Evidence of artificial selection: are orchids in cultivation an effective ex situ conservation strategy?

Resumen: La selección artificial es el proceso por el cual los humanos cambian las características morfológicas y genéticas de las especies a través de la selección de caracteres ''favorecidos''. Aunque la literatura gris sugiere que el proceso está ocurriendo en orquídeas mantenidas en colecciones ex situ, existe evidencia limitada de esto en la literatura científica. Esta presente una perspectiva común entre los cultivadores de que las especies (no los híbridos) que se mantienen en las colecciones ex situ son fuentes potenciales de material para usar en el restablecimiento in situ, sin embargo, esto supone que no ha habido ninguna selección artificial de caracteres morfológicos, o rasgos funcionales mientras crecían y se propagaban ex situ. Aquí evaluamos si las plantas cultivadas en colecciones ex situ muestran cambios en las características morfológicas a lo largo del tiempo y si el rango de tamaño de los fenotipos está dentro del rango de las poblaciones in situ. Evaluamos plantas de la base de datos de la American Orchid Society de nueve géneros y 54 especies. Notamos que el 35% de los caracteres evaluados tenían evidencia de cambios significativos a lo largo del tiempo. Además, para la mayoría de las características evaluadas de las especies ex situ estaban frecuentemente (95%) fuera del rango de plantas de poblaciones naturales según las descripciones de las especies. Si la variación en el tamaño de las plantas en colecciones ex situ en comparación con las plantas in situ tiene una base genética, es posible que estas sean funcionalmente inadaptadas si se reintroducen en su entorno natural. Los protocolos para los programas de conservación ex situ deben centrarse en las interacciones ecológicas incluyendo la variación morfológica, bioquímicas y la diversidad genética que harían que la reintroducción de plantas ex situ a su entorno natural maximizara la probabilidad del restablecimiento efectivo. Por consecuencia, las especies que reciben reconocimiento en las exposiciones de orquídeas pueden ser potencialmente inapropiadas dentro de un programa de conservación de reintroducción in situ.

Palabras clave: conservación in situ; evolución; selección fenotípica; selección natural; variación morfológica

