The Families and Genera of Vascular Plants

62 J.M. Arrington and K.
Kubitzki
Cistaceae
J.M. Arrington and K. Kubitzki
Cistaceae Juss., Gen. Pl.: 294 (1789), nom. cons. Vegetative Morphology and Anatomy. Cis-
taceae are predominantly shrubs or subshrubs;
Shrubs, subshrubs or herbs, often with stellate annuals are restricted to Tuberaria sect. Scorpi-
indumentum. Leaves alternate or opposite, simple, oides and Helianthemum sect. Brachypetalum
entire, often revolute, petiolate to sessile, stipulate (sensu Grosser 1903). Perennial herbs are lacking
or exstipulate; leaf venation pinnate, palmate, 1- in the family, and those appearing as such, as
nerved, or 3 more or less parallel veins. Flowers species of Lechea or Tuberaria sect. Tuberaria,
solitary or in cymose inflorescences, hermaphro- have a persistent, woody tap root and therefore are
ditic, actinomorphic, hypogynous, chasmogamous considered as halfshrubs (Troll 1969). Stipules are
or cleistogamous; sepals 3 or (4)5, when 5, the outer reported to occur erratically in the family but it
2 often bract-like and narrower or shorter, or needs to be explored whether or not these struc-
connate to the inner 3 prominent sepals, convolute, tures correspond to prophylls or other leaves
often persistent; petals yellow, white or reddish developed on precocious axillary shoots.
(maroon, pink, purplish, orange), free, (3)5 or 0 The combination of diverse hair types on the
(cleistogamous flowers), convolute in bud in the calyx, stem, and leaf is a characteristic of the
opposite direction to that of the sepals, ephemeral family. Hairs are glandular (multicellular and cap-
or rarely marcescent; stamens numerous, or 3–10 itate or elongate-uniseriate, rarely peltate scales)
(cleistogamous flowers, some annuals, Lechea), or nonglandular (simple, tufted, stellate). Stellate
filaments distinct, sometimes sensitive; anthers hairs are actually clustered, “falsely bicellular”
basifixed, introrse, tetrasporangiate, 2-locular at unicellular trichomes (similar hairs occur only in
anthesis, longitudinally dehiscent (sometimes Combretaceae). The illusion of basal bicellularity
apically in cleistogamous flowers); gynoecium is due to a conical or concave cellulose membrane
(2)3–5(6–12)-carpellate; ovary 1- or imperfectly within the cell wall (Grosser 1903; Metcalfe and
(2)3–5(6–12)-locular due to intruded parietal Chalk 1950). The assemblage of this hair type is a
placentae; placentae filiform (shield-like); ovules distinctive trait of the Cistaceae.
2–many on each placenta, orthotropous (ana- The leaves of the Cistaceae often possess cys-
tropous, hemianatropous), bitegmic and crassinu- toliths. Stomates are anomocytic, present on one
cellar; funicles well-developed; style simple, long or both leaf surfaces, and without subsidiary cells.
to very short or wanting; stigma large and capitate In the minor leaf veins, phloem transfer cells
or discoid, often lobed, seldom minute, or rarely are present (e.g., Helianthemum) or absent (e.g.,
stigmas 3 and fimbriate-plumose. Capsule loculici- Cistus).
dal (septifragous), (2)3–5(6–12)-valved, upright Nodes have been reported to be unilacunar.
or pendent, few- to many-seeded; often enveloped However, a comprehensive survey of the taxa and
by persistent sepals. Seeds very small; seed coat of trace-type appears to be lacking. The xylem of the
2 integuments, the outer thin, sometimes gelati- Cistaceae consists of both vessels with simple end
nous when moist, the inner very hard; endosperm walls and tracheids. Fibre tracheids are present;
starchy, nuclear, thin-walled; embryo curved, bent vestured pits have been observed in the secondary
into a hook or ring, more or less circinately coiled, xylem of Cistus (Baas and Werker 1981). Wood
plicate or biplicate,rarely almost straight,central or rays are low and heterocellular, mostly uniseriate.
more or less peripheral; cotyledons narrow to ellip- Wood parenchyma is wanting or very scanty. Sec-
tical-orbicular,straight to strongly bent. ondary thickening of the stem develops from a
The family comprises 8 genera and about 180 typical cambial ring. Sieve elements of the phloem
species mostly in temperate and subtropical are of the Ss-type (Behnke 1991); internal phloem
regions of the northern hemisphere, especially the is absent. Stems in cross section are cylindrical
western Mediterranean region, with a secondary or tetragonal. Root hairs are lacking; see under
center in the eastern United States. Distribution and Habitats below.
K. Kubitzki et al. (eds.), Flowering Plants · Dicotyledons

© Springer-Verlag Berlin Heidelberg 2003
Cistaceae 63
Inflorescence and Floral Structure. Cleistogamous flowers regularly occur in certain

