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Morphological Characterisation of Wild Rubus rosifolius (Rosaceae) Plants Growing in Jamaica Prior to Agricultural Pursuits

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Precise identification of plants is critical for informed agriculture where nutraceutical properties and productivity are simultaneously important. The Jamaican-grown West Indian raspberry (Rubus rosifolius) falls into this category. The first step to taking this wild-growing plant into agricultural production is morphological identification. Although there are no botanical reports of multiple varieties of this species in Jamaica, two morphotypes were found, which were named ‘Red' and ‘Wine Red’ based on fruit colour. Morphological methods were used to characterise these plants growing at Holywell (located over 900 m above sea level), in the Blue Mountain region of Jamaica. Morphological analyses revealed that growth form, leaf, flower, and fruit characteristics of the morphotypes were statistically distinctive between the two morphotypes for 48 of the 59 measured parameters. Several descriptors allowed the morphotypes to be distinguished before their fruits became visible. These findings support the hypothesis that there is a standard morphotype having a scrambling morphology, smaller leaves, and darker red, oblong, solitary fruits (Wine Red, WR), and a distinct morphotype with more upright stems, larger leaves, and lighter red, spherical, bunched fruits (Red, R). This information can now be used to facilitate molecular analyses and ramp-up clonal production of these morphotypes to determine the agricultural factors that are linked to yield and nutritionally relevant traits.
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Caribbean Journal of Science (2021), 51: pp. 1–13.
© Copyright 2021 by the College of Arts and Sciences of the University of Puerto Rico, Mayagüez
1
Morphological Characterisation of Wild Rubus rosifolius (Rosaceae) Plants Growing in
Jamaica Prior to Agricultural Pursuits
Theresa F. rambaran1,2,3,*, Joape GiniGini2, paTricia a. mcLenachan4, camiLLe s. bowen-Forbes1, and syLvia
adJoa miTcheLL5
1Department of Chemistry, Faculty of Science and Technology, The University of the West Indies, Mona,
Jamaica
2Institute of Applied Sciences, Faculty of Science and Technology, University of the South Pacic, Suva, Fiji
3Current address: Department of Public Health and Clinical Medicine, Section of Sustainable Health, Umeå
University, Umeå, Sweden
4Institute of Fundamental Sciences, Massey University, Palmerston North, New Zealand
5The Biotechnology Centre, Faculty of Science and Technology, The University of the West Indies, Mona,
Jamaica
*Corresponding author: theresa.rambaran@umu.se
AbstrAct—Precise identication of plants is critical for informed agriculture where nutraceutical properties
and productivity are simultaneously important. The Jamaican-grown West Indian raspberry (Rubus rosifolius) falls
into this category. The rst step to taking this wild-growing plant into agricultural production is morphological
identication. Although there are no botanical reports of multiple varieties of this species in Jamaica, two
morphotypes were found, which were named ‘Red’ and ‘Wine Red’ based on fruit colour. Morphological methods
were used to characterise these plants growing at Holywell (located over 900 m above sea level), in the Blue Mountain
region of Jamaica. Morphological analyses revealed that growth form, leaf, ower, and fruit characteristics of the
morphotypes were statistically distinctive between the two morphotypes for 48 of the 59 measured parameters.
Several descriptors allowed the morphotypes to be distinguished before their fruits became visible. These ndings
support the hypothesis that there is a standard morphotype having a scrambling morphology, smaller leaves, and
darker red, oblong, solitary fruits (Wine Red, WR), and a distinct morphotype with more upright stems, larger
leaves, and lighter red, spherical, bunched fruits (Red, R). This information can now be used to facilitate molecular
analyses and ramp-up clonal production of these morphotypes to determine the agricultural factors that are linked
to yield and nutritionally relevant traits.
Rubus L. (Rosaceae) is a fruitful genus that includes
raspberries, blackberries, and dewberries, and has over
700 species (Hummer 2010). Rubus species are highly
valued for their sweet and tangy fruit. Rubus rosifolius
Sm., also called West Indian raspberry, roseleaf
raspberry, and redberry, is widespread in Southeast
Asia, the Caribbean, Brazil, and Hawaii (Adams
1972). In addition to food applications, R. rosifolius
has several ecological functions, including its role in
helping return disturbed areas to forest vegetation, soil
protection, and providing food and cover for wildlife
and livestock (Francis 2004). Ornamental variants,
such as R. rosifolius var. commersonii (Poir.) Tirveng.
and R. rosifolius var. coronarius Sims., produce elegant
white owers, and are easily distinguished from
the typical variety as R. rosifolius var. commersonii
produces owers with 10 petals, while R. rosifolius var.
coronarius produces elaborate, rose-like owers and no
fruit (Brown 1999).
