Small Fruits Review

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Thornlessness in Blackberries
A Review

M. A. Coyner, Robert M. Skirvin, M. A. Norton & A. G. Otterbacher

To cite this article: M. A. Coyner, Robert M. Skirvin, M. A. Norton & A. G. Otterbacher (2005)
Thornlessness in Blackberries, Small Fruits Review, 4:2, 83-106, DOI: 10.1300/J301v04n02_09

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 Thornlessness in Blackberries:
A Review
M. A. Coyner
Robert M. Skirvin
M. A. Norton
A. G. Otterbacher

ABSTRACT. There are several known sources of thornlessness for

blackberries. Some of these have been used as parents and their genetics
are quite well understood; others remain relatively unknown. The source
and nature of thornlessness in blackberry has been reviewed by George
Darrow (1937), Jack Hull (1968), and Derek Jennings (1986). In this pa-
per we review the literature on thornlessness, discuss the status of the
thornless parents discussed in these older reviews, and we also discuss
more recent literature related to thornlessness, new sources of thorn-
lessness, and the genetic basis of thornlessness. [Article copies available
for a fee from The Haworth Document Delivery Service: 1-800-HAWORTH.
E-mail address: <docdelivery@haworthpress.com> Website: <http://www.
HaworthPress.com>  2005 by The Haworth Press, Inc. All rights reserved.]

KEYWORDS. Rubus, thorns, prickles, breeding

M. A. Coyner is Graduate Student, Robert M. Skirvin is Professor of Horticulture,

M. A. Norton is Post-Doctoral Researcher, and A. G. Otterbacher is Senior Researcher,
University of Illinois, Department of Natural Resources and Environmental Sciences,
258 Edward R. Madigan Biotechnology Laboratory, 1201 West Gregory, Urbana, IL
61801.
This research is supported in part by funds provided by the University of Illinois
Agricultural Experiment Station project number 65-0323 and funds from the Univer-
sity of Illinois Research Board. Any opinions, findings, conclusions, or recommenda-
tions expressed in this publication are those of the author(s) and do not necessarily
reflect the view of the US Department of Agriculture.
Small Fruits Review, Vol. 4(2) 2005
http://www.haworthpress.com/web/SFR
 2005 by The Haworth Press, Inc. All rights reserved.
Digital Object Identifier: 10.1300/J301v04n02_09 83
84 SMALL FRUITS REVIEW

INTRODUCTION

The stiff outgrowths commonly known as thorns on the canes of

blackberries and raspberries are technically known as prickles or spines.
Prickles form from the outer layer of tissue (epidermis) and divide tan-
gentially to form ridged multicellular appendages which later become
cutinized as hard sharp appendages known as prickles. Morphologically
there is no difference between glandular hairs and prickles (Peitersen,
1921), but prickles are stiffer (Figure 1). Epidermal spines or prickles
are commonly referred to as “thorns,” however, they are morphologi-
cally different from true thorns, which are modified branches with ad-
joining vascular tissue. Prickles lack internal vascular material and
result from multiple cellular divisions of the epidermis (Esau, 1977).1
Mode of Inheritance. Segregation analysis has shown that both reces-
sive and dominant inheritance systems for thornlessness are found
among various blackberry cultivars. The genetic nature of some thorn-
lessness genes is not understood; this has hindered breeding for thorn-
less blackberries. Differences in ploidy levels among blackberry cultivars
and species are other factors that have made it difficult to breed for
thornlessness in blackberries (Darrow, 1928). Most commercial black-
berries are polyploid; the majority of these are tetraploids, a few are
hexaploid, and other ploidy levels are less common (Table 1). Due to
differences in ploidy, segregation ratios are not always the same for all
crosses. For instance, for a recessive character to be expressed in a
tetraploid, four copies of the recessive gene must be present; only one
copy of a dominant gene is required for expression of the phenotype, re-
gardless of ploidy. Progress in blackberry breeding was first reported by
Darrow (1928; 1937) and by Stene and Odland (1937).
Chromosomal Diversity. The basic chromosome number of the Rubus
genus is x = 7 and ploidy ranges from 2x (diploid) to 12x (dodecaploid)
(Robertson, 1974). In order to successfully transfer genetic material the
chromosomes must pair with their homologous chromosomes or else
essential genes and chromosomes can be lost (see Table 2). Such losses
can result in embryo abortion (Daubeny, 1996). Since thornlessness is
expressed phenotypically it is possible to screen for thornlessness in a
single season, but it is not possible to screen for fruitfulness or make
crosses in that period. Blackberry canes are biennial. The primocanes
grow vegetatively, and following a cold period, the floricanes flower
and set fruit. Thus a seed generation in blackberry requires at least two
growing seasons or two years. A year can be saved by germinating the
seeds in vitro (Ke et al., 1985; Mian et al. 1995). A minimum of two
 Coyner et al. 85

FIGURE 1. Variations in prickle morphology over several Rubus species.

R. allegheniensis R. argutus R. pergratus R. frondosus

R. canadensis R. elegantulus R. vermontanus R. setosus

R. hispidus R. flagellaris R. enslenii R. baileyanus

generations (four years) is required to detect thornlessness in a recessive

inheritance system, since the recessive trait is not expressed in the first
generation hybrid. In a dominant system a thornless phenotype can be
detected after a single generation (two years).
Since blackberry cultivars are clonally propagated, the identification
86 SMALL FRUITS REVIEW

TABLE 1a. Thornless blackberry species and cultivars. Part I: Origin, ploidy,
gene action and growth habit.

Name Introduction Location of Gene Ploidy Gene Action Habit

date discovery
Merton Thornless-types
‘Apache’ 1999 Arkansas s 4x recessive erect
‘Arapaho’ 1992 Arkansas s 4x recessive erect
‘Black Satin’ 1974 Illinois s 4x recessive semi-erect
‘Burbank Thornless’ 1914 California s 2x recessive semi-erect
‘Chester Thornless’ 1985 Illinois s 4x recessive semi-erect
‘Dirksen Thornless’ 1974 Illinois s 4x recessive semi-erect
‘Ebano’ 1981 Brazil s 4x recessive semi-erect
‘Georgia Thornless’ 1967 Georgia - 4x recessive semi-erect
‘Hull Thornless’ 1981 Maryland s 4x recessive semi-erect
‘Loch Ness’ 1988 UK s 4x recessive erect
‘Merton Thornless’ 1956 UK s 4x recessive semi-erect
‘Navaho’ 1988 Arkansas s 4x recessive erect
‘Ouachita’ 2003 Arkansas s 4x recessive erect
Rubus rusticanus var. inermis - wild species s 2x recessive semi-erect
‘Santa Rosa’ 1900’s California s 4x recessive semi-erect
‘Smoothstem’ 1966 Maryland s 4x recessive semi-erect
‘Thornfree’ 1966 Maryland s 4x recessive semi-erect
‘Triple Crown’ 1996 Maryland s 4x recessive semi-erect

