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Cactus seed germination: A review



The present review tries to give a general overview of the available information on cactus seed germination. First, information about the family Cactaceae is discussed, concerning aspects such as distribution and general characteristics. Seed distinctive features are mentioned, such as colour, form, and size. Aspects of seed physiology, such as germination and dormancy, as well as seed dynamics including dispersal, predation, and soil seed bank formation, are included in the discussion. Techniques of propagation and some aspects of longevity and conservation are mentioned. The areas where there is scarce information available are highlighted, and, therefore, are important areas in which to continue research in order to generate data for immediate and future conservation efforts.
*Author for correspondence. E-mail:
Journal of Arid Environments (2000) 44: 85–104
Article No. jare.1999.0582
Available online at on
Cactus seed germination: a review
Mariana Rojas-AreHchiga* & Carlos VaHzquez-Yanes
Instituto de Ecologn&a, UNAM, Apartado Postal 70-275,
04510 Me&xico, D.F., Me&xico
(Received 8 June 1998, accepted 10 August 1999)
The present review tries to give a general overview of the available information
on cactus seed germination. First, information about the family Cactaceae is
discussed, concerning aspects such as distribution and general characteristics.
Seed distinctive features are mentioned, such as colour, form, and size. Aspects
of seed physiology, such as germination and dormancy, as well as seeddynam-
ics including dispersal, predation, and soil seed bank formation, are included in
the discussion. Techniques of propagation and some aspects of longevity and
conservation are mentioned. The areas where there is scarce information
available are highlighted, and, therefore, are important areas in which to
continue research in order to generate data for immediate and future conserva-
tion efforts.
(2000 Academic Press
Keywords: cactus seed; dormancy; germination; propagation; longevity;
dissemination; conservation
Arid and semi-arid regions cover about 30% of the world’s continental surface (Meigs,
1953 in Kigel, 1995). The best-represented plant families in these regions are
Asclepiadaceae, Aloaceae, Apiaceae, Asteraceae, Cactaceae, Chenopodiaceae, Euphor-
biaceae, Fabaceae, Malvaceae, Poaceae and Zygophyllaceae (Kigel, 1995).
Of these families, the Cactaceae are one of the most interestingdue to their extensive
set of peculiar adaptations to water scarcity, which allow them to be perennial and
evergreen despite the sometimes extreme dry conditions of their environment. The
Cactaceae are exclusively restricted to America (with the exception of the genus
Rhipsalis and some introduced species of Opuntia). The family consists of approxim-
ately 1500 to 2000 species distributed from the north of Canada to Patagonia in
Argentina. They mainly inhabit arid and semi-arid regions sometimes constituting
a conspicuous vegetation known as xerophilous scrubland. They also grow in temperate
forests, subtropical and tropical zones where some show an epiphytic form, and in cold
regions with winter snowfalls as in Canada or Argentina. The xerophilous family
is large and diverse, including broadleaf plants (Pereskia), giant arborescent plants
(Carnegiea gigantea, Pachycereus pringlei, P.schottii), columnar species (Neobuxbaumia
polylopha,Cephalocereus columna-trajani), candelabriform species (Myrtillocactus
0140-1963/00/010085#20 $35.00/0 (2000 Academic Press
geometrizans,Pachycereus weberi), globose forms (Mammillaria, Coryphantha), and
epiphyte forms (Hylocereus,Rhipsalis) (Bravo-Hollis & Scheinvar, 1995).
Cactus seed characteristics
Cactus seeds present considerable variations in form, size, structure, embryo character-
istics, and colour of the testa. Some external characteristics, such as form, size, number
of seeds produced by fruit, and colour of the seeds are of interest to this review. In
general, mature cactus seeds have the following parts: testa, embryo, endosperm,
perisperm (in most primitive groups), arillus cover (which characterizes the seeds of the
subfamily Opuntioideae; Buxbaum, 1951), funicle (Opuntia), and hilum (Elizondo-
Elizondo et al., 1994). Some species have a caruncle (Pereskia) (Dau & Labouriau,
1974) and a strophiole or raphe (Mammillaria erectacantha) (Fittkau, 1968). The
number of seeds produced by a single fruit can be enormous, sometimes more than 1000
seeds per fruit (Pilosocereus chrysacanthus; pers. obs.), or just a few (1 to 5 seeds per fruit in
Epithelantha and Pereskia aculeata)(LodeH, 1995; Pedroni & SaHnchez, 1997). Even within
one species the number of seeds per fruit can vary greatly. Zimmer (1966) reported that
some Epiphyllum anguliger fruits contain 1500 seeds, while others contain 5500, depend-
ing on the age of the plant, the number of flowers per plant, and the size of the plant. Del
Castillo (1988) reported that some Ferocactus histrix fruits contain up to 2200 seeds, while
other fruits have only 300. Otero & MeyraHn (1966) reported that Echinocereus pulchellus
had 40 to 100 seeds per fruit, LeoHndelaLuz&DommHnguez-Cadena (1991) reported 52
to 1566 seeds per fruit for Stenocereus gummosus,andWeisset al. (1995) reported 100 to
500 seeds per fruit for Selenicereus megalanthus. Cactus seeds can show diverse forms,
colours, appearances, and sizes which are shown in Table 1.
Seed predation
In desert areas, it is well known that there is a massive consumption of seeds (Gutter-
man, 1994). Most studies have been essentially concerned with annual plant seeds
(Morton, 1985; Boeken & Gutterman, 1990), and studies of cactus seeds have been
carried out on some Opuntiaspp., mainly in the Chihuahuan Desert‘nopaleras’ (Janzen,
1971, 1986; Grenot & Serrano, 1981; GonzaHlez-Espinosa & Quintana-Ascencio, 1986;
Vargas-Mendoza & GonzaHlez-Espinosa, 1992). Frugivores are very important seed
dispersal agents (Howe & Smallwood, 1982), and in arid and semi-arid habitats they
constitute important cactus fruit and seed predators. These predators are mainly
rodents, and to a lesser extent, ants, birds, lizards, and some mammals (GonzaHlez-
Espinosa & Quintana-Ascencio, 1986; Wendelken & Martin, 1988; Cortes Figueira et
al., 1994; May, 1994; Silvius, 1995). Wendelken & Martin (1988) demonstrated that 18
avian species are potential dispersers of Stenocereus eichlamii and Pilosocereus leucocephalus
seeds, but subsequent germination trials were not made. Cortes Figueira et al. (1994)
reported that Melocactus violaceus fruits are consumed by a specific lizard species and seed
dormancy was broken when the seeds passed through the lizard’s digestive tract.
The damage produced by the consumer to the seed is variable, for example, while
ants do not kill the embryo and only remove the pulp, funiculus rests, or mucilaginous
layer adhering to the testa (Del Castillo, 1988), rodents completely destroy the seed
(Vargas-Mendoza & GonzaHlez-Espinosa, 1992). Silvius (1995) worked with 14 avian
consumers of Stenocereus griseus and showed that while some birds destroyed the seeds
prior to or during digestion of the fruits, others defecated intact seeds viable for
germination. In all cases of non-damaged seeds, a portion can be incorporated into a soil
seed bank and can germinate if they are deposited onto a safe micrositewhere conditions
are suitable.
Table 1. Some cactus seed characteristics
Characteristic Examples References
Form Reniform Mammillaria meyranii var. michoacana,Neobux-
baumia macrocephala,Ferocactus flavovirens Buchenau, 1969; Bravo-Hollis et al.,
Globular Echinocereus grandis,Turbinicarpus lophophoroides Lindsay, 1967; Bravo-Hollis &
SaHnchez-Mejorada, 1991
Piriform Echinocereus pulchellus,Mammillaria varieaculeata,
M.nana,Melocactus macracanthos Buchenau, 1964, 1966; Otero &
MeyraHn, 1966; Antesberger, 1991
Hat-like Lophophora williamsii,Astrophytum capricorne,
Leuchtenbergia principis Bravo-Hollis, 1967; Elizondo-
Elizondo et al., 1994; Bravo-Hollis
&SaHnchez-Mejorada, 1991
Ovoid Ariocarpus kotschoubeyanus,Ferocactus
haematacanthus,Disocactus kimnachii Kimnach, 1984; Bravo-Hollis &
SaHnchez-Mejorada, 1991
Mussel-shaped Selenicereus wittii Barthlott et al., 1997
Lens-shaped Pereskia Dau & Labouriau, 1974
Colour and
appearance Black to brown colour Most seeds, Neobuxbaumia spp,
Peniocereus castellae Bravo-Hollis et al., 1970, 1971, 1973;
SaHnchez-Mejorada, 1973
Reddish black Carnegiea gigantea Gibson & Nobel, 1986
Reddish brown Pelecyphora strobiliformis Elizondo-Elizondo et al., 1994
White (with aril) Opuntia Gibson & Nobel, 1986
Tan Pterocactus Gibson & Nobel, 1986
Shiny Bergerocactus,Neobuxbaumia,
Disocactus kimnachii Moran, 1965; Bravo-Hollis et al., 1971;
Kimnach, 1984
Opaque Stenocereus chrysocarpus,Matucana formosa SaHnchez-Mejorada,
1972; Bregman et al., 1987
Size 40)5mm Blossfeldia,Strombocactus Gibson & Nobel, 1986
55)0mm Nyctocereus,Opuntia Bravo-Hollis & Scheinvar, 1995
Intermediate sizes Mammillaria magnimamma,Epiphyllum
phyllanthus,Selenicereus megalanthus Bravo-Hollis & Scheinvar, 1995;
Weiss et al., 1995
Neotropical arid and semi-arid habitats contain abundant, large columnar cacti, which
produce a considerable biomass of fleshy fruits(Silvius, 1995). The mode of dispersal of
certain cacti is strongly associated with the structural characteristics of both its fruit and
seed (Bregman, 1988). Cactus seeds are mainly transported in two ways by differ-
ent vectors: wind and animals. Water should be considered as a third vector of seed
Anemochory constitutes the least common mode of dispersal in the Cactaceae. The
winged seeds of the genus Pterocactus are dispersed by this method (Kiesling, 1968;
Bregman, 1988). Zoochory (dispersal by animals) can be divided into three categories:
(1) Endozoochory. Many cactus species produce fleshy fruits (berries) with bright
colours which are an attraction mechanism, encouraging consumption by
many frugivorous animals (birds, small mammals such as rabbits and other
rodents, reptiles, and bats). All these animals can disperse seeds away from the
parent plant by means of regurgitation or defecation, sometimes to safe sites for
germination and establishment. They function as a dynamic link between the
fruiting plant and the seed or seedling bank in their community (Jordano, 1992).
Most seeds that are dispersed via animals are characterized by a very thick or
resistant testa which can withstand stomachal acids and enzymes. For example,
Opuntia,Epiphyllum, Hylocereus, Pachycereus, Ferocactus, Melocactus,Carnegiea,
Sclerocactus polyancistrus,Pereskia aculeata and Stenocereus griseus (Bregman, 1988;
Del Castillo, 1988; Cortes Figueira et al., 1994; May, 1994; Silvius, 1995; Pedroni
&SaHnchez, 1997).
(2) Synzoochory. This is probably the most common mode of seed dispersal. Ants are
the main vectors, but other insects can also participate. Seed dispersal by ants
(myrmecochory) provides an additional advantage for the plant, in that a suitable
site for the seed to germinate is created and the seed is protected from predation.
