Chapter 4
Wild Beans (Phaseolus L.)
of North America
Sarah Dohle, Jorge Carlos Berny Mier y Teran, Ashley Egan,
Theodore Kisha, and Colin K. Khoury
Abstract The wild relatives of the five domesticated species of bean (Phaseolus
L.) are widely distributed across the tropics and subtropics of the New World,
with taxa extending from the Canadian border to Argentina, and on the Caribbean
Islands, Bermuda, and the Galapagos Islands. Mesoamerica holds the largest concentration of species, particularly in the highlands of central Mexico, northward
along the Sierra Madre Occidental, and south to Chiapas. The progenitors and
close relatives of all five domesticates are also concentrated in this region. Plant
breeding involving the use of wild relatives has almost entirely been directed
toward the improvement of common bean (Phaseolus vulgaris L.), the most
widely cultivated species, and successful contributions have mostly come from its
progenitor (Phaseolus vulgaris L.) and a few other taxa. Wild relatives are considered to possess novel useful genetic variation that has not yet been fully explored.
S. Dohle
Department of Plant Sciences, Delaware Valley University, Doylestown, PA, USA
e-mail: sarah.dohle@delval.edu
J. C. Berny Mier y Teran
Department of Plant Sciences, University of California, Davis, CA, USA
e-mail: jcberny@ucdavis.edu
A. Egan
Smithsonian Institution, National Museum of Natural History, Department of Botany,
Washington, DC, USA
e-mail: ashegan2@gmail.com
T. Kisha
USDA Agricultural Research Service, Western Regional Plant Introduction Station,
Pullman, WA, USA
e-mail: theodore.kisha@ars.usda.gov
C. K. Khoury (*)
USDA, Agricultural Research Service, Center for Agricultural Resources Research,
National Laboratory for Genetic Resources Preservation, Fort Collins, CO, USA
International Center for Tropical Agriculture (CIAT), Cali, Colombia
e-mail: colin.khoury@ars.usda.gov
© This is a U.S. Government work and not under copyright protection
in the U.S.; foreign copyright protection may apply 2019
S. L. Greene et al. (eds.), North American Crop Wild Relatives, Volume 2,
https://doi.org/10.1007/978-3-319-97121-6_4
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A number of wild Phaseolus are rare endemics that are threatened in their natural
habitats and are insufficiently protected in situ. Significant ex situ collections of
wild Phaseolus are maintained at the International Center for Tropical Agriculture
(CIAT), the USDA-ARS National Plant Germplasm System, within the Sistema
Nacional de Recursos Fitogenéticos para la Alimentación y la Agricultura
(SINAREFI) Conservation Centers Network in Mexico, and at the Botanic Garden
Meise, Belgium. Unfortunately, over 26% of Phaseolus taxa are not represented
at all in these ex situ conservation facilities, and another 29% are represented by
less than ten accessions, making over half of the species highly underrepresented
in genebanks. Further efforts to enhance the protection of vulnerable species in
their natural habitats, and further collecting to fill critical gaps in germplasm collections, are warranted.
Keywords Genetic resources · Ex situ conservation · In situ conservation ·
Grain legumes
4.1
Introduction
Beans (Phaseolus L.) have been cultivated for over 7000 years in the neotropics
and continue to be a critical source of plant protein, dietary fiber, iron, zinc, and
other micronutrients for human nutrition (Broughton et al. 2003; RamírezVillegas et al. 2010). The plants are also important to sustainable agricultural
production practices due to their ability to enrich soil nitrogen by way of their
symbiotic association with nitrogen-fixing rhizobia bacteria (Rubiales and
Mikic 2015).
The genus Phaseolus has five domesticated species; each of the product of independent domestication processes from different wild progenitor species: common
bean (Phaseolus vulgaris L), lima bean (P. lunatus L.), runner bean (P. coccineus L.),
tepary bean (P. acutifolius A. Gray), and year bean (P. dumosus Macfady) (Bitocchi
et al. 2017). Both common bean and lima bean were in fact domesticated more than
once from different populations of their wild progenitor taxa, in Mesoamerica and in
the Andes (Gepts 1998; Motta-Aldana et al. 2010; Bitocchi et al. 2013).
Common bean is the world’s most important legume for direct human consumption, comprising more than 50% of the grain legumes eaten worldwide, holding
primary importance in the staple diet of over 500 million people, and being cultivated on over 23 million hectares, on every inhabitable continent (Gepts 2001;
McClean et al. 2004; CGIAR 2017). Major production areas include Brazil, Mexico
and Central America, the Iberian Peninsula, Central-Eastern and Southern Africa,
China, and India (Bitocchi et al. 2017).
The other four domesticated species have more distinct and somewhat narrower ecological adaptations (Debouck and Smartt 1995; Bitocchi et al. 2017).
Lima bean grows in the warm humid tropics, from sea level to over 2000 m asl
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101
(Baudoin 1988; Bitocchi et al. 2017; Serrano-Serrano et al. 2012). Runner bean
is adapted to cool highland conditions and is now cultivated primarily in
climatically appropriate regions of Spain, Italy, the UK, the Netherlands, and
other northern European countries with cooler growing season temperatures
(Rodino et al. 2007). Tepary bean, the most drought tolerant of cultivated
Phaseolus, is adapted to arid and semiarid conditions, with high temperatures,
low and variable precipitation, and alkaline soils (Pratt and Nabhan 1988; Porch
et al. 2013a). Tepary bean is still predominantly grown only in its region of
origin in the warm deserts of Mexico and the USA, although it is the subject of
increasing interest in similar agroecological regions of Africa and South Asia
(Small 2014). Year bean is the domesticated form of a species that is the result
of a naturally occurring hybridization event between the progenitors of common
and runner bean (Mina-Vargas et al. 2016). Year bean is cultivated in Central
America (particularly in the highlands of Guatemala), the Caribbean, and in the
Northern Andes from Venezuela south to Peru, especially in climatic zones
where common bean is highly susceptible to anthracnose, Ascochyta blight, and
root rot (Mina-Vargas et al. 2016; USDA, ARS, National Plant Germplasm
System 2017b; D. Debouck personal communication 2017).
Even with the incredible expansion and success of Phaseolus beans worldwide,
a wide range of challenges to their cultivation exist. Common beans are vulnerable
to drought and heat, which are likely to be exacerbated by climate change (Beebe
et al. 2011; Jarvis et al. 2012). Major diseases for the crop include root rot,
Xanthomonas blight, downy mildew, rust, angular leaf spot, anthracnose, web
blight, white mold, halo blight, bean common mosaic virus (BCMV), and a variety
of other viruses. Major insect pests include red spider, mealybug, white fly, leafhoppers, weevils, various caterpillars, flea beetles, Diabrotica, and Mexican bean beetles (Beebe 2012).