Rubus laciniatus-types
‘Everthornless’ 1995 Illinois STE 4x Dominant trailing
‘Thornless Evergreen’ 1926 Oregon STE 4x Dominant trailing

Austin Thornless-types
‘Austin Thornless’ 1924 Oklahoma Sf 8x Quasi-dominant trailing
‘Waldo’ 1989 Oregon Sf 6x Dominant trailing
‘Douglass’ 1993 Oregon Sf - Dominant trailing

Whitford Thornless-types
‘Whitford Thornless’ 1967 Illinois - 2x recessive erect

Rubus canadensis-types
Rubus canadensis - wild species - 3x - erect
‘Perron’s Black’ 1987 Canada - 3x - -
‘Per Can’ 1990 Canada - 3x - trailing

Logan Thornless-types
‘Cory Thornless’ 1911 California SfL 6x Dominant -
‘Thornless Logan’ 1993 New Zealand SfL 6x Dominant trailing
(‘Bauer Thornless’)
‘Rosati-Jamieson Logan 1988 Italy SfL 6x Dominant trailing
Thornless’

Other Thornless-types
‘Bedford Thornless’ 1930 UK - 6x Dominant -
‘Bursa’ Series - Turkey - - - -
‘Cameron’ 1926 North Carolina - - - -
‘Cox’s Miracle Berry’ 1988 West Virginia - - - semi-erect
‘Doyle’s Blackberry’ 1970 Indiana - - - semi-erect
‘Jumbo’ - - - - - -
‘Pecos’ 2003 California - 4x - erect
‘Thornless Boysen’ 1938 California - 7x - trailing
‘Young Thornless’ 1930 California - - - trailing
 Coyner et al. 87

TABLE 1b. Thornless blackberry species and cultivars. Part II: Source and
reference.

Name Source References

Merton Thornless-types
‘Apache’ Arkansas 1007 ⫻ ‘Navaho’ Clark and Moore, 1999
‘Arapaho’ Arkansas 883 ⫻ Arkansas 631 Moore and Clark, 1993
‘Black Satin’ SIUS 47 ⫻ ‘Thornfree’ Hull, 1975
‘Burbank Thornless’ - Hull, 1968
‘Chester Thornless’ SIUS 47 ⫻ ‘Thornfree’ Galletta et al., 1998a
‘Dirksen Thornless’ SIUS 47 ⫻ ‘Thornfree’ Hull, 1975
‘Ebano’ F2 of ‘Comanche’ ⫻ (‘Thornfree’ ⫻ ‘Brazos’) Bassols and Moore, 1981
‘Georgia Thornless’ USDA 1445 ⫻ ‘Early June’ Moore and Skirvin, 1990
‘Hull Thornless’ SIUS 47 ⫻ ‘Thornfree’ Galletta et al., 1981
‘Loch Ness’ SCRI 74126RA8 ⫻ SCRI 75131D2 Jennings, 1989
‘Merton Thornless’ self of ‘John Innes’ ⫻ ‘John Innes’c Scott et al., 1957
‘Navaho’ Arkansas 583 ⫻ Arkansas 631 Moore and Clark, 1989
‘Ouachita’ ‘Navaho’ ⫻ Arkansas 1506 Clark and Moore, 2003
‘Pecos’ ‘Navaho’ ⫻ ‘Loch Ness’ Fear, 2003
Rubus rusticanus var. inermis species Daubeny, 1996
‘Santa Rosa’ - Jennings, 1988
‘Smoothstem’ US 1482 open pollinated Hull, 1975
‘Thornfree’ (‘Brainerd’ ⫻ ‘Merton Thornless’) ⫻ Zych et al., 1970
(‘Merton Thornless’ ⫻ ‘Eldorado’)
‘Triple Crown’ SIUS 68-2-5 ⫻ (‘Darrow' ⫻ 'Brazos’) Galletta et al., 1998b

Rubus laciniatus-types
‘Everthornless’ Histogenic isolation of ‘Thornless Evergreen’ McPheeters and Skirvin, 1995
‘Thornless Evergreen’ Clonal selection of R. laciniatus Darrow, 1931

Austin Thornless-types
‘Austin Thornless’ ‘Austin Mayes’ seedling selection Hull, 1961
‘Douglass’ “hand pollinated a hand-pollinated cross Douglass , 1993
of two non-patented parents, Sander
and Lawrence”
‘Waldo’ derived from ‘Austin Thornless’ Daubeny, 1996; Strik, 1996

Whitford Thornless-types
‘Whitford Thornless’ selection from the wild near Farina, Illinois Zych et al., 1967

Rubus canadensis-types
Rubus canadensis species Hull, 1961
‘Perron’s Black’ Selected from wild R. canadensis Huber, 1987
‘Per Can’ Selected from wild R. canadensis Huber, 1990

Logan Thornless-types
‘Cory Thornless’ chimeral variant of ‘Mammoth’a Jennings, 1988
‘Thornless Logan’ (‘Bauer meristem tip culture of ‘Logan’b Hall et al., 1986b
Thornless’)
‘Rosati-Jamieson Logan selection from ‘Logan Thornless’ seedlings Rosati, 1988
Thornless’

Other Thornless-types
‘Bedford Thornless’ self of Veitchberry (4x raspberry) Daubeny, 1996
‘Bursa’ series - Turemis et al., 2003
‘Cameron’ ‘Young’ ⫻ ‘Lucretia’ Hull, 1961
‘Cox's Miracle Berry’ chance seedling Cox, 1988
‘Doyle's Blackberry’ open pollination of unknown parentage Doyle, 1977
‘Jumbo’ - Turemis et al., 2003
‘Thornless Boysen’ - Hull, 1975
‘Young Thornless’ Found at the home of Helen M. Ovenell. Fischer et al., 1940
a ‘Mammoth’ = ‘Aughinbaugh’ ⫻ ‘Crandall’
b ‘Logan’ = ‘Antwerp’ (red raspberry) ⫻ ‘Aughinbaugh’ (blackberry)
c ‘John Innes’ (4x) spined cultivar with the recessive thornless gene from Rubus rusticanus var. inermis
88 SMALL FRUITS REVIEW

TABLE 2. Ploidy levels of selected blackberry cultivars.