Genera	Specific epithet	First year	Last year	Sample size	Reference for in situ
Aerangis	articulata (Rchb.f.) Schltr.	1971	2014	13	NA
Aerangis	biloba (Lindl.) Schltr.	1969	2017	15	NA
Aerangis	citrata (Thouars) Schltr.	1965	2018	15	NA
Aerangis	hariotiana (Kraenzl.) P.J.Cribb & Carlsward	1975	2014	11	NA
Aerangis	luteoalba (Kraenzl.) Schltr.	1965	2020	37	NA
Angraecum	distichum Lindl.	1965	2018	18	NA
Angraecum	eburneum Bory	1965	2018	28	NA
Angraecum	leonis (Rchb.f.) André	1965	2018	20	Notahianjanahary 2016
Angraecum	magdalenae Schltr. & H.Perrier	1972	2016	12	NA
Angraecum	sesquipedale Thouars	1972	2021	44	NA
Angraecum	viguieri Schltr.	1976	2019	20	NA
Brassavola	nodosa (L.) Lindl.	1965	2020	62	Ackerman 2014
Brassia	gireoudiana Rchb.f. & Warsz.	1977	2018	27	NA
Brassia	verrucosa Lindl.	1977	2015	24	NA
Cattleya	aclandiae Lindl.	1965	2019	64	van den Berg 2020
Cattleya	amethystoglossa Linden & Rchb.f. ex Warner	1965	2020	79	van den Berg 2020
Cattleya	aurantiaca (Bateman) P.N.Don	1965	2020	50	Salazar 1990
Cattleya	bicolor Lindl.	1965	2019	30	van den Berg 2020
Cattleya	brevipedunculata (Cogn.) Van den Berg	1981	2020	21	NA
Cattleya	cernua Beer	1965	2019	55	Buzatto 2010
Cattleya	coccinea Lindl.	1965	2020	126	Buzatto 2010
Cattleya	dormaniana Rchb.f.	1977	2020	25	NA
Cattleya	dowiana Bateman	1988	2019	39	Pupulin 2015
Cattleya	forbesii Lindl.	1965	2018	23	NA
Cattleya	granulosa Lindl.	1965	2019	37	van den Berg 2020
Cattleya	harpophylla (Rchb.f.) Van den Berg	1965	2018	19	NA
Cattleya	harrisoniana Bateman ex Lind.	1965	2018	28	van den Berg 2020
Cattleya	intermedia Graham	1977	2019	72	NA
Cattleya	jongheana (Rchb.f.) Van den Berg	1978	2019	48	NA
Cattleya	labiata Lindl.	1952	2018	29	NA
Dendrobium	cuthbertsonii F.Muell.	1984	2020	117	NA
Dendrobium	farmeri Paxton	1965	2021	46	NA
Dendrobium	lindleyi Steud.	1965	2021	73	NA
Laelia	anceps Lindl.	1974	2020	144	Jiménez et al. 1997
Paphiopedilum	armeniacum S.C.Chen & F.Y.Liu	1983	2019	152	NA
Paphiopedilum	bellatulum (Rchb.f.) Stein.	1965	2019	220	Cribb 1987
Paphiopedilum	charlesworthii (Rolfe) Pfizer	1977	2019	62	Cribb 1987
Paphiopedilum	concolor (Bateman) Pfitzer	1965	2016	60	Cribb 1987
Paphiopedilum	delenatii Guillaumin	1965	2019	129	NA
Paphiopedilum	fairieanum (Lindl.) Stein	1973	2019	110	NA
Paphiopedilum	godefroyae Stein	1973	2019	89	NA
Paphiopedilum	hirsutissimum (Lindl. ex Hook) Stein	1975	2002	56	NA
Paphiopedilum	lowii (Lindl.) Stein	1975	2014	141	Cribb 1987
Paphiopedilum	macranthum Z.J.Lui & S.C.Chen	1985	2018	247	Cribb 1987
Paphiopedilum	malipoense S.C.Chen & Z.H.Tsi	1986	2019	108	Cribb 1987
Paphiopedilum	niveum (Rchb.f.) Stein	1970	2019	99	Cribb 1987
Paphiopedilum	philippinense (Rchb.f.) Stein	1973	2019	81	Cribb 1987
Paphiopedilum	rothschildianum (Rchb.f.) Stein	1965	2019	233	NA
Paphiopedilum	sukhakulii Schoser & Senghas	1965	2019	208	NA
Paphiopedilum	venustum (Wall. ex Sims) Pfitzer	1982	2020	46	NA
Paphiopedilum	wardii Summerh.	1972	2016	103	NA
Phalaenopsis	amboinensis J.J.Sm.	1969	2020	84	NA
Phalaenopsis	bellina (Rchb.f.) Christenson	1969	2021	148	NA
Phalaenopsis	cornu-cervi (Hasselt ex Hassk) Blume & Rchb.f.	1972	2021	100	Sweet 1980
Phalaenopsis	equestris (Schauer) Rchb.f.	1971	2019	104	Sweet 1980
Phalaenopsis	lueddemanniana Rchb.f.	1970	2016	51	Sweet 1980
Phalaenopsis	pulcherrima (Lindl) J.J.Sm.	1984	2021	128	NA
Phalaenopsis	violacea hort. ex H.Witte	1969	2021	131	Sweet 1980

