Chapter 7

Prunus

Daniel Potter

7.1 Description and Distribution

Prunus L. (Table 7.1) comprises roughly 200 species,
including all of the economically important crop species known as stone fruits almonds, apricots,
cherries, peaches, and plums as well as many ornamental species and species cultivated or harvested
from the wild for timber and medicinal purposes.
Morphological descriptions are provided by Rehder
(1940), Kalkman (1965), Wilken (1996), and Bortiri
et al. (2006). Members of the genus are deciduous or
evergreen trees or shrubs with alternate, simple leaves
with toothed or entire margins and deciduous stipules.
Nearly all species bear glands on the leaves, but the
details of their morphology vary considerably among
species. These are generally present in one to several
pairs, but are occasionally solitary or absent, they may
be found on the petiole or on the undersurface or the
margin of the blade, usually near the base, they range
from quite prominent to relatively inconspicuous, and
they may be flat, hollow, or cushion-shaped (Kalkman
1965). The function of these glands has not been
determined.
The inflorescence in Prunus varies from a solitary
flower to an umbel-like cluster or a raceme, which may
or may not bear leaves on the peduncle. The radially
symmetrical flowers have a well-developed hypanthium, whose shape varies from campanulate to tubular, with five sepals, five petals that vary in color from
white to pink or red, 15 or more stamens, and a single
simple pistil (composed of one carpel) with a superior

ovary. The fruit is a drupe. The base haploid chromosome number for Prunus is x 8 (Raven 1975). Like
many other members of Rosaceae, species of Prunus
produce significant amounts of both the sugar alcohol
sorbitol, which serves as the primary transport carbohydrate in these plants (Zimmermann and Ziegler
1975; Moing et al 1997), and cyanogenic glycosides,
which impart a characteristic acrid odor to crushed
vegetative portions and toxicity to the seeds of many
species (Wilken 1996). Members of the genus exhibit
a range of breeding systems; gametophytic selfincompatibility has been documented for several species, and polyploidy and interspecific hybridization
are both common.
Prunus occurs in a variety of habitats, from forests
to deserts, and across altitudinal ranges from sea level
to alpine zones. The genus is most abundant in the
temperate zone of the Northern Hemisphere and is
widely distributed in North America, Europe, and
northern Asia. This, combined with the fact that all
of the cultivated species of global economic importance originated and are primarily grown in temperate
regions, has led to the perception, even among many
botanists, that Prunus is an exclusively north temperate genus. In fact, however, about 75 species have
tropical and subtropical distributions, including about
4550 species in South and Southeast Asia, about 25
in Central and South America, and one or two in subSaharan Africa (Kalkman 1965).

7.2 Classification and Phylogeny

D. Potter (*)
Department of Plant Sciences MS2, University of California,
1 Shields Avenue, Davis, CA 95616, USA
e-mail: dpotter@ucdavis.edu

Prunus has been variously lumped and split by different taxonomists over the last several centuries
(reviewed by Wen et al. 2008), and as many as seven

C. Kole (ed.), Wild Crop Relatives: Genomic and Breeding Resources, Temperate Fruits,
DOI 10.1007/978-3-642-16057-8_7, # Springer-Verlag Berlin Heidelberg 2011

129

130

D. Potter

Table 7.1 Summary of Rehders (1940) classification of the

genus Prunus L. and subsequent modifications thereof, with
placements of representative cultivated and wild species, including all species discussed in this chapter, indicated
Subgenus Prunus (treated as Prunophora Focke by Rehder):
plums and apricots
Section Prunus (treated as Euprunus Koehne by Rehder):
Eurasian plums
Representative species: P. cerasifera Ehrh., P. domestica
L., P. insititia L., P. salicina Lindl., P. simonii Carr.,
P. spinosa L.
Section Piloprunus Masona
Representative species: P. texana Dietr.
Section Prunocerasus Koehne: North American plums
Representative species: P. alleghaniensis Porter,
P. americana Marshall, P. geniculata R. M. Harper,
P. maritima Marshall
Section Armeniaca (Lam.) Koch: apricots
Representative species: P. armeniaca L., P. mandshurica
(Maxim.) Koehne, P. mume Siebold & Zucc.
Section Penarmeniaca Masona
Representative species: P. andersonii Gray
Subgenus Amygdalus (L.) Focke: peaches and almonds
Representative species: P. davidiana (Carr.) Franch.,
P. dulcis (Mill.) D. A. Webb., P. fenzliana Fritsch, Prunus
ferganensis (Kost. & Rjab.) Y.Y.Yao, P. kansuensis Rehder,
P. mira Koehne, P. persica (L.) Batsch, P. tenella Batsch,
P.webbii (Spach) Vierh.
Subgenus Cerasus Pers.: cherries
Section Cerasus (treated as Eucerasus Koehne by Rehder)
Representative species: P. avium L., P. cerasus L.,
P. fruticosa Pall.
Section Microcerasus Webb.
Representative species: P. glandulosa Thunb.,
P. tomentosa Thunb.
Section Pseudocerasus Koehne: flowering cherries
Representative species: P. canescens Bois., P. incisa
Thunb., P. lannesiana E. H. Wilson, P. nipponica
Matsum., P. serrulata Lindl., P. yedoensis Matsum.
Section Lobopetalum Koehne
Representative species: P. dielsiana Schneid.
Section Phyllocerasus Koehne
Representative species: P. pilosiuscula Koehne
Section Mahaleb Focke
Representative species: P. mahaleb L., P. pennsylvanica L.
Section Phyllomahaleb Koehne
Representative species: P. maximowiczii Rupr.
Subgenus Emplectocladus (Torr.) Sargenta
Representative species: P. fasciulata Gray
Subgenus Padus (Moench) Koehne: bird-cherries
Representative species: P. maackii Rupr., P. napaulensis
(Ser.) Steud., P. padus L., P. serotina Ehrh., P. virginiana L.
Subgenus Laurocerasus Koehne: laurel-cherries
Section Laurocerasusb
(continued)

Table 7.1 (continued)

Representative species: P. africana (Hook. f.) Kalkm.,
P. laurocerasus L., P. lusitanica L
Section Mesopygeum (Koehne) Kalkm.b
Representative species: P. arborea (Bl.) Kalkm.
Unnamed section primarily tropical America, some
North Americab
Representative species: P. ilicifolia (Nutt.) Walp.,
P. integrifolia (C. Presl) Walp
a

Following Mason (1913)