Lima bean insect pests include thrips, aphids, leaf miners, Heliothis, and
chrysomelid beetles. They are also susceptible to mycoplasmas (Freytag and
Debouck 2002) and white mold (Boland and Hall 1994). Runner beans are primarily affected by rust, as well as angular leaf spot, anthracnose, and Ascochyta
leaf spot, and by Apion pod weevil, chrysomelid beetles, thrips, flea beetle, and
leafhoppers. Tepary beans are susceptible to powdery mildew, root rots, white
mold, rust, and viruses including alfalfa mosaic, bean yellow mosaic, BCMV,
bean golden mosaic, curly top, pod mottle, and four whitefly-transmitted viruses
(Freytag and Debouck 2002). Insect pests of tepary bean include leafhopper, leafeating beetles, leaf miners, flea beetles, and leaf-eating caterpillars (Debouck
1999; Freytag and Debouck 2002).
This chapter provides an overview of the identities and relationships, distributions, genetic resource potential, and conservation status of the North American
wild relatives of Phaseolus beans. We focus on the progenitors and close relatives
of the domesticated species, as they are considered the most promising taxa with
regard to successful present and future uses as genetic resources (Harlan and de
Wet 1971). We also provide details on Phaseolus polystachios (L.) Britton et al.,
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a distant relative of lima bean which is the most northerly occurring wild species
in the genus, known by the authors to be declining in portions of its natural habitat.
We conclude with a discussion of the key steps needed to improve the conservation of North American Phaseolus crop wild relatives.
4.2
Crop Wild Relatives of Phaseolus Beans
and the Relationships Between Wild Taxa
and the Domesticated Species
Phaseolus is a monophyletic genus native to the New World, comprising some
70–85 species and ca. 30 infraspecific taxa, which are currently parsed into eight
phylogenetic groups within two clades (Freytag and Debouck 2002; DelgadoSalinas et al. 2006; Ramírez-Villegas et al. 2010) (Fig. 4.1). The total number of
species may reach 100 or more with thorough fieldwork in understudied areas of the
neotropics in combination with phylogenetic analyses, in further search of narrow
endemic taxa (D. Debouck personal communication 2017).
The vulgaris group includes all cultivated species except P. lunatus, along with
four other noncultivated species, including the newly described Phaseolus debouckii
A. Delgado, segregated from P. vulgaris based on genome-wide single nucleotide
polymorphisms (Rendón-Anaya et al. 2017a, b). The lunatus group includes at least
five wild species, while its sister clade, the polystachios group, includes nine or more
taxa (Fig. 4.1).
Phaseolus species have been organized into genetic relative categories, or “genepools,” based on crossing studies and phylogenetic analyses (Smartt 1981; Singh
and Jauhar 2005; Rendón-Anaya et al. 2017a, b; The Harlan and de Wet Crop Wild
Relative Inventory 2017; USDA, ARS, National Plant Germplasm System 2017b).
The primary genepool of domesticated species includes both cultivated forms and
wild populations, hybrids of which are generally fully fertile with no major reproductive barriers (Harlan and de Wet 1971) (Table 4.1).
The secondary genepool is comprised of related species that may be crossable
with the cultivated forms, but with extra effort needed to overcome reproductive
barriers and to eliminate linkage drag (Prohens et al. 2017; Miller and Khoury
2018). Common bean has been successfully crossed with species in its secondary genepool; lima and tepary bean may be less capable of gene exchange with
their secondary relatives, although their potential has not been as comprehensively
studied (Debouck 1999; D. Debouck personal communication 2017). Tertiary relatives and other distantly related taxa within the genus are not considered in detail
here, as their potential for utilization as genetic resources is constrained by major
biological barriers to interfertility, although advancing techniques may make utilization of distant relatives more feasible in the future (Zhang et al. 2014; Miller
and Khoury 2018).
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Fig. 4.1 Phylogenetic tree of Phaseolus groups, modified from the trnK-ITS maximum parsimony analysis of Delgado-Salinas et al. (2006), with the nodal supports value noted above key
clades. Groups defined as in Delgado-Salinas et al. (1999). Domesticated species are in bold
4.3
Distribution of Wild Phaseolus in North America
The genus Phaseolus originated in the tropics and subtropics of the New World.
Wild species belonging to the genus are naturally distributed from the USA south to
Argentina (Ramírez -Villegas et al. 2010). Taxa are not known from Chile nor from
Canada (Freytag and Debouck 2002), although Sousa and Delgado-Salinas (1993)
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Table 4.1 Genepools of Phaseolus domesticated species
Common bean – Phaseolus vulgaris L., Phaseolus vulgaris L. var. vulgaris
Primary relatives
Secondary relatives
Tertiary relatives
Phaseolus acutifolius
Phaseolus albescens
Phaseolus vulgaris L. (including
Phaseolus vulgaris L. var. aborigineus McVaugh ex R. Ramírez A. Gray
& A. Delgado
(Burkart) Baudet and Phaseolus
vulgaris L. var. mexicanus A. Delgado)
Phaseolus coccineus L.
Phaseolus acutifolius
A. Gray var. acutifolius
Phaseolus costaricensis Phaseolus acutifolius
Freytag & Debouck
A. Gray var. tenuifolius
A. Gray
Phaseolus dumosus
Phaseolus angustissimus
Macfady
A. Gray
Phaseolus debouckii
Phaseolus carteri Freytag &
A. Delgado
Debouck
Phaseolus persistentus
Phaseolus filiformis Benth.
Freytag & Debouck
Phaseolus maculatus
Scheele subsp. ritensis
(M. E. Jones) Freytag
Phaseolus parvifolius
Freytag
Runner bean – Phaseolus coccineus L.
Primary
Secondary
Tertiary
Phaseolus acutifolius
Phaseolus coccineus L.
Phaseolus albescens
McVaugh ex R. Ramírez A. Gray
& A. Delgado
Phaseolus costaricensis
Freytag & Debouck
Phaseolus dumosus
Macfady
Phaseolus vulgaris L.
Year bean – Phaseolus dumosus Macfady
Primary
Secondary
Tertiary
Phaseolus parvifolius
Phaseolus dumosus Macfady
Phaseolus albescens
McVaugh ex R. Ramírez Freytag
& A. Delgado
Phaseolus coccineus L.
Phaseolus costaricensis
Freytag & Debouck
Phaseolus vulgaris L.
Tepary bean – Phaseolus acutifolius A. Gray
Primary
Secondary
Tertiary
Phaseolus acutifolius A. Gray var.
Phaseolus parvifolius
Phaseolus albescens
acutifolius
Freytag
McVaugh ex R. Ramírez &
A. Delgado
(continued)
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Table 4.1 (continued)
Phaseolus acutifolius A. Gray var.
tenuifolius A. Gray
Lima bean – Phaseolus lunatus L.
