-state or federal protected status depends on both the ed to prescribed burns to keep the habitat prime fornumber and distribution of plants on protected land the smaller, herbaceous species that would beand also on the knowledge of the species’ biology otherwise crowded out by succession.needed to determine whether protection is adequate. In contrast, other woodland species, such asA number of protected reserves in the United States Cypripedium acaule Aiton, C. fasciculatum Kelloggand Canada focus to some degree on orchids or on ex S. Watson, C. montanum Douglas ex Lindl., andhabitats with disproportionately many orchids. One C. reginae Walter, have ecological strategies andsuch example is the Bruce Peninsula National Park dependencies based both on edaphic constancy andin Ontario, which supports 43 of the orchid species occasional disturbance of their environments. Fornative to Canada that will be threatened as a result of example, even though C. fasciculatum and C.climate change (Suffling & Scott, 2002). The heart of montanum grow sympatrically in old growth forest,the Niagara Escarpment Biosphere Reserve includes there is evidence that C. montanum needs occasionalthe Bruce Peninsula National Park, a Nature disturbances, such as fire or tree-felling and thinning,Conservancy Preserve, and First Nations lands, all to create open, sunnier areas in which they will bloomof which share knowledge about species at risk. The and set seed more freely. Its sympatric relative, C.stated park goal is to maintain viable populations of fasciculatum, conversely seems to be inhibited byall native species in situ, and there is a program burns and other such disturbances. Therefore, itunderway to report on the condition of all SARA becomes increasingly important to investigate thespecies, including trends in populations and the individual ecological complexities of each individualfactors that contribute to their condition. species if comprehensive management plans are to beAttributing population trends to particular factors created.
highlights the importance of in-depth understanding Understanding what factors influence populationof species biology for accomplishing effective con- dynamics is critical for understanding how speciesservation. Most endangered orchids in North America will respond to warming global temperatures. Withhave had at least some investigation of population orchids, this uncertainty may be compounded bygenetic structure, and this has been used to considering the other species on which they depend.understand connectivity between populations and to Some orchids may be limited by the availability ofdetermine the contribution of outlying populations to pollinators. For example, the deceptive orchidspecies integrity. For example, Wallace (2003) found Cyrtopodium punctatum (L.) Lindl. depends on oil-that Platanthera leucophaea was a predominantly gathering Centris bees for pollination. These bees relyoutcrossing species and that inbreeding depression, on other flowering plants, especially Byrsonimaespecially in small populations, suppressed seed lucida (Sw.) DC. (Malpighiaceae), for the oils theyviability. These results suggested that larger, more collect. This led Pemberton and Liu (2008) to suggestdiverse and outcrossing populations were needed to that B. lucida, which is almost completely absentsupport population genetic variability in the current from the areas where the few remaining native orchidfragmented landscape. This white-fringed orchid populations of C. punctatum persist, be planted in thehistorically was distributed from Missouri and Iowa vicinity of orchids to attract and support the bees thatto Ontario with disjunct populations in Maine, New pollinate both taxa. Such activities may also beJersey, and Virginia (Bowles et al., 2005). The needed to support the interaction between orchidscurrent distribution is, however, much reduced and and their pollinators in the face of climate change. Infew populations are self-sustaining. particular, Liu et al. (2010) suggested that warmingAlso important to consider are critical issues of temperatures in southwestern China are differentially
land management. Many terrestrial orchids are affecting pollinator and orchid phenologies such thatpioneer species and cannot compete with overgrown availability of pollinators and orchid flowering may behabitat. One prime example is the Green Swamp increasingly out of synchronization. Effects of climatePreserve of southeastern North Carolina in Brunswick change on orchid mycorrhizal fungi are completelyand Columbus counties. Originally established to unknown, reflecting that very little is known aboutprotect the habitat of the Venus fly trap (Dionaea what factors drive the distribution and abundance ofmuscipula J. Ellis, Droseraceae), a great variety of nearly all fungi. Because of this, conservation of fungiorchids, including species of Calopogon R. Br., must be largely accomplished through habitatPlatanthera, and Cleistes Rich. ex Lindl., exist in the conservation.