Following Kalkman (1965)

different genera have been recognized in this group

(e.g., Takhtajan 1997). Currently, however, the most
widely accepted classification of Prunus is that of
Rehder (1940), who adopted a broad interpretation of
the genus and divided it into five subgenera that are
further split into sections, most of which correspond to
Old World and New World groups. Additional infrageneric taxa proposed by Mason (1913) and Kalkman
(1965) have also been widely accepted (Table 7.1).
Many of the tropical Old World species are sometimes
classified under the genus Pygeum, but these were
all transferred to Prunus subgenus Laurocerasus by
Kalkman (1965; see Table 7.1).
To date, infrageneric classifications have emphasized morphological characters such as presence or
absence of a sulcus on the fruit, the number of axillary
buds on twigs, and features of the inflorescence.
Molecular phylogenetic analyses of the genus conducted over the last decade (e.g., Bortiri et al. 2001,
2006; Lee and Wen 2001; Wen et al. 2008), however,
have revealed that many previously recognized subgenera and sections are not supported as monophyletic
(Fig. 7.1) and many of the taxonomically important
characters exhibit considerable homoplasy. In other
words, traits considered diagnostic for particular
groups have evolved more than once within the
genus and some of them likely arose multiple times
as adaptations to special habitats, such as the presence
of dry fruits in species of arid regions (Bortiri et al.
2006). These findings suggest that a new infrageneric
classification for Prunus is needed.
The overall consensus is that there are several
major clades within Prunus: one comprising species
of Maddenia and subgenera Padus and Laurocerasus,
another comprising the most members of subgenera
Cerasus, and yet another comprising primarily members of subgenera Prunus and Amygdalus (Fig. 7.1).

Fig. 7.1 Schematic representation

of current understanding of
phylogenetic relationships within
Prunus, based on several recently
published studies (Lee and Wen
2001; Bortiri et al. 2001, 2006; Wen
et al. 2008). Polytomies indicate
cases in which analyses to date have
not been able to resolve the
branching order among lineages.
Subgeneric names refer to those
listed in Table 7.1

131
Other
Rosaceae

7 Prunus
Maddenia,
some Laurocerasus some Laurocerasus
(incl. some Pygeum), (incl. some Pygeum),
some Padus
some Padus

Many relationships within the genus remain poorly

resolved to date, however, due to a combination of
limited taxon sampling, especially for the tropical
species, and the lack of strong support for some
nodes (Fig. 7.1). The last phenomenon, in turn, results
from a combination of lack of sufficient variation in
sequence data, homoplasy within individual data sets,
and conflict among data sets, especially nuclear ribosomal DNA internal transcribed spacer (ITS) vs. chloroplast DNA (cpDNA) regions, which suggest
different placements for most members of subgenus
Cerasus (Bortiri et al. 2001, 2006; Lee and Wen 2001;
Wen et al. 2008). Some analyses have demonstrated
support for particular infrageneric groups, including
subgenera Amygdalus and Emplectocladus and section
Prunocerasus (Shaw and Small 2005), but differences
in taxon and sampling and relationships among the
different studies conducted so far preclude definitive
decisions about the status of these taxa. Because of its
widespread distribution, Prunus provides an excellent
system in which to examine historical biogeography of
temperate and tropical regions of both the Old and
New Worlds. Analyses to date (e.g., Bortiri et al.
2006; Wen et al. 2008) indicate multiple New World
Old World disjunctions within the genus, but, again,
more thorough sampling and better resolved phylogenies are needed to provide a full understanding of
these patterns.

Most Cerasus

Prunus, Amygdalus,
Emplectocladus,
sect. Microcerasus

The position of Prunus within Rosaceae has varied

among taxonomic treatments over the last 50 years
(reviewed in Potter et al. 2007). In what was until
recently the most widely used classification of the
family, Schulze-Menz (1964) recognized four subfamilies; Prunus sensu lato was treated as the largest
genus in subfamily Amygdaloideae, which also
included the genera Maddenia Hook. f. Thomson,
with 45 Asian species, Prinsepia Royle, with 34
Asian species, and the monotypic Oemleria Reichenb,
from western North America, the members of all of
which produce drupes. All but the last of these were
also included in tribe Pruneae by Hutchinson (1964),
who did not recognize subfamilies within Rosaceae.
Takhtajan (1997) recognized 12 subfamilies in Rosaceae; his Amygdaloideae included the aforementioned genera plus Exochorda Lindley, with 15
Asian species that produce capsules, but, like the
other genera mentioned, have a base chromosome
number of x 8 (Raven 1975).
Recent phylogenetic studies at both the generic (see
Fig. 7.1) and familial (see Fig. 7.2) levels have
resulted in modifications of these schemes, however.
The combination of a unicarpellate gynoecium that
develops into a drupe and the base chromosome number
of x 8 are synapomorphies for Prunus (Bortiri et al.
2006). Both Maddenia and Pygeum are nested within
Prunus and, while Exochorda, Oemleria, and Prinsepia

132
Rosaceae

form a clade, it is not the sister clade to Prunus sensu

lato. As a result, in the latest infrafamilial classification for Rosaceae (Potter et al. 2007; Table 7.2), based
on phylogenetic analyses of sequences from multiple
chloroplast and nuclear genes and incorporating nonmolecular characters, Prunus (including Maddenia)
was placed by itself in tribe Amygdaleae, while the
other three aforementioned genera were classified in
tribe Osmaronieae; both of these tribes were, in turn,
classified within an expanded subfamily Spiraeoideae.
In summary, while recent phylogenetic analyses
support Rehders (1940) broad circumscription of
Prunus indeed, they favor an even broader concept
that includes Maddenia they have not supported
monophyly of all of the currently recognized infrageneric taxa. Because a robust and thoroughly sampled
phylogeny for the genus is not yet available, however,
it is premature to propose a new, phylogenetically
based, infrageneric classification for Prunus. Such a
phylogeny will also be required to gain a complete
understanding of patterns of historical biogeography
and character evolution across the genus (Bortiri et al.
2006). The thorough and careful morphological studies of past workers and the resulting classifications
(e.g., Mason 1913; Rehder 1940; Kalkman 1965)
provide an excellent framework and a solid foundation
for future classifications, in which modifications can
be made to recognize only groups strongly supported
as monophyletic.

Pyrus
Malus

Gillenia

Sorbaria

Spiraea

Exochorda

Kerria

Prunus

Neillia

Spiraeoideae
Lyonothamnus

Rosa
Fragaria

Rubus

Rosoideae

Dryadoideae

Dryas

Other
Rosales
(Moraceae,
Rhamnaceae,
etc.)