Notwithstanding their numerous benecial
properties, some Rubus species are regarded as
being invasive, for example Rubus alceifolius Poir.
on Réunion Island (Baret et al. 2003). According
to Randell et al. (2004), R. rosifolius Sm. occupies
habitats on all the islands and in most ecosystems
of Hawaii, but is considered the least invasive of
their alien Rubus species. This species, which is an
underutilised fruit-bearing plant in Jamaica, is recorded
as being introduced to Hawaii from Jamaica in 1880
(CABI 2019). When it was introduced to Jamaica is
not known, nor has it become a major crop there. With
the push towards agricultural diversication, research
aimed at nutritional and phytochemical characterisation
of Jamaican berries (including R. ellipticus Sm., R.
rosifolius, R. jamaicensis L., and R. racemosus Roxb.)
is needed to determine their potential for productive
utilisation and commercialisation.
2 caribbean JournaL oF science [Volume 51
Research on Rubus raspberries growing in
Jamaica that began in 2007 has resulted in a better
understanding of their phytochemistry and health-
promoting properties, including non-phenolic
functional food components, anthocyanin content, and
their antioxidant, anti-inammatory, and anticancer
effects (Bowen-Forbes et al. 2009, 2010). Several
Rubus species in other countries have also been
found to have benecial properties, and have been
used in the manufacture of nutraceuticals. These
include R. ulmifolius Schott, which has been used as
a lipid oxidation inhibitor in burger patties and as an
antioxidant ingredient in yoghurt. Additionally, Rubus
coreanus Miq. has been used in vinegar for its bone-
health enhancing properties (Ganhão et al. 2013;
Martins et al. 2014; Lee et al. 2017).
Further agricultural development was stymied
when it became evident that the Rubus rosifolius berries
provided by local farmers were of two types, with
two different shapes and colours; thus, research was
undertaken to see if traits of these morphotypes were
stable and dependable. As an initial test, colourimetric
analysis revealed that while both berries had the same
redness value, the berries of one morphotype (labelled
‘Red’ or ‘R’) were signicantly lighter and had a higher
yellowness value than the other (labelled ‘Wine Red’ or
‘WR’) (Rambaran and Bowen-Forbes 2020).
From an agro-processing perspective, it is
important for the sake of product consistency to know
if the variation noted in fruit colour, and possibly other
attributes, is random or due to intrinsic differences.
Proling of their phenolic contents, chemopreventative
properties (via cytochrome P450 inhibitory analysis),
antioxidant, antimicrobial, and hypoglycemic activities
revealed notable differences between these two
morphotypes (Campbell et al. 2017; Rambaran and
Ginigini 2020; Rambaran et al. 2020). The same was
true for their anthocyanins, essential oils, and several
sensory parameters (Rambaran and Ginigini 2020;
Rambaran and Bowen-Forbes 2020).
Of similar importance is being able to compare
plant morphotypes in one country with those found
elsewhere, to discern if they are endemic, native, or
introduced, and therefore what level of protection is
needed. Morphological and molecular methods are
important and reliable tools that can be used for the
characterisation and identication of plants (Tilneya et
al. 2009; Kostamo et al. 2013; Šarhanová et al. 2017;
Boonsrangsom 2020). Although molecular methods
are more denitive, morphological studies are very
important for physical characterisation and eld
identication of plants for which genetic ngerprinting
can then be carried out.
In this work, we used morphological characterisation
to determine the extent of the differences between
these two sympatric morphological forms of plants
resembling R. rosifolius. Other aims of this research are
to report on descriptors which can be used to describe
other Rubus plants, to identify parameters to be used
for eld identication, especially before fruit set, and
to determine which parameters to use for quantity
and quality analysis at harvest. Yet another aim is to
establish base-line data of wild-growing R. rosifolius to
compare with planned agricultural trials to determine
factors responsible for important phytochemical and
yield traits.
maTeriaLs and meThods
Plant Material and Sampling Protocol
Rubus rosifolius plants were found growing wild at
Holywell Park located in the forests of the Blue and John
Crow Mountains National Park, St. Andrew, Jamaica,
at over 900 meters above sea level. Park ofcials,
who reported that they had been working at the park
for several years, said they only removed wayward
stems but otherwise did not disturb growth within
each cluster of stems. They also indicated that two
morphotypes were distinguishable by them. These two
morphotypes were rst noticed by our research group
in 2011, with bright red fruits always occurring on one
morphotype, which we labelled R, and slightly darker
red fruits on the other morphotype, which we called
WR. The morphotypes were collected from different
areas of Holywell, St. Andrew in 2011, 2012, and 2015
(voucher specimen numbers 35596, 35652, and 36280
for the WR morphotype and 35595, 35653, and 36278
for the R morphotype), which indicated to us that the
two morphotypes were stable. They were identied
as R. rosifolius at the herbarium in the Department
of Life Sciences, The University of the West Indies,
Mona Campus. Although both morphotypes appear to
have been collected in Jamaica before 2011 (Table 1),
they were both identied on the herbarium sheets as
R. rosifolius, and were not further differentiated. The
next step was to determine if there were signicant
2021] rambaran eT aL.: morphoLoGy oF Rubus Rosifolius pLanTs 3
morphological differences between the two plants.
