R. Rutishauser et al., THELIGONUM CYNOCRAMBE - DEVELOPMENTAL MORPHOLOGY OF A PECULIAR RUBIACEOUS HERB, Plant systematics and evolution, 210(1-2), 1998, pp. 1-24
The annual Mediterranean herb Theligonum cynocrambe shows a peculiar c
ombination of morphological characters, e.g., switch from decussate to
spiral phyllotaxis with 90-100 degrees divergence, combined with a ch
ange from interpetiolar to lateral stipules, anemophily, lack of calyx
, flowers often dimerous to trimerous, corolla fused in both male and
female flowers, male flowers extra-axillary, with 2-19 stamens per flo
wer, female flowers axillary, with inferior uniovulate ovary, basilate
ral style and perianth, nut-like fruits with elaiosome. In male flower
s the androecium emerges as an (uneven) elliptical rim with a central
depression. This common girdling primordium is divided up into several
stamen primordia. In male flowers with low stamen number the stamen p
rimordia may occupy the corners alternating with the corolla lobes. Th
ere are no epipetalous androecial primordia that secondarily divide in
to stamens. Male flowers occasionally show a hemispherical base that m
ay be interpreted as remnant of the inferior ovary. In female flowers
a ring primordium grows into a tube on which the petal lobes arise. Th
e perianth and style become displaced adaxially by uneven growth of th
e inferior ovary. The ovary is basically bilocular. The lower region o
f the ovary is provided with a septum that is overtopped and hidden by
the single curved ovule. Theligonum is referred to the Rubiaceae-Rubi
oideae, with the Anthospermeae and Paederieae as most closely related
tribes.