Fig. 7.2 Schematic representation of

current understanding of phylogenetic
relationships in Rosaceae, based on
results presented by Potter et al. (2007)
with the circumscriptions of the three
subfamilies included in their
infrafamilial classification indicated.
Polytomies indicate cases in which
analyses to date have not been able to
resolve the branching order among
lineages

D. Potter

7.3 Diversity of Wild and Cultivated

Species of Prunus
Not surprisingly, given the large size and wide distribution of the genus and the fact that many of the
species exhibit one or more features of potential
value to people (i.e., high quality timber, beautiful
flowers, and/or edible fruit), Prunus includes species
with varying degrees of economic importance, from
exclusively wild species that are not used by people
through wild species that are sometimes cultivated and
are currently, or were historically, locally important as
sources of food, timber, or medicine, to true domesticates that are major crop plants. In addition to their
uses for food and timber and as ornamentals, medicinal uses are reported for a number of Prunus species.
The major cultivated species of Prunus are almond
(P. dulcis), peach (P. persica), sweet cherry (P. avium),
sour cherry (P. cerasus), European plum (P. domestica),
Japanese plum (P. salicina), and apricot (P. armeniaca).
Most ornamental flowering cherries belong to section
Pseudocerasus. Together, these species represent a
broad cross-section of the phylogenetic diversity of
Prunus (Table 7.1; Fig. 7.1). Each of the major domesticated species of Prunus shares its basic common
name with a number of wild and minor cultivated
species e.g., desert almond (P. fasciculata);
desert peach (P. andersonii), Manchurian apricot

7 Prunus
Table 7.2 Summary of infrafamilial classification of
Rosaceae by Potter et al. (2007)
Subfamily Rosoideae
No tribal placement:
Filipendula Adans., Rubus L., Rosa L.
Tribe Colurieae
Representative genera: Fallugia Endl., Geum L.
Tribe Potentilleae
Representative genera: Fragaria L., Potentilla L.
Tribe Sanguisorbeae
Representative genera: Agrimonia L., Sanguisorba L.
Subfamily Dryadoideae
Representative genera: Cercocarpus H. B. & K., Dryas L.,
Purshia DC.
Subfamily Spiraeoideae
No tribal placement:
Gillenia Moehcn., Lyonothamnus A. Gray
Tribe Amygdaleae
Representative genus: Prunus L.
Tribe Kerrieae
Representative genera: Kerria DC., Rhodotypos Sieb. &
Zucc.
Tribe Osmaronieae
Representative genera: Exochorda Lindl., Oemleria
Reichenb.
Tribe Neillieae
Representative genera: Neillia D. Don, Physocarpus
Maxim.
Tribe Pyreae
Representative genera: Lindleya H. B. & K., Malus Mill.,
Pyrus L.
Tribe Sorbarieae
Representative genera: Adenostoma Hook. & Arn.,
Sorbaria A. Braun
Tribe Spiraeeae
Representative genera: Aruncus Adans., Spiraea L.

(P. mandshurica), ground cherry (P. fruticosa),

black cherry (P. serotina), beach plum (P. maritima) which may or may not be closely related to
the major crop (Table 7.1).
Although they may be very clear in the case of
some major domesticates, the distinctions among
wild, cultivated, and domesticated taxa are often at
least somewhat ambiguous, and Prunus exhibits several features that make it especially challenging to
draw these distinctions. First, in woody perennials
with long generation times, the effects of human selection are not always as dramatic and obvious as they are
in many annual crop plants. Second, as noted above,
due to the large number of species in the genus, many
of which share basic features that are of interest to
humans, and its widespread distribution, there exists a

133

continuum of conditions from fully wild populations

to fully domesticated forms, not just across the genus,
but sometimes within a single species; the use of many
species and hybrids as rootstocks also contributes to
this phenomenon. Third, cross-compatibility among
species, especially closely related ones, has allowed
interspecific hybridization to play an important role in
breeding efforts, such that some cultivars include
genetic contributions from more than one naturally
occurring species.
Pandey et al. (2008) surveyed the wild and
cultivated species of Prunus available in India, where
considerable genetic diversity of the genus is found in
the Himalayan region and, to a lesser extent, at higher
elevations farther south (peninsular India). They documented the presence of 29 species used for food, 12
used as rootstocks, and 14 used medicinally; they also
mentioned the uses of several species as ornamentals.
These lists included a large number of native and
introduced wild species as well as the major cultivated
species of Prunus. They concluded that valuable
genetic diversity was present in all of the following
categories of material: cultivated species with high
regional and local diversity (e.g., P. persica), native
species that already exist in semi-domesticated forms
in some areas (e.g., P. napaulensis), and native wild
species with potential for domestication and worthy of
further investigation (e.g., P. tomentosa).
In some cases, multiple stages of domestication
may be observed within a single species. For example,
cultivated sweet cherry is Prunus avium, a species that
also occurs wild in Europe and North Africa and is
highly valued as a timber tree (Vaughan et al. 2007).
Browicz and Zohary (1996) explored the effects of
domestication on species of Amygdalus, using morphologically based taxonomic studies. They noted, as
many other authors have, the high frequency of naturally occurring hybrids within the group and pointed
out the potential value this has for transferring valuable traits from wild to cultivated species via classical
breeding methods. They further examined infraspecific variability in P. dulcis (which they treated as
A. communis L.), which they designated a crop complex because it comprises multiple categories of materials, including truly wild populations in the eastern
Mediterranean region, domesticated forms distinguished by sweet, non-poisonous seeds, and fruits
that are larger and have thinner endocarps than wild
forms and related wild species, and escapes from

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cultivation from the Mediterranean into Southwest and

central Asia.
As discussed above, there have been several recent
molecular phylogenetic studies aimed at resolving
relationships across Prunus (e.g., Aradhya et al.
2004; Bortiri et al. 2006; Wen et al. 2008). These
studies have, in general, confirmed ideas based on
morphology and crossing studies about the wild relatives of the major cultivated species. Lack of thorough
sampling, on the one hand, and the lack of phylogenetic resolution, on the other, have, however, precluded definitive tests of hypotheses about specific
wild progenitors to individual domesticated species.
Thus, because the aforementioned studies sampled
only one accession each of five or fewer species
other than P. dulcis and P. persica, and none of
them included P. fenzliana, it was not possible to test
Ladizinskys (1999) hypothesis that P. fenzliana is the
most likely wild ancestor to cultivated almond. However, a recent study based on nuclear and chloroplast
simple sequence repeat (SSR) markers (Zeinalabedini
et al. 2009), which focused on subgenus Amygdalus,
did point towards a close relationship between
P. fenzliana, represented by four accessions, and
P. dulcis, represented by 39.
Zohary (1992) hypothesized, based on cytogenetic
and morphological evidence, that the hexaploid
P. domestica is an autopolyploid derived solely from
P. cerasifera, which exhibits several ploidy levels,
rather than an allopolyploid derived from hybridization between diploid P. cerasifera and tetraploid
P. spinosa (Weinberger 1975). Subsequently, however, restriction site analyses of ribosomal RNA
genes suggested that P. spinosa itself is a hybrid, of
which P. cerasifera is one parent (Reynders-Aloisi
and Grellet 1994; Okie and Hancock 2008). Phylogenetic analyses of nuclear and cpDNA sequence data
indicate a very close relationship among all three
species, also supporting the allopolyploid origin
hypothesis (Bortiri et al. 2001). Interspecific hybrids,
including a variety of simple and complex hybrids
between wild and cultivated Prunus species, have
also been important as rootstocks for the various Prunus fruit crops (Bouhadida et al. 2007). Bouhadida
et al. (2007) used a polymerase chain reaction
restriction fragment length polymorphism (PCRRFLP) approach with several regions of cpDNA to
confirm the identity of the maternal parents of many
of these hybrids.