Cultivar Ploidy Reference

‘Burbank Thornless’ 2n = 2x = 14 Hull, 1968
Rubus rusticanus var. inermis 2n = 2x = 14 Daubeny, 1996
‘Whitford Thornless’ 2n = 2x = 14 Zych et al., 1967
Rubus canadensis 2n = 3x = 21 Hull, 1961
‘Merton Thornless’ 2n = 4x = 28 Hull, 1968
‘Cory Thornless’ 2n = 6x = 42 Jennings, 1988
‘Lincoln Logan’ 2n = 6x = 42 Hall et al., 1986c
‘Thornless Young’ 2n = 6x = 42 Fischer et al., 1940
‘Thornless Boysen’ 2n = 7x = 49 Jennings, 1988
‘Austin Thornless’ 2n = 8x = 56 Hull, 1961
‘Cascade’ 2n = 9x = 63 Hull, 1968

of a thornless plant that could become a new cultivar (or a parent for fur-
ther crosses) might be found in a single generation. Theoretically, a su-
perior blackberry cultivar could be produced in two seasons (one
generation) with a dominant inheritance, whereas a recessive inheri-
tance would require at least four seasons (two generations). However, it
is more likely that repeated cycles of backcrossing will be required be-
fore an acceptable genotype is isolated.

TYPES OF GENETIC SYSTEMS

IN THORNLESS BLACKBERRIES

Recessive Genes. ‘Merton Thornless’ (MT) originated from the Malling

Merton Experiment Station in 1943 (Scott et al., 1957). MT was se-
lected among progeny resulting from a cross between ‘John Innes’ and
a thornless seedling from ‘John Innes’.2 MT has a recessive thornless
gene, s. Since MT is tetraploid, four copies of the gene are needed to
produce the thornless phenotype. In a segregating population from an
MT cross, thorny phenotypes can be screened and eliminated by the
presence of stalked glands or glandular hairs on the cotyledons and/or
leaf petioles of germinating seedlings (Hull, 1968; Scott et al., 1957).
Although seedling screen is useful, Scott et al. (1957) reported that
seedlings needed to reach a height of 8-12 inches before thornlessness
could be reliably determined. Using these methods, thorny segregants
are eliminated, the percentage of thornless plants is increased, and the
amount of space required to grow the generation to fruiting is reduced.
Several undesirable characters are linked to thornlessness in MT. These
include susceptibility to cold, semi-erect growth habit, lateness of fruit-
 Coyner et al. 89

ing season, peculiar flavor, and partial sterility (Hull, 1968). The stems
of thornless segregants are free of thorns but the petioles can produce
rudimentary prickles (Jennings, 1988).
‘Burbank Thornless’ (BT) is a thornless blackberry genetically simi-
lar to ‘Merton Thornless’ that also shows stalked glands or glandular
hairs on thorny segregants (Hull, 1968). BT was found in the collection
of Luther Burbank after his death in 1923. Little is known of its origin or
genetics. It has a diploid chromosome number (2n = 2x = 14).
Dominant Thornless Genes. ‘Austin Thornless’ (AT; syn. ‘Austin
Thawnless’) (Butterfield, 1928) has a “quasi-dominant” thornless in-
heritance controlled by the thornless gene Sf (Hull, 1961). Thornlessness
in its segregating progeny ranges from moderately thorny to rudimen-
tary prickles on the petiole and leaf veins to completely thornless. Vig-
orous canes of AT may develop thorns near the lower nodes (Hull,
1961). Although seedlings of AT vary in their degree of expression of
thornlessness, it is more useful than recessive types since it is possible
to obtain a completely thornless segregant in a single generation. AT is an
octoploid blackberry. Crosses using the thornlessness of AT are limited
to blackberries with higher ploidy levels (Daubeny, 1996). A dominant
gene is more useful in breeding higher ploidy blackberry since only one
copy of the gene is necessary to produce the thornless phenotype
(Jennings, 1988).
‘Everthornless’ (ET) is a trailing type tetraploid blackberry devel-
oped at the University of Illinois and patented in 1995 (McPheeters and
Skirvin, 1995, 2000; Plant Patent No. 9407, December 26, 1995). It was
derived from ‘Evergreen’ via the chimeral ‘Thornless Evergreen’ (TE)
(Darrow, 1931) parent using tissue culture methods. The mode of inher-
itance is a dominant system (Hall et al., 1986a) that does not entirely
prevent thorns from developing; it appears to behave as a suppressor of
their development. Several classes of thorns from small bumps to stiff,
recurved thorns are found among thornless progeny (Norton and Skirvin,
1997). Several traits appear to be linked with the genetic thornlessness
of ET, including bent petioles lacking a clasping sheath around the stem
and dwarfism.
Less Understood Thornless Genes. ‘Whitford Thornless’ (WT) has
very erect, thornless canes with a few small thorns on leaf petioles and
midribs (Zych et al., 1967). Stalked glands, as observed in thorny
segregants of ‘Merton Thornless’ crosses, are present on both thorny
and thornless seedlings (Hull, 1968), so the character is of no value for
screening for thornlessness. Little is known of the genetics of thorn-
lessness in WT.
90 SMALL FRUITS REVIEW

Chimeras and Thornlessness in Blackberries. Since the thorns of

blackberries are epidermal in origin, it is possible for a thornless plant to
be chimeral, possessing both thorny and thornless tissue (Dermen,
1960). TE is such a plant. It has a thornless (mutant) epidermis, which
surrounds an inner core of wild-type (thorny) tissue. There are several
problems associated with a plant possessing dual genetic compositions.
The character may not pass through the sexual system and the pheno-
type may not be stable.
Chimeras in Rubus. The shoot meristem of a blackberry has three
distinct histogenic layers (Figure 2); each of which is superimposed on
top of the other and differentiates into specific regions and organs of the
plant. The outermost layer of cells in the apical dome (LI) develops into
the outer covering (epidermis) of the plant. LI cells divide anticlinally,
perpendicular to the surface. The layer immediately underneath the LI
layer is called the LII; this region develops from anticlinal and periclinal
(tangential) divisions into a thick, transversing layer that constitutes the

FIGURE 2. A diagram of a cross section through the apical meristem of a chi-

mera showing the various types of chimeras that could result from adventitious
shoot formation. Moving clockwise from the upper right hand diagram: 1. No
chimera (no mutations); 2. Sectorial chimera; 3. Fully mutated; 4. Periclinal chi-
mera; 5. Mericlinal chimera.