Primary
Phaseolus lunatus L.
Phaseolus carteri Freytag &
Debouck
Phaseolus coccineus L.
Phaseolus costaricensis
Freytag & Debouck
Phaseolus dumosus Macfady
Phaseolus filiformis Benth.
Phaseolus persistentus
Freytag & Debouck
Phaseolus vulgaris L.
Secondary
Tertiary
Phaseolus augusti Harms Phaseolus acinaciformis
Freytag & Debouck
Phaseolus
Phaseolus albinervus
longiplacentifer Freytag Freytag & Debouck
Phaseolus mollis Hook. f. Phaseolus jaliscanus Piper
Phaseolus
Phaseolus juquilensis
pachyrrhizoides Harms
A. Delgado
Phaseolus viridis Piper
Phaseolus lignosus Britton
Phaseolus maculatifolius
Freytag & Debouck
Phaseolus maculatus
Scheele
Phaseolus marechalii
A. Delgado
Phaseolus nodosus Freytag
& Debouck
Phaseolus novoleonensis
Debouck
Phaseolus polystachios (L.)
Britton et al.
Phaseolus reticulatus
Freytag & Debouck
Phaseolus rotundatus
Freytag & Debouck
Phaseolus salicifolius Piper
Phaseolus scrobiculatifolius
Freytag
Phaseolus sonorensis Standl.
Phaseolus venosus Piper
Phaseolus xolocotzii
A. Delgado
Adapted from USDA-ARS National Plant Germplasm System (2017b), The Harlan and de Wet
Crop Wild Relative Inventory (2017), Rendón-Anaya et al. (2017a, b), and D. Debouck personal
communication (2017)
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report wild populations in southern Quebec and Ontario. Members also occur on the
major Caribbean Islands (Phaseolus dumosus Macfady, Phaseolus lunatus L., and possibly P. polystachios [Liogier (1988)]), Bermuda (Phaseolus lignosus Britton), and the
Galapagos Islands (Phaseolus mollis Hook. f.) (Ramírez -Villegas et al. 2010; Arani
et al. 2017; Debouck 2017; USDA, ARS, National Plant Germplasm System 2017b).
The main areas of richness of species are Mesoamerica (from southern USA,
Mexico, and Central America down to Panama), the northern Andean region
(Colombia to northern Peru), and the central Andes (northern Peru, Bolivia, and
northwest Argentina). Of these, the Mesoamerican center is the richest in number of
taxa, reaching the highest concentrations in the highlands of central Mexico, northward along the Sierra Madre Occidental, and south to Chiapas (Nabhan 1990;
Debouck 2000; Freytag and Debouck 2002; Ramírez-Villegas et al. 2010) (Fig. 4.2).
Numerous rare endemic species are distributed in North America, especially in the
Sierra Madre Occidental and Sierra Madre Oriental of Mexico.
The distributions of the primary and secondary relatives of the cultivated species
are detailed below.
Fig. 4.2 Species richness map of modeled potential distributions of 57 North American Phaseolus
taxa. The map displays overlapping potential distribution models for assessed taxa. Potential distribution models are based on climatic and edaphic similarities with herbarium and genebank reference localities and reflect potential for occurrence rather than confirmed presence. Warmer colors
indicate areas where greater numbers of taxa potentially occur in the same geographic localities.
Full methods for generation of maps and occurrence data providers are listed in Appendix 1
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Wild Beans (Phaseolus L.) of North America
4.3.1
107
Distributions of Close Relatives of Common Bean
Wild common bean (P. vulgaris L.) occurs from southern Chihuahua, Mexico, south
through Central America and the Andes to Argentina (Nabhan 1985; USDA, ARS,
National Plant Germplasm System 2017b) (Fig. 4.3). Populations are found from
very arid conditions to over 100 cm annual precipitation (Gentry 1969; Freytag and
Debouck 2002). The plant is often found growing over or in shrubs on the edges
and within open pine-oak forests and woodlands, usually from 800 to 2000 m asl
(Freytag and Debouck 2002). Wild common bean is usually found in shallow, often
friable but sometimes heavy, soils, derived from limestone, schist, granite, lava, or
tufa (Freytag and Debouck 2002). The plant is a short-lived perennial, generally producing seeds during its first year of growth on indeterminate climbing vines reaching
over 3 meters long (Gentry 1969). The species produces vegetative growth during
the rainy season, developing pods during the dry season, which varies by region,
indicating a strong population-level adaptation to local climates (Gentry 1969). Wild
common bean has been separated into multiple varieties, including P. vulgaris L. var.
aborigineus [Burkart] Baudet and Phaseolus vulgaris L. var. mexicanus A. Delgado
based on geographic and morphological differences. Here we follow Freytag and
Debouck (2002) in treating the plants as one continuum within the species.
Fig. 4.3 Occurrence localities of North American Phaseolus primary and secondary wild relatives
within the common bean (P. vulgaris L.), runner bean (P. coccineus L.), and year bean (P. dumosus
Macfad.) genepools. Full methods for generation of maps and occurrence data providers are listed
in Appendix 1
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Secondary relative Phaseolus albescens McVaugh ex R. Ramírez & A. Delgado
is known from 16 populations distributed in montane forests in central Mexico, in
the states of Colima, Guerrero, Jalisco, and Michoacan (Ramírez-Delgadillo and
Delgado-Salinas 1999; Debouck 2017; USDA, ARS, National Plant Germplasm
System 2017b) (Fig. 4.3). Phaseolus costaricensis Freytag & Debouck occurs in
Costa Rica and Panama (USDA, ARS, National Plant Germplasm System 2017b),
the newly described Phaseolus debouckii A. Delgado is distributed in Ecuador
and northern Peru (Rendón-Anaya et al. 2017a), and Phaseolus persistentus
Freytag & Debouck occurs in Guatemala (USDA, ARS, National Plant Germplasm
System 2017b). Wild Phaseolus coccineus L. and P. dumosus are covered in the
sections below. North American occurrences of these species are mapped in
Fig. 4.3.
4.3.2
Distributions of Close Relatives of Runner Bean
Wild runner bean (P. coccineus) occurs from southern Chihuahua, Mexico south
to Matagalpa, Nicaragua; reports outside that range are likely to be escapes from
cultivation or misidentifications (Nabhan 1985; Freytag and Debouck 2002; USDA,
ARS, National Plant Germplasm System 2017b; D. Debouck personal communication 2017) (Fig. 4.3). Wild runner bean is a climbing perennial with large red flowers
that can vine up to 5 m. The species has been assigned several subspecies, some of
which have purple flowers; white flowers are generally restricted to cultivated types
(Freytag and Debouck 2002). Wild runner bean is normally found from 1500 to
2500 m asl in mixed forests of pine, oak, juniper, liquidambar, and hawthorn, growing over shrubs and herbs (Freytag and Debouck 2002). The various subspecies and
varieties of wild coccineus appear to be adapted to specific climates and habitats,
from cloud forest to desert and from rich to poor soils (Freytag and Debouck 2002).