Species	DSL	DSW	PL	PW	LSL	LL	NumF
A. articulata	0.008 (0.018)	0.002 (0.002)	0.010 (0.014)	0.003 (0.005)	-0.009 (0.007)	0.011 (0.014)	0.022a (0.009)*
A. biloba	0.028 (0.016)	0.010 (0.005)	0.025 (0.014)	0.009 (0.004)	0.011 (0.004)*	0.020 (0.020)	0.026a (0.011)*
A. citrata	0.012 (0.006)*	0.008 (0.007)	0.010 (0.005)	0.008 (0.003)*	0.003 (0.006)	0.008 (0.005)	-0.008a (0.014)
A. hariotiana	0.002 (0.003)	< 0.001 (0.002)	0.002 (0.002)	< 0.001 (0.002)	0.001 (0.002)	< 0.001 (0.003)	NA
A. luteoalba	0.020 (0.005)	0.001 (0.003)**	0.018 (0.006)**	0.013 (0.004)	0.009 (0.003)**	0.020 (0.008)*	0.004a (0.010)

Species	DSL	DSW	PL	PW	LSL	LL	NumF
A. distichum	-0.003 (0.002)	-0.001 (0.001)	-0.004 (0.002)	-0.001 (0.002)	0.002 (0.002)	-0.003 (0.002)	0.002a (0.025)
A. eburneum	-0.005 (0.009)	-0.003 (0.004)	0.123 (0.161)	-0.016 (0.011)	-0.005 (0.004)	-0.006 (0.011)	0.010a (0.020)
A. leonis	0.005 (0.015)	0.003 (0.003)	0.003 (0.014)	0.004 (0.004)	-0.013 (0.014)	0.056 (0.038)	< -0.0001a (0.013)
A. magdalenae	0.010 (0.028)	-0.011 (0.012)	-0.009 (0.017)	-0.005 (0.008)	-0.004 (0.007)	0.009 (0.019)	0.065 (0.008)
A. sesquipedale	-0.022 (0.013)	0.013 (0.006)*	-0.011 (0.012)	0.018 (0.006)**	0.013 (0.005)*	0.061 (0.062)	-0.017a (0.010)
A. viguieri	-0.006 (0.011)	0.003 (0.003)	0.014 (0.008)	-0.002 (0.002)	0.002 (0.002)	0.022 (0.033)	0.035 (0.010)

Species	DSL	DSW	PL	PW	LSL	LL	NumF
Bv. nodosa	0.03 (0.16)	0.004 (0.002)*	0.011 (0.013)	0.002 (0.001)*	0.012 (0.015)	0.002 (0.013)	-0.08a (0.18)
B. gireoudiana	-0.066 (0.036)	-0.001 (0.002)	0.022 (0.032)	0.001 (0.001)	-0.100 (0.053)	-0.006 (0.013)	0.32a (0.15)*
B. verrucosa	-0.20 (0.06)*	-0.005 (0.003)	-0.29 (0.097)*	-0.006 (0.003)	-0.17 (0.07)*	-0.076 (0.021)**	0.11 (0.20)
D. cuthbertsonii	0.004 (0.004)	0.003 (0.003)	-0.005 (0.004)	< 0.001 (0.003)	0.006 (0.004)	-0.035 ( 0.021)	-0.016a (0.013)
D. farmeri	-0.009 (0.006)	-0.002 (0.004)	-0.01 (0.005)	0.008 (0.003)*	-0.002 (0.004)	-0.006 (0.004)	0.005a (0.008)
D. lindleyi	< -0.001 (0.004)	-0.003 (0.003)	< -0.001 (0.003)	0.002 (0.003)	0.003 (0.003)	< 0.001 (0.004)	0.010a (0.013)
L. anceps	-0.004 (0.007)	0.008 (0.003)*	-0.005 (0.006)	0.015 (0.006)*	NA	0.005 (0.006)	0.029 (0.009)**