2
Mutated

Wild-type
3
 Coyner et al. 91

majority of the leaf tissue including mesophyll and palisade layers. The
mature sexual reproductive cells (gametes), ovules and pollen are de-
rived from the LII. The LII is positioned as a dome of cells in juxtaposi-
tion to a core histogenic region called the LIII. The LIII region contains
the bulk of the center of the stem, leaf midrib region, and roots. Black-
berries often form adventitious shoots from roots and crowns. These are
called root suckers. Root suckers develop from progenitor LIII cells and
reflect the genotype of the LIII (Dermen, 1960; Esau, 1977; Hartmann et
al., 1997).
Genetic changes in blackberry cells can occur in a number of ways
including point mutations, segmental inversions, rearrangements, and/
or transposable elements. Some mutations alter the plant’s phenotype.
For instance, a modification of the DNA in an LI cell can alter the ex-
pression of a gene in the epidermis. If the mutation affects the pheno-
type, the new phenotype (e.g., thornlessness) from the mutation can
mask the wild-type phenotype of the underlying LII and LIII cells. The
mutant phenotype is called a “sport.” Cells arising from mutant cells
maintain their identity by cellular division and continue to express the
mutant (thornless) trait while the unaltered cells of the LII and LIII layers
retain their thorny genotype. Depending on the location of a mutation
event in the apical meristem, various types of chimeras3 can result
(Tilney-Bassett, 1986).
The chimeral arrangements of thorny and thornless tissues are as-
signed specific names. (1) Sectorial. Thornless and thorny genetic types
are situated side-by-side. (2) Periclinal. Thornless (or thorny) tissue
completely encompasses a core of the opposite phenotype. (3) Mericlinal.
A mutated single histogenic layer partially encircles the internal tissue,
so it is not periclinal (Figure 2).
Chimeral analysis can be used to explain TE’s instability for the
thornless character (Darrow, 1955; Hall et al., 1986a) and its tendency
to revert back to the thorny condition as well as its failure to breed for
the thornless condition. First, TE is thornless. Prickles form from the LI
(thornless) layer; thus all epidermal tissues derived from that layer will
be thornless. Second, TE produces only thorny offspring. Gametes form
from the LII (thorny genotype); thus all offspring are thorny. Third, TE
forms thorny root suckers. Blackberry roots form adventitious shoots
(suckers) readily. Roots originate from the LIII (thorny genotype); thus
root suckers from a chimeral plant such as TE are thorny (McPheeters
and Skirvin, 1983).
TE arose as a chance mutation on the wild-type thorny parent ‘Ever-
green’ around 1926 (Darrow, 1931). TE is a periclinal chimera in which
92 SMALL FRUITS REVIEW

a layer of thornless tissue (LI) completely surrounds a core of thorny

cells in the LII and LIII. The thornless condition is not stable. The
periclinal chimeral arrangement is commonly known as a “hand-in-
glove” chimera. Although TE is phenotypically thornless, it is not a
pure thornless blackberry. As a result TE has had little use in commer-
cial breeding programs. Gametes arise from the LII histogenic layer,
which has the thorny, wild-type genotype. Since gametes form from the
LII layer TE will transmit the thorny character to the offspring, and so
only thorny offspring will be seen. In addition TE readily produces root
suckers. The roots arise from the LIII-derived internal tissue, which is
thorny; the root suckers will be thorny. Thus TE will revert back to the
thorny type if not maintained properly. In addition root suckers cannot
be used to propagate TE or maintain thornlessness. Furthermore, be-
cause thorny root suckers are more vigorous than thornless canes, the
plants can easily lose their beneficial thornless condition. Propagation
of TE must involve the use of preformed axillary buds to maintain the
chimeral state.
McPheeters and Skirvin (1983) were successful in causing cells from
external thornless layers to formulate whole plants in tissue culture to
produce a genetically pure thornless form of TE they called ‘Ever-
thornless’ (ET). A similar procedure was applied to chimeral ‘Thorn-
less Logan’ to produce a genetically pure form called ‘Lincoln Logan’
(Hall et al., 1986c; Rosati et al., 1988).
Genetically Pure (Non-Chimeral) Blackberries. In a genetically pure
blackberry all cells have the necessary information to express the
thornless phenotype. Roots suckers produced from a genetically pure
thornless blackberry will be thornless and gametes in the egg or sperm
will carry thornless genes into the next generation. Barring mutation,
genetically pure thornlessness is stable.

THORNLESS BLACKBERRY CULTIVARS

Merton Thornless-Types. ‘Merton Thornless’ (MT, Figure 3) MT was

the first commercial tetraploid genetically thornless blackberry cultivar.
M. B. Crane in England developed MT in 1948 from a backcross be-
tween ‘John Innes’ to a partially thornless seedling of ‘John Innes’.
‘John Innes’ is a thorny tetraploid blackberry, which is heterozygous for
a recessive thornless gene from Rubus rusticanus (ulmifolius) var.
inermis (Daubeny, 1996). ‘Merton Thornless’ was found to be self-fer-
tile and thus became an excellent source of thornlessness for breeding
 Coyner et al. 93

FIGURE 3. A comparison of primocanes from four different types of thornless

blackberries.

Austin Thornless-type Whitford Thornless-type

Rubus laciniatus-type Merton Thornless-type

tetraploid thornless blackberries. The thornless canes of MT are easily

managed and require less labor than thorny cultivars. Undesirable char-
acteristics from MT are lack of winter hardiness, late maturity and pecu-
liar flavor of the fruits (Hull, 1968). The fruits of MT ripen late in the
season (July or early August) and have large seeds (Scott et al., 1957).
‘Burbank Thornless’. ‘Burbank Thornless’, a genetically thornless
blackberry similar in appearance to ‘Merton Thornless’ (Hull, 1968).
‘Burbank Thornless’ is a diploid blackberry that was found in the col-
lection of Luther Burbank after his death in 1923. Little is known of its
origin or genetics.
‘Dirksen Thornless’. ‘Dirksen Thornless’ (SIUS 64-21-8) was se-
lected in 1967 at Carbondale, Illinois, by J.W. Hull from a cross of SIUS
47 (US 1482 ⫻ ‘Darrow’) ⫻ ‘Thornfree’. The cultivar was named in
honor of the late Senator Everett McKinley Dirksen of Illinois (Hull,
1975; Moore and Skirvin, 1990). The fruits are medium-large in size
with a dull black color and good flavor. ‘Dirksen Thornless’ is more
vigorous and productive than ‘Thornfree’ but less vigorous than ‘Black
Satin’ (Hull, 1975). It has a semi-erect growth habit, so trellising is re-
94 SMALL FRUITS REVIEW

