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Inflorescence and Flower Bracts

by Bob Harms  email-here
I view the bracts of B. trifoliolata and B. swaseyi as consisting of four 3-merous structures – i.e., a 3 prophyll spiral plus 3 x 3 sepal whorls. The traditional view is that a 'bract' occurs at or toward the base of the pedicel accompanied by two prophylls higher on the pedicel or attached to the receptacle. The images below clearly demonstrate that for these two species the 'bract/prophyll' forms are morphologically identical, differing only in size. Other Berberis/Mahonia species may not support this interpretation. And the prophylls are quite distinct from the outermost sepaloid bracts (III), sometimes considered to be bracteoles, constituting an epicalyx or calyculus. I share the view that these are sepals, supported by their similarity in form and color to sepals of the median whorl (II).

Key to labels in the images.
1,2,3 = prophylls III = outermost sepals
II = median sepals I = innermost sepals

B. swaseyi B. trifoliolata
B. swaseyi B. trifoliolata, inflorescence bases emerging from bud scales.
Click here to open page with additional images

Bracts, Prophylls, Bracteoles, Sepals in the Berberis (Mahonia) Literature

cataphylls basal prophyll upper 2
prophylls
epicalyx sepals
Pursh 1814       3 x 3 leaves
calyx triplex
Nuttall 1818       Calyx 9-leaved, ... in 3 series,
exterior bracteiform and small
Bonpland et al 1821       Calyx with 3 bracts
tribracteatus
and 6 leaves
hexaphyllus
Payer 1857 bud scale
écaille de bourgeon
basal bracts of infl. (2)
bractée foliacées
basal bract of fl. (1)
bractée mère
prophylls (2 + 3)
bractées secondaires
sepals proper (2 x 3)
sépales
Wooten 1898*     bracts (2) sepaloid
bracts (3)
sepals proper
Fedde 1901 bud scales
Knospenschuppe
bract
Tragblatt
prophylls (2 or absent)
Vorblatt
sepals (3 x 3)
Kelchblatt
Ahrendt 1961 bracts of
inflorescence
bract prophylls (2 or absent) sepals (3 x 3)
Harms** bud scales prophylls (3) sepals (3 x 3)
Hitchcock &
Cronquist 1973
bract     bracts (3) sepals (2 x 3)
Munz & Keck 1959       bractlets (3) sepals (2 x 3)
Whittemore (FNA)   ? bracteoles
(of infl.)
bracteoles (3)
(of flower)
sepals (2 x 3)
 *Wooten describes B. haematocarpa.
**In comparing the interpretations of the various authors in this table, I hasten to note that I have examined only
two species in detail, both from the Section Horridae, and my views may not be appropriate for other species.

Payer 1857 was the first to provide a detailed examination of Berberis floral morphology. And with the exception of Fedde's and Ahrendt's monographs the structure of the Berberis/Mahonia inflorescence has received scant attention in the botanical literature beyond discussion of what to call various floral components, and perhaps this accounts for the lack of a standardized descriptive terminology. Ahrendt 1961 notes (p.16) that

"different authors, or even the same author at different times, have referred to the outer flower components of the same species indiscriminately as bracteoles and outer sepals. Thus ... described, at one time as possessing prophylls and two whorls of sepals, and elsewhere as having no prophylls and three whorls of sepals.
[and for Mahonia]
it is generally possible, after experience in examining the flowers to make a satisfactory distinction. Whereas the prophylls are nearly always 1 or 2 in number, and triangular and acuminate, the small outer sepals in a whorl of 3, are ovate, and acute.

Whetstone et al (FNA) clarify their usage:

The perianth of Berberidaceae is commonly composed of three distinct types of organs, but terminology for the organs varies from author to author. In our treatment, we refer to the small, outer parts as bracteoles (collectively forming a calyculus); the large, middle parts as sepals; and the innermost parts, which are commonly nectariferous, as petals. Some authors have referred to the bracteoles as outer sepals and to the petals as staminodes.
Whittemore, as in his FNA description of Berberis swaseyi, follows this usage of bracteole for the perianth, but also as prophyll of the inflorescence:
bracteoles leathery, apex spinose-acuminate, sometimes with proximal bracteoles as described, distal membranous and acuminate
Although in this later sense it is difficult to discern whether he counts 3 prophylls (bracteoles) or 2. The above description of B. swaseyi is appropriate only if 3, since the upper (distal) 2 prophylls of a spiral series commonly reduce in this manner - and at times even all three in racemes with more than 3 flowers.