192 Annals of theMissouri Botanical Garden
TARGET 8, AT LEAST 75% OF THREATENED PLANT SPECIES IN growth. While fungal requirements can sometimesEX SITU COLLECTIONS, PREFERABLY IN THE COUNTRY OF be overcome under specialized conditions in theORIGIN, AND AT LEAST 20% AVAILABLE FOR RECOVERY AND laboratory, there is little question that under naturalRESTORATION PROGRAMS conditions fungi are required for orchid recruitment
and long-term survival of populations. Many orchidsThe Botanical Gardens Conservation Internation-
require specific fungi at all life history stages (e.g.,al’s (BGCI; 2012) Plant Search database shows that
McCormick et al., 2004), while other more generalist66% (139 species) of native North American orchids
orchids use fungi from one or a few families (e.g.,are found in ex situ collections in botanical
Shefferson et al., 2010). As a result, effective ex situinstitutions around the world (Table 1). Each of
conservation of orchids will require not just seedthese species is reported from an average of sixbanking, but also maintenance of required mycor-botanic gardens, but more than 25% (36 species) arerhizal fungi. This goal is still far from beingreported from a single botanic garden only. Theaccomplished. One of the central difficulties hasspecies most prevalent is Prosthechea cochleata (L.)been to simply identify the needed fungi. MostW. E. Higgins [[ Encyclia cochleata (L.) Dressler], aorchid mycorrhizal fungi rarely produce spores andspecies easily propagated (Pugh-Jones, 2009), whichso were known only from the anamorphic stage foris reported from 65 botanic gardens. Only two of 11many years. Now they can be identified largely byspecies ranked G1 by NatureServe are found in anyDNA sequencing. This rarity of spore production,botanic garden collection, being Peristylus holochilacoupled with inconspicuous and morphologicallyat three gardens and Spiranthes delitescens at onedepauperate sporulating bodies, has resulted in ainstitution. Five of 13 species ranked G2 arepoorly defined taxonomy of these cryptic fungi (e.g.,represented in botanic gardens, but only two areSwarts & Dixon, 2009); DNA sequences rarelyreported from more than one garden (two and fivematch the taxonomically described species. Fur-gardens, Platanthera leucophaea and S. diluvialis,thermore, with few spores produced, the fungi thatrespectively). It is highly unlikely that any of these exan orchid needs can rarely be stored as spores andsitu collections represents a genetically representa-so must be maintained as active cultures. However,tive sample of the species.some researchers have been testing methods forThe major method likely to preserve speciesstoring fungal cultures in liquid nitrogen (Batty etgenetic variation is seed banking, as storing manyal., 2001). If these methods prove successful, thengenetically distinct orchid seeds requires very littleconservation of genetically representative collec-space. Accomplishing such genetically representativetions of mycorrhizal fungi may become moreconservation is a major goal of the project Orchidcommon.Seed Stores for Sustainable Use (OSSSU; Seaton et al.,
2010). This project is initially focusing on hotspots of As yet there are relatively few orchid mycorrhizal
orchid diversity but is also beginning to work with fungi in culture. The University of Alberta Mycolog-
groups within North America to organize locations and ical Herbarium’s (UAMH; ,http://www.uamh.
organizations for genetically representative orchid devonian.ualberta.ca.) culture collection currently
seed storage. The Center for Plant Conservation maintains 113 fungal cultures obtained from 37
(CPC; ,http://www.centerforplantconservation.org.) orchid species. The Smithsonian Environmental
maintains seed collections for eight orchid species, Research Center (SERC) currently maintains more
two of which are ranked G1 (Peristylus holochila and than 400 fungal isolates in culture from more than 40
Spiranthes delitescens), three G2 (Isotria medeoloides, native orchid species (D. F. Whigham, M. K.
