On distinctions of
cultivated black-fruited aronia from its wild ancestors
A. K. Skvortsov, Yu. K. Maitulina
On the differences between the cultivated
chokeberry and its wild progenitors. — Bulletin of the Central Botanical
Garden, AN SSSR 126 (1982): 35-40
Translation: Irina Kadis
The genus Aronia (sometimes treated as a subgenus or section of
Sorbus) is distributed in eastern North America from Newfoundland and
southern Quebec and Ontario in the north to the Florida Peninsula in the south.
or chokeberry mostly occurs in wet habitats: wetlands, wet forests,
along streams and around lakes, in sandy lowlands. It can be found in dunes, on
rocky slopes and steep cliffs, on overgrown or bare rocky outcrops [1
In North America, chokeberry is considered a low-quality ornamental
shrub and thus is seldom cultivated, sometimes even exterminated as a weed by
chemical applications [2
Most American authors [1
two species of chokeberry: red-fruited A. arbutifolia
(L.) Elliott and
black-fruited A. melanocarpa
(Michx.) Elliott (Fig. 1), emphasizing
significant variability in both species, and especially in the latter one.
Fig.1. [Not provided] Herbarium sample of A. melanocarpa with ripe fruit (northern New York
State, Adirondack Mountains, Huntington Forest, sphagnum bog, 24 Aug 1976. coll. A.
Both species of Aronia
were cultivated in the European gardens as early as
the beginning of the 19th century, their various forms often described as distinct
species differing from the ones named above by height, pubescence, fruit color,
]. Nearly all these "species"
Lindl., A. grandifolia
Lindl., A. pubens
and others) are currently treated as garden forms of either A. arbutifolia
or A. melanocarpa
]. The only exception is A.
Spach, which is considered either a garden hybrid between A.
and A. melanocarpa
] or distinct
Plants appearing intermediate between A. arbutifolia
also exist in the wild. These are as well treated either as hybrids
] or a third species, A. prunifolia
]. The wild A. prunifolia
has been found identical
with the cultivated A. floribunda
], which made it
possible to synonymize these names.
Another viewpoint on Aronia
] is that it includes only
a single, extremely variable species: A. arbutifolia
At the same time, the black-fruited aronia cultivated in this country and
considered to be A. melanocarpa
, is characterized by extremely low
variability. Due to that, any further selective improvement of the cultivated
aronia employing the traditional approach (selection—crosses—selection) has no
perspective. The only promising method is chemical mutagenesis [10
As a participant in the Russian-American Expedition of 1976 across the Eastern US,
A.K. Skvortsov observed A. melanocarpa in its natural habitats (rocks, open
sand, and bogs in Virginia and New York State) and noticed drastic differences
between these and those cultivated in Russia. This triggered a domestic study of
cultivated aronia headed in four different directions: 1. variability of cultivated
aronia; 2. characters distinguishing cultivated aronia from its wild ancestors; 3.
history of aronia introduction in this country; 4. flowering and fruiting biology.
The following is a report on the results for the former two research directions.
Cultivated black-fruited aronia variability survey
The goals of the survey included:
- assessment of viability and variability of some aronia populations in the center
and at the margins of its range in the USSR;
- a study of fruit morphological variability;
- comparisons of morphology, phenology, and growth dynamics of seedlings
originating from different geographic areas, grown in uniform soil and climatic
conditions; delineation of variability range for certain characters; description of
The work was completed in 1976-1980 at the Flora Department of the Central Botanic
Garden, USSR Acad. Sci (GBS AN SSSR) in Moscow.
Fruits were collected in 1977 along a latitudinal profile within the secondary
range of aronia in [European] Russia, from its northernmost to southernmost limit:
in Arkhangelsk, Petrozavodsk, Vologda, Moscow, and Astrakhan.
In order to obtain comparable material, we collected fruits at the same stage of
ripening from plants growing in similar situations and having a similar habit
(young shrubs up to 1.5 m tall). 10-15 samples of fruit were collected at each
geographic destination, each from a certain mother-plant.
Fruits were also obtained from the US (Washington D.C., Virginia, Massachusetts)
and from other regions of the USSR: Riga, Jaunkalsnava (Latvia); Girionys
(Lithuania); Volgograd, Saransk [European Russia];
Barnaul [southwestern West Siberia];
Sakhalin I. [Far East].
Each fruit sample was represented by a random trial of 50 fruits, each of which was
measured fresh (diameter), then dried and weighted (air mass determined) using a
torsion scale. Because the amount of fruits received from other botanic gardens
through Index Seminum was not large, we had to reduce those trials to 10 fruits.
The mean for each sample was calculated upon running three random trials; then the
mean for each geographic destination was computed as well as variability range,
mean square deviation, and representative error. Student's t-test was employed for
assessing the statistical significance of the differences between sample means.