Some types are quite rare, such as Phaseolus coccineus L. subsp. coccineus var.
griseus (Piper) Freytag, occurring in dry conditions of southern Oaxaca in scrub oak
and mesquite (Freytag and Debouck 2002).
4.3.3
Distributions of Close Relatives of Year Bean
Wild year bean (P. dumosus) is distributed in Chiapas in southern Mexico and in
Guatemala and is naturalized in other regions of Central America as well as in the
Caribbean (USDA, ARS, National Plant Germplasm System 2017b) (Fig. 4.3).
Only four populations are known in Mexico (Freytag and Debouck 2002; Debouck
2017), in humid pine-oak forests, growing on very steep slopes and along streams,
mostly in open places, often over weeds and small trees, in deep, moist, welldrained, often sandy or rocky, humic soil derived from volcanic ash (Freytag and
Debouck 2002).
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Wild Beans (Phaseolus L.) of North America
4.3.4
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Distributions of Close Relatives of Tepary Bean
Wild tepary bean (Phaseolus acutifolius A. Gray) mostly occurs at elevations of 335–
2000 m asl from Arizona and New Mexico, USA, and south to Jalisco in central
Mexico (Freytag and Debouck 2002; USDA, ARS, National Plant Germplasm System
2017b) (Fig. 4.4). Populations may extend into Texas and Baja California (Freytag
and Debouck 2002). Wild tepary beans are annuals, with fibrous roots and small slender climbing vines. Plants are generally found along dry washes, streambeds, canyon
bottoms, and floodplains in pine-oak forest and along trails, abandoned fields, and
fence rows in drier areas (Freytag and Debouck 2002). Phaseolus acutifolius A. Gray
var. acutifolius is somewhat sympatric with Phaseolus acutifolius A. Gray var. tenuifolius A. Gray, although variety acutifolius is thought to occur in drier conditions and
generally more westerly in the Sierra Madre and lowlands, whereas variety tenuifolius
occurs in the higher parts of the mountains and eastward (Freytag and Debouck 2002;
D. Debouck personal communication 2017). A wild Phaseolus acutifolius A. Gray
var. latifolius Freeman has been described as occurring from northwest Mexico and
the southwestern USA (Freytag and Debouck 2002). Here we follow USDA-ARS
National Plant Germplasm System (2017b) in considering it a synonym of var. acutifolius. Secondary relative Phaseolus parvifolius Freytag is distributed from Sonora
and Baja California Sur, Mexico, south to Guatemala (Fig. 4.4).
Fig. 4.4 Occurrence localities of North American Phaseolus primary and secondary wild relatives
within the tepary bean (P. acutifolius A. Gray) genepool. Full methods for generation of maps and
occurrence data providers are listed in Appendix 1
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4.3.5
S. Dohle et al.
Distributions of Close Relatives of Lima Bean
Wild lima bean (P. lunatus L.) is distributed from Sinaloa, Mexico, south through
Central America to Brazil and Argentina (Freytag and Debouck 2002; USDA, ARS,
National Plant Germplasm System 2017b) (Fig. 4.5). One population is known from
Baja California Sur, Mexico. Wild lima bean is a climber, with vines up to 8 m long
and with perennial, fibrous, and sometimes fleshy roots. The plant is mostly found
along stream banks or other moist areas of pine-oak forests between 0 and 1600 m asl
(Freytag and Debouck 2002). Wild lima bean often grows over shrubs and small trees
in areas cleared for coffee, sugarcane and other crops, and is more abundant in areas
inaccessible to grazing. The plant is adapted to a wide range of soil types derived
from volcanic rock and ash, metamorphic schists, limestones, and basalt, from black
clay to brown friable rock, and even in sand dunes (Freytag and Debouck 2002).
Phaseolus longiplacentifer Freytag is a narrow endemic species described by the
senior author in Freytag and Debouck (2002), known only from its type collection
north of Orizaba, Veracruz, Mexico. Delgado-Salinas et al. (2006) assigned longiplacentifer as a synonym of Phaseolus viridis Piper (1926), which is found in scattered places in Veracruz, Oaxaca, and Chiapas, Mexico, and in Alta Verapaz,
Guatemala (Debouck 2017), while the senior author in Freytag and Debouck (2002)
Fig. 4.5 Occurrence localities of North American Phaseolus primary and secondary wild relatives
within the lima bean (P. lunatus L.) genepool. Full methods for generation of maps and occurrence
data providers are listed in Appendix 1
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Wild Beans (Phaseolus L.) of North America
111
considered P. viridis a synonym of (wild) P. lunatus. Phaseolus augusti Harms is
distributed in Ecuador, Peru, Bolivia, and Argentina; Phaseolus pachyrrhizoides
Harms occurs in Peru; and P. mollis is distributed only in the Galapagos Islands
(USDA, ARS, National Plant Germplasm System 2017b).
4.4
4.4.1
Utilization of Wild Phaseolus
Utilization of Wild Phaseolus as Genetic Resources
Considerable genetic variation in cultivated common bean has existed historically
as the result of multiple domestication events as well as further naturally occurring
gene flow between wild progenitors and early domesticates, resulting in six to seven
“races” within two main “genepools” with clear genetic structure (Acosta-Gallegos
et al. 2007). The diversity in major commercial common bean varieties has been considered to be relatively narrow, though, and numerous authors have recommended a
widening of this genetic base (e.g., Adams 1977; Singh 1992; Sonnante et al. 1994;
Beebe et al. 1995; Singh 1999; Métais et al. 2002; Rosales-Serna et al. 2004).
Interspecific crosses are possible between domesticated P. vulgaris, P. coccineus,
P. dumosus, and P. acutifolius (Fig. 4.6). Phaseolus coccineus and P. dumosus readily cross with P. vulgaris without embryo rescue, although the progeny have reduced
viability when using P. vulgaris as the male parent (Al-Yasiri and Coyne 1966;
Debouck 1999). Phaseolus acutifolius has been utilized in interspecific crosses with
common bean, mostly through congruity crosses (alternate backcrossing), using
embryo rescue with P. vulgaris as the cytoplasm source (Waines et al. 1988; MejíaJiménez et al. 1994; Anderson et al. 1996).