Strombocactus, Aztekium, Opuntia, and Parodia are a few genera that show this type
of dispersal. The seeds of Blossfeldia liliputana, which are arillate and hairy (a
characteristic unique within this family), represent an adaptation to facilitate ant
dispersal (Barthlott & Porembski, 1996).
(3) Epizoochory. Seeds are transported passively on the outside of the animal. Seeds of
these species commonly have a thin testa.
Valiente-Banuet & Arizmendi (1997) mention two groups of dispersors among cacti:
(1) primary dispersors which take the fruit directly from the plant. They can be diurnal
(some birds and lizards), or nocturnal animals (certain species of bats); (2) secondary
dispersors (mainly rodents and ants) which take the fruit from the ground. Animals in
the first group of dispersors eat the fruits and defecate sometimes in shaded places,
where seeds can find good conditions for establishment. Animals in the second group of
dispersors carry the seeds to their nests where, if the seeds escape predation, they can
find suitable conditions for germination.
Hydrochory has been demonstrated to occur in species which are prevalent in river
valleys, such as the Peruvian genus Matucana (Anon., 1997b) and Selenicereus wittii,
which has a seed structure that functions as a floating device (Barthlott et al., 1997).
Seeds dispersed by water have floating mechanisms provided mainly by a large hilum,
deep hilum cup, thin seed coat, and small embryo (Bregman, 1988).
Germination and dormancy
Dormancy in plants is a process where physiological activities cease in a reversible
manner, even when conditions are suitable for germination (i.e. enough moisture, air,
and suitable temperatures). Dormancy has a survivalvalue when conditions for germi-
nation and establishment are not suitable. Cactus seeds, which inhabit places where
water is in short supply, are said to be quiescent. Three types of dormancy have been
described: innate, enforced and induced (Harper, 1957 in Murdoch & Ellis, 1992;
Roberts, 1972). Innate dormancy (or primary dormancy) prevents seed germination on
the mother plant and for a time after dispersal; enforced dormancy is regulated by
environmental conditions such as light and/or temperature, and seeds are ready to
germinate immediately upon removal of the environmental limitation; and induced
dormancy (or secondary dormancy) is characterized by the persistence of the dormant
condition even when the seeds are returned to favourable conditions for germination
(Tran & Cavanagh, 1984). For cactus seeds, innate and enforced dormancy have been
Information concerning the dormancy and germination behaviour of cactus seeds
comes from the pioneering studies of Alcorn & Kurtz (1959) and McDonough (1964),
which demonstrated that light has a stimulating effect on the germination of
Carnegiea gigantea and Stenocereus thurberi. Zimmer (1969c) worked with several species
of cactus and found that some germinated in the dark, while others needed varying light
intensities for germination. He also worked with different light qualities and found
that red light stimulated germination for all species studied (Zimmer, 1969d). More
recently, in other investigations, it was demonstrated that either light stimulated
germination, or seeds were strictly positively photoblastic (Fearn, 1981; MartmHnez-
HolgumHn, 1983;Arias &Lemus, 1984;Del Castillo,1986; Nobel,1988; LoHpez-GoHmez&
SaHnchez-Romero, 1989). In other studies, it was shown that the light requirement was
limited only within certain temperature ranges, such as in Cereus jamacaru (Arias
& Lemus, 1984) and Melocactusviolaceus(Dau & Labouriau, 1974). In some species the
light requirement is conditioned to seed washing prior to sowing, as in Melocactus
curvispinussp. caesius, whose seeds which were sown just after harvest did not germinate
under light or dark conditions, but washing before sowing stimulated germination giving
a 100% germination rate under light conditions (Arias & Lemus, 1984). In contrast,
seeds of Pereskia aculeata are indifferent to light under a wide range of temperatures
(Dau & Labouriau, 1974; Pedroni & SaHnchez, 1997).
Some studies have been carried out to assess if the addition of the gibberellic acid
(GA) substitutes for the light requirements for germination on some species. Alcorn
& Kurtz (1959) demonstrated that germination in either light or darkness increased
when the seeds of Carnegiea gigantea were imbibed in a 500 to 1000 p.p.m. of GA
solution. Later, McDonough (1964) proved that GA was effective in the promotion
of the dark germination of Carnegiea gigantea and Stenocereus thurberi seeds at 25 and
303C. Brencher et al. (1978) and Zimmer & BuKttner (1982) demonstrated that some
species showed a significant increase in dark germination with 500, 1000, and
2000 p.p.m. of GA. Under light conditions, GA at a high concentration may inhibit
germination for some species or increase the germination rate for others. Despite these
results, germination under light conditions gave the best germination rate for most
Rojas-AreHchiga et al. (1997) suggest that the light requirement could be associated
with the cactus life-form (i.e. barrel cacti or columnar cacti) due to a maternal effect
induced by temperature. Other studies support this hypothesis (Arias & Lemus, 1984;
Romero-Schmidt et al., 1992; Maiti et al., 1994; Nolasco et al., 1996; Vega-Villasante
et al., 1996; Nolasco et al., 1997).
Cacti have a wide range of responses to temperature, as do most species of tropical
habitats. For cactus seeds, the favourable temperature ranges between 17 and 343C,
with optimal values frequently at 253C (Nobel, 1988). Some of the first investigations
into the effects of temperature on cactus seed germination were made by Zimmer.
He applied 25 diurnal and nocturnal temperature combinations to Astrophytum myrios-
tigma with a thermoperiod of 12 h. From the results he concluded that at 103C there is
Table 2. Optimum temperature to get over 90% germination for some cactus
species. Species marked with an asterisk showed above 75% germination. References
cited more than once are assigned a letter the first time they occur, using this letter for
subsequent citations
Species Optimum
temperature Reference
Oreocereus trollii 15–203C Zimmer, 1967 A
Pachycereus hollianus* Rojas-AreHchiga et al., 1998 B
Rebutia marsoneri A
Rebutia minuscula Zimmer, 1969aC
Oreocereus celsianus 15–253CC
Cereus peruvianus A
Astrophytum myriostigma A
Oreocereus erectocylindrica Zimmer, 1970 D
Parodia maassii C
Echinopsis pasacana 15–303CC
Echinopsis huascha C
Mammillaria polythele C
Cleistocactus strausii A
Eulychnia castanea D
Mammillaria longimamma C
Ferocactus glaucescens C
Parodia leninghausii 15–353C Zimmer, 1971 E
Rebutia xanthocarpa v.salmonea &16–193C Fearn, 1974 F
Parodia chrysacanthion &17–253CF
Echinopsis pasacana &18–273CF
Cleistocactus hyalacanthus 20–253CC
Mammillaria zeilmanniana C
Gymnocalycium saglionis C
Parodia chrysacanthion*E
Thelocactus setispinus 20–303CC
Mammillaria muehlenpfordtii C
Coryphanta gladispina 20–353CC
Mammillaria fuauxiana C
Haageocereus multangularis 25–303CD
Echinocactus grusonii 25–353CC
Opuntia phaecantha var. discata Potter et al., 1984 G
Opuntia lindheimeri G
Espostoa lanata 203CC
Mammillaria durispina C
Cleistocactus strausii C
Stenocereus griseus 213C MartmHnez-HolgumHn, 1983
Ferocactus histrix 24$1·53C Del Castillo, 1986
Carnegiea gigantea 253C Alcorn & Kurtz, 1959
Astrophytum myriostigma Arredondo-GoHmez &
Camacho-MorfmHn, 1995
Neobuxbaumia tetetzo var. tetetzo Rojas-AreHchiga,
unpublished data
Ferocactus latispinus var. spiralis*B
Gymnocalycium mihanovichii E
Pereskia aculeata 333C Dau & Labouriau, 1974
no germination, between 15 and 253C there is a high germination rate, and above 303C
germination is reduced considerably (Zimmer, 1965). Later, the same author worked
with 25 cactus species in a 10}303C temperature range, and found that some showed
optimum germination at low temperatures, while other groups showed optimum
germination at high temperatures (Zimmer 1969a, 1973a)(see Table 2). The latter
group also demonstrated that as temperature increases the time to complete germination
is reduced, so at 303C, 11 cactus species completed their germination in 10 days
(Zimmer, 1970). In Table 2, optimum germination temperatures for some species are
Cota SaHnchez (1984), obtained high germination percentages at 403C for Ferocactus
latispinus var. spiralis; Fearn (1974) found that Frailea pumila germinated at a rate of
more than 50% at 39·53C; and Vega-Villasante et al. (1996) exposed seeds of Pachy-
cereus pecten-aboriginum to temperatures of 453C for prolonged times and obtained good
germination rates. From the studies of Zimmer (1967, 1968a, 1969a,b, 1970, 1971,
1973a,b), Fearn (1974, 1981), Kwack & Zimmer (1978) and Nobel (1988), some
observations can be made of the temperature effect on cacti seed germination:
(1) Temperature extremes do not favour germination, that is, below 123C and above
283C. A temperature of 20$23C gives a good germination rate in a wide range of
(2) Different species have different responses to temperature.
(3) Time to complete germination decreases as temperature increases (i.e. Astrophytum
myriostigma and Gymnocalycium mihanovichii).
(4) Response to temperature depends on seed age (i.e. the optimum temperature for
1 year old seeds is lower than for the 7 year old seeds of Parodia chrysacanthion).
(5) Aged seeds take longer to germinate than young ones (e.g. Maihuenia poeppigii).
(6) Alternating temperatures give better germination results than constant temper-
Despite this last statement, alternating temperature effects on cactus seed germi-
nation are not very clear, because in most investigationsonly constant temperatures have
been used, and any experiments that have incorporated alternating temperatures do not
show any significantly different effects on germination compared to constant
temperatures, or simply do not favour germination in Rebutia minuscula,Parodia
maassii,Oreocereus celsianus, and Mammillaria zeilmanniana (Zimmer, 1968b); Opuntia
sp., O.phaecantha var. discata, and O.lindheimeri (Potter et al., 1984); Neobuxbaumia
tetetzo,F.flavovirens, and Pachycereus hollianus, (GodmHnez-Alvarez & Valiente-Banuet,
1998; Rojas-AreHchiga et al., 1998); Ferocactus latispinus var. spiralis, and Echinocactus
platyacanthus fa. grandis (Rojas-AreHchiga et al., 1998). Contrary to these last results,
GodmHnez-Alvarez & Valiente-Banuet (1998) found significant differences between
constant and fluctuating temperatures for the two latter species, probably because the
constant temperature used was too low (173C).
With respect to the effect of temperature on the germination of seeds of dif-
ferent ages, Zimmer & Schultz (1975) mentioned that the species studied showed
different responses, mainly between low and high temperatures. For example, fresh
and 4 year old seeds of Neoporteria subgibbosa and Eulychnia castanea had a significantly
reduced germination percentage at 303C, and at 10 and 153C, respectively.