Species	DSL	DSW	PL	PW	LSL	LL	NumF
C. aclandiae	-0.003 (0.006)	0.01 (0.003)**	-0.006 (0.005)	0.006 (0.003)	-0.008 (0.006)	0.0005 (0.006)	0.012 (0.007) NB
C. amethystoglossa	0.009 (0.007)	0.007 (0.004)	0.01 (0.007)*	0.02 (0.006)**	0.009 (0.005)	0.005 (0.005)	0.0005 (0.005)** NB
C. aurantiaca	-0.01 (0.007)	0.009 (0.002)	-0.01 (0.007)	0.005 (0.003)	0.003 (0.007)	0.0002 (0.006)	-0.017 (0.01)** NB
C. bicolor	0.005 (0.01)	-0.0006 (0.003)	0.004 (0.009)	0.002 (0.008)	0.002 (0.01)	0.004 (0.006)	-0.007 (0.007)** NB
C. brevipedunculata	0.03 (0.01)**	0.01 (0.007)	0.04 (0.009)	0.02 (0.01)	0.02 (0.01)	0.02 (0.007)*	0.03 (0.02) NB
C. cernua	0.008 (0.003)*	0.005 (0.002)**	0.004 (0.003)	0.002 (0.003)	0.007 (0.003 )*	-0.0002 (0.002)	-0.0002 (0.007)** NB
C. coccinea	0.003 (0.003)	0.008 (0.002)	0.001 (0.003)	0.02 (0.004)	0.002 (0.003)	-0.001 (0.002)	0.008 (0.008)* NB
C. dormaniana	0.02 (0.008)	0.001 (0.002)	0.01 (0.007)	0.002 (0.003)	0.009 (0.01)	0.008 (0.006)	0.07 (0.02) NB
C. dowiana	0.05 (0.02)**	0.01 (0.006)	0.01 (0.02)	0.02 (0.01)	0.008 (0.01)	0.009 (0.02)	-0.015 (0.012)** NB
C. forbesii	0.002 (0.01)	0.004 (0.006)	-0.002 (0.01)	0.006 (0.007)	0.004 (0.01)	0.007 (0.02)	0.04 (0.02) NB
C. granulosa	-0.01 (0.01)	0.003 (0.003)	-0.004 (0.008)	-0.0002 (0.004)	-0.004 (0.009)	0.009 (0.009)	-0.02 (0.006)** NB
C. harpophylla	0.002 (0.01)	-0.002 (0.003)	0.005 (0.01)	0.001 (0.002)	-0.01 (0.01)	0.02 (0.009)*	0.02 (0.01) NB
C. harrisoniana	0.005 (0.009)	0.007 (0.004)	0.001 (0.01)	0.02 (0.01)	0.007 (0.01)	0.01 (0.008)	0.004 (0.009) **NB
C. intermedia	-0.004 (0.01)	0.01 (0.004)*	-0.003 (0.01)	0.02 (0.009)**	0.02 (0.01)	-0.004 (0.008)	-0.0004 (0.02)** NB
C. jongheana	0.01 (0.01)	0.006 (0.002)**	0.01 (0.008)	0.02 (0.007)*	0.01 (0.009)	0.01 (0.007)	0.01 (0.01) NB
C. labiata	-0.04 (0.02)*	-0.02 (0.008)*	-0.04 (0.01)**	-0.04 (0.02)**	-0.05 (0.03)*	-0.03 (0.02)	0.007 (0.02)* NB