D. Potter

7.4 Use of Wild Species in Crop

Improvement Efforts
There is considerable variation in the degree to which
wild species have been important in the history of
breeding of particular crop species within Prunus.
Wild species have probably been most important
in the history of development of plums (Okie and
Hancock 2008). Okie and Hancock (2008) described
Luther Burbanks use of the Chinese species P. simonii
and several native North American plum species
(section Prunocerasus) to develop new cultivars of
diploid Japanese plum (P. salicina) for the California
plum industry and subsequent use of local species
native to the northeastern and southeastern US, respectively, to develop varieties adapted to growth in those
areas, though they note that most of those are no
longer available due to the demise of industries outside
of California. Okie and Hancock (2008) also discuss
the limited use of diploid P. cerasifera, a progenitor of
hexaploid European plum (P. domestica), in genetic
improvement of the latter.
At the other extreme, nearly all modern peach cultivars were derived from a small number of P. persica
cultivars, all of which trace their parentage back to
Chinese Cling, introduced to the United States from
China via England in 1850 (Hancock et al. 2008). This
narrow genetic base has stimulated interest in modern
use of wild species in peach breeding (Foulongne et al.
2003a).
In the case of cherries (subgenus Cerasus), there are
two major species that are cultivated for fruit: diploid
sweet cherry, P. avium, and tetraploid sour cherry,
P. cerasus L. (Iezzoni 2008). A third closely related
species, the tetraploid ground cherry P. fruticosa Pall.,
hybridizes freely with P. cerasus, contributing to
genetic and morphological diversity, as well as reduced
fertility, in sour cherry (Iezzoni 2008).
Dirlewanger et al. (2004) discussed the potential
value of several wild species closely related to
P. persica namely P. davidiana, P. kansuensis, and
P. mira as possible sources of resistance to several
important pests and diseases of peach. They further
pointed out that, although there is great potential for
transfer of traits among the many intercompatible species within Prunus, realization of that potential has
been limited, due primarily to the slowness of traditional breeding methods, but that recently developed

7 Prunus

genomic methods may allow greater use of the genetic

variability available in the genus and already conserved in the many germplasm collections for Prunus.
At the same time, Gradziel (2003) has demonstrated
that interspecific hybridization and backcrossing can
provide an effective means of introgressing desirable
genes from wild species into both peach and almond
cultivars. Wild species have been important in a few
cases as sources of disease resistance for Prunus
crop species, e.g., resistance to plum pox virus in
P. armeniaca derived from P. mandshurica (Ledbetter
2008), and resistance to cherry leaf spot in P. cerasus
derived from P. maackii and P. canescens (Iezzoni
2008). Hybrids resulting from a cross between
P. persica and P. davidiana were used to select for
resistance to powdery mildew, peach green aphid,
and plum pox virus (Kervella et al. 1998).
As a contrast to their usefulness as sources of disease and pest resistance, wild species may also serve
as hosts for diseases of cultivated species and thus
contribute to the continued presence of diseases in
areas where crops are grown. Carraro et al. (2002)
reported that several wild species of Prunus
P. spinosa, P. cerasifera, and P. domestica serve as
hosts for the phytoplasma that causes European stone
fruit yellows (EFSY), a disease of many cultivated
Prunus species, and the psyllid vector (Cacopsylla
pruni) that transmits the disease. Similarly, P. virginiana
serves as an alternate host for X-disease, a phytoplasma disease of cherries transmitted by leafhoppers
(Iezzoni 2008). Damsteegt et al. (2007) showed that 40
species and varieties of Prunus, including many wild
species native to the US, were susceptible to infection
by plum pox virus, which causes Shakra disease,
considered the most important viral disease of stone
fruits in Europe, and maintained infections through
repeated cycles of cold-induced dormancy over 4
years. The results suggested that many native and
introduced species may serve as reservoirs for this
serious disease, presenting a significant challenge to
eradication efforts.
The efficiency of transferring desirable traits can be
greatly enhanced by modern genetic and genomic
methods, including comparative mapping and
marker-assisted selection. Hybrids with wild species
have been important for genetic mapping studies in
Prunus, including mapping studies of cherry
(Emperor Francis  P. nipponica and  P. incisa;
also P. avium cv. Napoleon and P. nipponica;

135

Iezzoni 2008), peach (P. persica  P. ferganensis),

and peach and almond (P. cerasifera  (P. dulcis 
P. persica); Dirlewanger et al. 2003).
Dirlewanger et al. (2004) reviewed the status of
Prunus genetic maps as of 2004. At that time, the
Prunus reference map, constructed in a peachalmond
hybrid, included 562 markers covering 519 cM.
Using anchor markers from the reference map, 13
additional maps were constructed for other species of
Prunus, including two additional cultivated species
(P. armeniaca and P. avium) and three wild species
(P. cerasifera, P. davidiana, and P. ferganensis). The
availability of these maps has allowed mapping of
28 major genes affecting horticulturally important
characters in the different species, including genes
involved in the determination of fruit quality, phenological traits, and pest and disease resistance traits.
Comparison of these maps (Dirlewanger et al. 2004)
revealed essential colinearity among these diploid
Prunus species, all of which are members of one of
the three subgenera Amygdalus, Cerasus, or Prunus
(Table 7.1). The fact that the degree of synteny
observed between the Prunus genome and both component genomes of Malus (apple), a member of the
polyploid tribe Pyreae, was quite high, albeit lower
than within Prunus (Dirlewanger et al. 2004), suggests
that, not surprisingly, gene order and overall chromosomal structure has been conserved within subfamily
Spiraeoideae (Table 7.2), with the degree of rearrangement correlated to phylogenetic distance (Fig. 7.2).
A general problem with the use of wild species
as sources of desirable traits in breeding programs is
the concomitant introgression of unfavorable traits
(Quilot et al. 2004). Thus, in Prunus, as in other fruit
crops, transfer of disease resistance genes from
wild relatives may result in decreased fruit quality.
Foulongne et al. (2003a) demonstrated the potential
value of the Chinese species Prunus davidiana as
a source of genes that could be introgressed into
the peach genome using comparative genetic mapping of RFLP, SSR, and amplified fragment length
polymorphism (AFLP) markers in F1, F2, and BC2
generations resulting from a cross between the two
species. Subsequently, Foulongne et al. (2003b)
found quantitative trait loci (QTL) for resistance to
powdery mildew in hybrid and backcross generations
derived from a cross between the commercial peach
variety Summergrand and a member of the closely
related wild species P. davidiana. For nine of the 13

136

QTLs detected, the favorable allele was derived from

the wild species.
Quilot et al. (2004) reported the results of QTL analyses of fruit quality in P. persica (peach) based on an
advanced backcross population derived from a cross
between the commercial peach variety Summergrand
and a member of the closely related wild species
P. davidiana. They found QTLs for 24 physical and
biochemical fruit quality traits and identified some horticulturally desirable alleles in the wild species. They
identified three primary genomic regions where QTLs
with negative effects are located. They proposed that
future breeding efforts using P. davidiana should focus
on suppressing those chromosomal regions and on finemapping of regions in which QTLs with beneficial resistance and negative fruit quality effects are colocated.