Innate dormancy produced by the presence of endogenous inhibitory compounds in
the testa has been detected in the seeds of Opuntia phaecantha var. discata and
O.lindheimeri (Pilcher, 1970), Stenocereus griseus (Williams & Arias, 1978), Melocactus
curvispinussp. caesius (Arias & Lemus, 1984), Opuntia sp. O.phaecantha var. discata,O.
lindheimeri (Pilcher, 1970; Potter et al., 1984), Stenocereus gummosus (LeoHn de la Luz
& DommHnguez-Cadena, 1991), and Opuntia joconostle (SaHnchez-Venegas, 1997), which
all need to be washed or imbibed for certain periods of time after sowing to get high
germination rates, meaning that in natural conditions the seeds need long periods of
moisture to wash out soluble germination inhibitors from their testa. In contrast,
GodmHnez-Alvarez & Valiente-Banuet (1998) suggested that imbibition periods did not
enhance seed germination in the species studied, while Alvarez-Aguirre & Montana
(1997) mentioned that an imbibition period of 36h prior to sowing gave good germina-
tion results in the species studied, but this is not necessary to get high germination
Several works have demonstrated the effect that fruit ingestion by animals can
have on the seed testa, allowing the seeds to germinate if placed in suitable conditions.
For example, seeds of Pereskia aculeata collected from the faeces of two species of
monkeys germinated much better than seeds sown from intact fruits (Pedroni &
SaHnchez, 1997). Also, seeds of Melocactus violaceus collected from lizard faeces ger-
minated well, while seeds collected directly from ripe fruits did not germinate (Cortes
Figueira et al., 1994). Alternatively, Silvius (1995) showed that the seeds of Stenocereus
griseus collected from ripe fruits and from the faeces of some birds failed to germinate
until they were provided with large amounts of water, suggesting an enforced dormancy
of survival value. Some studies have been carried out on the effect of chemical or
mechanical scarification, to simulate the requirements of seeds with impermeable seed
coats that are satisfied when passing through animal digestive tracts. A brief immersion
in a low concentration of acid solutionincreased germination in some species of Opuntia
(Potter et al., 1984), Echinocactus grusonii, E.platyacanthus fa. visnaga (De la Rosa-
Ibarra & GarcmHa, 1994), Ferocactus peninsulae (Romero-Schmidt et al., 1992), and
Pachycereus pringlei (Nolasco et al., 1996). The two latter species were also tested for
salinity effects, and from the results obtained it was demonstrated that as the
concentration of NaCl increased, the germination percentage decreased (Romero-
Schmidt et al., 1992; Nolasco et al., 1996). SaHnchez-Venegas (1997) reports that the
seeds of Opuntia joconostle need mechanical scarification to germinate, and the seeds of
Sclerocactus mariposensis germinate better with mechanical scarification (Moreno et al.,
1992). Seeds that are dispersed by water need a mechanical scarification that simulates
the natural abrasion of the soil when they are transported by water runoffs in their
habitat (Kemp, 1989). Some other species, such as Opuntia compressa and O.macro-
rhiza, need a stratification period to obtain high germination percentages (Baskin
& Baskin, 1977) and the germination of the seeds of Maihueniapoeppigiitakes place only
after a moist low temperature treatment (between 5 and 103C) is applied before sowing
(Zimmer, 1972; Zimmer, 1973c; Kwack & Zimmer, 1978). This might be related to the
removal of dormancy, or to the mobilization of nutrients from the perisperm, and this
has not been reported for seeds of tropical cactus species.
Embryo immaturity is another factor that can cause innate dormancy, so the seeds
need an after-ripening period to germinate, and this varies for each species. This has
been reported for Eriocereus bonplandii and Mammillaria zeilmanniana, whose seed
germination rate increased with age (Zimmer, 1967, 1969b). Young seeds of Ferocactus
latispinus var. spiralis germinated at under 50% and did not germinate at 20 and 353C,
whereas 45-month old seeds germinated at above 80% in a 20}353C range (Zimmer,
1980). Similar responses were found withseeds of Ferocactus wislizenii(Zimmer, 1980).
This has also been demonstrated in Opuntia rastrera seeds (Mandujano et al., 1997) and
for Echinocactus platyacanthus fa. grandis (pers. obs.). Also, May (1994) mentions that
seeds of Sclerocactus polyancistrus need to be ‘aged’ before germination can occur.
Soil seed bank
The seed bank composition in deserts mainly consists of annual plants, considering both
biomass and numbers of seeds (Brown etal., 1979;Inouye,1991)as they constitute95%
of the total number of seeds. Seeds in desert soils are distributed mostly near the surface
(Kemp, 1989).
There is no available information in the reviewed literature about the seed
bank of cacti species, but it is possible that many species showing any kind of dormancy
could form at least a short-term seed bank if they are able to avoid predation. Silvius
(1995) mentions that seeds of Stenocereus griseus from a semi-arid area of Venezuela
remained in the soil for 4 months before germinating. Mandujano et al. (1997) state that
as Opuntia rastrera seeds gave better germination results with ageing, implying the
presence of primary dormancy, they could also form a seed bank. Also, Potter et al.
(1984) found that seeds of Opuntia lindheimeri and Opuntia spp. which were stored for
some time showed a higher germination percentage than recently collected seeds,
meaning that they need an after-ripening period allowing them to stay in the soil for
some time.
Ecology and germination
Many studies that have assessed cactus seed germination have focused on their estab-
lishment and survival. Studies have been carried out on germination and recruitment
under shade provided by associate plants (nurse plants) or rocks (nurse rocks). These
elements provide extra moisture and protection against excessive radiation during the
early stages of growth, which have been shown to be a basic requirement for seedling
survival. This has been well studied in the Saguaro (Turner et al., 1966; Steenbergh
& Lowe, 1969; Brum, 1973; McAuliffe, 1984; Franco & Nobel, 1989). Other
studies on the effect of nurse plants on species of cactus seedlings (Nobel, 1988;
Valiente-Banuet et al., 1991a,b; Valiente-Banuet & Ezcurra, 1991), and on species in
which a shaded microsite or a specific microhabitat is needed for seedling survival
(Del Castillo, 1986; Nobel, 1989; LeoHn de la Luz & DommHnguez-Cadena, 1991; Silvius,
1995; Nolasco et al., 1996; Vega-Villasante et al., 1996; Nolasco et al., 1997). Alterna-
tively, Cortes Figueira et al. (1994) suggest that for Melocactus violaceus, shady habitats
may pose restrictions on survivorship because most plants were observed in open
habitats. Nurse plants are mostly perennial shrubs that, in some cases, can be replaced
by the grown columnar cacti (Valiente-Banuet et al., 1991a).
The role of inhibitory substances in the germination of cactus seeds seems to have
ecological implications as it provides information on the environmental conditions in
which they inhabit. Soluble inhibitors present in the testa or in the fruit flesh seem to
control seed germination by maintaining the dormant state until environmental condi-
tions for growth are suitable. Rainfall patterns may affect the germination of some
cactus seeds, as in the case of Melocactus curvispinus sp. caesius and Stenocereus griseus
seeds. The former species germinates immediately after a brief rinse, but the latter
requires prolonged leaching (Williams & Arias, 1978; Arias & Lemus, 1984), suggesting
that ample soil water may be necessary for varying periods of time in order to trigger
germination. Pilcher (1970) and Potter et al. (1984) demonstrated that Opuntia species
require different periods of time of soaking to promote germination.
Dubrovsky (1996, 1998) evaluated the promotive effect of discontinuous
hydration (hydropriming) on seed germination in Pachycereus pecten-aboriginum,
Ferocactus peninsulae, and Stenocereus gummosus, and found improved seedling survival
during a drought. It seems that cactus seeds have a memory of the internal changes
induced by a hydration treatment, which can allow them to resist periods of desiccation
and germinate faster after rehydration (Dubrovsky, 1996, 1998).
Bregman & Bouman (1983) studied patterns of seed germination in relation to
morphological and phylogenetical lines of development. Most cactus seeds germinate
within a week, but in the subfamily Opuntiodeae it may take a few months. These seeds
are inoperculate, but have a sclerified arillus, whichis necessary for passage through the
digestive tract of a bird. Thus, operculum formation could be correlated with ecological
conditions. Bregman & Graven (1997) demonstrated that in some genera of the tribu
Notocacteae, the ripe seed is surrounded by a protein-enriched layer that appears to be
hydrophilous, and improves germination under relatively dry conditions.
It has also been suggested that a rapid and high percentage of germination is
associated with a thin testa and with the presence of starch granules (Maiti et al., 1994).
Propagation and conservation
The Cactaceae family has always been subjected to intensive exploitation due to its great
diversity and value, mainly as ornamental plants, and as a result their populations have
been drastically affected due to overcollection and severe perturbation of their
habitat. Today, 35% of Mexican cactus speciesare endangered, many of which are listed
in the Appendix. Some of these species are Aztekium ritterii, Strombocactus disciformis,
Mammillaria pectinifera, Cephalocereus senilis, and Astrophytum asterias (CITES, 1995).
Propagation studies constitute an alternative to the conservation of this natural
resource, because the possibility of obtaining valuable plants through artificial methods
could decrease the demand for wild-collected material. Cactus propagationcan be done
in three ways: through in vitro tissue culture, by means of vegetative propagation, or by
seed germination.
Tissue culture propagation of Cactaceae has been extensively studied (Mauseth,
1977; Johnson & Emino, 1979a,b; Lazarte et al., 1982; Starling & Hutson, 1984;
Starling, 1985; Ault & Blackmon, 1987; RodrmHguez-Garay & Rubluo, 1992; Bonness
et al., 1993; Ortiz-Montiel & Vargas-Figueroa, 1995). This technique could be success-
fully used to propagate threatened or endangered species, with two main problems:
(1) genetic diversity is reduced, and (2) it is an expensive technique; and two ad-
vantages: (1) you can produce many plants from only one specimen and from various
parts of the plant, and (2) growth rate is increased.
Vegetative propagation can be achieved via shoots and cuttings, by division of
caespitose specimens, and by grafting. All of these are asexual methods without genetic
recombination, and have been used extensively for many species (Cattabriga, 1994;
Reyes-Santiago & Arias-Montes; 1995; Reyes-Santiago, 1997).
Propagation by seed is an important method because it allows genetic diversity of
populations and species to be maintained.Yet, little is known about cacti seed germina-
tion requirements, and about their viability and longevity. Studies on germination and
seedling establishment are important for understanding reproductive strategies, and for
artificial propagation and conservation. Most information concerning cactus seed
propagation has come from horticulturists, nursery workers, and amateurs (SchuKtz,
1973; Dunlap, 1976; Rivas, 1978, 1986, 1993; Morgan, 1983; Braunton, 1984; Kewen,
1984; Gerber, 1987; Fitz Maurice, 1989; Johnstone, 1990; Simerda, 1990; Moss, 1993;
Pilbeam, 1993; Anon., 1995a,b; Du Plooy, 1995; Grandet, 1995; Panarotto, 1995;
Anon., 1996a,b,c; Jolly & Lockert, 1996; Moakes, 1996; Quail, 1997; Bach, 1998;
Dimmitt, 1998; Kohlschreiber, 1998; Peters,1998; and Internet pages), and mainly only
suggests which type of substrate to usefor germination and growth, and some tips (types
of containers, sterilizing system, etc.) about seed sowing and transplanting.
There is an urgent need for more studies on propagation for the conservationist
effort to progress.
