23 CACTUS AND SUCCULENT JOURNAL
H. DAVID JIMENO SEVILLA1,2, ANGÉLICA M. HERNÁNDEZ RAMÍREZ1,3 & THORSTEN KRÖMER1,4
Echeveria rosea Lindley
(Crassulaceae): a hummingbird-
dependent succulent epiphyte
The stonecrop family Crassulace-
ae has a worldwide distribution
with exception of Australia and
the Polynesian area. There are
about 1,500 species of Crassu-
laceae distributed in 33 genera,
which are characterized by their
rosetted succulent leaves. The family is often viewed
as a typical northern temperate element, although its
highest species richness is concentrated in Mexico
(about 325 species) and South Africa (about 250 spe-
cies). Most species grow saxicolous and/or terrestri-
al in dry areas and show Crassulacean Acid Metabo-
lism (CAM) as photosynthetic pathway, which is an
adaptation to arid conditions. In contrast, only seven
species in three genera (Echeveria, Kalanchoe, Sedum)
have been reported to grow mainly epiphytic (Kress
1989). Furthermore, Crassulaceae species are known
for a variable floral morphology, which is attractive for
a diverse spectrum of pollinators.1
A general approach to specify the pollination biol-
ogy of the Crassulaceae family includes the establish-
ment of five major pollination syndromes based on
1. Melittophily or bee pollination is assumed for
species with short tubular corollas, which is
the most frequent and least specialized floral
characteristic in Crassulaceae (Figs. 1 & 2),
2. Psychophily or pollination by butterflies is
associated with long-tubed salver-shaped
flowers or flowers with petals forming a tube-
like structure, which have an intensive color-
ation (red, yellow) and a perfume-like scent
production over day (Fig. 3),
3. Sphingophily or pollination by hawkmoths
corresponds to long, whitish corolla tubes and
nocturnal strong sweet scent,
1Centro de Investigaciones Tropicales, Universidad
Veracruzana, Xalapa, Veracruz, México.
1. Sedum praealtum and Bombus sp. Tlacolulan,
2. Villadia elongata and Apis melifera. Huayacocotla,
2013 VOLUME 85 NUMBER 3 24
4. Ornithophily or bird pollination is assumed to
red, long-tube corollas, abundant nectar pro-
duction and exserted anthers, and
5. Myophily or pollination by flies corresponds to
open flowers with darkish colors, freely acces-
sible nectar produced by large nectary scales
and foetid flowers (Thiede & Eggli 2006),
Nonetheless, studies on the pollination biology of
Crassulaceae family members are scarce, and only a
few species of the genera Aeonium (Esfeld et al. 2009,
Nelson 2010), Crassula ( Johnson et al. 1993), Diamor-
pha (Wyatt 1997, 1981), Dudleya (Aigner 2005, Jones
et al. 2010), Echeveria (Parra et al. 1993, 1998, Vargas
& Parra-Tabla 2002), and Sedum (Shahani 2007) have
been investigated in detail. Thus, for the great major-
ity of the species, there is no information on this sub-
Mexico as a biodiversity
center of Echeveria
Echeveria is represented by about 140 species dis-
tributed from southwest Texas to Central and South
America. Of these, about 110 species occur in Mexico
and 107 species (97.2%) are endemic to the country
(Pérez-Calix & Franco-Martínez 2004). Furthermore,
Echeveria is the most speciose genus of the Crassu-
laceae family in Mexico. The name of the genus was
honored to the Mexican illustrator and botanist Ata-
nasio Echeverría y Godoy, who made several scientific
illustrations for the Flora of Mexico in collaboration
with the Spanish botanist Martín de Sessé y Lacas-
tra and the Mexican naturalist José Mariano Moci-
ño during the “Real Expedición Botánica a la Nueva
España” in the XVIII century.
3. Crassula coccinea and butterfly. Drawing by H. David Jimeno Sevilla.
25 CACTUS AND SUCCULENT JOURNAL
The Echeveria genus seems to include a small spe-
cialized group of epiphytic succulent plants, which is
an exception in the Crassulaceae family. At least in
this genus, the epiphytic habit appears to be associat-
ed with humid montane forest, whereas most species
grow saxicolous in zonal arid habitats such as rocky
slopes and rock fissures under otherwise more humid
climatic conditions, or in mountain regions of moder-
ately arid areas.
Echeveria rosea an endemic
plant of Mexico
Echeveria rosea Lindley is a succulent epiphyte that
occurs in cloud and pine-oak forests from Tamauli-
pas to Chiapas, with a disjunct distribution in Guer-
rero. The species usually flowers from November to
February and has showy inflorescences with terminal
and lax racemes. The length of the inflorescences is
between 21-40 cm, each bearing 12-41 flowers, with
one to four open flowers per day. The yellow flower is
hermaphroditic and subtended by a red colored flo-
ral bract. The petals are fused at the base, providing
a receptacle for nectar. The flowers of E. rosea have
attributes that are related to hummingbird-pollination
(ornithophily), i.e., long-tubed corollas, lack of odour,
abundant nectar production and exserted anthers
(Parra et al. 1993).
Pollinators and floral visitors
Nectar is the most common form of floral reward
provided by animal-pollinated plants to their mutual-
istic partners. The preliminary results of our research
on the pollination biology of Echeveria rosea realized
in a pine-oak forest at “Cerro de la Magdalena”, cen-
tral Veracruz, indicate that Garnet-Troated Hum-
mingbird (Lamprolaima rhami) is the most frequent
floral visitor (Fig. 5), followed by White-Eared
Hummingbird (Basilinna leucotis = Hylocharis leu-
cotis), whereas Magnificent (Rivoli’s) hummingbird
(Eugenes fulgens) only occasionally visits E. rosea. Al l
mentioned hummingbirds forage for nectar and con-
tact the anthers and/or stigmas of the flowers, sug-
gesting that these are the main pollinators of E. rosea.
Interestingly, the geographic distributions of E.
rosea and Garnet-Troated Hummingbird in Mexico
are coincident. Moreover, the breeding season of Gar-
net-Troated Hummingbird in Mexico occurs when
E. rosea blooms. In this scenario, the relation between
Echeveria rosea and Lamprolaima rhami seems to be a
keystone mutualistic interaction with potential eco-
logical and evolutionary processes involved.
Other floral visitors are mites, however, flower
mites are considered as nectar robbers, because they
live and reproduce in flowers of the plant species, and
feed on their nectar and pollen without a pollinator
service for the plant.
The cloud forest habitat of both species is threat-
ened, because it is strongly disturbed by illegal defor-
estation and agropecuary activities. As Echeveria rosea
is an epiphyte and thus dependent on the availabil-
ity of adequate host trees, it is very susceptible to dis-
appear by immoderate land use changes. Furthermore,
the Garnet-Troated Hummingbird is included in the
IUCN Red List of threatened species (BirdLife Inter-
national, 2009) and protected under Mexican law
(SEMARNAT, 2010). Thus, it is important to get a
better knowledge about the relationship of E. rosea
and their pollinators to devise adequate conservation
strategies in the future.
This study was funded by the Consejo Nacional
de Ciencia y Tecnología (CONACyT, grant number
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