Rubus rosifolius plants growing at Holywell Park
and at farmer’s elds located in the forests of the Blue
and John Crow Mountains National Park were analysed
in April 2015. Plants from both morphotypes were then
established in the Mona Heights area of St. Andrew
parish, at 179 metres above sea level. Their gross
morphology was compared to those plants studied
in detail from Holywell Park to indicate if the main
features of these two morphotypes were maintained
and therefore independent of location, especially in
terms of elevation.
Morphological Data
The gross morphology (appearance of brambling,
ower colour, ower type [single or bunched, terminal
or sub-terminal], fruit colour, and fruit placement) of
over 50 plants was observed at Holywell Park and
neighbouring farmer’s elds. Next, ten owers, ripe
fruits, green fruits, and leaves (including all leaets of
each leaf) of each morphotype were taken from three
separate plant clumps and measured. The petal and
sepal length, ower width and mid-height, and fruit
characteristics (ripe or green, length, and width) were
measured. Leaves were randomly selected from the
main and side axes. For these analyses, leaets were
numbered from the leaf base to the tip on both sides,
with the terminal leaet having the largest number. The
length and width of the leaves, and component leaets,
as well as leaf area, were determined and analysed.
Finally, the morphology of ve representative
stems taken from each of the two Rubus morphotypes
were analysed. Stem height and total number of nodes
present on the main stem axis, and percentage of
nodes bearing fruit was tabulated. Then, in order to
decrease variability, upper stems at similar stages of
growth (tip and next ten nodes) were also analysed for
each morphotype. The descriptors chosen here were:
internode length and width, average number of prickles
per internode, number of branches (primary, secondary,
and higher branch order if present); percentage of nodes
bearing owers and/or fruits (mature, green, or dried),
and stem placement of owers and fruit. Morphological
descriptions given for R. rosifolius and other Rubus
species by Adams (1972), Whistler (1995), and Wagner
et al. (1999) were then compared with those obtained
during this morphological analysis.
Statistical Analysis
Plants selected for morphological analysis were of a
similar level of maturity, with only stems from patches
of the two morphotype having both mature and unripe
berries being selected for analysis (eg. oricanes). In
all, mean values and associated statistical signicances
were calculated for 59 parameters. All data from the
morphological analysis was subjected to statistical
analysis using IBM SPSS® v. 22.0 where paired t-tests
were conducted. All values are reported as mean ±
standard deviation and p < 0.05 was considered to be
statistically signicant.
resuLTs
Morphological Traits: Gross Morphology
TabLe 1. Records of dried specimens of Rubus rosifolius from Jamaica at The University of the West Indies, Mona
Campus, Jamaica Herbarium (UWI). Morphotypes labelled ‘a were identied by the herbarium curator, Mr. Patrick
Lewis, from pressed, dried specimens on 29 Sep 2015.
Herbarium # Locality Parish Date Morphotype
31182 Orange River St. Mary 1 Apr 1974 Reda
33211 Clydesdale St. Andrew 19 Mar 1982 Reda
33405 Hardware Gap St. Andrew 1989 Wine Reda
34759 Christiana Manchester 20 Nov 1999 Wine Reda
35352 Below Gap Cafe St. Andrew 28 Apr 2007 Wine Reda
35595 Holywell St. Andrew 5 Nov 2011 Red
35596 Holywell St. Andrew 5 Nov 2011 Wine Red
35653 Holywell St. Andrew 16 Oct 2012 Red
35652 Holywell St. Andrew 16 Oct 2012 Wine Red
36278 Holywell St. Andrew 8 Oct 2015 Red
36280 Holywell St. Andrew 8 Oct 2015 Wine Red
4 caribbean JournaL oF science [Volume 51
Gross morphology was observed for over 50 Rubus
rosifolius plants growing at four separate locations
(two farmer’s elds near Holywell Park and two
locations within the Park). On all occasions, the gross
morphology as reported here was the same for each
plant of the respective morphotype, without exception.
For both morphotypes, white owers were found. The
WR morphotype was decidedly more scrambling than
the R counterpart.