Cultivated material used for seedling comparison was started from seeds obtained in
1977 through own collecting as well as from other botanic gardens. All seeds were
stratified in peat moss at 0-2°C for 4 months. The sample plots were started in the
Experimental Nursery of the Central Botanical Garden in the spring of 1978. 10
samples, 30-50 g each, were sown for each of the four geographic destinations where
the seed had been collected; additional sample plots were started with seed
obtained by mail (a single sample plot for each destination). Every sample had two
replications sown separately. Soil and light conditions were uniform for all plots.
Care for the plots included weeding, tillage, and a one-time thinning of seedlings.
Watering was only allowed during exceptionally long drought periods.
Growth rate and dynamics were assessed during the first two vegetation periods. At
the end of each growing season height and stem diameter were measured in all
seedlings (ca. 50 plants from each of the four major geographic destinations, in
two replications). In mid-summer, seedling dry biomass was assessed (30 plants per
geographic destination, in two replications).
The studied parameters were measured separately for each sample plot. Because
visual observations suggested very little variability among sample plots from a
certain geographic destination, the mean value for each character (height, stem
diameter, and biomass) was calculated for a geographic destination. For statistical
analysis, mean square deviation and representation error were calculated. Student's
t-test was used for assessment of differences in respective characters for each
pair of geographic destinations.
Morphology of vegetative organs was studied in live plants and herbarium samples
representing aronia in its natural range and in cultivation. Herbarium specimens
from cultivated plants were obtained from Arkhangelsk, Petrozavodsk, Moscow,
Volgograd, and Astrakhan. Collections were studied in the following depositories:
Moscow State University; Central Botanical Garden, USSR Acad. Sci.; V.L. Komarov
Botanical Institute, USSR Acad. Sci.; N.G. Kholodny Institute of Botany, Ukrainian
Acad. Sci.; Tartu University (Estonia); Nikita Botanic Garden. The authors are
especially grateful to Dr. Raimond Tsinovskis (Riga), who kindly shared his large
Size and mass of fruits collected in different regions of the USSR vary
insignificantly: a single fruit weighs 115-125 mg (Table 1). This might be
attributed not just to the biological attributes of the species, but also to
uniformity in cultivation situations, which may level differences in soil and
Table 1. Dry mass of black-fruited aronia from different
||USSR European Part
|Place of collection
||Mean mass of 10 fruits, g
||Place of collection
||Mean mass of 10 fruits, g
||Place of collection
||Mean mass of 10 fruits, g
||0.45 ± 0.05
||0.34 ± 0.08
||1.19 ± 0.02
||0.52 ± 0.04
||0.62 ± 0.03
||1.23 ± 0.03
||0.43 ± 0.03
||0.82 ± 0.03
||1.21 ± 0.02
||0.70 ± 0.00
||0.95 ± 0.05
||1.22 ± 0.01
||0.38 ± 0.00
||0.98 ± 0.03
||1.22 ± 0.0
||0.84 ± 0.03
||0.69 ± 0.04
||1.20 ± 0.03
||0.81 ± 0.11
||1.21 ± 0.02
||0.70 ± 0.06
||0.46 ± 0.02
Table 2. Leaf blade size in black-fruited aronia
|Herbarium Specimens Provenance
| Riga (Latvia)
||41.7 ± 0.4
||23.8 ± 0.3
||1.83 ± 0.21
||51.7 ± 0.9
||34.0 ± 0.8
||1.54 ± 0.01
||55.7 ± 1.6
||34.3 ± 1.1
||1.58 ± 0.03
| Virginia 1*
||56.2 ± 2.8
||19.9 ± 1.1
||2.89 ± 0.12
| Virginia 2
||59.1 ± 5.8
||20.1 ± 2.4
||3.09 ± 0.21
| Virginia 3
||57.1 ± 3.2
||20.6 ± 1.1
||2.78 ± 0.11
| Virginia 4
||50.3 ± 7.4
||17.8 ± 3.0
||2.97 ± 0.25
||33.4 ± 3.0
||17.0 ± 2.7
||2.04 ± 0.13
| Quebec 1
||51.9 ± 3.1
||22.1 ± 2.0
||2.43 ± 0.19
| Quebec 2
||18.9 ± 0.9
||9.3 ± 0.6
||2.06 ± 0.04
| Quebec 3
||31.5 ± 1.5
||16.5 ± 0.8
||1.92 ± 0.05
|* collection number
Morphological characters of aronia vegetative organs have proved to be constant in
different regions of the USSR. Even the leaf shape and size show only insignificant
variability (Table 2).
Unfortunately, none of the seeds collected or received via mail from North America
germinated. Seeds from across this country germinated all at once, in 10 days upon
During their first year, the total biomass of the above-ground parts was measured
for plants of various provenances (Table 3). While it was found that the biomass of
a single plant can vary within 40-60 mg, mean values for plants of different
provenances were practically the same.
In the second-year plants, the biomass of stems and leaves was measured separately.