Fig. 4.6 Viability of interspecific crosses of the five domesticated species of Phaseolus based on
Al-Yasiri and Coyne (1966), Smartt (1970), and Debouck (1999). The arrow points toward the
female in the cross
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Runner bean accessions have been employed in common bean breeding as new
sources of resistance to bean golden yellow mosaic virus (Osorno et al. 2007), white
mold (Schwartz et al. 2006; Singh et al. 2009; Vasconcellos et al. 2017), and common bacterial blight (Freytag et al. 1982; Miklas et al. 1999). Year bean accessions
have been researched for resistance to anthracnosis (Mahuku et al. 2002), Ascochyta
leaf blight (Schmit and Baudoin 1992), and white mold (Hunter et al. 1982) as
well as to enhance nutritional quality (Blair 2013). Tepary bean accessions have
contributed new sources of resistance to common bacterial blight, leafhopper, and
bruchid (Singh et al. 1998), and the species has been explored for drought and heat
tolerance (Rao et al. 2013). Although attempts to cross P. vulgaris and P. lunatus
have mostly been unsuccessful (Al-Yasiri and Coyne 1966; Smartt 1970), viable
but infertile plants have been reported (Mok et al. 1978), although not replicated
(Debouck 1999).
Practical limitations constrain the use of wild germplasm for common bean cultivar improvement, as breeding is mainly restricted to the variation within market
classes (Kelly et al. 1998; Singh 2001). The complexity of inheritance and genetic
linkage of traits of commercial importance, such as seed color, seed size, and growth
habit, has hindered the introgression of novel variation (Koinange et al. 1996; Kelly
et al. 1999; Singh 2001; McClean et al. 2002; Blair et al. 2006; Moghaddam et al.
2016). The lack of useful characterization and evaluation data on wild accessions
has also been considered a constraint. Wild accessions are avoided because of the
limited power of evaluation of the germplasm for agronomically useful traits, due to
attributes such as vigorous growth habits, long growing cycles, photoperiod sensitivity, and dehiscence (Singh 2001; Acosta-Gallegos et al. 2007). Finally, a simple
lack of readily available wild germplasm may have constrained its use in bean
breeding, particularly prior to the 1970s (Freytag and Debouck 2002).
That said, Phaseolus wild relatives are considered to possess valuable traits
that may be difficult to find in domesticated materials (Porch et al. 2013b; Rao
et al. 2013). Target traits in common bean research with focus on wild vulgaris
germplasm have included resistance to storage insects, leafhoppers, Ascochyta
blight, common bacterial blight, white mold, BCMV, Fusarium root rot, and rust,
and tolerance to abiotic stresses and low soil fertility, as well as early maturity,
adaptation to higher latitudes, upright plant type, pod quality, seed size, seed
yield, protein digestibility, and nodulation traits (Kornegay and Cardona 1991;
Shellie-Dessert and Bliss 1991; Kipe-Nolt et al. 1992; Kornegay et al. 1993;
Delgado-Salinas et al. 1999; Singh 2001; Acevedo et al. 2006; Blair et al. 2006;
Acosta-Gallegos et al. 2007; Cortés et al. 2013; De Ron et al. 2015; Blair et al.
2016; Rodriguez et al. 2016).
Wild P. vulgaris has been successfully used in common bean breeding as a
source of resistance to bruchids (Osborn et al. 1988, 2003; Kornegay et al. 1993),
common bacterial and web blight (Beaver et al. 2012), white mold (Mkwaila et al.
2011), and for yield (Acosta-Gallegos et al. 2007; Wright and Kelly 2011; Porch
et al. 2013b). In addition, wild P. acutifolius has been utilized in common bean
breeding as a source of resistance to bruchids (Singh et al. 1998; Kusolwa et al. 2016),
4
Wild Beans (Phaseolus L.) of North America
113
Table 4.2 Accessions of wild Phaseolus published as utilized in common bean breeding. For
accession codes, G refers to International Center for Tropical Agriculture (CIAT) collections and
PI to USDA-ARS National Plant Germplasm System accessions
Trait
Bruchids
Bruchids
Cold and
drought
Common
bacterial
blight
Web blight
White
mold
White
mold
Yield
Source
species
P. vulgaris L.
PI 638833
References
Osborn et al. (1988, 2003) and
Kornegay et al. (1993)
Singh et al. (1998) and
Kusolwa et al. (2016)
Souter et al. (2017)
PI 417662
Beaver et al. (2012)
P. vulgaris L.
P. vulgaris L.
PI 417662
PI 318695
Beaver et al. (2012)
Mkwaila et al. (2011)
P.
costaricensis
Freytag &
Debouck
P. vulgaris L.
G40604
Singh et al. (2009), Schwartz
and Singh (2013)
G24423
Acosta-Gallegos et al. (2007)
P. acutifolius
A. Gray
P. acutifolius
A. Gray
P. vulgaris L.
Accession
G12882, G12866, G12952,
G02771
G40199
low temperatures, and drought tolerance (Souter et al. 2017). Phaseolus costaricensis has contributed white mold resistance (Singh et al. 2009; Schwartz and
Singh 2013). Wild Phaseolus accessions published as sources of traits utilized in
common bean plant breeding programs are listed in Table 4.2.
Secondary relative P. costaricensis has also been noted as belonging to a germplasm group potentially having resistance to angular leaf spot, anthracnose, Ascochyta
blight, bean golden mosaic virus, bean yellow mosaic virus, common bacterial blight,
and Fusarium root rot, as well as cold tolerance (Singh 2001). Some distant relatives
of common bean have been recorded as possessing interesting characteristics of
potential value to breeding, although successful introgression would be difficult.
Examples include Phaseolus leptostachyus Benth. for its determinate growth habit
(Freytag and Debouck 2002), Phaseolus angustissimus A. Gray for cold tolerance
(Balasubramanian et al. 2004), and the Mexican restricted endemic Phaseolus leptophyllus G. Don, whose unique leaf morphology may provide protection from water
loss during drought stress (Freytag and Debouck 2002).
Widening of genetic diversity in the other Phaseolus crop species may also prove
important, as species such as tepary bean are thought to possess relatively low levels
of genetic variation in cultivated forms (Schinkel and Gepts 1988; Garvin and
Weeden 1994; Munoz et al. 2006). With regard to lima bean, tertiary relatives
Phaseolus jaliscanus Piper, Phaseolus maculatus Scheele, Phaseolus maculatus
Scheele subsp. ritensis (M. E. Jones) Freytag, P. polystachios, and Phaseolus salicifolius Piper have been noted for their disease resistance (van der Maesen and
Somaatmadja 1992).