quired to support the canes. Harvesting of ‘Dirksen Thornless’ occurs

three weeks earlier than ‘Thornfree’ (Hull, 1975).
‘Thornfree’. ‘Thornfree’ is a semi-erect thornless blackberry cultivar
that was introduced in 1966 by Scott and Ink from the USDA in
Beltsville, Maryland (Zych et al., 1970). ‘Thornfree’ was selected from
a cross between two thorny plants, US 1410 and US 1414, which were
heterozygous for the recessive thornless ‘Merton Thornless’ gene. A
thornless offspring with four copies of the recessive gene was selected
and field-tested. The selected blackberry plant had large, firm berries
and tart flavor. ‘Thornfree’ has a semi-erect growth habit that requires
additional support to assist in fruit development and harvesting. Staking
the canes to a single pole or attaching them to a two-wire vertical trellis
is commonly practiced. The fruits of ‘Thornfree’ mature over a 35-40
day period. The ripe fruits are recommended for home gardens, local
markets, and pick-your-own operations. ‘Thornfree’ is marginally hardy
in Central Illinois and is better suited for the milder winters in the south-
ern part of Illinois (Zych et al., 1970).
‘Smoothstem’. ‘Smoothstem’ was selected from the progeny of an
open pollination of US 1482. The medium-large fruit of ‘Smoothstem’
are black in color with good flavor, but secondary in overall quality to
‘Thornfree’. ‘Smoothstem’ is more vigorous than ‘Thornfree’ but lacks
winter hardiness (Hull, 1975; Moore and Skirvin, 1990).
‘Black Satin’. ‘Black Satin’ (SIUS 64-21-11) is a sibling to ‘Dirksen
Thornless’. In 1967 J.W. Hull selected ‘Black Satin’ from the SIUS 47
(US 1482 ⫻ ‘Darrow’) ⫻ ‘Thornfree’ cross at Carbondale (Moore and
Skirvin, 1990). The dull-black fruit is larger and the plant is more vigor-
ous and winter hardy than ‘Thornfree’ (Hull, 1975). ‘Black Satin’ yields
a greater quantity of fruit than ‘Dirksen Thornless’. ‘Black Satin’ is
semi-erect and trellising is required. The fruit of ‘Black Satin’ ripen one
week later than ‘Dirksen Thornless’ and two weeks earlier than ‘Thorn-
free’ (Hull, 1975).
‘Navaho’ (Plant Patent No. 6679). ‘Navaho’ (Arkansas 1172) was
selected in 1977 from a cross of two thorny blackberry selections het-
erozygous for the recessive ‘Merton Thornless’ gene for thornlessness
(s) from ‘Thornfree’. ‘Navaho’ was the first ‘Merton Thornless’ de-
rived thornless blackberry with an erect stature. The erect growth habit
of ‘Navaho’ reduces the need for trellis support. The thornless character
of ‘Merton Thornless’ is closely linked to trailing cane growth habit.
Several generations of crosses were required to break the linkage and
introduce the erect phenotype into thornless plants. The medium-sized
fruits of ‘Navaho’ are glossy black with good yields. ‘Navaho’ has pro-
 Coyner et al. 95

vided great utility for breeding the erect habit to other thornless tetraploid
cultivars (Moore and Clark, 1989) and is a parent of the ‘Arapaho’,
‘Ouachita’ and ‘Pecos’ cultivars.
‘Ebano’. ‘Ebano’ is a thornless blackberry originally crossed at Ar-
kansas Agricultural Experiment station. The seeds were transported to
Brazil for germination and selection. The selected blackberry plant was
thornless and semi-erect with numerous large clusters of medium-sized
fruit. Thornlessness in ‘Ebano’ resulted from a cross from an offspring
of ‘Thornfree’ and subsequent selection of a thornless plant with four
copies of the recessive thornless gene in a segregating F2 generation.
The cultivar was named after the Portuguese word for ebony, referring
to the dark black fruit color. ‘Ebano’ has been popular in Brazil because
of its high quality fruits that ripen after the other major crops have been
harvested. The fruits of ‘Ebano’ are commonly used in jams, jellies,
canned and frozen packs and used as flavor and coloring in dairy prod-
ucts such as yogurt and ice cream (Bassols and Moore, 1981).
‘Hull Thornless’. ‘Hull Thornless’ (SIUS 68-6-6) is a product of the
same cross that produced ‘Dirksen Thornless’ and ‘Black Satin’; it was
the fifth thornless blackberry cultivar produced from the USDA and
collaborating organizations including the University of Illinois and
Southern Illinois University. The plant was selected by J.W. Hull in
1972 at Carbondale, Illinois and later named ‘Hull Thornless’ in his
honor. Hull was a blackberry and raspberry breeder at the University of
Maryland, the University of Arkansas, and the US Department of Agri-
culture in Carbondale, IL. ‘Hull Thornless’ is a semi-erect vigorous
blackberry with fruits similar to ‘Black Satin’. The fruits are large, firm
and sweet in flavor. The plants are winter hardy south of Urbana, IL.
‘Hull Thornless’ is recommended as an alternative thornless blackberry
cultivar to ‘Black Satin’ in USDA hardiness zones 6-8 (Galletta et al.,
1981).
‘Arapaho’ (Plant Patent No. 8510). ‘Arapaho’ (Arkansas 1536) was
introduced in 1992 as an early ripening, erect-caned thornless black-
berry. ‘Arapaho’ is a half-sibling to ‘Navaho’ from the cross Arkansas
631 ⫻ Arkansas 883 made in 1982. ‘Arapaho’ offers several improve-
ments over ‘Navaho’. The roots of ‘Arapaho’ produce more primocanes
which increases its ability to establish a dense planting quickly. The
fruits ripen earlier so the extreme summer temperatures are avoided; it
is probably the earliest ripening thornless cultivar thus far introduced.
‘Arapaho’ has a very erect growth habit derived from thorny cultivars
such as ‘Darrow’, ‘Hillquist’, ‘Brazos’, and ‘Cherokee’, and it derived
its thornlessness via ‘Thornfree’. Since it is an erect cultivar, trellis sup-
96 SMALL FRUITS REVIEW