The inorescences examined (Fig. 1) were
cincinnus and subsolitary in the WR morphotype,
and corymbose and cymose in the R morphotype,
respectively. While fruiting of both morphotypes was
terminal, the fruits were solitary (only one per node)
in the WR morphotype, but appeared in bunches in the
R morphotype. The fruits also had distinct differences
in colour and shape (Fig. 2). The WR morphotype had
fruits which were deeper red and were more oblong-
ellipsoid in shape (longer fruits) while the fruits of the
R morphotype had a brighter red hue and were spherical
(wider fruits). These characteristics were stable, as the
WR morphotype never had R-type fruits nor vice versa.
There were also no intermediate-looking fruits on either
morphotype.
Morphological Traits: Fruit, Flower, and Leaf
Characteristics
For the ower and fruit characteristics measured
for WR and R Rubus rosifolius morphotypes, all ower
parameters, with the exception of petal length, differed
signicantly (p < 0.05) between the two morphotypes,
and in each instance, that of WR was larger than R
(Table 2). The ripe and unripe/green WR fruits were
signicantly longer than those of its R counterpart, and
vice versa for fruit width. As expected, the length and
width of the ripe fruits were greater than those of the
green fruits for both WR and R morphotypes (Fig. 2;
Table 2).
Compound leaves were measured due to perceived
differences between the two morphotypes (Table 3).
Since eld identication was the aim, leaf/leaet
size, area, shape, and structure were considered. The
leaves from the main stem, in most cases, consisted of
seven leaets; however, leaves bearing ve and (less
frequently) nine leaets, were also found on each
plant sampled. The leaf area (total leaet area) of six
randomly selected typical leaves from the main stem
and side branches of both morphotypes was measured
(Table 3). All leaves chosen possessed either ve or
seven leaets, and the leaf area (total area of leaets)
FiG. 1. Fruiting pattern of the R (cymose and corymbose) and WR (subsolitary) Rubus rosifolius morphotypes. Photos
by T. Rambaran.
2021] rambaran eT aL.: morphoLoGy oF Rubus Rosifolius pLanTs 5
FiG. 2. The R and WR Rubus rosifolius morphotypes. Photos by T. Rambaran.
TabLe 2. Flower and fruit characteristics of two Rubus rosifolius morphotypes, R and WR. SS (statistical signicance)
at the 95 % condence level, n = 10.
Parameter (mm) Red (R) Wine Red (WR) SS
Flowers
Width 2.54 ± 0.3 3.25 ± 0.7 yes
Petal Length 14.00 ± 0.1 13.60 ± 0.2 no
Sepal Length 11.00 ± 0.1 17.00 ± 0.3 yes
Mid-height 6.91 ± 1.0 8.51 ± 2.0 yes
Fruits
Ripe
Length 14.50 ± 0.2 18.30 ± 0.3 yes
Width 20.90 ± 0.4 14.90 ± 0.3 yes
Green
Length 11.40 ± 0.2 14.60 ± 0.2 yes
Width 11.70 ± 0.2 8.40 ± 0.1 yes
Length
Ripe vs. Green 14.50 ± 0.2 vs. 11.40 ± 0.2 yes
18.30 ± 0.3 vs. 14.60 ± 0.2 yes
Width
Ripe vs. Green 20.90 ± 0.4 vs. 11.70 ± 0.2 yes
14.90 ± 0.3 vs. 8.40 ± 0.1 yes
6 caribbean JournaL oF science [Volume 51
TabLe 3. Mean leaet area for main stem and side branch leaves for two Rubus rosifolius morphotypes, R and WR. SS
(statistical signicance) at the 95 % condence level, n = 10.
Parameter (cm2) Red (R) Wine Red (WR) SS
Leaet Area
Main Stem
Leaets (1–6) 411.0 ± 145.6 154.5 ± 36.5 yes
Leaet 7 481.0 ± 150.2 145.0 ± 42.4 yes
Side Branch
Leaets (1–4) 217.0 ± 78.3 127.7 ± 31.4 yes
Leaet 5 261.6 ± 181.7 133.0 ± 44.6 yes
Avg. Leaet Area
Comparisons
(1–6) vs. 7 411.0 ± 145.6 vs. 481.0 ± 150.2 no
154.5 ± 36.5 vs. 145.0 ± 42.4 no
(1–4) vs. 5 217.0 ± 78.3 vs. 261.6 ± 181.7 yes
127.7 ± 31.4 vs. 133.0 ± 44.6 no
5 vs. 7 261.6 ± 181.7 vs. 481.0 ± 150.2 yes
133.0 ± 44.6 vs. 145.0 ± 42.4 no
FiG. 3. Leaves from comparable R and WR Rubus rosifolius morphotypes with seven leaets. Photos by S. Mitchell.
2021] rambaran eT aL.: morphoLoGy oF Rubus Rosifolius pLanTs 7
of both the main stem and side branch leaves was
signicantly greater for those of the R morphotype than
for the WR morphotype.