There are few works that evaluate the loss of viability over time, although data suggest
that most cactus seeds have an orthodox storage behaviour (Roberts, 1972). The
optimal conditions for long-term storage are not known for most cactus seeds (Alcorn
& Martin, 1974). Myers (1939 in Fearn, 1977), carried out some pioneering studies on
cactus seed viability when he demonstrated that seeds of some Opuntia species remain
viable for years. Later, Zimmer (1970) mentioned that Haageocereus turbidus and
Oreocereus erectocylindrica seeds sown 4 years after they were harvested had good
germination rates, while Melocactus peruvianus,Echinopsis tegeleriana, and
Samaipaticereus corroanus lost their viability quickly. Fearn (1977) mentioned that seeds
of Ferocactus spp. and Echinocereus spp. show a slow initial loss of viability for the first
4 years, and that Neoporteria shows a relatively slow decline in viability in contrast with
Rebutia. A 7-year seed sample of Coryphanta odorata gave 100% germination, but when
tested later their viability was lost rapidly (Fearn, 1977).
Ten-year old seeds of Ferocactus herrerae and F.emoryi still showed 80 and 90%
germination, respectively (Fearn, 1977). In contrast, seeds of two speciesof Brasilicactus
did not remain viable for longer than 4 years, and their viability declined rapidly after
1 year (Fearn, 1977). Alcorn & Martin (1974) proved that the germination rate of 10
year old seeds of Carnegiea gigantea of various crops stored dry at laboratory temper-
atures ranged from 4 to 51%. In some genera, such as Frailea (Jolly & Lockert, 1996)
and some members of the genus Gymnocalycium (Fearn, 1981), viability is lost very
quickly, while in other genera like Echinocereus, Ferocactus, Neoporteria,Eulychnia, and
Haageocereus, viability is maintained for longer periods (Fearn, 1981). Zimmer
& Schultz (1975) demonstrated that 3-year old seeds of Oreocereus erectocylindrica gave
over 80% germination within a 15}303C range, and 4-year old seeds of Eulychnia
castaneagave over 70% germination within the same range. Del Castillo (1986) reported
that a high viability is maintained for 2 years in Ferocactus histrix seeds, for 5 years or
more in Melocactus seeds, for 4 or 5 years in Matucana seeds, and for approximately
9 years in Parodia seeds (Anon., 1997a,b,c). Unfortunately, the conditions in which
these seeds were preserved to keep their viability are not mentioned. Six-year old seeds
ofFerocactus latispinus var. spiralisand Echinocactus platyacanthusfa. grandis kept in glass
containers at ambient temperature (20$23C) still gave above 50% germination (pers.
Longevity of seeds under natural or controlled storage conditions depend on many
factors, including type of seed, stage of maturity, viability and moisture content when
stored, temperature, degree of infection by fungi and bacteria, etc. (Roberts, 1972; Stein
et al., 1974; Fearn, 1981).
An effective way to conserve endangered species by ex situ methods is by creating
a germplasm or seed bank which could maintain viable seeds if placed under carefully
controlled conditions (the seeds are placed inside a sealed chamber at a low temperature
(!183C) and a low relative humidity (c. 5%). In this way, viability could be maintained
for a long period of time (sometimes up to 100 years).
The information that is available on cactus seeds is already of a considerable volume,
however, valuable information on important ecological and conservation aspects is
lacking. Almost nothing is known about the presence and dynamics of cactus seeds in
the desert soil seed bank. Some information exists on seed longevity in nature or under
controlled storage conditions. Aspects of seed recollection, handling, storage, and
certification needs to be extensively studiedin order to develop successful techniques of
seed management and ex situ conservation.
We greatly appreciate assistance from Ulises GuzmaHn who helped us with taxonomic aspects and
gave us some valuable references. We also thank Rosalba Silva and Teresa Zamora for gathering
bibliographic references, and Pedro PeHrez Granados for helping us with the translation of German
papers. This work was partially supported by CONACYT (The Mexican Council of Science and
Technology) research grant G0011-N9607.
Antesberger, H. (1991). Melocactus laui, a new species from Aruba, Netherland Antillas. Cactus
and Succulent Journal (U.S.), 63: 241–243.
Alcorn, S.M. & Kurtz, E.B. (1959). Some factors affecting the germination of seed of the
saguaro cactus (Carnegiea gigantea). American Journal of Botany,46: 526–529.
Alcorn, S.M. & Martin, S.C. (1974). Cereus giganteus Engelm. In: Seeds of Woody Plants in the
UnitedStates, pp. 313–314. Washington, D.C.: Forest Service U. S. Departmentof Agriculture.
883 pp.
Alvarez-Aguirre, M.G. & Montan8a, C. (1997). GerminacioHn y supervivencia de cinco especies de
cactaHceas del Valle de TehuacaHn: Implicaciones para su conservacioHn. Acta Bota&nica Mexicana,
40: 43–58.
Anonymous (1995a). Culture de: Copiapoa.Cactus-Aventures International,25: 16.
Anonymous (1995b). Culture de: Epithelantha.Cactus-Aventures International,28: 16.
Anonymous (1996a). Culture de: Neoporteria.Cactus-Aventures International,29: 17.
Anonymous (1996b). Culture de Lobivia.Cactus-Aventures International,30: 16.
Anonymous (1996c). Growing: Frailea.Cactus-Aventures International,31: 17.
Anonymous (1997a). Growing: Melocactus.Cactus-Aventures International,33: 16.
Anonymous (1997b). Growing: Matucana.Cactus-Aventures International,34: 16.
Anonymous (1997c). Growing: Parodia.Cactus-Aventures International,35: 17.
Arias, I. & Lemus, L. (1984). Interaction of light, temperature and plant hormones in the
germination of seeds of Melocactus caesius Went (Cactaceae). Acta Cientn&fica Venezolana,35:
Arredondo-GoHmez, A. & Camacho-MorfmHn, F. (1995). GerminacioHndeAstrophytum myriostigma
en relacioHn con la procedencia de las semillas y la temperatura de incubacioHn. Cacta&ceas
y Suculentas Mexicanas,40: 34–38.
Ault, J.R. & Blackmon, W.J. (1987). In vitro propagation of Ferocactus acanthodes (Cactaceae).
HortScience,22: 126–127.
Bach, D. (1998). Soils for succulents—A nurseryman’s view. Cactus and Succulent Journal (U.S.),
Barthlott, W. & Porembski, S. (1996). Ecology and morphology of Blossfeldia liliputana
(Cactaceae): a poikilohydric and almost astomate succulent. Botanica Acta,109: 161–166.
Barthlott, W., Porembski, S., Kluge, M., Hopke, J. & Schmidt, L. (1997). Selenicereus wittii
(Cactaceae): an epiphyte adapted to Amazonian IgapoHinundationforests. Plant Systematics and
Evolution,206: 175–185.
Baskin, J.M. & Baskin, C.C. (1977). Seed and seedling ecology of OpuntiacompressainTennessee
cedar glades. Journal of the Tennessee Academy of Science,52: 118–122.
Boeken, B. & Gutterman, Y. (1990).The effect of temperature on seed germination in three
common bulbous plants of different habitats in the Central Negev Desert of Israel. Journal
of Arid Environments,18: 175–184.
Bonness, M.S., PareH, P.W. & Mabry, T.J. (1993). Novel callus and suspension cultures of the
‘old man’ cactus (Cephalocereus senilis). Cactus and Succulent Journal (U.S.),65: 144–147.
Braunton, E. (1984). Best soil for cactus. Cactus and Succulent Journal (U.S.),57:8.
Bravo-Hollis, H. (1967). Una revisioHn del geHnero Lophophora.Cacta&ceas y Suculentas Mexicanas,
12: 8–17.
Bravo-Hollis, H. & SaHnchez-Mejorada, H. (1991). Las Cacta&ceas de Me&xico, Vol. 2. MeHxico, D. F.:
UNAM. 404 pp.
Bravo-Hollis, H. & Scheinvar, L. (1995). El Interesante Mundo de las Cacta&ceas.MeHxico, D. F.:
CONACyT-FCE. 233 pp.
Bravo-Hollis, H., Scheinvar, L. & SaHnchez-Mejorada, H. (1970). Neobuxbaumia polylopha
Backeberg. Cacta&ceas y Suculentas Mexicanas,15: 75–82.
Bravo-Hollis, H., Scheinvar, L. & SaHnchez-Mejorada, H. (1971). Estudio comparativo del geHnero
Neobuxbaumia (Backeberg) II. N.macrocephala (Web.) Daws. Cacta&ceas y Suculentas
Mexicanas,16: 3–14.
Bravo-Hollis, H., Scheinvar, L. & SaHnchez-Mejorada, H. (1973). Estudio comparativo del geHnero
Neobuxbaumia IV. N.multiareolata (Dawson) Bravo, Scheinvar and SaHnchez-Mejorada.
Cacta&ceas y Suculentas Mexicanas,18: 59–67.
Bregman, R. (1988). Forms of seed dispersal in Cactaceae. Acta Botanica Neerlandesa,37:
Bregman, R. & Bouman, F. (1983). Seed germination in Cactaceae. Botanical Journal of the
Linnean Society,86: 357–374.
Bregman, R. & Graven, P. (1997). Subcuticular secretion by cactus seeds improves germination
by means of rapid uptake and distribution of water. Annals of Botany,80: 525–531.
Bregman, R., Hensel, H.B., Laney, P., Meerstadt, A., Melis, P., Pullen, A.B. & VanTillborg, A.
(1987). Het geslacht Matucana. Br. & R. (VII). Succulenta,66: 56–59.
Brencher, W., Stange, L. & Zimmer, K. (1978). Ersatz des Lichts der Keimung von Kakteensa-
men durch GibberellinsaKure. Gartenbauwissenschaft,43: 91–94.
Brown, J.H., Reichman, O.J. & Davidson, D.W. (1979). Granivory indesert ecosystems. Annual
Review of Ecology and Systematics,10: 210–227.
Brum, G.D. (1973). Ecology of the saguaro (Carnegiea gigantea): phenology and establishment in
marginal populations. MadronJo,22: 195–204.
Buchenau, F.G. (1964). Mammillaria nana Backeberg. Cacta&ceas y Suculentas Mexicanas,10:
Buchenau, F.G. (1966). Mammillaria varieaculeata (Buchenau). Cacta&ceas y Suculentas
Mexicanas,11: 79–81.
Buchenau, F.G. (1969). Mammillaria meyranii var. michoacana.Cacta&ceas y Suculentas Mexicanas,
14: 75–76.
Buxbaum, F. (1951). Morphology of cacti. Pasadena, CA: Abbey Garden Press. 223 pp.
Cattabriga, A. (1994). Propagation of threatened Mexican cacti. In: Anderson, E.F., Arias
Montes, S. & Taylor, N.P. (Eds), Threatened Cacti of Mexico, pp. 117–133. Kew: Royal Botanic
Gardens. 135 pp.
CITES (1995). Appendices I, II and III to the Convention on International Trade in Endangered
Species of Wild Fauna and Flora. Washington, D.C.: U.S. Department of the Interior. U.S. Fish
and Wildlife Service. 23 pp.
Cortes Figueira, J.E., Vasconcellos-Nieto, J., Alice GarcmHa, M.& Teixeira de Souza, A.L. (1994).
Saurocory in Melocactus violaceus (Cactaceae). Biotropica,26: 295–301.
Cota SaHnchez, J.H. (1984). Influencia de la luz, temperatura y sustancias qumHmicas sobre la
germinacioHn de semillas de Ferocactus latispinus (Haw.) Br. and Rose (Cactaceae). Bachelor
Thesis, ENCB-Instituto PoliteHcnico Nacional, MeHxico, D.F. 82 pp.