7.5 Population and Evolutionary

Genetic Studies of Wild Species
While wild species have been valuable in the improvement of cultivated species, on one hand, the existence
of genetic tools for characterizing cultivated species
has facilitated evolutionary and population genetic
studies of wild species, on the other. Cross-species
transportability of molecular markers, such as SSR
primers, within Prunus, including both cultivated and
wild species, has been reported by multiple workers.
These include Vendramin et al.s (2007) report of 21
expressed sequence tag SSRs (EST-SSRs) isolated
from the peach fruit transcriptome that successfully
amplified PCR products in six other Prunus species,
five cultivated (P. dulcis, P. armeniaca, P. avium,
P. salicina, P. domestica) and one wild (P. ferganensis),
Rohrer et al.s (2004) use of SSR markers from 15
primer pairs originally developed in P. persica and
P. avium to examine phylogenetic relationships among
13 known wild species and several undetermined
wild accessions of North American plums (subgenus
Prunus, section Prunocerasus), and Pairon et al.s
(2008) use of microsatellite markers originally developed for various cultivated species to identify
genome-specific markers for the allotetraploid wild
species P. serotina.
Wild species of Prunus have been the subject of
numerous studies aimed at understanding the evolutionary histories and dynamics of populations. Jordano

D. Potter

and Godoy (2000) used random amplified polymorphic DNA (RAPD) markers to study population
genetic structure in Prunus mahaleb among seven
populations across an area of about 100 km2 in Parque
Natural de las Sierras de Cazorla in southeastern
Spain. They found evidence both for extensive gene
flow among populations and for a degree of isolation
by distance, which they attributed to the combined
effects of efficient long-distance dispersal by frugivorous birds and mammals and local fragmentation
resulting from vicariant factors including demographic bottlenecks due to high post-dispersal seed
and seedling mortality.
Mohanty et al. (2002) examined cpDNA diversity,
using a PCR-RFLP approach, among 25 wild populations of P. spinosa from forests across Europe. They
found 32 haplotypes, of which 10 were shared by
multiple populations and 22 were private. Overall, no
clear phylogeographic structure was detected, but
higher haplotype diversity in southern than northern
Europe was attributed to glacial refugia in the more
southerly locations.
Roh et al. (2007) used inter-SSR (ISSR) markers
and sequences of two cpDNA regions to clarify the
distinction between wild Korean plants referred to as
P. yedoensis and cultivated hybrid ornamental Yoshino
cherries from Japan, referred to as P.  yedoensis. They
concluded that the two are sufficiently distinct that
they should be treated as separate taxa.
Several studies have focused on wild populations of
P. avium in Europe. Frascaria et al. (1993) examined
isozyme variation among four populations of the species in France. They found no significant genetic structure within the populations and no significant
differentiation among them. They attributed these
results to the effects of human dispersal, perhaps combined with the limited time since the last glaciation in
the areas studied.
Mohanty et al. (2001) surveyed variation PCRRFLP patterns of cpDNA among 23 wild populations
of Prunus avium from across Europe and found a total
of 16 haplotypes, six of which were shared by two or
more populations and ten of which were unique. They
found no genetic structure among wild populations,
which they attributed to long-distance gene flow
among populations mediated by birds, mammals, and
humans. Subsequently, Panda et al. (2003) expanded
upon this study by surveying a total of 96 cultivars. In
their study, they found 16 haplotypes among wild

7 Prunus

populations and only three among cultivars, which

represented the most common three in the wild populations, indicating higher cpDNA diversity in wild
populations than in the cultivars, and providing information useful for developing germplasm conservation
strategies for the species.
Schueler et al. (2003), using seven microsatellite
markers originally developed in peach (Prunus persica), examined genetic structure in a natural population of wild P. avium in Germany, and found sufficient
variability in the markers to allow identification of
individual trees. They also demonstrated that genotyping of endocarps with their markers could be used to
identify the mother tree of dispersed seeds. Vaughan
and Russell (2004) developed primers for 14 microsatellite loci in cultivated P. avium. Genetic mapping
studies of their seven most polymorphic loci with
four from a previous study (Clarke and Tobutt 2003)
revealed that the 11 loci are genetically unlinked,
providing powerful tools for use in studies of population structure of wild forms of the species. Subsequently, Vaughan et al. (2007) used 13 of these loci
to examine patterns of spatial-genetic structure in two
wild populations of P. avium, one managed and one
unmanaged, in Britain. They found evidence of significant clonal reproduction and restricted gene dispersal
via both pollen and seed, leading to two recommendations that should help maintain genetic diversity of the
species: selective removal of mature trees from particular areas and establishment of minimum distances
(they suggested 100 m) between trees to be used as
sources of seeds for propagation.

7.6 Evolutionary Studies of SelfIncompatibility Genes

Prunus is one of the several genera in Rosaceae that
exhibits gametophytic self-incompatibility (GSI), in
which specificity of self-pollen rejection is determined
by a stylar component known to be an S-RNase (Ushijima et al. 1998) and its genetically linked pollen
component known to be an F-box protein, which, in
Prunus, has been named SFB (Ushijima et al. 2003).
Because of its importance in breeding and production
of fruit crops, considerable attention has been directed
to understanding in detail the mechanism and genetics
of GSI. Numerous S-RNase/pollenSF-box protein gene