114
4.4.2
S. Dohle et al.
Direct Uses of Wild Phaseolus Species
The seeds of wild common and lima beans are used as food by indigenous peoples
in times of scarcity but are not regularly consumed (Gentry 1969; Freytag and
Debouck 2002; Zizumbo-Villareal et al. 2012). Kaplan (1965) noted that the roots
of wild runner bean are poisonous, but that tubers which form on the roots are
occasionally boiled for consumption. Wild vulgaris, coccineus, and acutifolius var.
tenuifolius are known to be grazed by cattle and goats (Freytag and Debouck 2002).
A number of additional wild Phaseolus species were probably occasionally consumed and possibly even cultivated, including P. polystachios (possibly cultivated)
(Kaplan 1965), P. leptostachyus, and Phaseolus glabellus Piper (Freytag and
Debouck 2002).
4.5
4.5.1
Conservation of Wild Phaseolus in North America
In Situ Conservation of Wild Phaseolus in North
America
NatureServe has published conservation assessments for 17 currently accepted
Phaseolus taxa (NatureServe 2017). Of these only Phaseolus texensis A. Delgado &
W. R. Carr is listed as a threatened species (i.e., G1 or G2), assigned a G2 (imperiled) status. The species is narrowly endemic to rocky canyons in the eastern and
southern parts of the Edwards Plateau of Texas and is known from only a few localities (Delgado-Salinas and Carr 2007). Phaseolus supinus Wiggins & Rollins is also
listed imperiled and is also present (although not listed) in the US Fish and Wildlife
Service Environmental Conservation Online System (USFWS 2017), but this species is no longer an accepted Phaseolus taxon, having been reassigned as
Macroptilium supinum (Wiggins & Rollins) A. Delgado et al. (USDA, ARS,
National Plant Germplasm System 2017b).
The IUCN Red List of Threatened Species lists P. lignosus, the Bermuda
endemic, as Critically Endangered (Copeland et al. 2014). The species is restricted
to six populations due to habitat loss and invasive species, and surveys completed
in 2004 counted a total of only 29 mature individuals, which may be all that
are left of naturally occurring plants. Surveys conducted in 2014 were successful in finding populations only within Walsingham Natural Reserve, although it
may also occur in three other protected areas in Bermuda. Phaseolus lignosus is
listed as Critical Risk B1, C, at Level 3 in Bermuda, under the Protected Species
Order 2012, and benefits from a recovery plan led by Bermuda Department of
Environment Protection, including habitat protection and restoration as well as ex
situ conservation. Seeds are in long-term storage at the International Center for
Tropical Agriculture and at Kew’s Millennium Seed Bank, and propagation efforts
4
Wild Beans (Phaseolus L.) of North America
115
have translocated the species to five sites, where populations appear to be viable
(Copeland et al. 2014; Debouck 2015).
The Red List also covers P. polystachios and Phaseolus xanthotrichus Piper, both
as species of Least Concern, assessed as widespread species with generally stable
populations (Contu 2012; Groom 2012). Recent efforts by authors Kisha and Egan
to document the extant range of P. polystachios relative to historical collections
have revealed a striking decline in range and commonality therein, particularly
along northern and western range boundaries (T. Kisha & A.N. Egan personal
observations 2017). Furthermore, while P. polystachios has a wide range within
North America, it is now declared extirpated from Michigan (MI DNR 2017),
Connecticut (CT DEEP 2015), and Pennsylvania (PA Natural Heritage Program
Species Lists 2014) and imperiled in several others (NatureServe 2017).
The IUCN Red List and NatureServe programs currently offer no information on
a large number of endemic and otherwise rare North American Phaseolus taxa,
including Phaseolus polystachios (L.) Britton et al. subsp. smilacifolius (Pollard)
Freytag occurring in Florida; Phaseolus carteri Freytag & Debouck in Baja
California Sur, Phaseolus amabilis Standl. in Chihuahua; Phaseolus reticulatus
Freytag & Debouck in Durango; Phaseolus albiflorus Freytag & Debouck,
Phaseolus albiviolaceus Freytag & Debouck, Phaseolus altimontanus Freytag &
Debouck, Phaseolus maculatifolius Freytag & Debouck, Phaseolus novoleonensis
Debouck, Phaseolus neglectus F. J. Herm., and Phaseolus trifidus Freytag in Nuevo
Leon, Tamaulipas, and/or Coahuila; Phaseolus purpusii Brandegee in San Luis
Potosi; P. jaliscanus, Phaseolus esperanzae Seaton, Phaseolus magnilobatus
Freytag & Debouck, Phaseolus marechalii A. Delgado, and Phaseolus rotundatus
Freytag & Debouck in central Mexico; Phaseolus leptophyllus G. Don in Guerrero;
and Phaseolus chiapasanus Piper in southern Mexico. Conservation status and
threat assessment information are needed for such species to be able to prioritize
conservation action.
Aside potentially from the populations of wild Phaseolus monitored in the Sierra
de Manantlan Biosphere Reserve in Jalisco and Colima, Mexico (Vázquez-García
1995), no active management of wild Phaseolus in protected areas is known by the
authors to be ongoing in North America. And even in the Biosphere Reserve, the
mention by Vázquez-García et al. (1995) of two unidentified species clearly indicates
that the first limiting factor for in situ conservation is that protected areas lack a
complete inventory of plants (D. Debouck personal communication 2017).
In situ conservation is certainly occurring in federal, state, provincial, Native
American, Indigenous Peoples, nongovernmental, and privately managed protected areas without inventories, active management plans, or regularly scheduled monitoring. But information on these populations is not available. Outside
of these areas, wild Phaseolus populations are considered to be vulnerable to
habitat destruction (Freytag and Debouck 2002) and potentially (for progenitor
species) to genetic contamination due to gene flow with cultivated forms (Papa
and Gepts 2003).
116
4.5.1.1
S. Dohle et al.
Remarks on Phaseolus polystachios (L.) Britton et al.
Phaseolus polystachios, the North American wild kidney bean or thicket bean, is the
only Phaseolus species native to temperate North America, with a distribution from
Texas to Connecticut, USA. As such, it has evolutionarily adapted to different climatic, ecological, and pest conditions compared to most of its congeners and may
thus harbor unique genetic traits that could prove useful for Phaseolus breeding
efforts. The thicket bean is an herbaceous, perennial vine with tuberous roots that
overwinter. Thicket bean is known to have hypogeal germination wherein the seed
and cotyledons can stay below ground through germination, a trait that may have
enabled P. polystachios to flourish in colder latitudes (Dhaliwal and Pollard 1962).
Phaseolus polystachios has a documented distribution reaching from New
England to Florida, west to the eastern edges of Texas, and north to Michigan. The
plant favors the edges of forests, roads, and streams where it has garnered its name
by twining into thickets in rather open areas, particularly on slopes with southwestern, southern, and southeastern aspects. The thicket bean can establish colonies on
open forest slopes, but it much prefers well-drained soils and full-sun conditions
(Allard 1947). The thicket bean prefers an intermediate day length, with too-short
or too-long days hampering its growth. Depending on latitude, P. polystachios will
flower between June and September and fruit between August and November.