The terminal leaet area was compared to the
area of the other leaets of that particular leaf (Fig.
3). There were no signicant differences amongst the
leaet areas for the WR morphotype. However, for the
R morphotype, the terminal leaet of ve-leaet leaves
was signicantly larger than the other leaets. Also, for
the R morphotype, the terminal leaet of leaves bearing
seven leaets was signicantly larger than those of
leaves bearing ve leaets.
The leaets of both the main stem and side branches
were further examined in terms of their length and
width (Table 4). The overall length and width of the
R morphotype leaves was signicantly greater than the
WR leaves. Comparisons were made between the leaet
length and width (Table 4) in a similar manner as done
for leaet area (Table 3). In each case, the leaets were
signicantly longer and wider for the R morphotype
than for the WR morphotype. The only exceptions
were the average of leaets one to six and leaets one
to eight compared to their respective terminal leaet of
the R morphotype.
TabLe 4. Mean leaf and leaet lengths and widths for two Rubus rosifolius morphotypes, R and WR. SS (statistical
signicance) at the 95 % condence level, n = 10.
Parameter (mm) Red (R) Wine Red (WR) SS
Leaf Length 9.60 ± 0.3 5.40 ± 0.1 yes
Leaf Width 13.80 ± 0.4 7.80 ± 0.2 yes
Leaet Length (LL)
(1–4) 5.10 ± 0.1 3.70 ± 0.1 yes
5 6.20 ± 0.1 4.90 ± 0.1 yes
(1–6) 6.90 ± 0.1 4.10 ± 0.0 yes
7 7.30 ± 0.2 5.90 ± 0.0 yes
(1–8) 8.60 ± 0.1 5.00 ± 0.1 yes
9 8.50 ± 0.0 none found
LL Comparison
(1–4) vs. 5 5.10 ± 0.1 vs. 6.20 ± 0.1 yes
3.70 ± 0.1 vs. 4.90 ± 0.1 yes
(1–6) vs. 7 6.90 ± 0.1 vs. 7.30 ± 0.2 no
4.10 ± 0.0 vs. 5.90 ± 0.0 yes
(1–8) vs. 9 8.60 ± 0.1 vs. 8.50 ± 0.0 no
none found
Leaet Width (LW)
(1–4) 2.10 ± 0.0 1.80 ± 0.0 yes
5 2.70 ± 0.1 2.30 ± 0.0 yes
(1–6) 2.90 ± 0.1 1.70 ± 0.0 yes
7 4.30 ± 0.2 2.30 ± 0.0 yes
(1–8) 3.60 ± 0.1 2.30 ± 0.0 yes
9 3.50 ± 0.0 none found
LW Comparison
(1–4) vs. 5 2.10 ± 0.0 vs. 2.70 ± 0.1 yes
1.80 ± 0.0 vs. 2.30 ± 0.0 yes
(1–6) vs. 7 2.90 ± 0.1 vs. 4.30 ± 0.2 yes
1.70 ± 0.0 vs. 2.30 ± 0.0 yes
(1–8) vs. 9 3.60 ± 0.1 vs. 3.50 ± 0.0 yes
none found
8 caribbean JournaL oF science [Volume 51
Morphological Traits: Stem Morphology
Stems support the leaves, owers and, by extension,
the fruits. Special emphasis was given to the placement
and occurrence of owers, fruits, and prickles on a
typical stem; the average number of nodes per stem,
along with internode length and width, and stem branch
order (Table 5). The main stem length was comparable
between both morphotypes; however, the WR stems
were bent (scrambling) as a result of them being more
branched and thinner, while the R morphotype stems
were thicker, less branched, and more upright (Table
5; Fig. 5). The percentage of nodes bearing owers or
fruits was comparable between the two morphotypes
along with the total number of nodes present per equal
length of stem. The R morphotype had signicantly
more fruits and owers as many nodes had more
than one fruit (bunched). The number of prickles per
internode length was signicantly different, with the R
morphotype having fewer prickles (Table 5).
Although the WR and R Rubus morphotypes had
different branching patterns, with WR having up to four
branch order causing it to scramble (Table 5), the lengths
of the internodes of the main stem were not found to be
signicantly different in most cases (Fig. 4). Differences
were only noted for the youngest internodes, with
the R internodes being signicantly longer than their
WR counterparts (Fig. 4). For internode width (with
increasing distance from the tips), the internodes of the
R Rubus morphotype were signicantly wider, giving
thicker stems (Fig. 5).
Summary of Morphological Traits
Morphological analysis revealed similarities and
differences between the two morphotypes. They were
similar in respect to growth type, since both bore
stems which arose from underground runners, and
they both had white owers with similar petal length,
and with similar leaet shape (lanceolate). For both
morphotypes, the terminal leaet tips were acuminate.