Platanthera leucophaea, and S. diluvialis), and three McCormick & J. P. O’Neill, pers. comm.). TheG3 (Cypripedium kentuckiense C. F. Reed, Platan- American Type Culture Collection (ATCC) maintainsthera praeclara, and S. parksii). These species are 30 isolates from 13 orchid species (,http://www.atcc.specifically directed to be available for conservation org.). Other scattered fungal cultures exist inand restoration activities, so this would suggest that laboratories around the country where orchid re-seeds of 16% (eight of 50) of threatened North search is conducted, but the maintenance of theseAmerican orchid species are available for conserva- collections is often uncertain. Most of the existingtion and restoration, though the extent to which their collections focus on saprotrophic Tulasnella J.collections are genetically representative is unclear. Schrot. and Ceratobasidium D. P. Rogers fungi, whileOne shortcoming of seed storage for orchids is many orchids rely on fungi that form ectomycorrhizal
that all orchids are dependent on or benefit from associations with other plants. These fungi are oftenmycorrhizal fungi for seed germination and all difficult to culture and many cannot be grown withoutspecies are dependent on fungi for protocorm a photosynthetic host, often a tree, making them very
Volume 99, Number 2 Krupnick et al. 1932013 Status and Future of Orchid Conservation in
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difficult to establish or maintain in culture. Even or certificates to import, export, or re-export orchidwhen fungi are available in culture, however, their species across international lines. However, therole in facilitating seed germination is far from smuggling of wild orchids remains a problemcertain. (Phelps et al., 2010).Germination requirements for seven of 11 G1-
ranked and five of 13 G2-ranked orchids have yet to TARGET 14, THE IMPORTANCE OF PLANT DIVERSITY AND THE
be studied (Stewart & Hicks, 2010). One threatened NEED FOR ITS CONSERVATION INCORPORATED INTO
orchid (Isotria medeoloides) has so far proven COMMUNICATION, EDUCATION, AND PUBLIC AWARENESScompletely recalcitrant in culture or in the field. Of PROGRAMthe 11 G1- and G2-ranked orchids for which somelevel of germination success has been obtained, two As environmental awareness matures and enters
have had only limited asymbiotic success and only the digital age, several organizations previously
five have been germinated symbiotically. Symbiotic limited to spreading their messages slowly thoughgermination has not been reported for the other six of periodicals and mailings have found much broaderthese 11 species. Symbiotic germination is expected and enthusiastic younger audiences. One example isto be the method that leads to seedlings best able to the Native Orchid Conference (,http://tech.groups.survive reintroduction and also provides a method for yahoo.com/group/nativeorchidconference/.) that hasco-introducing needed fungi and orchids (e.g., become the ‘‘go to’’ place for information about nativeStewart, 2008). orchid species and access to experts, photographs,Few well-documented orchid reintroductions exist. ecology, phenology, and field information to any
Often reintroductions occur in one location at one interested party. It is particularly useful to the public,time and either succeed or fail without providing who may not otherwise have access to or might beinformation about what may have been suboptimal or intimidated by scientific publications.may increase success in the future (Stewart, 2008). Botanical institutions nationwide, such as theTwo examples of reintroductions designed as studies Smithsonian Institution and the U.S. Botanic Gardenare outlined by Stewart (2008). Elements of another (USBG), provide a type of outreach through conser-reintroduction are described by Zettler and Piskin vation messages that regularly appear in exhibits,(2011). All three studies were designed to assess particularly annual orchid exhibits, and are viewedeffectiveness of introduction into different habitats, by their many visitors. In addition, speakers frombut only one utilized symbiotic seedlings because these institutions regularly travel around the countryfungi were not available for the other two species. In discussing conservation values to likely interestedtwo of the three studies, survival was highest in sites parties at orchid societies and special events acrossthat already had the target orchid species. In the third
North America. These efforts, however, are notstudy, reintroduction was only attempted in sites with
sufficient. Considerably more could, and should, bethe target species so existing plants could act as
done to raise public awareness about the importancesources for mycorrhizal colonization of the trans-
of North American native orchid species.plants. This suggests that mycorrhizal colonizationmay be critical for reintroduction success. No
15,mention of pollinator availability was made in these TARGET THE NUMBER OF TRAINED PEOPLE WORKING WITH
three studies. APPROPRIATE FACILITIES SUFFICIENT ACCORDING TO NATIONAL
NEEDS, TO ACHIEVE THE TARGETS OF THIS STRATEGY
TARGET 11, NO SPECIES OF WILD FLORA ENDANGERED BY Without a centralized organization focusing onINTERNATIONAL TRADE North American orchids, it is difficult to assess how
many trained individuals work on the in situ and exThe United States and Canada are both parties tositu conservation of orchid species. Kramer et al.The Convention on International Trade in Endan-
gered Species of Wild Fauna and Flora (CITES; (2010) report on a U.S. survey that revealed a major
,http://www.cites.org.), the lead coordinating decline in botanical courses and degree programs at
agency for the implementation, monitoring, and universities and colleges nationwide, as well as a
review of Target 11. All orchid species are listed in deficiency of botanists at U.S. government agencies.