Thicket bean’s wide range may have been historically augmented by indigenous gathering, caching, or even cultivation. Seeds or phytoliths of P. polystachios
have been identified at the Parkin site in Arkansas, dating from the fifteenth to
sixteenth centuries (Scarry and Reitz 2005) and much earlier from the Hunter’s
Home site in New York, with potential dates as early as 2500 B.C. (Hart et al.
2008). That said, even with its wide distribution, the thicket bean is not commonly
found within its range and seems to be on the decline. The plant is considered
extirpated in Michigan, where it was historically documented from the Detroit
River International Wildlife Refuge, Pennsylvania, where author A.N. Egan failed
to find any extant populations at 21 historical localities, and in Connecticut, listed
as special concern by state agencies as of November 2017 (see above). Thicket
bean’s decline may be due to competitive exclusion from nonnative, introduced
vines, habitat destruction from urban development (e.g., there are old herbarium
records of populations located within what are now highly urbanized New York
City and Washington DC [D. Debouck personal communication 2017]), quarrying, mowing, herbicide use, and fire suppression around human-built structures
(A.N. Egan, personal observation 2017). Authors A.N. Egan and T. Kisha have
completed extensive locality studies across its range, visiting over 200 locations in
efforts to collect plants, but averaged about a 25% success rate of finding extant
populations from historical collection locality information or firsthand knowledge
of state and federal botanists.
The closest cultivated relative of P. polystachios is lima bean, which is susceptible to white mold (Sclerotinia sclerotiorum (Lib.) de Bary), leading to flower drop,
emergence issues, and pod set failure. Through coevolution in its natural habitat,
P. polystachios may have acquired true resistance to the ubiquitous pathogen.
4
Wild Beans (Phaseolus L.) of North America
117
Further, transfer of the hypogeal germination trait to lima bean may be useful.
Several decades ago, successful crosses between the two species were made, but
hybrids showed high pollen sterility, likely due to meiotic irregularities due to chromosomal and genetic differences (Dhaliwal and Pollard 1962). With advances in
plant breeding and genetics techniques, renewed efforts to tap into the P. polystachios
genome may be fruitful for lima bean improvement. One P. lunatus x P. polystachios
hybrid accession (G40503) is represented in the CIAT collection, originating
from the work of Albert P. Lorz of the University of Florida (D. Debouck personal
communication 2017).
The USDA-ARS National Plant Germplasm System (NPGS) safeguarded until
recently only 13 accessions of P. polystachios, six of which were collected in
2013 in Florida. Recent explorations supported by the NPGS have augmented the
holdings. A trip throughout Ohio in 2015 by authors Egan and Kisha increased
holdings by 11 accessions, and a 2016 trip to Virginia, North Carolina, and South
Carolina by Egan increased holdings by 19 accessions. In 2017, trips by Egan to
Alabama, Arkansas, Mississippi, and Louisiana yielded 25 more accessions, while
a trip by Kisha to Kentucky and Tennessee yielded an additional 10. Enrichment of
germplasm collections from across the range of P. polystachios will enable a much
more comprehensive assessment of genetic diversity within the species, providing
valuable knowledge to in situ conservation efforts, and may in addition help to
reveal further traits of value for crop improvement. Genetic diversity studies are
currently underway by Kisha and Egan.
4.5.2
Ex Situ Conservation of Wild Phaseolus in North
America
Significant ex situ collections of wild Phaseolus are maintained at the International
Center for Tropical Agriculture (CIAT), Colombia (ca. 2000 accessions); the
USDA-ARS National Plant Germplasm System (ca. 900 accessions), within the
Sistema Nacional de Recursos Fitogenéticos para la Alimentación y la Agricultura
(SINAREFI) Conservation Centers Network in Mexico (ca. 400 accessions); and
the Botanic Garden Meise, Belgium (ca. 400 accessions).
Counts of accessions of Phaseolus taxa are provided in Table 4.3. Of 94 taxa, 25
(26.6%) are not represented at all in these ex situ conservation facilities, and another
27 (28.7%) are represented by less than ten accessions, making over half of Phaseolus
taxa highly underrepresented in these genebanks. Given that some duplication exists
between the major collections as a result of collaborative collecting missions and
recent repatriation (e.g., repatriation of 915 wild Mexican Phaseolus accessions
maintained at CIAT to the Centro Nacional de Recursos Genéticos [CNRG] of the
Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias [INIFAP])
(F. de la Torre personal communication 2017), the number of unique accessions
held in these facilities is likely even less than the total counts for most taxa.
118
S. Dohle et al.
Table 4.3 Counts of accessions in major wild Phaseolus collections databases
USDAARS
Taxon
CIAT NPGS
Phaseolus acinaciformis 1
Phaseolus acutifolius
53 (33)
Phaseolus acutifolius var. 81
36 (16)
acutifolius
Phaseolus acutifolius var. 84
55 (50)
tenuifolius
Phaseolus albescens
1
Phaseolus albicarminus 1
Phaseolus albiflorus
7
1 (1)
Phaseolus albinervus
Phaseolus albiviolaceus 2
Phaseolus altimontanus 1
1 (0)
Phaseolus amabilis
Phaseolus amblyosepalus
Phaseolus angustissimus 5
4 (2)
Phaseolus anisophyllus
Phaseolus augusti
29
16 (1)
Phaseolus campanulatus
Phaseolus carteri
1
2
Phaseolus chiapasanus
4
1
Phaseolus coccineus
168 74 (18)
Phaseolus coccineus
5 (0)
subsp. coccineus
Phaseolus coccineus var.
17 (1)
coccineus
Phaseolus coccineus var.
2 (0)
griseus
Phaseolus coccineus var.