The attachment and arrangement of their leaves were
TabLe 5. Assessment of the stem (length and width of internodes), number of nodes, prickles, fruits, and owers on
typical stems of two Rubus rosifolius morphotypes, R and WR. SS (statistical signicance) at the 95 % condence level,
n = 5. Full Stem: Main stem from tip to ground level; Upper Stem: Main stem 10 nodes long and measured from the tip;
*Number of branches per primary stem with 10 nodes.
Parameter Red (R) Wine Red (WR) SS
Full Stem
Main Stem Length (cm) 171.30 ± 65.7 103.80 ± 54.2 no
Total No. of Nodes 26.00 ± 5.2 17.20 ± 3.7 yes
Main Stem Length / Total
No. of Nodes (cm) 6.59 6.03 no
No. of Nodes Bearing Fruits
or owers 22.00 ± 5.2 (85 %) 16.40 ± 3.8 (95 %) no
Supper Stem
Total Nodes on 1° Branch 10 10 no
No. of Prickles per
Internode 2.58 ± 0.9 3.26 ± 0.8 yes
Avg. No. of Branches 1.22 ± 0.3 4.60 ± 3.7 yes
No. of 2° Branches*1.4 3.6 yes
No. of 3° Branches 0 8.6 yes
No. of 4° Branches 0 1.6 yes
Flowers & Fruits per Stem
No. of Mature Fruits 0.24 ± 0.3 0.14 ± 0.1 yes
No. of Green Fruits 4.34 ± 1.4 1.24 ± 1.0 yes
No. of Dried Fruits 0.80 ± 0.9 1.57 ± 1.3 yes
Total Fruits 5.38 ± 2.2 2.95 ± 0.7 yes
Total No. of Flowers 1.68 ± 0.6 0.58 ± 0.8 yes
2021] rambaran eT aL.: morphoLoGy oF Rubus Rosifolius pLanTs 9
also found to be similar, being petiolate and bipinnate,
respectively. The most distinctive morphological
differences distinguishable in the eld between the two
morphotypes was the scrambling nature and smaller
leaves of the WR morphotype, fruit colour and shape,
and fruiting pattern.
discussion
Rubus rosifolius berries are mostly picked from the
wild (Surya et al. 2018) and are considered famine food
(www.purdue.edu), but after being proven edible and of
nutraceutical value, this species is starting to be cultivated
in several countries. Rubus rosifolius is not yet grown as
an agricultural crop in Jamaica, although some farmers
have shown interest in doing so. Rubus rosifolius has
been identied as a ‘plant for a future’ (https://pfaf.org/
user/Plant.aspx?LatinName=Rubus+rosifolius) and is
being cultivated in some countries (Campagnolo and
Pio 2012; Patto et al. 2013).
Prior to agricultural pursuits, it is important to
know what parameters one will be measuring, and
for what morphotype. While literature could be found
on R. rosifolius phytochemistry, taxonomy, genetics,
even mutant breeding (Surya et al. 2016), a detailed
list of morphological descriptors to be used for eld
identication could not be found. The descriptors
published for Rubus idaeus L. (UPOV 2003) were the
most useful, but not in all respects. Also, in taxonomic
descriptions reported, including for Jamaican species
in Adams (1972), the level of detail was insufcient
to separate the two morphotypes. We decided to
undertake this task by using 59 different descriptors
to determine which could be used to separate the two
Rubus morphotypes in the eld, and thus to recommend
the most useful ones for further studies. Phytochemical
analysis reported in Campbell et al. (2017), Rambaran
et al. (2020), Rambaran and Ginigini (2020), and
Rambaran and Bowen-Forbes (2020) were also
undertaken to see if statistically relevant variations
could be found between these two morphotypes.
Morphological Analysis
Morphological differences between the two Rubus
morphotypes were observed for all major plant parts.
Most parameters measured were statistically different
between the two morphotypes. The WR morphotype was
scrambling (due to thinner stems, and more branches
with a higher branch order) while the R morphotype
had upright stems (due to them being thicker and less
branched). The WR morphotype had more prickles on
the stems, smaller and shorter leaves, longer sepals,
and fruits with deeper red pigmentation than the R
morphotype. Differences also existed in fruit shape and
inorescence type, the WR fruit being oblong-ellipsoid
and cincinnus, and its counterpart (R) fruits being
spherical and inorescence corymbose, respectively,
in these two characters. Also, the WR fruits were only
found singly per node while the R fruits were clumped
at the nodes. This was an obviously distinctive feature
(Fig. 1).