Species	DSL	DSW	PL	PW	LSL	LL	NumF
P. armeniacum	0.027 (0.006)	0.019 (0.005)	0.023 (0.005)	0.023 (0.006)	NA	0.005 (0.013)	0.004a (0.009)
P. bellatulum	0.019 (0.003)	0.019 (0.002)	0.019 (0.003)	0.017 (0.002)	NA	0.026 (0.016)	-0.007b (0.003)*
P. charlesworthii	-0.016 (0.007)*	-0.003 (0.010)	0.005 (0.006)	0.012 (0.005)*	NA	NA	-0.004a (0.009)
P. concolor	-0.004 (0.005)	0.016 (0.005)	0.003 (0.005)	0.006 (0.004)	NA	NA	-0.318a (0.010)**
P. delenatii	0.016 (0.003)	0.020 (0.003)	0.019 (0.003)	0.027 (0.004)	NA	NA	< -0.001a (0.007)
P. fairieanum	0.007 (0.004)	0.019 (0.005)	<-0.001 (0.006)	0.006 (0.004)	NA	0.011 (0.015)	-0.045a (0.007)
P. godefroyae	0.016 (0.004)	0.025 (0.004)	0.004 (0.004)	0.013 (0.005)**	NA	NA	0.001b (0.003)
P. hirsutissimum	0.003 (0.012)	-0.009 (0.009)	0.015 (0.015)	-0.003 (0.011)	NA	NA	0.015a (0.019)
P. lowii	0.046 (0.041)	0.022 (0.005)	0.028 (0.013)*	0.017 (0.010)	NA	0.038 (0.005)	-0.012a (0.007)**
P. macranthum	0.013 (0.003)	0.005 (0.015)	0.012 (0.003)	0.015 (0.004)	NA	0.023 (0.021)	0.017a (0.007)*
P. malipoense	0.020 (0.10)*	0.018 (0.007)*	0.023 (0.010)**	0.025 (0.008)**	NA	0.077 (0.028)*	0.027a (0.010)**
P. niveum	0.012 (0.003)**	0.021 (0.004)	0.015 (0.004)**	0.015 (0.003)	NA	0.018 (0.015)	-0.005a (0.007)
P. philippinense	0.021 (0.012)	0.006 (0.004)	0.025 (0.035)	0.012 (0.012)	NA	-0.028 (0.017)	-0.010a (0.005)
P. rothschildianum	0.011 (0.006)	0.037 (0.004)	0.049 (0.015)**	0.003 (0.012)	NA	0.003 (0.011)	0.002a (0.003)
P. sukhakulii	0.015 (0.003)	0.009 (0.002)	-0.004 (0.008)	0.008 (0.005)	NA	0.005 (0.012)	-0.012a (0.005) NB
P. venustum	-0.003 (0.007)	0.013 (0.006)*	-0.003 (0.007)	0.008 (0.005)	NA	NA	-0.003a (0.017)*
P. wardii	0.001 (0.006)	0.019 (0.004)	0.016 (0.009)	0.006 (0.008)	NA	0.006 (0.028)	0.002 (0.010)

Species	DSL	DSW	PL	PW	LSL	LL	NumF
P. amboinensis	0.004 (0.003)	0.007 (0.002)**	0.004 (0.002)*	0.002 (0.002)	0.004 (0.003)	0.002 (0.003)	-0.009a (0.009)*
P. bellina	-0.003 (0.003)	0.009 (0.001)	-0.0005 (0.002)	0.008 (0.002)	0.002 (0.003)	0.004 (0.002)	-0.022a (0.004)
P. cornu-cervi	-0.004 (0.004)	0.003 (0.002)	-0.003 (0.003)	-0.002 (0.002)	-0.002 (0.004)	< -0.001 (0.003)	-0.020a (0.010)
P. equestris	0.0008 (0.003)	-0.0004 (0.002)	0.0005 (0.005)	-0.002 (0.003)	-0.0005 (0.003)	-0.004 (0.003)	0.018a (0.009)*
P. lueddemanniana	-0.005 (0.004)	0.003 (0.002)	-0,008 (0.003)*	0.007 (0.003)*	-0.008 (0.004)*	-0.002 (0.002)	-0.008a (0.010)
P. pulcherrima	0.008 (0.003)*	0.011 (0.002)	0.011 (0.003)	0.027 (0.005)	0.011 (0.003)**	0.011 (0.004)**	0.022a (0.007)**
P. violacea	-0.009 (0.002)	0.0008 (0.001)	-0.012 (0.002)	0.0004 (0.0017)	0.011 (0.06)	-0.004 (0.003)	-0.017a (0.006)**

Species	DSL	DSW	PL	PW	LSL	LL	NumF
Positive effect	0.24 (14)	0.45 (26)	0.21 (12)	0.36 (21)	0.12 (5)	0.10 (5)	0.35 (20)
Negative effect	0.07 (4)	0.02 (1)	0.07 (4)	0.02 (1)	0.08 (3)	0.02 (1)	0.26 (15)
No significant selection	0.69 (40)	0.53 (31)	0.72 (42)	0.62 (36)	0.80 (32)	0.88 (46)	0.39 (22)