Dau, L. & Labouriau, L.G. (1974). Temperature control of seed germination in Pereskia aculeata
Mill. Anais Academia Brasileira de Ciencias,46: 311–322.
Del Castillo, R.F. (1986). Semillas, germinacioHn y establecimiento de Ferocactus histrix.Cacta&ceas
y Suculentas Mexicanas,31: 5–11.
Del Castillo, R.F. (1988). FenologmHa y remocioHn de semillas en Ferocactus histrix.Cacta&ceas
y Suculentas Mexicanas,33: 5–14.
De la Rosa-Ibarra, M. & GarcmHa, H. (1994). EstimulacioHn de la germinacioHn de cinco especies de
cactaHceas consideradas en peligro de extincioHn. Phyton,56: 147–150.
Dimmitt, M.A. (1998). Potting media for succulents in hot, dry climates. Cactus and Succulent
Journal (U.S.),70: 14–17.
Dubrovsky, J.G. (1996). Seed hydration memory in Sonoran Desert cacti and its ecological
implication. American Journal of Botany,83: 624632.
Dubrovsky, J.G. (1998). Discontinuous hydration as a facultative requirement for seed germina-
tion in two cactus species of the Sonoran Desert. Journal of the Torrey Botanical Society,125:
Dunlap, M.S. (1976). Germination of cactus seed. Cactusand Succulent Journal (U.S.),48: 83–86.
Du Plooy, F. (1995). Growing succulents and cacti from seed. Aloe,32: 6–7.
Elizondo-Elizondo, J., ValdeHs Reyna, J., Arias Montes, S. & Hatch, S.L. (1994). MicromorfologmHa
de las semillas de algunas especies de la tribu Cacteae. Cacta&ceas y Suculentas Mexicanas,39:
Fearn, B. (1974). An investigation into the effect of temperature on the seed germination of
nine species of cacti using thermal gradient bars. Cactus and Succulent Journal (U.S.),46:
Fearn, B. (1977). An investigation into the effect of age on the germination potential of seeds
of 600 species of cacti, together with a note on the viability of Lithopsseeds. Excelsa,7:103–108.
Fearn, B. (1981). Seed germination: the modern approach. Cactus and Succulent Journal (G.B.),
43: 13–16.
Fittkau, H.W. (1968). Mammillaria erectacantha Foerster. Cacta&caeas y Suculentas Mexicanas,13:
Fitz Maurice, W.A. (1989). A system of seed propagation for cacti. Cactus and Succulent Journal
(U.S.),61: 14–16.
Franco, A.C. & Nobel, P.S. (1989). Effect of nurse plants on the microhabitat and growth of
cacti. Journal of Ecology,77: 870–886.
Gerber, H. (1987). Trials and tribulations when growing cacti from seed. Cactus and Succulent
Journal (U.S.),59: 104–105.
Gibson, A.C. & Nobel, P.S. (1986). The Cactus Primer. Cambridge, MA: Harvard University
Press. 286 pp.
GodmHnez-Alvarez, H. & Valiente-Banuet, A. (1998). Germination and early seedling growth of
Tehuacan Valley cacti species: the role of soils and seed ingestion by dispersers on seedling
growth. Journal of Arid Environments,39: 21–31.
GonzaHlez-Espinosa, M. & Quintana-Ascencio, P.F. (1986). Seed predation and dispersal in
a dominant desert plant: Opuntia, ants, birds and mammals. In: Estrada, A. & Fleming, T.H.
(Eds), Frugivores and Seed Dispersal, pp. 273–284. Dordrecht: Dr. W. Junk Publishers. 392 pp.
Grandet, G. (1995). Quelques observations sur des semis comparatifs de CacteHes. Cactus-
Aventures International,26: 7–10.
Grenot, C. & Serrano, V. (1981). Ecological organization of small mammal communities at the
BolsoHn de MapimmH(MeHxico). In: Barbault, R. & Halffter, G. (Eds), Ecology of the
Chihuahuan Desert, pp. 89–100. MeHxico, D.F.: Instituto de EcologmHa, a.c. 167 pp.
Gutterman, Y. (1994). Strategies of seed dispersal and germination in plants inhabiting deserts.
The Botanical Review,60: 373–425.
Howe, H.F. & Smallwood, J. (1982). Ecology of seed dispersal. Annual Review of Ecology and
Systematics,13: 201–228.
Hunt, D. (comp.) (1992). CITES Cactaceae Checklist. Kew: Royal Botanic Gardens. 190 pp.
Inouye, R.S. (1991). Population biology of desert annual plants. In: Polis, G.A. (Ed.), The Ecology
of Desert Communities, pp. 27–54. Tucson: The University of Arizona Press. 456 pp.
Janzen, D.H. (1971). Seed predation by animals. Annual Review of Ecology and Systematics,2:
Janzen, D.H. (1986). Chihuahuan desert nopaleras: defaunated big mammal vegetation. Annual
Review of Ecology and Systematics,17: 595–636.
Jolly, J. & Lockert, X. (1996). Sowing: germination and substrates. Cactus-AventuresInternational,
31: 22–25.
Johnson, J.L. & Emino, E.R. (1979a). In vitro propagation of Mammillaria elongata.HortScience,
14: 605–606.
Johnson, J.L. & Emino, E.R. (1979b). Tissue culture propagation in the Cactaceae. Cactus and
Succulent Journal (U.S.),51: 275–277.
Johnstone, R. (1990). Propagation by seed. Cactus and Succulent Journal (Australian),17: 92–95.
Jordano, P. (1992). Fruits and Frugivory. In: Fenner, M. (Ed.), The Ecology of Regeneration in
Plant Communities, pp. 105–156. U.K.: C.A.B. International. 373 pp.
Kewen, P. (1984). Growing cacti. Cactus and Succulent Journal (U.S.),57: 11–13.
Kemp, P.R. (1989). Seed banks and vegetation processes in deserts. In: Leck, M.A.,
Parker, V.Th. & Simpson, R.L. (Eds), Ecology of Soil Seed Banks, pp. 257–281. San Diego:
Academic Press, Inc. 462 pp.
Kiesling, R. (1968). Pterocactus.Cacta&ceas y Suculentas Mexicanas,13: 36–38.
Kigel, J. (1995). Seed germination in arid and semi-arid regions. In: Kigel, J. & Galili, G. (Eds),
Seed Development and Germination, pp. 645–699. New York: Marcel Dekker, Inc. 853 pp.
Kimnach, M. (1984). Nopalxochia horrichii Kimn., a new cactus from Costa Rica. Cactus and
Succulent Journal (U.S.),56: 4–8.
Kohlschreiber, D. (1998). Soil mixes for epiphytic cacti. Cactus and Succulent Journal (U.S.),70:
Kwack, B.H. & Zimmer, K. (1978). Germination of aged seeds of Maihuenia poeppigii.Garten-
bauwissenschaft,43: 188–191.
Lazarte, J.E., Gaiser, M.S. & Brown, O.R. (1982). In vitro propagation of Epiphyllum chrysocar-
dium.HortScience,17: 84.
LeoHn de la Luz, J.L. & DommHnguez-Cadena,R. (1991). EvaluacioHn de la reproduccioHn por semilla
de la pitaya agria (Stenocereus gummosus) en Baja California Sur, MeHxico. Acta Bota&nica
Mexicana,14: 75–87.
Lindsay, G.E. (1967). Los Echinocereus de Baja California. Cacta&ceas y Suculentas Mexicanas,12:
LodeH, J. (1995). Le genre: Epithelantha Weber ex Britton & Rose. Cactus-Aventures International,
28: 12–15.
LoHpez-GoHmez, R. & SaHnchez-Romero, P. (1989). GerminacioHn de dos variedades de pitaya
Stenocereus griseus (Haworth) Buxbaum. Cacta&ceas y Suculentas Mexicanas,34: 35–40.
May, R.W. (1994). The ecology of Sclerocactus polyancistrus (Cactaceae) in California and
Nevada. Desert Plants,11: 6–22.
McAuliffe, J.R. (1984). Sahuaro-nurse tree associations in the Sonoran Desert: competitive
effects of sahuaros. Oecologia,64: 319–321.
McDonough, W. (1964). Germination responses of Carnegiea gigantea and Lemairocereusthurberi.
Ecology,45: 155–159.
Maiti, R.K., HernaHndez-Pin8ero, J.L. & ValdeHz-MarroqumHn, M. (1994). Seed ultrastructure and
germination of some species of Cactaceae. Phyton,55: 97–105.
Mandujano, M.C., Golubov, J. & Montan8a, C. (1997). Dormancy and endozoochorous dispersal
of Opuntia rastrera in the Southern Chihuahuan Desert. Journal of Arid Environments,36:
MartmHnez-HolgumHn, E. (1983). GerminacioHn de semillas de Stenocereus griseus (Haw) Buxbaum
(Pitayo de mayo). Cacta&ceas y Suculentas Mexicanas,28: 51–57.
Mauseth, J.D. (1977). Cactus tissue culture: a potential method for propagation. Cactus and
Succulent Journal (U.S.),49: 80–81.
Moakes, P. (1996). Successful seed raising. British Cactus and Succulent Journal,14: 24–25.
Moran, R. (1965). RevisioHn del geHnero Bergerocactus.Cacta&ceas y Suculentas Mexicanas,10:
Moreno, N., LoHpez, J.J. & Arce, L. (1992). Aspectos sobre las semillas y su germinacioHnde
Echinomastus mariposensis Hester. Cacta&ceas y Suculentas Mexicanas,37: 21–27.
Morgan, L. (1983). Cacti from seed. Cactus and Succulent Information Exchange,16: 102–104.
Morton, S.R. (1985). Granivory in arid regions: Comparison of Australia with North and South
America. Ecology,66: 1859–1866.
Moss, B. (1993). Ariocarpus from seed. British Cactus and Succulent Journal,2: 2–3.
Murdoch, A.J. & Ellis, R.H. (1992). Longevity, viability and dormancy. In: Fenner, M. (Ed.), The
Ecology of Regeneration in Plant Communities, pp. 193}229. U.K.: C.A.B. International. 373 pp.
Nobel, P.S. (1988). Environmental Biology of Agaves and Cacti. New York, NY: Cambridge
University Press. 270 pp.
Nobel, P.S. (1989). Temperature, water availability, and nutrient levels at various soil depths
— consequences for shallow-rooted desert succulents, including nurse plant effects. Ameri-
can Journal of Botany,76: 1486–1492.
Nolasco, H., Vega-Villasante, F. & DmHaz-Rondero, A. (1997). Seed germination of Stenocereus
thurberi (Cactaceae) under different solar irradiation levels. Journal of Arid Environments,
36: 123–132.
Nolasco, H., Vega-Villasante, F., Romero-Schmidt, H.L. & DmHaz-Rondero, A. (1996). The
effects of salinity, acidity, light and temperature on the germination of seeds of cardoHn
(Pachycereus pringlei) (S.Wats) Britton & Rose (Cactaceae). Journal of Arid Environments,33:
Ortiz-Montiel, J.G. & Vargas-Figueroa, M. (1995). PropagacioHnin vitro de Heliocereus elegantis-
simus (Britton y Rose) var. elegantissimus (Cactaceae). Cacta&ceas y Suculentas Mexicanas,40:
Otero, F. & MeyraHn, J. (1966). Echinocereus pulchellus.Cacta&ceas y Suculentas Mexicanas,11:
Panarotto, P. (1995). I Turbinicarpus che crescono sul gesso: note di coltivazione. Piante Grasse,
15: 46–48.