137

pairs have been identified and sequenced from species

of Prunus, including both cultivated (P. armeniaca,
P. avium, P. cerasifera, P. cerasus, P. domestica,
P. dulcis, P. mume, P. salicina) and wild (P. lannesiana
var. speciosa Makino, P. spinosa, P tenella, P. webbii)
taxa. As a result of these efforts, Prunus has become
a model system in which to examine the evolution of
self-incompatibility at the molecular level.
Surbanovski et al. (2007) examined sequences of
SFB and S-RNase alleles in wild Prunus tenella, native
to the Balkan Peninsula. They found evidence for
positive selection on the sequences of S-RNase alleles
of this species, in contrast to results obtained for
P. lannesiana (Kato and Mukai 2004), P. dulcis, and
P. avium (Ma and Oliveira 2002). In addition, they
found that the amino acid sequence of the S-RNase
encoded by one of the alleles from P. tenella was
identical to one from P. avium, but that the
corresponding SFB alleles showed many differences
between the two species. They discussed their results
in terms of the models for evolution of GSI specificities in Prunus. Specifically, their results show that
the same pistil determinant (S-RNase) can tolerate
variability in the pollen determinant (SFB), suggesting
that the evolution of new GSI specificities is initiated
by mutations in the pollen-determinant genes.
Vieira et al. (2008) used phylogenetic analyses coupled with models of sequence evolution and estimates
of the age of Prunus based on calibrated molecular
phylogenies (Wikstron et al. 2001) to develop hypotheses about the evolution of GSI in Prunus. Their results
suggested that extant Prunus harbor only about a
third of the GSI specificities that would have been
present in their common ancestor, suggesting one or
more evolutionary bottlenecks during the evolution
of the genus, perhaps resulting from processes associated with speciation and/or domestication.
Several models have been proposed to explain
the generation of new alleles and, correspondingly,
new specificities, in a two-gene system of self-incompatibility. These include models that require selfcompatible intermediates (Uyenoyama et al. 2001),
dual-specificity intermediates (Matton et al. 1999),
and gradual accumulation of mutations while maintaining self-incompatibility (Chookajorn et al. 2004).
Implicit in all of these models is the tight linkage and
coevolution between the two loci involved, such that
mutations in one must be followed by compensatory
mutations in the other in order to restore or maintain

138

D. Potter

self-incompatibility. Intriguingly, none of these models has been completely supported by empirical data
from Prunus. In particular, lack of correspondence
between the phylogenies for the pistil and pollen determinants has suggested a role for recombination in
the evolution of new specificities (Nunes et al. 2006;
Tsukamoto et al. 2008).
The phylogenies presented in Fig. 7.3 illustrate
several striking and related features that have been
noted in recent studies of the evolution of S-RNase
and SFB genes in Prunus (Nunes et al. 2006; Suther-

S-RNase

land et al. 2008; Tsukamoto et al. 2008; Vieira

et al. 2008). First, the two phylogenies have some
similar characteristics: homoplasy is high (consistency
indices are low) in both data sets, support for the
relationships among lineages of both genes, especially
the deeper internal branches, is generally weak, and
many of those branches are quite short, suggesting that
early diversification of these genes may have occurred
rapidly in ancestral species, with subsequent lineage
sorting and/or recombination giving rise to the extant
alleles, as suggested by Tsukamoto et al. (2008).

PaS1
100
99
PdS11
PtS8
56
PaS13
62
PspS12
PdSb
PaS5
63
PmS1
PweS1
PspS10
PsSe
ParS4
PsSh
71
PaS2
PspS12
ParS1
59
ParS1
66
PdSd
PaS2
100
PsSg
100 PaS4
PcsfS10
PdoS9
76
PaS5
100 PsSd
64
PweS1
PcsfS9
PaS7
52
PspS7 1
100
PsSa
100 ParS17
100
PdoS6
PspS3 1
99
PspS7 1
86 PspS3 2
100
PaS4
PsS7
PdoS9
PaS6
100
PsSd
PaS7
PcsfS9
100 100 PaS12
PsSf
PdSk
77
PaS6
PdS12
76
PspS8
79 PcsfS3
PspS9
PaS13
74
PaS12
PdSd
PdSk 97 100 100
96
PcS26
PdS12
PmS7
PcsfS3
PsSf
PcS26
ParS4
PmS7
PspS8
PaS3
PsSc
100
PcS34
PspS9
PdSb
PdoS5
100 PsSa
PsSb
ParS17
100
PdoS6
PspS3 1
PdSc
51
PaS3
100
PspS3
2
94
PcS33
PsS7
53
100
PmS1
PcS33
ParS2
PspS10
ParS2
PsSe
100 PsSg
PsSh
PdSc
PcsfS10
PsSc
PsSb
PcS35
PdSa
50 changes
PaS1
100
71 PdS11
98
PtS8
100
PcS34
PdoS5

SFB

100

PcS35
PdSa
10 changes

Fig. 7.3 Comparison of relationships among S-RNase and SFB

alleles from wild and cultivated species of Prunus. Left: relationships among S-RNase alleles. Single most parsimonious
tree (2002 steps, ci excluding autapomorphies 0.3860,
ri 0.5065); based on alignment of 50 published sequences
with 747 characters, of which 239 were constant, 129 variable
but uninformative, and 379 parsimony-informative. Right: relationships among SFB alleles. One of five most parsimonious
trees (2,816 steps, ci excluding autapomorphies 0.4047,
ri 0.4886); based on alignment of 50 published sequences
with 1,158 characters, of which 343 were constant, 227 vari-

100

able but uninformative, and 588 parsimony-informative. Numbers on branches represent bootstrap support values. In the tree
at right, nodes marked with an asterisk were not present in the
strict consensus tree for the five most parsimonious trees.
Analyses were conducted as in Tsukamoto et al. (2008),
where Genbank accession numbers are listed for all sequences
except Pwe S1 RNase (DQ993660) and Pwe S1 SFB
(DQ993667). Pa P. avium; Par P. armeniaca; Pc P. cerasus;
P csf P. cerasifera; Pd P. dulcis; Pdo P. domestica; Pm P.
mume; Ps P. salicina; Psp P. spinosa; Pt P. tenella; Pwe P.
webbii

7 Prunus

Second, both genes show a pattern known as transspecific evolution (Richman et al. 1996), in which
alleles from individual species do not form monophyletic groups; i.e., the closest relatives of many alleles
are alleles from other species. This pattern may reflect
the role of balancing selection in the evolution of selfincompatibility specificities (Richman and Kohn
2000), resulting in retention of alleles through evolution over long periods of time and multiple speciation
events, although it has been shown that in Prunus, in
contrast to Solanaceae, the pattern of trans-specific
evolution may not be interpretable as evidence for
great age of specificities (Vieira et al. 2008). A related
phenomenon is that neither genes phylogeny is congruent with species phylogenies in Prunus (e.g.,
Fig. 7.1), and this lack of congruence has been
shown to be significant at all taxonomic levels within
the genus (Tsukamoto et al. 2008), while at least for
the S-RNase locus, alleles of Malus and Pyrus (both
members of tribe Pyreae) are phylogenetically distinct
from those of Prunus (Igic and Kohn 2001). This
pattern, like that of trans-specific evolution discussed
above, results from incomplete lineage sorting (Lu
2001) and indicates that, for the members of Rosaceae
sampled to date, coalescence of alleles has not
occurred below the level of the genus at either locus.
The third notable pattern is that the phylogenies of the
two genes are incongruent with one another, which
may reflect a role of intragenic recombination in the
evolutionary histories of the two genes, which otherwise would be expected to show congruent patterns of
relationship (Tsukamoto et al. 2008). Future studies
incorporating S-RNase and SFB sequences from additional wild species of Prunus, especially members of
subgenera Laurocerasus and Padus (Table 7.1;
Fig. 7.1) and representatives of other tribes in Rosaceae (Table 7.2; Fig. 7.2), are required to gain a more
thorough understanding of patterns and processes of
evolution of self-incompatibility in the genus and the
family.