port is not required. It has shown good resistance to the disease rosette
(double blossom), a serious disease of blackberry in southern areas. The
fruit is a firm, glossy black, similar in flavor to ‘Navaho’ but with
smaller seeds. Mixed plantings of ‘Arapaho’ and ‘Navaho’ will produce
a long fruitful growing season (Moore and Clark, 1993).
‘Loch Ness’ (Plant Patent No. 6,782). ‘Loch Ness’ (Jennings, 1988)
was a selection derived from crossing two unnamed blackberry selec-
tions at the Scottish Crop Research Institute (SCRI) Dundee, United
Kingdom, in 1978 and was patented in 1989. Sturdy, erect turning
semi-erect canes bear large, slightly glossy black fruit with a pleasant
sharp flavor. The greatest attributes of ‘Loch Ness’ are winter hardiness
and early ripening.
‘Chester Thornless’. ‘Chester Thornless’ (SIUS 68-6-17) is another
thornless blackberry selection from the USDA program. SIUS 68-6-17
was selected from a SIUS 47 ⫻ ‘Thornfree’ cross made by J.W. Hull in
1968. One the authors of this review (Skirvin) actually made the cross
and planted the cultivar while a student of Hull working in Carbondale,
Illinois. It was named ‘Chester Thornless’ by two of the authors of this
chapter (Skirvin and Otterbacher) in recognition of Dr. Chester Zych, a
former bramble researcher at the University of Illinois (Galletta et al.,
1998a). ‘Chester Thornless’ is semi-erect like the other cultivars se-
lected from the SIUS 47 ⫻ ‘Thornfree’ cross. It has large, firm fruits
and has an improved flavor. ‘Chester Thornless’ is similar to ‘Hull
Thornless’ in fruit qualities but is more productive and is more winter
hardy. Because of its semi-erect growth habit ‘Chester Thornless’ needs
trellis support like the other semi-erect cultivars. The fruits have excel-
lent shelf life and its bold color provides a fresh appearance for frozen
desserts. The fruits are firm enough to be shipped and sold as a fresh
product. ‘Chester Thornless’ is one of the hardiest thornless blackberry
cultivars and is recommended as a replacement for ‘Thornfree’ in
UDSA hardiness zones 5-7 (Galletta et al., 1998a). ‘Chester Thornless’
is a parent of the hardy (and very thorny) cultivar named ‘Illini Hardy’
(Plant Patent 8333; ‘Chester Thornless’ ⫻ NY 95) that is reported to
survive temperatures as low as ⫺17 to ⫺23° F (Skirvin and Otterbacher,
1993).
‘Triple Crown’. ‘Triple Crown’ was selected in 1983 at Beltsville,
Maryland, by G.J. Galletta from the cross of SIUS 68-2-5 and Arkansas
545 (‘Darrow’ ⫻ ‘Brazos’). SIUS 68-2-5 is a sibling of ‘Black Satin’,
‘Chester Thornless’, ‘Dirksen Thornless’ and ‘Hull Thornless’. In 1996
the selection was named ‘Triple Crown’ for its three crowning attrib-
utes: vigor, aroma and sweetness, and high productivity. The overall
 Coyner et al. 97

performance of the plant coupled with its superior attributes was sug-
gestive of winning a “Triple Crown” in horse racing. ‘Triple Crown’
produces large fruits with a uniform distribution along the cane contrib-
uting to easier harvesting. The aroma is second to ‘Marion’, the most
widely cultivated blackberry. Unfortunately its fruits are too soft for
shipping, so it will remain a pick-your-own or backyard cultivar (Galletta
et al., 1998b).
‘Apache’ (Plant Patent 11,865). ‘Apache’ resulted from a cross of
Arkansas 1007 ⫻ ‘Navaho’ made in 1988 (Clark and Moore, 1999). The
original plant was selected in 1991. It is the third thornless erect cultivar
released from the Arkansas program. ‘Apache’ produces larger fruit
and higher yields than the previously released Arkansas thornless, erect
cultivars ‘Navaho’ (Moore and Clark, 1989) and ‘Arapaho’ (Moore and
Clark, 1993).
‘Ouachita’ (Plant Patent Pending). ‘Ouachita’ is a thornless, erect,
firm-fruited blackberry (Clark and James N. Moore, 2003) resulting
from a cross between ‘Navaho’ ⫻ Ark. 1506. The fruit is medium-large,
productive, moderately vigorous with a hardiness down to ⫺17.4 °C. It
is reported to be resistant to double blossom/rosette (Cercosporella
rubi); moderately resistant to anthracnose (Elsinoë veneta).

Rubus canadensis-Types

‘Perron’s Black’. ‘Perron’s Black’ is a thornless blackberry selection

from Rubus canadensis. It originated from W.H. Perron & Co. Ltd.,
Laval, Quebec, Canada. ‘Perron’s Black’ was a clone selected for ex-
treme cold hardiness from a Rubus canadensis population. The plant is
well adapted to most soil types but prefers neutral to acidic soils. The
canes are vigorous, producing an aromatic black fruit when mature. The
canes can grow up to 12 feet in length so trellising or wire support is
needed. ‘Perron’s Black’ is well suited for pick-your-own operations,
home gardens and road-side markets (Huber, 1987).
‘Per Can’ (Plant Patent No. 7,251). ‘Per Can’ is a selection of Rubus
canadensis made by Tony Huber in Laval, Canada (Huber, 1990). ‘Per
Can’ was selected for its exceptional cold hardiness in Canada, resistant
to temperature as low as ⫺40°C. The canes are vigorous but rarely pro-
duce root suckers. The fruits turn dark black at maturity and have an ex-
tended harvest season (60 days). The trailing growth habit of ‘Per Can’
requires trellis support.
98 SMALL FRUITS REVIEW