A lingering question was whether these observed
differences were stable, and thus would be maintained
at different locations. Representative plants from both
morphotypes taken from Holywell Park (~ 900 m)
were established at Mona Heights, St. Andrew (179
m); the latter experiencing higher temperatures than
FiG. 4. Internode Number vs. Internode Length of
the R and WR Rubus rosifolius morphotypes. (1.0–2.0 =
internode between nodes 1 and 2 numbered from the tip of
the plant). n = 5.
FiG. 5. Internode Number vs. Internode Width of the
R and WR Rubus rosifolius morphotypes. (1.0–2.0 =
internode between nodes 1 and 2 numbered from the tip of
the plant). n = 5.
10 caribbean JournaL oF science [Volume 51
the former. At Mona Heights, the two morphotypes
could be distinguished as the WR stems scrambled
and had smaller leaves while the R morphotype stems
shot upwards and had larger leaves, although it was
noted that for both morphotypes, owering and fruiting
occurred inconsistently and less abundantly compared
to the plants at Holywell Park. When they did ower,
they showed the same pattern, shape, and colour as
reported here for those at Holywell Park.
The differences identied between the two
morphotypes suggest that they are either different
varieties of the same species, are two different
species, or one is a hybrid between R. rosifolius and
a yet to be determined Rubus taxon. While the present
morphological study alone cannot answer which one of
these is correct, data presented here indicate that they
are two distinct morphotypes.
Of the Jamaican-grown Rubus species described
by Adams (1972), in addition to R. rosifolius, there are
two wild raspberry species, R. racemosus Roxb. and R.
ellipticus Sm. (with the latter growing in the same area
where the plants of interest were collected). Comparing
these three Rubus species (Table 6) indicates that the
WR and R Rubus morphotypes most closely t the
description for R. rosifolius. The other two species were
obviously different, one bearing purplish-black fruits
(R. racemosus), and the other having reddish bristles
on the stems and yellow fruits (R. ellipticus).
There has been no botanical report of R. rosifolius
varieties matching all the morphological characteristics
outlined in this paper. Different descriptions of R.
rosifolius Sm. have been reported, and three were
closest to the morphotypes considered in this paper;
Adams (1972) for plants found in Jamaica, Whistler
(1995) for plants grown in Hawaii, and Wagner et al.
(1999) for plants grown in Hawaii. Those of Whistler
(1995) and Wagner et al. (1999) being almost identical.
The descriptions of R. rosifolius, as outlined by these
authors, were therefore compared with the WR and R
Rubus morphotypes (Table 7). The WR morphotype
was found to agree more closely with Adams (1972)
for R. rosifolius based on three descriptors: scrambling
shrub, subsolitary owers, and oblong-ellipsoidal fruits.
Comparing the WR and R morphotypes (Tables 2–6;
TabLe 6. Comparison of Rubus rosifolius, R. ellipticus, and R. racemosus using descriptors employed by Adams
(1972), NA = Not available in Adams (1972).
Descriptors R. rosifolius R. ellipticus R. racemosus
Height (cm) Max of 200 250–500 200
Growth type Scrambling Shrub Shrub Shrub
Thorns per internode Sparsely
More Thorned (with reddish
bristles and white shaggy
hairs)
NA
Leaet shape Ovate-lanceolate Elliptical Broadly Ovate
Position of lateral leaets Opposite Opposite Opposite
Leaets Doubly Serrated Serrated NA
Number of Leaets 5, 7, and 9 3 3, 5, and 7
Largest leaet Terminal Terminal Terminal
Terminal leaet length
(main stem) (cm) 7 710 6
Flowers Subsolitary Axillary and Terminal Axillary Corymbose Raceme
Sepals Attenuate-aristate Deltate NA
Sepal length (mm) 15 8 NA
Petal colour White White Purple
Petal length (mm) 10 10 8
Ripe fruit shape Oblong-ellipsoid NA NA
Ripe fruit length (mm) 15 10 NA
Ripe fruit colour Red Yellow Purplish-black
2021] rambaran eT aL.: morphoLoGy oF Rubus Rosifolius pLanTs 11
Figs. 4–5) with the description given by Wagner et al.
(1999) was inconclusive, as Wagner et al. (1999) refers
to erect stems (more similar to R), solitary, axillary,
or terminal owers (more similar to WR), subglobose
fruit (more closely ts WR), and long, attenuate sepals
(which ts neither) (Table 7). While the three closest
taxonomic descriptions (Adams 1972, Wagner et al.
1999, or Whistler 1995) and several others were not
sufcient to distinguish between these two morphotypes,
they lean towards the WR morphotype, especially that
Table 7. Comparison of the R and WR Rubus rosifolius morphotypes using descriptors previously used to describe
Rubus rosifolius as outlined by Adams (1972) and Wagner et al. (1999). NA = Not Available. Different superscript letters
across a row indicate that the corresponding value is signicantly different at the 95 % condence level. *Whistler’s
(1995) account when it deviated from that of Wagner et al. (1999).