CITES appendix II, thus preventing the endanger- With fewer college graduates entering the botanicalment of North American orchids by over-exploitation workforce and many government botanists retiring incaused by international trade. Artificially propagat- the coming years, it will be difficult to increase theed plants, hybrids, plant parts, products, or number of trained people working on orchidderivatives, with a few exceptions, require permits taxonomy and conservation.
TARGET 16, INSTITUTIONS, NETWORKS, AND PARTNERSHIPS propagation, and the maintenance of evolutionaryFOR PLANT CONSERVATION ESTABLISHED OR STRENGTHENED processes influencing population distribution pat-AT NATIONAL, REGIONAL, AND INTERNATIONAL LEVELS TO terns. In order to do this, conservation must combineACHIEVE THE TARGETS OF THIS STRATEGY detailed experimentation directed at continued
survival of the species both in situ and ex situThe CPC is a successful network of botanic
(Ramsay & Dixon, 2003). The integrated conserva-institutions dedicated to preventing the extinction of
tion strategy emphasizes the study of interactionsU.S. native plants by ensuring that ex situ material isamong land conservation, biological management,available for restoration and recovery efforts. Orchids,research and propagation and reintroduction andhowever, are unique in that effective ex situhabitat restoration (Hopper, 1997).conservation of orchids requires not just seed
banking, but also maintenance of cultures of required1mycorrhizal fungi. A network of partnerships focusing CASE
exclusively on the needs of orchids is necessary. Platanthera leucophaea (Fig. 1A) is currentlylisted as federally threatened, under the ESA, and
THREE CASE STUDIES has declined in the United States by more than 70%from estimates provided by original county records.As described throughout this manuscript, effectiveThis decline has mainly been due to habitat loss.conservation of any orchid species is a complexMost of the remaining 79 populations are small, theseundertaking that includes a wide range of variableswith fewer than 50 plants, and only 28 havefrom long-term storage and maintenance of collec- %adequate protection and management (U.S. Fish andtions of materials (e.g., seeds and mycorrhizal fungi)
that would be required for cultivation and propaga- Wildlife Service, 2007). Investigation of the orchid’s
tion to the establishment and maintenance of habitats genetics indicates that the species is primarily
that support the long-term success of orchid outcrossing and demonstrates significant inbreeding
populations. For many of the reasons described depression particularly prevalent in small popula-
above, we are unaware of any effort that has been tions (Wallace, 2003). Despite the low number of
fully successful in assuring the long-term survival of protected populations, the outlook for conservation of
any native orchid species. There are, however, a few P. leucophaea is relatively good. Well-coordinatedexamples of the efforts that are necessary to build an efforts to recover this species are currently in placeinformation base, which can support further attempts and involve a network of scientists, private landown-to conserve all native orchid species. ers, and volunteers (Zettler & Piskin, 2011). FungiEffective orchid conservation must integrate the needed by P. leucophaea are known, and symbiotic
understanding of existing and future environmental germination is regularly accomplished in laboratorythreats, taxonomic distinctiveness, numbers of indi- cultures, although successful transplantation ofviduals in populations, reproductive biology, ex situ seedlings into natural populations has met with
Figure 1. Three endangered orchid species. —A. Platanthera leucophaea (Nutt.) Lindl. (photo by Timothy Bell). —B. Isotriamedeoloides (Pursh) Raf. (photo by Melissa McCormick). —C. Tolumnia bahamensis (Nash) Braem. (photo by Matt Richards).