‘Logan’ Derived Cultivars

‘Cory Thornless’. ‘Cory Thornless’ originated in California and is a

thornless cultivar of ‘Mammoth’ blackberry. The fruits of ‘Cory Thorn-
less’ are large and black without a characteristic flavor. Although fairly
productive, the plant is marginally hardy and not grown in Oregon
(Hull, 1968). ‘Cory Thornless’ is not grown in cultivation (Waldo and
Hartman, 1946).
‘Thornless Logan’. ‘Thornless Logan’ is a thornless sport of ‘Lo-
gan’. ‘Thornless Logan’ was discovered in 1962 and distributed as
‘Bauer Thornless’ (Jennings, 1981). The thornlessness is due to a
periclinal chimera. The outer layer (LI) of mutant cells composing the
epidermis is genetically different from the wild-type internal layers (LII
and LIII). The thornless condition of ‘Thornless Logan’ is not stable.
Root suckers, which originate from internal tissue, will express the
wild-type thorny phenotype. As a result a chimeral ‘Thornless Logan’
plant can revert from thornless to a thorny condition (Rosati et al., 1988).
‘Lincoln Logan’. In 1986 the histogenic layers of chimeral ‘Thornless
Logan’ were separated in tissue culture to form a pure type (Hall et al.,
1986b). The new plant was believed to be derived from epidermal cells
and genetically pure thornless. ‘Thornless Logan’ was determined to be
pure thornless by assessing the phenotype of root suckers; if the root
suckers were thorny then the chimeral status had been maintained.
Thornless plants indicated segregation. The pure thornless ‘Thornless
Logan’ was named ‘Lincoln Logan’. The features of ‘Thornless Logan’
are identical to ‘Logan’. The fruits are medium to large with a dark red
color and firm texture. The attractive flavor makes ‘Lincoln Logan’
ideal for juice and wine making (Waldo and Hartman, 1946; Hull,
1975). ‘Thornless Logan’ is grown on the Pacific coast of the United
States.

Other Sources of Thornlessnes

‘Austin Thornless’. ‘Austin Thornless’ (a.k.a. ‘Austin Thawnless’) is

a genetically pure blackberry (Butterfield, 1928; Hull, 1961) (Figures 3
and 4). Unlike ‘Merton Thornless’ and ‘Whitford Thornless’, ‘Austin
Thornless’ has a dominant thornless character and is octoploid. Crosses
between ‘Austin Thornless’ and thorny compatible cultivars produce
thornless hybrid plants in the F1 generation indicating dominance. Hull
(1961) observed variability in the segregation patterns, and as a result
he called the inheritance in ‘Austin Thornless’ “quasi-dominant.” Re-
 Coyner et al. 99

FIGURE 4. A comparison of primocanes from ‘Austin Thornless’ (2n = 6x = 42)

and ‘Whitford Thornless’ (2n = 2x = 14) blackberries.

‘Austin Thornless’ ‘Whitford Thornless’

Austin Thornless-type Whitford Thornless-type

gardless of its mode of inheritance, using ‘Austin Thornless’ in breed-

ing programs offers an advantage over the recessive because a reduced
number of generations are required to detect thornlessness. A severe
disadvantage associated with thornlessness in ‘Austin Thornless’ is
varying degrees of female sterility in thornless segregants (Hull, 1961).
In addition the increased chromosome number of ‘Austin Thornless’
makes it difficult to breed with tetraploid or diploid blackberries. ‘Aus-
tin Thornless’ is a vigorous plant with large black berries with a pleasing
flavor. In cultivation ‘Austin Thornless’ is only moderately productive
and is restricted to growth in the south (Hull, 1961; 1975).
‘Waldo’ Blackberry. ‘Waldo’ was a joint release of the Oregon State
University and the USDA in 1989 (Strik, 1996). It was the first geneti-
cally thornless trailing blackberry cultivar released. Despite this major
advantage in addition to a relatively small seed size and firm fruit, this
cultivar is not widely planted. Plants lack sufficient vigor making man-
agement more difficult and fruit lack the “Marion” flavor (Strik, 1996).
‘Douglass’ Blackberry (Plant Patent 8423). ‘Douglass’ (Douglass,
1993) is a thornless trailing blackberry developed in Oregon and pat-
100 SMALL FRUITS REVIEW

ented in 1993. The cultivar is reported to be thornless on most regions

except the base of canes where “incipient thorns like those of raspberry”
develop.

‘Evergreen’-Derived Cultivars

‘Evergreen’. ‘Evergreen’ (EG, Rubus laciniatus Willd.) is a thorny

member of the common European blackberry Rubus laciniatus Willd
(Figure 5). EG is widely distributed in the wilds of Oregon but its exact
origin is questionable. Rubus laciniatus’ seeds were believed to be car-
ried on ships from England to Australia and New Zealand, then depos-
ited on the American West Coast. EG is highly adapted to Oregon’s
climate and has subsequently become naturalized in the wild (Waldo,
1977). Despite the heavy presence of thorns EG, was revered for its fruit

FIGURE 5. A comparison of primocanes from three different types of R.

laciniatus. Right to left: ‘Evergreen’ = original thorny type; ‘Thornless Ever-
green’ = chimeral thornless type; and ‘Everthornless’ = pure thornless type de-
veloped through tissue culture.

‘Thornless Evergreen’
‘Everthornless’ ‘Evergreen’

Rubus laciniatus-types
 Coyner et al. 101

quality and processing ability and was once the most-planted black-
berry cultivar in Oregon (Jennings, 1988).
‘Thornless Evergreen’. ‘Thornless Evergreen’ was discovered in
1926 by Frank Siegmond growing along a fence on his farm in Oregon
(Waldo, 1977). ‘Thornless Evergreen’ is a thornless sport of the ‘Ever-
green’, a thorny blackberry plant of uncertain origin (discussed earlier).
Mr. Siegmond was a grain farmer and had no interest in his thornless
version of ‘Evergreen’ so he offered the plant to Phillip Steffes for prop-
agation. Mr. Steffes planted several acres of the thornless blackberry.
The planting attracted the attention of George Darrow of the USDA. At
the Steffes farm Darrow, evaluated the planting and found that the
thornless plants were as vigorous and hardy as the parental ‘Evergreen’.
The thornless sport was later named ‘Thornless Evergreen’ (TE, Darrow,
1931). The yield and quality of ‘Thornless Evergreen’ was equal to ‘Ev-
ergreen’ but without the hazards of the thorns. As the thornless plants
propagated themselves from root suckers thorny canes were observed.
Although the thornless phenotype was unstable, it was still a great ad-
vantage for growers.
It was determined that the thornless instability of TE was a result of
being a chimera, where the outer layer (epidermis) had mutated to the
thornless state which produced a different phenotype than the internal
regions which retained their thorny genotypes (discussed earlier). Since
the outer layer is genetically different from the internal tissue, root
suckers arising from internal tissue were derived from the wild-type
thorny phenotype. ‘Thornless Evergreen’ is not a genetically pure
thornless blackberry. Gametes, produced from internal tissue, will propa-
gate the thorny characteristic and not transmit thornlessness to hybrid
plants. ‘Thornless Evergreen’ detracts commercial production value
due to the chimerism (Waldo, 1977).
‘Thornless Evergreen’ is a fully fruitful thornless version of ‘Ever-
green’. The fruits are sweet, large and black and grow vigorously. The
growth habit is trailing so trellising and supports are needed. TE is
adapted for growth in Pacific Northwest (Waldo and Hartman, 1946;
Hull, 1975; Skirvin, 1983).
‘Everthornless’ (Plant Patent No. 9407). ‘Everthornless’ (ET) is a
genetically pure form of ‘Thornless Evergreen’ (McPheeters and Skirvin,
1995, 2000). McPheeters and Skirvin separated the histogenic layers of
the periclinal chimera into pure types using tissue culture (McPheeters
and Skirvin, 1983). Thornless plants were selected and evaluated in
field studies. The result was a trailing type blackberry without thorns.
To test genetic purity, adventitious buds were derived from roots and
102 SMALL FRUITS REVIEW

used to determine the thornless phenotype, pure thornless or chimeral.