Genus (number of species)	Number of characters evaluated	Positive	Negative	No selection
Aerangis (5)	33	0.30	0	0.70
Angraecum (6)	42	0.12	0	0.88
Brassavola (1)	7	0.29	0	0.71
Brassia (2)	14	0.14	0.29	0.57
Cattleya (16)	112	0.24	0.10	0.66
Dendrobium (3)	21	0.05	0	0.95
Laelia (1)	6	0.5	0	0.5
Paphiopedilum (17)	96	0.41	0.07	0.52
Phalaenopsis (7)	49	0.27	0.14	0.59

Species	DSL	DSW	PL	PW	LSL	LL	NumF
Angreacum leonis	--	--	--	--	--	--	+O
Brassavola nodosa	+O	+O	W	+O	+O	+O	+O
Cattleya aclandiae	+O	+O	+O	+O	+O	+O	+O
Cattleya amethystoglossa	+O	+O	+O	+O	+O	+O	+O
Cattleya aurantiaca	+O	+O	±O	+O	+O	±O	+O
Cattleya bicolor	±O	+O	±O	+O	+O	±O	+O
Cattleya cernua	+O	+O	+O	+O	--	--	+O
Cattleya coccinea	+O	+O	+O	+O	+O	--	+O
Cattleya dowiana	--	--	±O	+O	--	+O	+O
Cattleya granulosa	±O	+O	±O	+O	±O	±O	+O
Cattleya harrisoniana	--	--	±O	+O	--	±O	+O
Paphiopedilum bellatulum	+O	+O	+O	+O	--	W	+O
Paphiopedilum charlesworthii	±O	+O	±O	+O	--	--	+O
Paphiopedilum concolor	+O	+O	+O	+O	--	--	+O
Paphiopedilum lowii	+O	+O	+O	W	--	--	+O
Paphiopedilum macranthum	W	+O	+O	+O	--	±O	+O
Paphiopedilum malipoensis	+O	+O	+O	±O	--	W	+O
Paphiopedilum niveum	+O	+O	+O	+O	--	--	+O
Paphiopedilum philippinense	+O	+O	+O	--	--	--	±O
Phalaenopsis cornu-cervi	±O	±O	+O	±O	±O	--	--
Phalaenopsis equestris	+O	+O	+O	+O	±O	+O	--
Phalaenopsis lueddemanniana	+O	+O	--	--	+O	±O	--
Phalaenopsis violacea	+O	+O	±O	+O	-O	-O	--

Genus	+	-	±	W
Angreacum	1.00 (1)	0.0 (0)	0.0 (0)	0.0 (0)
Brassavola	0.86 (6)	0.0 (0)	0.0 (0)	0.14 (1)
Cattleya	0.78 (42)	0.0 (0)	0.22 (12)	0.0 (0)
Paphiopedilum	0.79 (33)	0.0 (0)	0.12 (5)	0.09 (4)
Phalaenopsis	0.57 (12)	0.10 (2)	0.33 (7)	0.0 (0)

Categories	+	-	±	W
DSL	0.75 (15)	0.0 (0)	0.20 (4)	0.5 (1)
DSW	0.95 (19)	0.0 (0)	0.05 (1)	0.0 (0)
PL	0.62 (13)	0.0 (0)	0.33 (7)	0.05 (1)
PW	0.85 (17)	0.0 (0)	0.10 (2)	0.05 (1)
LSL	0.64 (7)	0.09 (1)	0.27 (3)	0.0 (0)
LL	0.36 (5)	0.07 (1)	0.43 (6)	0.14 (2)
NumF	0.95 (18)	0.0 (0)	0.05 (1)	0.0 (0)

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Lankester Botanical Garden, University of Costa Rica Jardín Botánico Lankester, Cartago, Cartago, Costa Rica, CR, 302-7050, 25117949, 25117931 - E-mail: franco.pupulin@ucr.ac.cr

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