25. Rubus L. (raspberry, blackberry, bramble)
Contributed by Mark P. Widrlechner
Plants shrubs
(perennial herbs elsewhere), sometimes suckering or reproducing vegetatively
from rooted stem tips. Stems (referred to as canes) biennial or (in sect. Rubus)
occasionally persisting longer, prostrate, climbing, arched, or erect, all
those found in Missouri armed with prickles, sometimes also with long, stiff
bristles (except for horticultural selections); first year’s stems generally
vegetative (called primocanes), unbranched or few-branched (moderately branched
in a few species); second year’s stems fertile (called floricanes), usually
dying back to the rootstock at the end of the second growing season. Leaves
rarely evergreen. Stipules small to conspicuous, persistent, leaflike, fused to
the petiole laterally or attached at the junction of the stem and petiole. Leaf
blades palmately or pinnately compound, the margins toothed and sometimes
lobed, the surfaces often hairy, sometimes with gland-tipped hairs or bristles,
the main veins sometimes also with prickles. Inflorescences typically axillary
on floricanes, of branched or simple clusters, racemes, panicles, or
occasionally solitary flowers. Flowers perfect (imperfect elsewhere), with a
noticeable hypanthium. Sepals 5, fused basally, persistent at fruiting, erect,
horizontally spreading, or reflexed. Petals usually 5 (rarely more in
horticultural selections), white or less commonly pale pink to rose pink,
glabrous or hairy, the margins usually entire, rarely lobed (in R.
laciniatus). Stamens numerous, the filaments attached at the mouth of the
hypanthium. Pistils many, attached to an enlarged receptacle, this hemispheric,
or becoming conic or cylindric as the fruits mature, each with 1 carpel and 2
ovules, 1 of these aborting as the fruit develops. Style 1, threadlike or
occasionally narrowly club-shaped, glabrous or hairy. Fruits drupelets
positioned on the enlarged receptacle to form an aggregate, which either
separates freely from the receptacle when ripe (raspberries) or is shed with
the receptacle attached (blackberries). About 800 species (although species
concepts and the resulting estimates differ widely), worldwide, particularly
abundant in temperate regions of the Northern Hemisphere.
Rubus is a large genus that is diverse in the
midwestern United States. Like Crataegus, its reputation for
hybridization, polyploidy, and apomixis has given it a well-deserved reputation
as a taxonomically difficult genus. Additionally the prickly canes that change
appearance from the first to second growing seasons have caused many collectors
to avoid the plants. Rubus is thus underrepresented in herbaria and many
of the specimens that are present are relatively incomplete. Plants often
thrive in disturbed, successional habitats, sometimes in mixed-species
thickets. A number of mixed collections in various herbaria involve the
primocanes of one species and the floricanes of a second species growing at the
same site.
The genus
produces a number of important fruit crops, including raspberries,
blackberries, and loganberries. In addition to being consumed as fresh fruits,
the fruits are used extensively in baked goods, jams, jellies, preserves, and
as juices and flavorants. The fruits also are an important food source for
wildlife. A few species are cultivated as ornamentals.
Whenever
possible, users of this treatment should examine primocane leaves for
identification characters involving leaves. References to primocane leaves and
leaflets in the descriptions that follow are based on fully expanded leaves. It
should be noted that in leaves with five leaflets, the apical or middle leaflet
is called the central leaflet, the adjoining pair of lateral leaflets are
termed the middle leaflets, and the pair positioned closest to the petiole are
known as basal leaflets. Floricane foliage is primarily found on inflorescence
branches, but can also be found on sterile branches. Large, sterile branches
that sometimes form near the base of floricanes are called parcifronds and can
easily be confused with primocanes. Floricane leaves on sterile branches
generally resemble those of primocanes, but often are smaller and have only 3
leaflets. Floricane foliage on inflorescence branches is denoted in the species
descriptions below as bracts or leafy bracts.
In the present
treatment, inflorescences are described and measured from well-developed
examples on floricanes. Inflorescences can also emerge directly from the crown,
generally when a primocane has been damaged or winter-killed. These atypical
inflorescences are called novirames, can flower out of season, and are
extremely difficult to identify to species.
North American
herbaria hold thousands of fragmentary Rubus specimens that cannot be
determined to species. When vouchering Rubus, it is important to collect
pieces of both the primocane and floricane, with the primocane displaying both
sides of fully-expanded leaves and the floricane displaying well-developed
inflorescences, collected at any time between the start of flowering and the
formation of ripe fruits. These pieces should be kept together, either by
mounting on the same sheet or through appropriate labeling. On labels, it is
also useful to note the overall plant habit and the presence or absence of
rooting at the tips of the canes.
The genus Rubus
generally is divided into five subgenera and numerous sections. The
relationships among the sections are poorly understood, and it is not clear if
the traditional classification circumscribes natural subgroups. The
nomenclature of the sections is also still somewhat controversial, as Liberty
Hyde Bailey (1858–1954; the leading specialist on the genus in the first half
of the twentieth century) initially treated these as unranked groups and by the
time that he validated the sectional names, some might already have been
published by other authors. Thus the names applied to the sections should be
treated as provisional, pending further review of the literature on Rubus.
Grouping the species into morphologically based subgenera and sections, as is
done in the present treatment, has the advantage of saving space by requiring
that characters shared among species within a section be listed only once in
the section description. Users are cautioned to read the subgeneric and
sectional descriptions carefully in addition to the species descriptions for
help in confirming the identity of plants being determined. The following key
to the subgenera and sections of Rubus uses a combination of characters
from primocanes and floricanes, but can be navigated with only one type of
cane, with the exception of couplet 4, which requires inflorescences.