7.7 Issues of Concern: Conservation

As is to be expected for such a large, diverse, and
widely distributed genus, Prunus species exhibit a
range of conservation statuses, from widely distributed
taxa that have become invasive following human dis-

139

persal to new environments to those with very

restricted distributions that are considered rare or
threatened, including one that is now endangered due
to overharvesting.
Due to their considerable economic importance,
many collections of cultivated and wild Prunus germplasm exist throughout the world. A search of the
Biodiversity Internationals Biodiversity Directory of
Germplasm Collections (Biodiversity International
2009) on June 30, 2009, retrieved 3,982 accessions
of Prunus classified as wild species at 40 institutions. When the type of germplasm was not restricted
to accessions classified as wild, the numbers were
60,168 accessions and 168 institutions. Inspection of
some of these records revealed that many wild taxa
were not explicitly designated as such and so were not
recovered by the first search. The largest single institution housing Prunus germplasm is the United States
Department of Agriculture (USDA) National Clonal
Germplasm Repository at Davis, CA; the Germplasm
Resources Information Network (GRIN) database
lists 108 taxa of Prunus for which accessions are
preserved there.
The GRIN database (USDA, ARS, National
Genetic Resources Program 2009) lists four taxa of
Prunus as rare and endangered. They are: P. africana
(African cherry), widely distributed in sub-Saharan
Africa, listed in CITES Appendix II and with one
accession preserved in the US National Germplasm
System, P. alleghaniensis (Allegheny plum), distributed
in the eastern US (Rehder 1940), listed by the Center
for Plant Conservation (CPC) and with four accessions
preserved in the US National Germplasm System,
P. geniculata (scrub plum), with a limited distribution
in Florida, listed by the CPC and on the Endangered list
of the US Fish and Wildlife Service and with one
accession preserved in the US National Germplasm
System, and P. maritima var. gravesii (Small) G. J.
Anderson, with a very limited distribution in Connecticut, listed by the CPC and with no accessions preserved
in the US National Germplasm System.
In addition to the species listed above, concern has
been raised about the conservation status of some
other taxa, including wild species as well as local
varieties of cultivated species, in certain regions.
Vivero et al. (2001) described ecology and ethnobotany of six species of Prunus that occur wild in
Andalusia, Spain, and proposed strategies to conserve
germplasm of wild populations and local varieties for

140

the three most economically important of those species, P. avium, P. mahaleb, and P. insititia. Among
their recommendations were designation of an area in
the Sierra Nevada for in situ germplasm conservation
and raising awareness of the importance of these
species and their conservation among forest workers
and managers and the general public. P. lusitanica ssp.
azorica was one of three taxa identified by Ferreira
and Eriksson (2006) as a target for conservation in
their proposed plan for conservation of forest tree
genetic resources in the Azores. This species was
selected due to its status as one of the most threatened
in the archipelago.
Perhaps of greatest concern purely from the point of
view of biodiversity conservation are the tropical
species of Prunus, which have received relatively little
attention from researchers to date and are poorly represented in germplasm collections, many of which occur
in areas where their habitats are threatened by anthropogenic factors such as logging, expansion of agriculture, and/or urbanization. In western New Guinea
(Papua Province, Indonesia), rapid deforestation is
threatening the habitats of several of the endemic species of Prunus (D. Potter, pers observ) and it is likely
that the same situation exists for many of the paleoand neotropical species. A recently initiated taxonomic
revision of Prunus for Colombia has so far revealed
three new species (Perez-Zabala 2007), all considered
by the author to merit conservation concern, two as
endangered and one as near threatened following
IUCN criteria (International Union for Conservation
of Nature and Natural Resources 2001).
One of the most interesting cases of an endangered
species of Prunus is P. africana, which is widely
distributed in montane regions of sub-Saharan Africa,
has been used traditionally by people throughout its
range for multiple purposes (Stewart 2003), and, at
least in some areas, is an important food source for
wildlife, including some rare and endangered species of
primates and birds (Fashing 2004). The discovery, in the
late 1960s, that bark extracts from this species were
effective in treating benign prostatic hyperplasia (Bombardelli and Morazzoni 1997) led to extensive international trade of the bark and herbal remedies prepared
from it, which in turn resulted in overharvesting of wild
trees (Cunningham and Mbenkum 1993), ultimately leading to the listing of the species in CITES Appendix II.
Several recent studies (e.g., Dawson and Powell 1999)

D. Potter

have examined the distribution of genetic diversity in this

species throughout its range, resulting in recommendations for conservation strategies. In addition, several studies (e.g., Cunningham et al. 2002; Stewart 2003; Fashing
2004) have called attention to the need for establishing
plantations of the species in order to reduce pressure on
wild populations.
P. africana presents an extremely challenging
problem for conservation. The medicinal value of the
species, the economic situations of local human inhabitants throughout much of its range, and the large gap
between the price paid for raw bark and that paid for
the final medicinal preparations tend to encourage
unsustainable wild-harvesting by local people, even
where this practice is in violation of local regulations
(Stewart 2003). Synthesis of the therapeutically active
components of the bark extracts has not been
attempted and is likely to be complicated and expensive (Stewart 2003), due to the fact that synergistic
interactions of multiple compounds are indicated in
the effectiveness of the extracts (Bombardelli and
Morazzoni 1997). Prospects for cultivation are poor
in many areas due to limited availability of appropriate
land (Stewart 2003). Studies of genetic diversity,
based on RAPD (Dawson and Powell 1999; Muchugi
et al. 2006) and SSR (Farwig et al. 2008) markers,
have revealed significant variation within populations
and have provided tools for identifying especially
diverse populations that should be prioritized for conservation, but in situ conservation efforts may be
undermined by a paradox pointed out by Fashing
(2004): while P. africana appears to require disturbance for successful regeneration (Kiama and Kiyiapi
2001), disturbance can be detrimental to the species,
either directly because of overharvesting that often
occurs when disturbance causes or results from
increased human access to an area, or indirectly due
to reduced genetic diversity resulting from forest fragmentation caused by human activity (Farwig et al.
2008). Recent efforts to include P. africana as a
model agroforestry species for participatory domestication (reviewed by Simons and Leakey 2004), in
which local small-scale farmers are engaged in the
process of identifying, cultivating, and improving
valuable germplasm selected from wild trees, thereby
alleviating pressure on wild populations, are encouraging and may represent the best hope for conservation of genetic diversity of this species.