194 Annals of theMissouri Botanical Garden
Volume 99, Number 2 Krupnick et al. 1952013 Status and Future of Orchid Conservation in
North America
limited success (Zettler et al., 2005; Zettler & Piskin, population persistence and also the founding of new2011). Genetic surveys have found that even small populations. This suggests that this species, many ofpopulations of P. leucophaea may retain relatively whose extant populations are relatively well protect-high levels of genetic diversity (Holsinger & Wallace, ed, is sufficiently poorly understood and that its2004), potentially resulting in significant seed maintenance in the face of a changing climate is aproduction and recruitment where habitat is avail- serious concern.able. This level of information about the species andthe network of public and private agencies cooper- CASE 3ating toward the orchid’s conservation suggest that P.
An additional case study involved returning aleucophaea has a strong potential for recovery.semiepiphytic species, Tolumnia bahamensis (Nashex Britton & Millsp.) Braem (Fig. 1C), to reasonably
CASE 2pristine habitat, where ostensibly all the other pieces
In contrast to the favorable outlook for Platanthera of the ecological puzzle remain. In this case, suitableleucophaea, Isotria medeoloides (Fig. 1B) is listed by unspoiled, historical habitat in Jonathan DickinsonNatureServe as imperiled (G2) in 14 (78%) of the 18 State Park in southern Florida was assessed and thestates and provinces in which it is still known to few extant plants of T. bahamensis were cross-occur; the orchid is thought to be historical or pollinated and grown ex situ at Atlanta Botanicalextirpated in five states. Nowhere across its distribu- Garden. With most of the host plants and presumablytional range in eastern North America is I. mycorrhizal fungi intact in the orchid’s preferredmedeoloides considered secure or common. The environment, reintroduction has initially been suc-primary threat to its existence is destruction of its cessful. These types of enrichment reintroductionswoodland habitat for development or forestry. The should be attempted whenever suitable protectedmajority of its populations number fewer than 25 habitat is available. This underscores the need for aplants and are thus vulnerable to local extinction holistic approach to orchid conservation in which(U.S. Fish and Wildlife Service, 1994). Many of the entire habitats and ecosystems are sought to beextant populations, including some of the largest preserved whenever possible (Jonathan Dickinsonpopulations, occur on land protected by federal or State Park, 2011).state agencies or the U.S. military, and on this basisthe plant might be considered well protected (U.S. THE FUTURE OF ORCHID CONSERVATIONFish and Wildlife Service, 1994). Its preferredhabitat conditions have also been identified (Sperdu- International efforts such as CITES have focused
to & Congalton, 1996), and searching these habitat on the illegal trade of orchids and many organizations
types allowed researchers to locate many new have been established to cultivate, market, and enjoy
populations in the late 1990s. However, the plant’s orchids, but there is no one national organization that
biology throughout much of its range is defined by focuses on their conservation and restoration. Neither
many small, ephemeral populations that make it quite is there one entity devoted to educating the public
difficult to target areas to protect. Additionally, long about the evolutionary and ecological importance of
periods of dormancy, common in many terrestrial orchids. Organizations (e.g., federal agencies and theorchids, are characteristic of the lifecycle of this U.S. military) that are mandated to identify andspecies, making it difficult to assess population sizes protect orchids on public lands have been involved inor even plant presence (Mehrhoff, 1989). research on relatively few species (see examplesOngoing management experiments of Isotria me- above), and they rely mostly on habitat conservation
deoloides are beginning to reveal a management for management. While habitat management istechnique, tree thinning, which can benefit local important, ecological attributes of orchids (e.g., thepopulations (e.g., Brumback et al., 2011). The fungi obligatory relationships between orchids and fungi atneeded by this orchid have recently been identified critical life history stages) dictate that habitat(M. K. McCormick, unpubl. data), but they have so management alone will not result in orchid conser-far been resistant to culture in the laboratory. Seed vation or restoration. Every U.S. state lists at least onegermination, either symbiotic or asymbiotic, has orchid species that is rare or threatened and mostnever been accomplished either in the field or in states list multiple orchid species. There is, however,the laboratory, yet based on population demographic little coordination among states and no one organi-studies (Mehrhoff, 1989) and preliminary genetic zation that can provide answers to basic questionsanalyses (M. K. McCormick, unpubl. data), recruit- that would guide effective management plans. Privatement from seed is critically important to both land-management conservation groups (e.g., The
196 Annals of theMissouri Botanical Garden
Nature Conservancy) face a similar dilemma. Perhaps roles. NAOCC administration and research are basedmost important, the public has little recognition of the at the SERC, which provides research services todiversity and importance of orchids and there is no private and public organizations, and collaboratescentral organization that focuses on orchids as an with the CPC and other affiliated organizations (e.g.,important aspect of education and outreach to the Kew Gardens and the U.S. Bureau of Landpublic. Management’s Seeds of Success program) to developWe propose a possible solution to the lack of a genetically diverse seed bank for all native orchids.