Pedroni, F. & SaHnchez, M. (1997). Dispersao de sementes de Pereskia aculeata Muller
(Cactaceae) num fragmento florestal no sudeste do Brasil. Revista Brasileira de Biologia,57:
Peters, R.R. (1998). Growing plants in low-organicsoil-based mixes. CactusandSucculentJournal
(U.S.),70: 10–11.
Pilbeam, J. (1993). Seed-raising. Cactus and Succulent Journal (G.B.),11: 55–56.
Pilcher, B.L. (1970). Germination of seeds of four species of Opuntia.Cactus and Succulent
Journal,42: 281–282.
Potter, R.L., Petersen, J.L. & Ueckert, D.N. (1984). Germination responses of Opuntia spp. to
temperature, scarification and other seed treatments. Weed Science,32: 106–110.
Quail, D.E. (1997). Slow-growing cacti from seed. British Cactus and Succulent Journal,15:
Reyes-Santiago, J. (1997). Cultivo y propagacioHn como plantas de ornato. In: Valles SeptieHn, C.
(Ed.), Suculentas Mexicanas/Cacta&ceas, pp. 69–77. MeHxico, D.F.: CVS Publicaciones. 143 pp.
Reyes-Santiago, J. & Arias-Montes, S. (1995). CactaHceas de MeHxico: conservacioHn y produccioHn.
Revista Chapingo/Horticultura,3: 85–92.
Rivas, M. (1978). Una forma sencilla de sembrar semillas de cactos. Cacta&ceas y Suculentas
Mexicanas,23: 94–96.
Rivas, M. (1986). Observaciones sobre el cultivo de cactos. Cacta&ceas y Suculentas Mexicanas,31:103.
Rivas, M. (1993). Notas sobre el cultivo de cactaHceas por semilla. Cacta&ceas y Suculentas
Mexicanas,38: 93–94.
Roberts, E.H. (1972). Dormancy: a factor affecting seed survival in the soil. In: Roberts, E.H.
(Ed.), Viability of Seeds, pp. 321–359. London: Chapman & Hall. 448 pp.
RodrmHguez-Garay, B. & Rubluo, A. (1992). In vitro morphogenetic responses of the endangered
cactus Aztekium ritteri (Boedeker). Cactus and Succulent Journal (U.S.),64: 116–119.
Rojas-AreHchiga, M., Orozco-Segovia, A. & VaHzquez-Yanes, C. (1997). Effect of light on the
germination of seven species of cacti from the ZapotitlaHn Valley in Puebla, Mexico. Journal of
Arid Environments,36: 571–578.
Rojas-AreHchiga, M., VaHzquez-Yanes, C. & Orozco-Segovia, A. (1998). Seed response to tempe-
rature of two life forms of Mexican cacti species: an ecophysiological interpretation. Plant
Ecology,135: 207–214.
Romero-Schmidt, H.L., Vega-Villasante, F., Nolasco, H. & Montan8o, C. (1992). The effect
of darkness, freezing, acidity and salinity on seed germination of Ferocactus peninsulae
(Cactaceae). Journal of Arid Environments,23: 389–395.
SaHnchez-Mejorada, H. (1972). Stenocereus chrysocarpus, una nueva especie de MichoacaHn.
Cacta&ceas y Suculentas Mexicanas,17: 95–98.
SaHnchez-Mejorada, H. (1973). Nuevas cactaHceas de la Nueva Galicia. Cacta&ceas y Suculentas
Mexicanas,18: 12–18.
SaHnchez-Venegas, G. (1997). GerminacioHn, viabilidad y caractermHsticas distintivas de la semilla
de Opuntia joconostle Weber, forma cuaresmero. Cacta&ceas y Suculentas Mexicanas,42:
SchuKtz, B. (1973). Einige Erfahrungen bei der erdelosen Kakteenkultur. Kakteen und andere
Sukkulenten,24: 62–3.
Silvius, K.M. (1995). Avian consumers of cardoHn fruits (Stenocereus griseus: Cactaceae) on
Margarita Island, Venezuela. Biotropica,27: 96–105.
Simerda, B. (1990). Effective ways of propagating endangered cacti. British Cactus and
Succulent Journal,8: 9–12.
Starling, R. (1985). In vitro propagation of Leuchtenbergia principis.Cactus and Succulent Journal
(U.S),57: 114–115.
Starling, R.J. & Hutson, R. (1984). Sterile culture of succulent plants. British Cactus and Succulent
Journal,2: 69–70.
Steenbergh, W.F. & Lowe, Ch.L. (1969). Critical factors during the first years of life of the
saguaro (Cereus giganteus) at Saguaro National Monument, Arizona. Ecology,50: 825–834.
Stein,W.F.,Slabaugh,P.E.&Plummer, A.P.(1974).Harvesting,processing,andstorage of fruits
and seeds. In: Schopmeyer, C.S. (Ed.), Seeds of Woody Plants in the United States, pp. 98–126.
Washington, D.C.: Forest Service U.S. Department of Agriculture. 883 pp.
Tran, V.N. & Cavanagh, A.K. (1984). Structural aspects of dormancy. In: Murray, D.E. (Ed.),
Seed Physiology, Vol. 2, pp. 1–44. Australia: Academic Press. 295 pp.
Turner, R.M., Alcorn, S.M., Olin, G. & Booth, J.A.(1966). The influence of shade, soil and water
on saguaro seedling establishment. Botanical Gazette,127: 95–102.
Valiente-Banuet, A. & Arizmendi, M. del C. (1997). InteraccioHn entre cactaHceas y animales:
polinizacioHn, dispersioHn de semillas y nuevos individuos. In: Valles SeptieHn, C. (Ed.), Suculentas
Mexicanas/Cacta&ceas, pp. 61–67. MeHxico, D.F.: CVS Publicaciones. 143 pp.
Valiente-Banuet, A. & Ezcurra, E. (1991). Shade as a cause of the association between the cactus
Neobuxbaumia tetetzo and the nurse plant Mimosa luisana in the Tehuacan Valley, Mexico.
Journal of Ecology,79: 961–971.
Valiente-Banuet, A., Bolongaro-Crevenna, A., Briones, O., Ezcurra, E., Rosas, M., NuHn8ez, H.,
Barnard, G. & VaHzquez, E. (1991a). Spatial relationships between cacti and nurse shrubs in
a semi-arid environment in central Mexico. Journal of Vegetation Science,2: 15–20.
Valiente-Banuet, A., Vite, F. & Zavala-Hurtado, J. A. (1991b). Interaction between the cactus
Neobuxbaumia tetetzo and the nurse shrub Mimosa luisana.Journal of Vegetation Science,2:
Vargas-Mendoza, M.C. & GonzaHlez-Espinosa, M. (1992). Habitat heterogeneity and seed disper-
sal of Opuntia streptacantha (Cactaceae) in nopaleras of Central Mexico. The Southwestern
Naturalist,37: 379–385.
Vega-Villasante, F., Nolasco, H., Montan8o, C., Romero-Schmidt, H. & Vega-Villasante, E.
(1996). Efecto de la temperatura, acidez, iluminacioHn, salinidad, irradiacioHn solar y humedad
sobre la germinacioHn de semillas de Pachycereus pecten-aboriginum ‘‘cardoHn barboHn’’
(Cactaceae). Cacta&ceas y Suculentas Mexicanas,41: 51–61.
Weiss, J., Scheinvar, L. & Mizrahi, Y. (1995). Selenicereus megalanthus (the yellow pitaya),
a climbing cactus from Colombia and Peru. Cactus and Succulent Journal (U.S.),67: 280–283.
Wendelken, P.W. & Martin, R.F. (1988). Avian consumption of the fruit of the cacti Stenocereus
eichlamii and Pilosocereus maxonii in Guatemala. The American Midland Naturalist,119:
Williams, P.M. & Arias, I. (1978). Physio-ecological studies of plant species from the arid and
semiarid regions of Venezuela. I. The role of endogenous inhibitors in the germination of the
seeds of Cereus griseus (Haw.) Br. and R: (Cactaceae). Acta CientnHfica Venezolana,29: 93–97.
Zimmer, K. (1965). Untersuchungen uKber den Einflubder Temperatur auf die Keimung von
Kakteen-Kultur-Saatgut.II. UGber die Keimung von Astrophytum myriostigma Lem. Die Garten-
bausissenschaft,30: 381–387.
Zimmer, K. (1966). UGber Samengewichte und Samenproduktion einiger Kakteenarten. Kakteen
und andere Sukkulenten,17: 153–154.
Zimmer, K. (1967). Temperatur und Keimung bei verschiedenen Kakteen. Kakteen und andere
Sukkulenten,18: 31–33.
Zimmer, K. (1968a). Untersuchungen uKber den Einflubder Temperatur auf die Keimung von
Kakteen-Saatgut. VI. UGber die Keimung einiger mexikanischer Arten. Gartenbauwissenschaft,
33: 167–175.
Zimmer, K. (1968b). Untersuchungen uKber den Einflubder Temperatur auf die Keimung von
Kakteen-Saatgut. VII. UGber die Keimung einiger argentinischer Arten. Gartenbauwissenschaft,
33: 335–344.
Zimmer, K. (1969a). UGber die Keimung von Kakteensamen I. Quellung und Keimung bei
konstanten Temperaturen. Kakteen und andere Sukkulenten,20: 105–107.
Zimmer, K. (1969b). UGber die Keimung von Kakteensamen II. Wechseltemperaturen-nachreife-
wirkung niedriger temperatur. Kakteen und andere Sukkulenten,20: 125–128.
Zimmer, K. (1969c). UGber die Keimung von Kakteensamen III. Die bedeutung des Lichtes.
Kakteen und andere Sukkulenten,20: 144–147.
Zimmer, K. (1969d). UGber die Keimung von Kakteensamen (Schlub). Bedeutung des Lichts und
Zusammenfassung. Kakteen und andere Sukkulenten,20: 178–180.
Zimmer, K. (1970). UGber die Keimung einiger bolivianischer und peruanischer Kakteenarten.
Kakteen und andere Sukkulenten,21: 177–178.
Zimmer, K. (1971). Ein weiterer Beitrag zur Keimung von Kakteensamen. Kakteen und andere
Sukkulenten,22: 153–155.
Zimmer, K. (1972). Untersuchungen uKber den Einflubder Temperatur auf die Keimung von
Kakteen-Saatgut. VIII. Zum KaKltebedurfnis bei der Keimung von Maihuenia poeppiggi (Otto)
Web. Gartenbauwissenschaft,37: 109–121.
Zimmer, K. (1973a). UGber die Keimung einiger chilenischer Kakteen-Arten. Kakteen und andere
Sukkulenten,24: 58–59.
Zimmer, K. (1973b). Zur Keimung von Selenicereus grandiflorus (LineeH) Britton & Rose. Kakteen
und andere Sukkulenten,24: 227–228.
Zimmer, K. (1973c). Zur Keimung von Maihuenia poeppiggi.Kakteen und andere Sukkulenten,24:
Zimmer, K. (1980). Einflubder Temperatur auf die Keimung von Kakteensaatgut. X. Keimung
einiger Ferocactus-Arten. Gartenbauwissenschaft,45: 121–123.