Flowers are solitary or grouped in unilateral, scor- species of Helianthemum, Tuberaria and Crocan-
pioid, or more or less symmetrical cymose inflo- themum, and rarely (M. Sanchez and J. Güemes,
rescences. Flowers usually open only in full pers. comm.; Grosser 1903) in Fumana. Cleis-
sunlight for a few hours, and the petals are typi- togamy tends to develop subsequent to chas-
cally ephemeral. Most flowers have five conspicu- mogamy in late summer and autumn. However,
ous petals, exceptions include cleistogamous chasmogamous and cleistogamous flowers regu-
flowers and Lechea (three minute petals). The larly co-occur in Helianthemum sect. Eriocarpum
petals have a multipapillate epidermis with about (arid regions of the Old World) and Crocanthe-
10 micro-papillae per cell which project from an mum sect. Lecheoides (New World). In cleistoga-
elongate epidermal cell (Barthlott 1981). Petals mous flowers, the petals (sometimes absent) and
with such an epidermis are rare among sepals are smaller, the stamens are usually reduced
angiosperms (Kay et al. 1981), and the cistaceous in number and size, and the capsules are often
type is a potential apomorphy for the family. smaller and contain fewer seeds. The anthers of
Stigmas are dry and possess a papillate surface. cleistogamous flowers in Crocanthemum sect.
The multicellular and multiseriate papillae are Lecheoides and Helianthemum sect. Eriocarpum
exclusive to the Cistaceae among other malvalean are usually fused to the stigma and apically
families (Nandi 1998a), and are a rare feature dehisce at the point of adherence (Grosser 1903;
among the angiosperms (Heslop-Harrison and Daoud and Wilbur 1965). The occurrence of cleis-
Shivanna 1977). togamy, reported in Halimium (Lord 1981) and
Floral ontogeny has been studied by several Lechea (Darwin 1877), should be further investi-
authors, most recently by Nandi (1998b). The gated, especially in the flowers of Lechea which
irregular calyx (unequal and/or variable in sepal rarely fully expand.
number) is a distinguishing characteristic for the
family. The basic sepal number is five. Sepals arise Embryology. Anthers are tetrasporangiate; the
sequentially; in some species the inner three tapetum is glandular. Microsporogenesis is simul-
develop from a ring primordium. Early in devel- taneous, and the initial microspore tetrads are
opment five sepal primordia appear but later on in tetrahedral, isobilateral or decussate. Ovules are
some species sepals 1 and 4, and/or sepals 2 and 5 bitegmic, crassinucellar, and orthotropous, except
become partially or wholly fused (Saunders 1936). for Fumana where the ovules are anatropous to
The androecium develops centrifugally and hemianatropous. The micropyle is formed by both
usually from a ring wall primordium (in taxa with integuments, mostly the outer (Nandi 1999b). The
a higher number of stamens). The first stamens to embryo sac formation follows the Polygonum
be formed on the ring wall tend to appear in type. The endosperm is nuclear (Kapil and
alternipetalous position (Saunders 1936; Nandi Maheswari 1964; Davis 1966).
1998b). Further stamen primordia appear in the
gaps between the previously formed primordia. In Pollen Morphology. (Heydacker 1963;
the genera examined by Nandi (1998b), Lechea is Ukraintseva 1993; J.M. Arrington, pers. obs.).
exceptional in producing single and compound Pollen grains are spheroid, oblate or prolate and
stamen primordia. The epidermis of cistaceous tricolpor(oid)ate monads; two Crocanthemum
anthers is persistent and the endothecium devel- spp. are reported as having loose tetrads (Tomb
ops fibrous thickenings. 1999; J.W. Horn, pers. comm.). Exine sculpture is
In Cistus, the gynoecium is 5(6–12)-carpellate, reticulate, rugulose, striate, or striate-reticulate.
but in all other Cistaceae it is 3-carpellate. Placen- Pollen morphology is largely in accord with
tation is parietal, but the placentae can be deeply generic, subgeneric and sectional classification.
intruded (e.g., Cistus) and then appear axile. The Six major pollen types can be recognized: (1)
shape is generally filiform with the exception of the Fumana type is tricolpor(oid)ate, oblate-
Lechea in which the placentae are shield-like. spheroidal to spheroidal with short, broad colpi
Funicles are well-developed and vary from filiform and large, unequal lumina; (2) the Lechea type is
(e.g., Cistus, Halimium, Crocanthemum, Hudsonia, tricolporate, oblate-spheroidal to slightly prolate-
a few Helianthemum) to robust (Lechea) and spheroidal with irregular long colpi and a distinc-
swollen in the center (e.g., Tuberaria) or towards tive infra-reticulate texture; (3) the Helianthemum
the end (e.g., most Helianthemum). They are type is tricolporate, prolate-spheroidal to prolate
ascending, somewhat straight or resupinate, and with long colpi; subtypes correspond to sub-
elongate or short. generic taxonomy; (4) the Cistus type is tricol-

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62 J.M. Arrington and K.

K. Kubitzki et al. (eds.), Flowering Plants · Dicotyledons

Inﬂorescence and Floral Structure. Cleistogamous ﬂowers regularly occur in certain

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