Thornless root suckers indicated pure thornlessness; thorny suckers in-
dicated a chimera. The thornless condition now has the potential for
commercial breeding. The fruit quality characteristics of ‘Everthorn-
less’ are similar to ‘Thornless Evergreen’ but ‘Everthornless’ is less
acidic with more soluble solids than its predecessor. ‘Everthornless’ has
great promise as a source of pure genetic thornlessness (McPheeters and
Skirvin, 2000). Hall et al. (1986a) showed that the gene controlling
thornlessness in ET is dominant.
‘Thornless Boysen’. ‘Thornless Boysen’ originated in California and
is a thornless sport of ‘Boysen’ quite similar to ‘Thornless Young’
(Moore and Skirvin, 1990). Like ‘Thornless Logan’ and ‘Thornless Ev-
ergreen’, the thornlessness of ‘Thornless Boysen’ is only expressed on
the external layers of the chimera and does not pass its thornlessness
through the sexual cycle. The fruits are large and dark red with an excel-
lent flavor. The plant grows vigorously and is fairly productive. ‘Thorn-
less Boysen’ is ideal for canning or freezing (Hull, 1975; Waldo and
Hartman, 1946).
‘Thornless Young’. ‘Thornless Young’ is a relatively thornless sport
of ‘Youngberry’ reported by Darrow (1929). ‘Thornless Young’ origi-
nated in California and was found at the home of Helen M. Ovenell
(Darrow, 1929). Its large fruits are dark red in color with an excellent
sweet flavor. ‘Thornless Young’ is moderately hardy and is common in
Southern and Pacific states (Hull, 1975). Like ‘Thornless Boysen’ the
fruits of ‘Thornless Young’ are canned and frozen (Waldo and Hartman,
1946).

‘Whitford Thornless’ and Other Unique Thornless Cultivars

‘Whitford Thornless’. ‘Whitford Thornless’ is a diploid thornless

blackberry discovered in the wild near Farina, Illinois, east of St. Louis
(Zych et al., 1967). ‘Whitford Thornless’ is believed to be a clone of a
wild blackberry, Rubus argutus Link, and was named after its discov-
erer, A. M. Whitford. The canes of ‘Whitford Thornless’ are thornless
but the occurrence of an occasional rudimentary prickle on the midrib
and petiole is not uncommon (Hull, 1967). ‘Whitford Thornless’ has a
very erect growth habit and produces many suckers from its roots.
‘Whitford Thornless’ is best suited for climates similar to that found in
Southern Illinois. Severe winter dieback was reported in Urbana, IL.
‘Whitford Thornless’ shows a recessive inheritance of its thorn-
lessness (Hull, 1968). Since ‘Whitford Thornless’ is a diploid species,
 Coyner et al. 103

thornlessness is not easily transferred to most eastern tetraploid black-

berries. ‘Whitford Thornless’ has not been a successful parent of pro-
ductive thornless blackberry cultivars and is not suited for immediate
commercial blackberry production (Zych et al., 1967).
‘Cox’s Miracle Berry’ (Plant Patent 6105). ‘Cox’s Miracle Berry’
(Cox, 1988) was found growing in Mr. Cox’s fields. It is thornless and
is reported to produce two crops of fruits per season.
‘Doyle’s Blackberry’ (Plant Patent 4094). ‘Doyle’s Blackberry’
(Doyle, 1977) was observed by Thomas Doyle growing in his berry
patch in 1970. ‘Doyle’s Blackberry’ was developed from an open polli-
nation of unknown parentage. The growth habit of ‘Doyle’s Black-
berry’ is semi-erect with a few trailing canes. Root suckers rarely
develop. The fruit of ‘Doyle’s Blackberry’ is blue-black in color and is
produced abundantly. The fruit is reported to have an excellent shelf life
and stays firm for several days following harvest (Doyle, 1977).
‘Pecos’ (Plant Patent 13525). ‘Pecos’ (Fear, 2003) is a thornless
cultivar that arose from a cross between ‘Navaho’ and ‘Loch Ness’. It is
a proprietary cultivar developed for the Driscoll’s Company®. The
cultivar is reported to be erect and produce firm fruits that ship well.
There is a report (Turemis et al., 2003) of thornless blackberry
cultivars selections from the Marmara and Black Sear regions of Turkey
called ‘Bursa-1’, ‘Bursa-2’, ‘Bursa-3’ and ‘Bartin’. They also report a
cultivar called ‘Jumbo’ which may be from Switzerland. The authors of
this review know nothing else about these cultivars.
In conclusion, in this article we have attempted to address the status
of thornlessness blackberry breeding. Although the genetic nature of
thornlessness is best understood for ‘Merton Thornless’ and ‘Austin
Thornless’, the usefulness of the other sources of blackberry thorn-
lessness remain largely unexplored. The authors of this paper hope that
this review will encourage geneticists to investigate the nature of these
other genetic systems through traditional breeding systems as well as
consider using biotechnology methods to identify and isolate specific
thornlessness genes (DNA sequences) and use them to transform thorny
cultivars directly to the thornless state.

NOTES
1. The terms prickles, spines and thorns are used interchangeably in most texts but
for ease of usage and to cohere with previous bramble research, throughout this manu-
script we will use the term “thorn” to refer to the multicellular epidermal appendages
present in blackberries.
104 SMALL FRUITS REVIEW

2. ‘John Innes’ is a thorny tetraploid hybrid blackberry from Rubus thysiger ⫻

Rubus rusticanus var. inermis (4x), also known as Rubus ulmifolius var. inermis. The
tetraploid R. rusticanus var. inermis formed from an unreduced germ cell. ‘John Innes’
is a heterozygous carrier for the recessive thornless gene, s.
3. “Chimera” is derived from Greek mythology, it is a creature with a lion’s head, a
goat’s body, and a dragon’s tail.

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