Zimmer, K. & BuKttner, P. (1982). Ersatz des LichtbeduKrfnisses bei der Keimung von Kakteensa-
men durch GibberellinsaKure. Gartenbauwissenschaft,47: 97–101.
Zimmer, K. & Schultz, A. (1975). Untersuchungen uKber den Einflubder Temperatur auf die
Keimung von Kakteensaatgut. IX. Zur VeraKnderung des Temperaturbereichs fuKr die Keimung
in AbhaKngigkeit vom Samenalter. Gartenbauwissenschaft,3: 120–125.
Authority (ies) of species mentioned in the text. Species names according to
Hunt (1992)
Species Authority (ies)
Ariocarpus kotschoubeyanus (Lem.) K. Schum.
Astrophytum asterias (Zucc.) Lem.
Astrophytum capricorne (Dietrich) Britton & Rose
Astrophytum myriostigma Lem.
Aztekium ritterii Boed. ex A. Berger
Blossfeldia liliputana Werderm.
Carnegiea gigantea (Engelm.) Britton & Rose
Cephalocereus columna-trajani (Karw.) K. Schum.
Cephalocereus senilis (Haw.) Pfeiff.
Cereus jamacaru DC.
Cereus peruvianus (L.) Mill.
Cleistocactus hyalacanthus (Backeb.) Backeb.
Cleistocactus strausii (Heese) Backeb.
Coryphanta gladispina (Boed.) A. Berger
Coryphanta odorata Boed.
Disocactus kimnachii G. D. Rowley
Echinocactus grusonii Hildm.
Echinocactus platyacanthus fa. grandis (Rose) Bravo
Echinocactus platyacanthus fa. visnaga (Hook.) Bravo
Echinocereus grandis Britton & Rose
Echinocereus pulchellus (Mart.) K. Schum.
Echinopsis pasacana (F.A.C. Weber) K. Friedrich &
G. D. Rowley
Echinopsis huascha (F.A.C. Weber) K. Friedrich &
G. D. Rowley
Echinopsis tegeleriana (Backeb.) D. R. Hunt
Epiphyllum anguliger (Lem.) G. Don
Epiphyllum phyllanthus (L.) Haw.
Eriocereus bonplandii (Parment.) Riccob.
Espostoa lanata (HBK) Britton & Rose
Eulychnia castanea Phil.
Ferocactus emoryi (Engelm.) Backeb.
Ferocactus flavovirens (Scheidw.) Britton & Rose
Ferocactus glaucescens (DC.) Britton & Rose
Ferocactus haematacanthus (Salm-Dyck) Bravo
Ferocactus herrerae J.G. Ortega
Ferocactus histrix (DC.) G. Linds.
Ferocactus latispinus var. spiralis (Karw. ex Pfeiff.) N. P. Taylor
Ferocactus peninsulae (Engelm. ex Weber) Britton & Rose
Ferocactus wislizeni G. Linds.
Frailea pumila (Lem.) Britton & Rose
Gymnocalycium mihanovichii (Fric & GuKrke) Britton & Rose
Gymnocalycium saglionis (Cels) Britton & Rose
Haageocereus multangularis (Willd.) Cullm.
Leuchtenbergia principis Hook.
Lophophora williamsii (Lem. ex Salm-Dyck) J. M. Coult.
Maihuenia poeppiggi (Otto) F. A. C. Weber
Mammillaria durispina Boed.
Mammillaria erectacantha C. F. Foerst.
Mammillaria fuauxiana Backeb.
Mammillaria longimamma DC.
Mammillaria magnimamma Haw.
Mammillaria meyranii var. michoacana Buchenau
Mammillaria muehelnpfordti C. F. Foerst.
Mammillaria nana Backeb. ex Mottram
Mammillaria pectinifera (Stein) F. A. C. Weber
Mammillaria polythele Mart.
Mammillaria varieaculeata Buchenau
Mammillaria zeilmanniana Boed.
Matucana formosa F. Ritter
Melocactus curvispinus ssp. caesius (H. L. Wendl.) N. P. Taylor
Melocactus macracanthos (Salm-Dyck) Link & Otto
Melocactus peruvianus Vaupel
Melocactus violaceus Pfeiff.
Myrtillocactus geometrizans (Mart.) Console
Neobuxbaumia macrocephala (F. A. C. Weber) E. Y. Dawson
Neobuxbaumia polylopha (DC.) Backeb.
Neobuxbaumia tetetzo (Coulter) Backeb.
Neoporteria subgibbosa (Haw.) Britton & Rose
Opuntia compressa (Salisb.) T. Macbr.
Opuntia joconostle F. A. C. Weber
Opuntia lindheimeri Engelm.
Opuntia macrorhiza Engelm.
Opuntia phaecantha var discata (Griffiths) L. D. Benson & Walk.
Opuntia rastrera F. A. C. Weber
Oreocereus celsianus (Lem.) Riccob.
Oreocereus erectocylindrica Rauh & Backeb.
Oreocereus trollii (Kupper) Backeb.
Pachycereus hollianus (F. A. C. Weber) Buxb.
Pachycereus pecten-aboriginum (Engelm.) Britton & Rose
Pachycereus pringlei (S. Watson) Britton & Rose
Pachycereus schottii (Engelm.) D. R. Hunt
Pachycereus weberi (J. M. Coult.) Backeb.
Parodia chrysacanthion (K. Schum.) Backeb.
Parodia leninghausii F. Haage
Parodia maassii (Heese) A. Berger
Parodia scopa (Spreng.) N. P. Taylor
Pelecyphora strobiliformis (Werderm.) Fric & Schelle
Peniocereus castellae SaHnchez-Mej.
Pereskia aculeata (Plum) Mill.
Pilosocereus chrysacanthus (F. A. C. Weber) Britton & Rose
Pilosocereus leucocephalus (Poselg.) Vyles & G. D. Rowley
Rebutia marsoneri Werderm.
Rebutia minuscula K. Schum.
Rebutia xanthocarpa v. salmonea Fric ex Backeb.
Samaipaticereus corroanus Card.
Sclerocactus mariposensis (Hester) N. P. Taylor
Sclerocactus polyancistrus (Engelm. & Bigelow) Britton & Rose
Selenicereus megalanthus (K. Schum.) Moran
Selenicereus wittii (K. Schum.) G. D. Rowley
Stenocereus chrysocarpus SaHnchez-Mej.
Stenocereus eichlamii (Britton & Rose) Bravo & A. S. Arias
Stenocereus griseus (Haw.) Buxb.
Stenocereus gummosus (Engelm.) A. C. Gibson & K. E. Horak
Stenocereus thurberi (Engelm.) Buxb.
Strombocactus disciformis (DC.) Britton & Rose
Thelocactus setispinus (Engelm.) E. F. Anderson
Turbinicarpus lophophoroides (Werderm.) Buxb. & Backeb.
... This has been well studied in desert annuals (Gutterman, 2000), but particularly for the family Cactaceae, there are few works that assess dormancy. In a review by Rojas-Aréchiga and V azquez-Yanes (2000) it is mentioned that among cacti two kinds of dormancy occurred: innate and enforced, following Harper (1957). According to Baskin and Baskin's (2014) seed dormancy classification, a physiological dormancy has been demonstrated for some Opuntia species (Mandujano et al., 2005;Orozco-Segovia et al., 2007;Reinhardt et al., 1999) and for Sclerocactus polyancistrus (May, 1994), and a physical dormancy has been demonstrated for Corryocactus melanotrichus (Larrea-Alc azar & L opez, 2008); however, most cacti seeds are nondormant (Baskin & Baskin, 2014). ...
... Seed longevity in Cactaceae has been poorly studied despite its importance for ex situ conservation research, as all species of this family are included in Appendix III and many of them in Appendix I of the Convention on International Trade in Endangered Species (CITES) (Hunt, 1999). Some studies on loss of viability through time under different conditions have shown variable results (Rojas-Aréchiga & V azquez-Yanes, 2000). Recently, Nascimento and Meiado (2017) demonstrated that germinability of seeds of Disocactus bahiensis after being stored inside a cold chamber for 24 months was high (70%). ...
... Primary dormancy, together with other morphological and physiological seed traits, such as small size and rounded shape and a light requirement for germination (positive photoblastism), have been associated with their potential to form a soil seed bank and their persistence in it (Thompson et al., 1993), and this is not an exception within cacti species (Bowers, 2000;Rojas-Aréchiga & Batis, 2001). Unfortunately, research on soil seed banks for cacti species is still scarce (Holland & Molina-Freaner, 2013;Rojas-Aréchiga & V azquez-Yanes, 2000), and seed bank formation in natural environments is difficult to assess. Current studies have demonstrated either no seed bank formation (Holland & Molina-Freaner, 2013) or its existence ( Alvarez-Espino et al., 2014;Bowers, 2005;Cano-Salgado et al., 2012;Lindow-L opez et al., 2018). ...
Full-text available
Knowledge on seed dormancy is crucial for the understanding of plant population dynamics, as it controls seed germination and seed bank formation. Dormant seeds have high potential to establish in soil seed banks, but such information within Cactaceae is scarce, although it is essential for conservation programs. The aim of this study was to determine if seeds of Ferocactus peninsulae showed any kind of dormancy and to test their germination capacity after storage. This was assessed with 15 seed sowing experiments done over 4 years with seeds stored under room conditions (20 ± 2°C). We demonstrated the existence of physiological dormancy in F. peninsulae seeds that is broken with an after‐ripening period. Germination was low during the first 3 months of storage (d = 0.206) but increased after 10 months of storage (d = 0.654), and seeds maintained their viability at 48 months (d = 0.707). Also, their speed of germination increased with storage time. Ferocactus peninsulae seeds are positively photoblastic, and the requirement for light for germination persisted over all experiments. The results provide crucial information for propagation and conservation research and may allow us to infer that F. peninsulae seeds are able to form a persistent soil seed bank, as they maintained their viability after dormancy is released. Dry after‐ripening breaks dormancy in Ferocactus peninsulae seeds. Ferocactus peninsulae seeds need at least 3 months under room storage to break dormancy. Seeds of F. peninsulae can maintain their viability for several years.
... El tamaño pequeño puede evitar la depredación por mamíferos (pequeños o medianos) e insectos (Rojas-Aréchiga & Batis, 2001). Esta característica también puede facilitar la dispersión por viento y agua (Rojas-Aréchiga & Vázquez-Yanes, 2000). El viento puede ser un vector importante en la dispersión debido al peso y en época de lluvias las corrientes de agua pueden trasladar las semillas lejos de las plantas madre. ...
Full-text available
Mammillaria parkinsonii is a mexican endemic species protected by NOM-059-SEMARNAT-2010 in the special category (Pr), thus the present study is a contribution to the knowledge of the morpho-anatomy, histochemistry and germination of Mammillaria parkinsonii seeds from a population in Tolimán, Querétaro which is characterized by a high diversity of cacti. The anatomical and histochemical descriptions were made on 20 randomly selected seeds, which were processed using standard protocols for scanning electron and light microscopy studies and to obtain permanent and semi-permanent slides. For germination study 210 seeds were sown in Petri dishes on agar, using six replicas; weight and size were obtained using 50 seeds randomly selected. The results show that the seed coat is formed by a non-striated cuticle with pectics and proteinaceous substances. The exotesta occupies 6.8% of the total volume of the seed, it has lignified-suberized and protein-containing anticline walls, forming U-type undulations with protrusions. The endotegmen is suberized and an embryonic membrane lacking ergastic contents surrounds the embryo. The endosperm is confined to a dorsal ridge. The embryo is globose, with two cotyledonous protrusions, and the hypocotyl assumes the reserve function with a "globoid crystals/protein bodies" association. The micropylar hilum region is small and oval, filled with parenchyma. Germination started on day 5th and ended on day 29th, with a mean germination time on day 8.9. The final germination percentage was 74.29%. The age, seed size, crystal and protein reserve as well as germination percentage of the M. parkinsonii suggest a potential orthodox or intermediate behavior to form soil seed banks in a desert habitat.