Descriptors Red (R) Wine Red (WR) Adams (1972) Wagner et al. (1999)
and Whistler (1995)
Height (cm) 171.3 103.8 Max of 200 100–200 (or > 200)*
Growth Type Shrub Scrambling Shrub Scrambling Shrub Shrub/subshrub (stems
erect)
Thorns per internode Sparse; (2.58 ± 0.9a)More Thorned; (3.26 ±
0.8b)Sparse; Prickly Sparse; Prickly
Phyllotaxy Alternate Alternate NA Alternate
Leaf Attachment Petiolate Petiolate Petiolate Petiolate
Leaf Arrangement Pinnate Pinnate NA Pinnate
Under-leaet Colour Green Green NA NA
Leaet number per leaf 5–9 5–9 NA 5–7 (3–7)
Leaet Tip Acuminate Acuminate NA Acuminate to acute
Leaet Base Unequally Rounded Unequally Rounded NA Rounded to truncate
Leaet Shape Ovate-lanceolate Ovate-lanceolate Ovate-lanceolate Broadly (Ovate)
lanceolate
Position of Lateral
Leaet Opposite Opposite Opposite NA
Leaet Margin Doubly Serrated Doubly Serrated Doubly Serrated Doubly Serrated
Largest Leaet Terminal Terminal Terminal Terminal
Terminal Leaet
Length(cm) Longer (7.3 ± 0.2a)
Shorter
(5.9 ± 0.0b)
Up to 7 cm NA
Inorescence Corymbose Cincinnus NA Solitary, Axillary or
Terminal
Flowers Cymose Subsolitary Subsolitary Solitary, Axillary or
Terminal
Sepals Attenuate-aristate Attenuate-aristate Attenuate-aristate Lanceolate/Long
Attenuate
Sepal Length (mm) Shorter (11.0 ± 0.1a) Longer (17.0 ± 0.3b) 15 815 (1425)*
Petal Colour White White White White
Petal Length (mm) Same; (14.0 ± 0.1a) Same; (13.6 ± 0.2a) ~10 1020
Ripe Fruit Shape Spherical Oblong-ellipsoid Oblong-ellipsoid Subglobose
Ripe Fruit Width (mm) Wider (20.90 ± 0.4a)More Narrow (14.90 ±
0.3b)NA 2035
Ripe Fruit Length (mm) Shorter (14.5 ± 0.2a) Longer (18.3 ± 0.3b) 15 NA
Fruits Multiple Singular NA Multiple
Fruit easily removed Ye s No NA Ye s
Ripe Fruit Colour Lighter Red (Red) Deeper Red (Wine Red) Red Red
12 caribbean JournaL oF science [Volume 51
of Adams (1972). There are several other botanical
reports of R. rosifolius Sm., some of which include
photos of leaves and fruits, and they are also generally
in line with the WR morphotype herein described
(CABI 2019; GBIF 2019; Francis 2004; Pao 1791;
Plants of the World online 2017). Tropicos (https://
www.tropicos.org) has 22 entries for R. rosifolius, with
12 named varieties, but unfortunately no pictures or
descriptions. TRAMIL (http://www.tramil.net/en) has
pictures of plants growing in the Caribbean region, but
only for those with identied folk medicinal uses, and
it has no Rubus species. Based on the foregoing, the
WR morphotype appears to be the typical R. rosifolius,
while the R morphotype appears to be taxonomically
undescribed.
Concluding Remarks
Two Rubus rosifolius morphotypes could be
distinguished in the eld based on statistically different
values of 48 out of 59 plant descriptors. Many of the
descriptors allowed morphological distinction in the
eld before fruits become visible. This study, therefore,
suggests that the WR and R plants are distinct and
stable R. rosifolius morphotypes. The R morphotype
produced more fruit and was more upright, which could
be important agricultural characteristics. Our studies
therefore show that morphological information is useful
for identifying new plants for breeding and developing
a raspberry industry. Future work in molecular analysis,
the development of clonal multiplication methods, and
carrying out agricultural trials to identify useful traits
can now utilise these morphological data to further
develop these morphotypes towards commercial
production while ensuring their continuity.
AcknowledgementsThe authors wish to express
gratitude to Mr. Patrick Lewis, the herbarium curator
from the Department of Life Sciences, The University
of the West Indies, Mona Campus, for his assistance in
the identication of plant materials. We are also grateful
to the Education, Audiovisual, and Culture Executive
Agency (EACEA) of the European Union for offering a
Caribbean-Pacic Island Mobility Scheme (CARPIMS)
scholarship to Dr. T. Rambaran.
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