1 (0)
tridentatus
Phaseolus costaricensis 27
6
Phaseolus dasycarpus
Phaseolus dumosus
9
6 (3)
Phaseolus esperanzae
7
Phaseolus esquincensis
Phaseolus filiformis
36
19 (15)
Phaseolus glabellus
8
5 (1)
Phaseolus gladiolatus
Phaseolus grayanus
2
18 (0)
Phaseolus hintonii
5
5 (2)
Phaseolus hygrophilus
1
Phaseolus jaliscanus
2 (0)
Botanic
Garden
Meise
BGCI GENESYS
34
10*
2
1130*
92
10
2
75
1
1
1
9
6
2
1
1
3
3
2
13
2
1
59
1
2
46
1
44*
4
5
2871*
154
Mexico
SINAREFI
101*
2
1
1
716*
243
1
2
1
1
1
18
24*
2
1
2*
604*
8
4
3
10
5
4
1
97
22
3
5
8
2
2
25
21
1
1
1
3
(continued)
4
Wild Beans (Phaseolus L.) of North America
119
Table 4.3 (continued)
Taxon
Phaseolus juquilensis
Phaseolus laxiflorus
Phaseolus leptophyllus
Phaseolus leptostachyus
Phaseolus leptostachyus
var. leptostachyus
Phaseolus lignosus
Phaseolus
longiplacentifer
Phaseolus lunatus
Phaseolus macrolepis
Phaseolus maculatifolius
Phaseolus maculatus
Phaseolus maculatus
subsp. maculatus
Phaseolus maculatus
subsp. ritensis
Phaseolus macvaughii
Phaseolus magnilobatus
Phaseolus marechalii
Phaseolus micranthus
Phaseolus microcarpus
Phaseolus mollis
Phaseolus neglectus
Phaseolus nelsonii
Phaseolus nodosus
Phaseolus novoleonensis
Phaseolus oaxacanus
Phaseolus oligospermus
Phaseolus opacus
Phaseolus
pachyrrhizoides
Phaseolus parvifolius
Phaseolus parvulus
Phaseolus pauciflorus
Phaseolus pedicellatus
Phaseolus perplexus
Phaseolus persistentus
Phaseolus plagiocylix
Phaseolus pluriflorus
Phaseolus polymorphus
USDAARS
CIAT NPGS
Mexico
SINAREFI
105
45
31 (23)
1 (0)
Botanic
Garden
Meise
BGCI GENESYS
43
2
1
144
43
1
1
62
25*
1
6223*
3
5
33
2
2
220
2
74 (46)
2 (0)
280*
1
18 (0)
1 (0)
10
23 (4)
8
3
2
4
5
1
29
1
1
2 (0)
1 (0)
17 (6)
4
1 (0)
32
2
19
2
1
3
2
15
1
1
2
1
3
5
3
12
4
76
2
2
1
5
2
1
1
1
1
13
2 (1)
6
1
21
21
8
3
1
31
20
2 (0)
12 (1)
9
1
2
7
3 (0)
1 (0)
12
1
3
2
3
46
15
2
12
1
1
1
3 (0)
1
3
1
2
1
1
7
1
(continued)
120
S. Dohle et al.
Table 4.3 (continued)
Taxon
Phaseolus polystachios
Phaseolus polystachios
subsp. polystachios
Phaseolus polystachios
subsp. sinuatus
Phaseolus purpusii
Phaseolus reticulatus
Phaseolus rotundatus
Phaseolus salicifolius
Phaseolus scabrellus
Phaseolus
scrobiculatifolius
Phaseolus smilacifolius
Phaseolus sonorensis
Phaseolus talamancensis
Phaseolus tenellus
Phaseolus teulensis
Phaseolus texensis
Phaseolus trifidus
Phaseolus tuerckheimii
Phaseolus venosus
Phaseolus vulgaris
Phaseolus vulgaris var.
aborigineus
Phaseolus xanthotrichus
Phaseolus xolocotzii
Phaseolus zimapanensis
USDAARS
CIAT NPGS
3
57 (1)
7 (0)
Mexico
SINAREFI
Botanic
Garden
Meise
3
3
2
1
1 (0)
2
1 (0)
2
1 (0)
2 (0)
1
1 (0)
13
2
1032 176 (156)
112 (81)
41
8 (3)
8
8 (4)
6
1
BGCI GENESYS
2
8
1
10
1
2
1
1
1
1
4
2
1
1
4
2
1
3995*
12
1
1
6
99
7
33*
2
83092*
225
11
1
7
1
55
1
24
2
Data from CIAT (2017); USDA-ARS National Plant Germplasm System (2017a), R. González
Santos personal communication (2017), Botanic Garden Meise (2017), BGCI (2017), and Data
Providers and the Crop Trust (2017)
CIAT, USDA-ARS NPGS, and Botanic Garden Meise databases were queried only for accessions
noted as wild; Mexico SINAREFI, BGCI, and GENESYS database counts for P. acutifolius, P.
coccineus, P. dumosus, P. lunatus, and P. vulgaris are likely to include cultivated forms and are
marked with an “*.” The GENESYS database includes information from over 400 institutes,
mainly in the CGIAR, USDA-ARS NPGS, and European genebank systems (Data Providers and
the Crop Trust 2017) and thus replicates the information from CIAT and USDA-ARS NPGS. For
USDA-ARS NPGS, counts of currently available accessions are given in parentheses
Moreover, many of these accessions are unavailable to researchers due to a lack of
adequate seed for distribution.
Ramírez-Villegas et al. (2010) assessed the comprehensiveness of major ex situ
collections with regard to taxonomic, geographic, and environmental or ecological
niche gaps (i.e., the degree of representation of the range of climates that wild
4
Wild Beans (Phaseolus L.) of North America
121
Phaseolus species occupy). Of 85 assessed taxa, they assigned 48 (56.5%) a high
priority for further collecting due to being completely absent or highly underrepresented in major genebanks. Seventeen taxa were assigned medium priority for further collecting, 15 low priority, and only 5 assessed as adequately represented ex
situ. Geographic hotspots for further collecting were concentrated in central Mexico,
although various narrow endemic species occurring in other parts of the country
were also prioritized (Ramírez-Villegas et al. 2010). Gap analyses for close relatives
of common and lima bean performed more recently corroborated these results
(Castañeda-Álvarez et al. 2016).
4.6
Final Remarks
The wild relatives of the five domesticated species of Phaseolus beans are clearly a
very diverse group of plants extending widely across the New World. The majority
of these species appear to be fairly well resolved and understood with regard to their
taxonomy and distributions. Further research is still needed to clarify the identities
and relationships among an additional dozen or more taxa, particularly with regard
to narrow endemic species.
Aside from the progenitor of common bean and a few studies involving other
relatives, the wild members of the bean crop genepools are still largely unexplored,
despite indications that novel useful diversity may be found within them. Given the
relative sensitivity of common bean to heat, drought, and other effects of climate
change, numerous promising wild species, as well as domesticates such as P. acutifolius, may play an increasing role as contributors of valuable genetic resources to
the crop. Contributions from wild relatives to improving agronomic and
market-related traits in the other domesticated species would also be valuable to
increasing their importance worldwide.
But these contributions can only be made if these resources are well conserved
and available to plant breeders and other researchers. A number of wild Phaseolus
are rare endemics that are threatened in their natural habitats and are insufficiently
protected in situ. Aside perhaps from the progenitors and a few other species, the
wild taxa are also largely underrepresented in major genebanks. Further efforts to
enhance protection of vulnerable species in their natural habitats, and further collecting to fill critical gaps in germplasm collections, are highly warranted.
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