7 Prunus

7.8 Issues of Concern: Invasive Species

At the other end of the spectrum from rare and
endangered species are those that are relatively common
in their native ranges and have also become invasive in
areas to which they are not native. Examples from
Prunus include P. laurocerasus, native to southeastern
Europe and Asia Minor, in western Europe
(Hattenschwiller and Korner 2003) and in the Pacific
Northwest of North America (Evergreen 2010) and
North American P. serotina in Europe (e.g., Godefroid
et al. 2005). P. cerasifera, native to southeastern Europe and naturalized in California, is included by the
California Invasive Plant Council in its Invasive Plant
Inventory but rated as limited because its ecological
impacts are considered minor on a statewide level (California Invasive Plant Council 2009). Concerns have
also been raised about northern European P. padus
in Alaska (Alaska Natural Heritage Program 2006).
Hattenschwiller and Korner (2003) studied the
effects of elevated CO2 levels on growth rates of
P. laurocerasus, whose abundance in the understoreys
of Swiss forests, where the species is not native, had
raised concern about its potential to become invasive.
They found that P. laurocerasus seedlings grown
in elevated CO2 concentrations for three growing
seasons showed an average of 56% greater biomass
than plants grown at ambient CO2 levels, while native
species showed a range of responses to elevated CO2.
They concluded that increases in atmospheric CO2
levels, an element of current and projected future
global change, may facilitate naturalization and spread
of this non-native species, thereby contributing to
another component of global change, biotic invasions.
P. serotina (black cherry), an allotetraploid species
native to North America, was introduced to Europe,
especially Germany, Belgium, and the Netherlands,
for various purposes beginning several hundred years
ago. Starfinger et al. (2003) provided a fascinating
account of the history of the introduction of this
species to Europe and how perceptions of it have
changed over the centuries. First introduced as an
ornamental in the seventeenth century, it was later
widely planted as a forest timber tree beginning
in the late eighteenth century; when its value as a
timber tree was shown to be questionable, it began to
be used for non-timber purposes, such as improving
litter due to the low C/N ratio in leaves of the species.

141

It had become naturalized in western Europe by

the mid-twentieth century, and by the late twentieth
century, it was widely considered an invasive forest
pest (Starfinger et al. 2003; Pairon et al. 2006; ClossetKopp et al. 2007).
Recent studies have been undertaken to investigate
and characterize the factors that contribute to the invasiveness of this species in European forests; these
studies have included investigations of the effects of
landscape structure (Deckers et al. 2005), ecological
variables (Godefroid et al. 2005; Verheyen et al.
2007), reproductive traits (Pairon et al. 2006), disturbance history (Chabrerie et al. 2008), and propagule
pressure (Vanhellemont et al. 2009) on the ability of P.
serotina to invade forests.
Godefroid et al. (2005) investigated the ecological
factors that affect the abundance of P. serotina in
forests in Belgium. Species richness in the herb
layer was negatively correlated with the abundance
of P. serotina in the shrub layer. Slope and light
intensity were the only abiotic factors measured that
explained significant portions of the variation in
P. serotina abundance. The light intensity results suggested that P. serotinas response to light intensity
changes as the tree matures: seedlings showed a positive response to 5880% of full light and a negative
response to lower light intensities, while saplings
showed the reverse trend. In addition, further growth
of saplings to maturity and seed production again
requires high light intensities, but saplings can adopt
a sit-and-weight strategy, forming a long-lived
seedling bank until a canopy light gap occurs (ClossetKopp et al. 2007). Thus, the establishment and persistence of P. serotina depends on opening of light
gaps in the canopy.
Studying the same system, Pairon et al. (2006)
investigated sexual regeneration traits of P. serotina
growing in a Belgian pine plantation, in order to gain a
better understanding of how those traits might affect
the invasiveness of the species. They found that fruit
production was high in spite of low fruit/flower ratio,
because of the large number of flowers produced per
tree. Seeds fell into two size classes: large (the majority) seeds, which are gravity-dispersed, and smaller,
bird-dispersed seeds, resulting in thorough coverage of
the area by seeds. While seed germination and seedling survival rates were low, the high seed density
means that each year, the entire forest floor is covered
with seedlings; the high survival rate of saplings helps

142

ensure maintenance of the population. Thus, the ecological and reproductive characteristics of P. serotina
seem to have pre-disposed it to be highly successful as
an invader in European forests.
Vanhellemont et al. (2009), noting that most studies
of the invasiveness of P. serotina in western Europe,
including those discussed above, had been conducted
in areas where the species had been intentionally
introduced, which were subject to considerable
anthropogenic disturbance, and where propagule pressure was high, undertook a study to address the question of whether or not P. serotina acts as an aggressive
invader in areas within its potential range that had not
yet been heavily invaded. They focused on a forest
reserve in central Belgium that met those criteria.
They found that the spread of P. serotina in this
reserve had slowed since the first establishment of
the species there around 1970s and subsequent further
spread in the 1980s, presumably from seedlings produced by the first arrivals. They assumed that the slowdown was due to lack of disturbance creating light
gaps needed for seedling establishment. At the same
time, they found no evidence that P. serotina was
inhibiting the regeneration of native understorey species in this forest. They concluded that P. serotina
could not be considered an aggressive invader in the
study area, but they pointed out that future disturbance
events opening up the canopy could result in accelerated spread and invasion of the species.

7.9 Summary and Conclusions

Prunus is a large genus of tremendous economic and
ecological importance worldwide. The group includes
several major fruit crop species, a large number of minor
cultigens and species collected from the wild for a range
of uses, and many wild species that have been used as
rootstocks for cultivated taxa and in their genetic
improvement. The economically important species represent several phylogenetic lineages within the genus.
Phylogenetic studies have confirmed some aspects of
past taxonomic treatments and hypotheses about the
origins and placement of particular crop species and
challenged others. To date, poor sampling of some
lineages, especially those including the approximately
75 species native to the New and Old World tropics, and
weak resolution of some relationships across the genus,
have precluded generation of a new phylogenetically

D. Potter

based classification. Nonetheless, ongoing efforts in

multiple labs throughout the world, some focusing on
relationships across the entire genus, others on particular
species and their closest relatives, are leading to a thorough understanding of phylogeny of Prunus, which will
allow robust investigations of the historical biogeography of the genus, the evolution of particular genes and
traits, and the interplay of natural and human selection in
shaping the extant variation in this group.
Wild species of Prunus have been important in the
histories of several cultivated species, and modern
methods such as comparative genetic mapping and
marker-assisted selection should help to facilitate the
transfer of desirable traits and to minimize the concomitant transfer of undesirable traits, from wild to
cultivated species. At the same time, there is considerable interest in wild species in their own right, and
Prunus provides an excellent example of a system in
which a complementary and synergistic relationship
exists between studies of cultivated species and those
of wild relatives. Tools developed for characterizing
cultivated taxa have been tremendously useful in ecological and evolutionary genetic studies of wild species,
while the results of the latter have provided valuable
information for crop improvement efforts, and for
understanding the economically significant issues associated with the spread of invasive species and the conservation of rare and potentially valuable taxa. Future
efforts in all of the aforementioned areas should continue. Particular attention should be paid to the tropical
species, which have received relatively little attention
from researchers to date and which may be among the
most threatened in terms of conservation status.

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