The leaf biomass was observed to be somewhat larger than that of stems. Due to some
variability in biomass among different cultivated plots, the Student's Criterion
indicated unreliable differences between mean biomass values of different
Comparisons of annual increments among samples of various provenance (Table 3) did
not reveal any significant differences.
Variability of the measured parameters turned out to be low (variability
coefficient was 10%). These data demonstrate high genotypic stability of characters
in aronia across all of its secondary area.
Table 3. Size and biomass of cultivated aronia seedlings
originating from different regions of the USSR
(biomass values per single plant)
(N to S, W to E)
||3.8 ± 0.2
||5.2 ± 0.8
||3.9 ± 0.2
||7.2 ± 0.4
||3.9 ± 0.1
||6.5 ± 0.5
||3.8 ± 0.1
||6.6 ± 0.4
||3.7 ± 0.1
||8.0 ± 0.5
||3.8 ± 0.2
||5.8 ± 0.5
||4.0 ± 0.2
||6.9 ± 0.6
||4.1 ± 0.2
||5.4 ± 0.8
||4.2 ± 0.1
||7.6 ± 0.6
||3.8 ± 0.1
||6.8 ± 0.3
Differences of cultivated aronia from its wild American ancestors
The measurements of fruit diameter and dry mass demonstrated reliable differences
between the plants from primary and secondary aronia range. Mass of fruits from
plants collected in a number of geographic destinations within the USSR is 2-3
times as large as that of fruits originating from North American plants (see Table
1). Plants grown in the West and Central European botanic gardens also have fruits
about 1.5-2 times as light as those of our cultivated aronia. V.I. Protsenko
] arrived at similar results.
In addition, the following morphological and qualitative differences were observed.
In the North American wild-collected plants, fruits are oval or slightly pyriform
(pear-shaped), shiny. In the cultivated aronia, they are globular, mostly somewhat
depressed (at least at the apex), and always opaque. Fruits from cultivated aronia
are more juicy than those wild-collected, hence they shrink much on drying, thus
partially alleviating drastic size differences between wild-collected and
cultivated specimens observed in fresh material.
Browsing through herbarium collections suggested differences in size and quantity
of flowers: North American plants have fewer and smaller flowers than specimens of
cultivated aronia. Reliable differences in the shape and size of leaf blades were
also revealed (see Table 2). We observed the significant infraspecific variability
in North American plants previously described by American researchers. North
American herbarium samples were greatly variable in leaf size and shape: from
small, round leaves to large, lanceolate ones with attenuate apex.
While North American aronia is only moderately hardy (Zone 4, according to
]), aronia cultivated in the USSR is extremely hardy, so that
it is possible to assign it to Zone 2.
Taking all these facts into account, one cannot identify plants cultivated in the
USSR with any of the forms occurring within aronia's natural area. The cultivated
aronia is reliably different from wild-collected in a number of quantitative and
Since the cultivated aronia exhibits distinct differences from its wild ancestors,
remains constant in its characteristics, and has acquired a very wide range across
Northern Eurasia, it appears to be quite reasonable to describe it as a new species
Fig. 2. [Not provided] Herbarium sample of cultivated aronia, A. mitschurinii (USSR,
Moscow, Moscow University Botanical Garden. Cultivated. 6 Aug 1973. coll. A.
Aronia mitschurinii Skvortsov et Maitulina sp. nova.— Frutex sat robustus,
ad 3 m (vulgo 1.25-2 m) altus, ramulis, foliis pedicellisque plus minusve
cinereo-tomentosis, foliis late ovalibus, corymbo multifloro, baccis sphaericis vel
apice ± applanatis, atrofuscis opacis in vivo 9-12 mm in diametro.
Habitat: in hortis numerosis-sintis in URSS culta, specimina silvestria abhuc
Typus: Mosqua, Hortus botanicus principalis, culta, 10.9.1980 leg. J. Maitulina,
A specie affini (et verosimiliter parentali), A. melanocarpa (Michx.)
Elliott habitu robustiore, pubescentia fere omnium partium, corymbo conspicue
majore necnon fructibus multo majoribus opacis manifeste discernitur.
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plants, which have flowered in the garden of the Horticultural Society at Chiswick.
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America containing abridged descriptions of all the known indigenous and
naturalized plants growing north of Mexico
. Vol. 1. N.Y., London.
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aronia introduction in Kaluga Oblast.] — In: Vtoraya krayevedcheskaya
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culture], Kaluga. P. 60-64. In Russian.
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selection in black-fruited aronia]. — Byull. Sibir. bot. sada
10: 58-60. In
10. Raudsepp, A.D. 1976. [Effect of chemical
mutagens on black-fruited aronia seedlings.] — In: Effektivnost khimicheskikh
mutagenov v selektsii
[Chemical mutagen efficiency in selection]. P. 297-301.
Moscow: Nauka Publ.
The Central Botanical Garden USSR Academy of Sciences
Translation I. Kadis
30 December 2011