coordination and the pooling of resources to focus on The seed collection will not only be used to assurethe more than 200 native orchids listed within the the long-term survival of the germplasm of eachUnited States and Canada. The NAOCC (http:// species, but will also serve as a resource for materialnorthamericanorchidcenter.org) is the first interna- to support efforts to grow and cultivate all nativetionally focused public–private effort to support the orchids. SERC is collaborating with the partners ofconservation, cultivation, and restoration of native NAOCC to expand its collection of orchid mycorrhi-orchid species. NAOCC began as a collaborative zal fungi to include fungi from all native orchids.effort between the SERC, Smithsonian Gardens, SERC is also playing a lead role in developing aDepartment of Botany at the National Museum of network of laboratories that provides services for theNatural History (NMNH), Exhibits and Park Man- molecular analysis of orchid fungi. The Smithsonian’sagement Department of the National Zoological Park Department of Botany at the NMNH will focus on the(NZP), and USBG. Other government agencies, development of a well-curated and complete herbar-botanic gardens, and public and private landowners ium-based orchid collection and will develop DNAare joining the collaboration. NAOCC launched in barcodes for all North American orchids. NMNH will
2012 and the network that will support the effort will also develop a digital library of all North American
be developed over approximately 10 years. orchids, including visual images of all species, and
The NAOCC’s mission is to conserve the native will actively partner with SERC and NAOCC to
orchid heritage of the United States and Canada develop web-based technologies to provide up-to-date
through preservation, restoration, and cultivation of public access to orchid information. Smithsonian
native orchids and to convey the importance of Gardens, the Exhibits and Park Management De-
NAOCC to the public through innovative educational partment of the NZP, and the USBG will coordinate
programs. The goals of NAOCC are to: efforts to cultivate all orchids within the Washingtonecoregion into their living collections, and they will
Develop an international seed bank collection, in collaborate with partner-gardens to develop and putcollaboration with the CPC, that will be represen- into effect a plan to cultivate all 210 native orchids intative of the genetic diversity of all North a range of gardens across the United States andAmerican orchid species. Canada. The Smithsonian and the USBG will alsoDevelop an international collection of fungi include exhibits about native orchids in theirrepresentative of the genetic diversity of mycor- biannual orchid show.rhizal fungi required by native orchids.Develop techniques to conserve the genetic CONCLUSION
diversity of all native orchids by cultivating themNative orchids occur in every state in the Unitedin an international network of botanic gardens and
States and every Canadian province, and one or morearboretums.species is listed as endangered or threatened in everyUse seed and mycorrhizal fungal banks to developstate and province. As described above, national andtechniques for restoring, conserving, cultivating,international efforts have provided a degree ofand restoring orchids in native habitats.protection for native orchids, and there have beenSupport efforts to conserve orchid populationsefforts to conserve and restore a small number ofthrough habitat conservation and restoration.species. It is our view that progress toward theDevelop web-based materials that will provide up-effective conservation of the numerous species thatto-date information on the ecology, conservationare listed as threatened or endangered will require astatus, and techniques for the cultivation of Northlarge-scale integrated effort to develop the knowledgeAmerican orchids.base required to develop effective management
An initial goal of the network of botanic gardens is strategies to assure the survival of the more thanto grow and display all native orchids in the United 200 species of native orchids. In establishing theStates and Canada using an ecoregional approach. NAOCC, our goal is to develop the resource base andThe primary partners play different yet integrated integration of public and private organizations
Volume 99, Number 2 Krupnick et al. 1972013 Status and Future of Orchid Conservation in
North America
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