... It is a very common adaptive plant strategy in unpredictable and harsh environments, such as arid and semiarid ones . The Cactaceae is a plant family having most of its species in unpredictable and harsh environments such as deserts, and several cactus species have dormant seeds (Rojas-Aréchiga and Vázquez-Yanes, 2000;Flores et al., 2005Flores et al., , 2006Flores et al., , 2008Barrios et al., 2020), especially in the genus Opuntia (Mandujano et al., 2005(Mandujano et al., , 2007Orozco-Segovia et al., 2007;Ochoa-Alfaro et al., 2008;González-Cortés et al., 2018). At the end of the XIX century, Ganong (1898) found that some O. echinocarpa seeds did not germinate for a year, and he suggested that it is a phenomenon that perhaps has an ecological meaning, but he did not mention the seed dormancy term. ...
Full-text available
Opuntia species from arid and semiarid environments have dormant seeds. The objective of this study was to evaluate how the soil influences seed germination of the cactus O. dejecta. We hypothesized that O. dejecta seeds in the basaltic rocky soil will show lower seed germination than seeds in a site with alluvial soil from the coastal plain. An experiment of partial reciprocal transplant was performed, placing seeds from basaltic soil (San Ignacio population) in pots containing alluvial soil from Puente Nacional population and in slabs from basaltic soil, and placing them on two greenhouses, in San Ignacio population and Puente Nacional population. We found that seed germination (%) was lower in San Ignacio site (with basaltic soil) than in Puente Nacional site (with alluvial soil), and that alluvial soil showed higher seed germination than basaltic soil, as well as higher seed germination in Puente Nacional than in San Ignacio. The results suggest that seed dormancy for the population situated in the rocky condition is a trait locally adapted, and the basaltic soil can be considered as a selection factor.
... El tamaño pequeño puede evitar la depredación por mamíferos (pequeños o medianos) e insectos (Rojas-Aréchiga & Batis, 2001). Esta característica también puede facilitar la dispersión por viento y agua (Rojas-Aréchiga & Vázquez-Yanes, 2000). El viento puede ser un vector importante en la dispersión debido al peso y en época de lluvias las corrientes de agua pueden trasladar las semillas lejos de las plantas madre. ...
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Mammillaria parkinsonii is a mexican endemic species protected by NOM-059-SEMARNAT-2010 in the special category (Pr), thus the present study is a contribution to the knowledge of the morpho-anatomy, histochemistry and germination of Mammillaria parkinsonii seeds from a population in Tolimán, Querétaro which is characterized by a high diversity of cacti. The anatomical and histochemical descriptions were made on 20 randomly selected seeds, which were processed using standard protocols for scanning electron and light microscopy studies and to obtain permanent and semi-permanent slides. For germination study 210 seeds were sown in Petri dishes on agar, using six replicas; weight and size were obtained using 50 seeds randomly selected. The results show that the seed coat is formed by a non-striated cuticle with pectics and proteinaceous substances. The exotesta occupies 6.8% of the total volume of the seed, it has lignified-suberized and protein-containing anticline walls, forming U-type undulations with protrusions. The endotegmen is suberized and an embryonic membrane lacking ergastic contents surrounds the embryo. The endosperm is confined to a dorsal ridge. The embryo is globose, with two cotyledonous protrusions, and the hypocotyl assumes the reserve function with a "globoid crystals/protein bodies" association. The micropylar hilum region is small and oval, filled with parenchyma. Germination started on day 5th and ended on day 29th, with a mean germination time on day 8.9. The final germination percentage was 74.29%. The age, seed size, crystal and protein reserve as well as germination percentage of the M. parkinsonii suggest a potential orthodox or intermediate behavior to form soil seed banks in a desert habitat.
... Schoenus nigricans is a major canary due to its strict association with water availability and to the time seeds take to sense the environment seedlings would grow in if they germinate Baskin 1971, 2014;Rojas-Aréchiga and Vázquez-Yanes 2000;Martínez-Villegas et al. 2012). Since it forms big clumps, changes in the riparian habitat they occupy are usually soon visible through the landscape modification along the water bodies. ...
Of the seemingly innumerable jewels of Cuatro Ciénegas, the Rio Churince system was singular, a microcosm of the basin’s ecological, evolutionary, and hydrogeological diversity. Recent drying of Churince extinguished this small, relatively unaltered desert spring ecosystem and with it an unmatched educational asset for all who could traverse its mere 2.5 km length. In Cuatro Ciénegas and beyond, there is urgent need to conserve desert spring systems that remain intact and to reignite the vibrancy of ones that can be restored. This chapter documents in photos the demise of Churince.
The Cactaceae family has a strong predisposition for hybridization, which is a mechanism suggested to be relevant to its evolution. However, a few studies evidence the consequences of hybridization in members of this family. A population of Cephalocereus columna-trajani × C. tetetzo cohabits with both parental species in the Tehuacán-Cuicatlán Reserve, Mexico. A comparative analysis was conducted between these three taxa with three principal aims: a) determine, through in situ experiments, the mating system of a hybrid taxon; b) compare seed morphology; and c) evaluate seed germination under controlled conditions. Our results show that the mating success was higher between hybrid individuals than between hybrids and C. tetetzo members. The hybrid seed showed the typical morphology of the tribe Pachycereeae but was statistically smaller than the seed of C. columna-trajani. Under white and red light the total germination percentage of hybrid seeds was higher than that of both parental species, which was interpreted as hybrid vigor. In summary, the successful mating between individuals preceding the formation of viable seeds with better germination capacity than both parental species is evidence of the reproductive success of hybrids, disproving their sterility and challenging the view that the hybrid taxon represents an “evolutionary endpoint”.
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Los Jardines Botánicos han jugado un papel importante en la protección de plantas, especialmente especies que se encuentran en peligro de extinción. La mayoría de estas especies viven en hábitats particulares; en su mayoría fragmentados y con baja densidad poblacional. Uno de los objetivos para la conservación de cactáceas, es la información relevante sobre la germinación, biología de sus poblaciones, y las principales condiciones ecológicas para la sobrevivencia, así como la creación de bancos de semillas. En el estado de Oaxaca, México, hay especies de cactáceas del género Mammillaria que han restringido su distribución y por lo tanto son altamente vulnerables. Entre las investigaciones que se llevan a cabo en la colección científica del Jardín Botánico Regional “Casiano Conzatti” destacan los realizados en el Valle de Tehuacán-Cuicatlán (VTC), donde se tienen reportadas 71 especies de cactáceas, de las cuales 18 son endémicas. Se han realizado estudios poblacionales de distintas especies de este Valle, sin embargo, en este trabajo se presenta información obtenida de estudios realizados con Mammillaria dixanthocentron, M. supertexta y M. huitzilopochtli, especies endémicas de Oaxaca y catalogadas en la NOM 059-2010. Se realizaron estudios para describir la germinación de las especies mencionadas. Los experimentos de realizaron bajo condiciones controladas con el fin de establecer las características apropiadas para la germinación de cada especie, así como el potencial de generar un banco de semillas del suelo. Se llevaron a cabo estudios sobre la biología de sus poblaciones. Ninguna de estas especies había sido estudiada desde un punto de vista demográfico, a pesar de que esta información sugiere importantes fluctuaciones espacio-temporales en sus poblaciones que sólo se pueden entender mediante análisis demográficos. Los resultados obtenidos indican que existe diferencia tanto en la demografía como en la dinamica poblacional de las especies en estudio. Para contribuir a la conservación de estas especies, se realizan proyectos de investigación, y se participa en la enseñanza sobre propagación de cactáceas, y ecología de poblaciones y comunidades. Palabras clave: conservación, dinamica poblacional, jardín botánico, Mammillaria
The seed anatomy of eight species of ornamental cacti from the Chihuahuan desert was described, germplasm in the process of long-term conservation in the Germplasm Bank of the National Center for Genetic Resources of INIFAP; different qualitative, quantitative and pseudo-qualitative aspects were tested for the characterization of the seed, likewise it was supported with an X-ray equipment. The shape of the seed was varied as well as the size, probably associated with the ecological conditions in which species are distributed; X-rays provide important information on the development and condition of the embryo, however, it is not indicative of the viability of the seed.
Miners used to take canaries with them when descending underground to work. They acted as an early warning for them, and many miners’ lives were saved. There are also environmental canaries, under and above ground, but unfortunately, they are usually ignored. In the Cuatro Ciénegas Basin (CCB), the mismanagement and overexploitation of water have led to a generalized disturbance of the hydrological systems that characterize the basin and, in recent years, to the desiccation of the Churince system. Several canaries have been tweeting, sighing and agonizing for a long time, but most stakeholders have not listened at all. Therefore, we analysed three riparian plant species that herein act as plant canaries of the overexploitation of the aquifer in the CCB wetland: Samolus ebracteatus var. coahuilensis, Flaveria chlorifolia and Schoenus nigricans. The distribution and behaviour of these species are related to their hydrophyte character, and their presence indicates water availability. As a consequence of the anthropogenic disturbance, the Churince system suffered as the original riparian habitat disappeared. As a result of the dry-out, many of the plant species colonized newly formed habitats such as a gypsum-based, highly saline, dry plain. Riparian species tried to follow the water in sinkholes formed due to abnormal sublevel water flow and the dry beds of the terminal lake and the river. Their behaviour accounts for the continual loss of ground and underground water. Canaries tweeted on time, but decision-makers did not listen opportunely. The loss of water in CCB is still ongoing, and now it is the time for everyone to respond before all this precious water is gone forever.
Soil conditions were evaluated over the rooting depths for Agave deserti and Ferocactus acanthodes from the northwestern Sonoran Desert. These succulents have mean root depths of only 10 cm when adults and an even shallower distribution when seedlings, which often occur in association with the nurse plant Hilaria rigida, which also has shallow roots. Maximum soil temperatures in the 2 cm beneath bare ground were predicted to exceed 65 C, which is lethal to the roots of A. deserti and F. acanthodes, whereas H. rigida reduced the maximum surface temperatures by over 10 C, providing a microhabitat suitable for seedling establishment. Water Availability was defined as the soil-to-plant drop in water potential, for periods when the plants could take up water, integrated over time. Below 4 cm under bare ground, simulated Water Availability increased slightly with depth (to 35 cm) for a wet year, was fairly constant for an average year, and decreased for a dry year, indicating that the shallow rooting habit is more advantageous in drier years. Water uptake by H. rigida substantially reduced Water Availability for seedlings associated with this nurse plant. On the other hand, a 66–90% higher soil nitrogen level occurred under H. rigida, possibly representing its harvesting of this macronutrient from a wide ground area. Phosphorus was slightly less abundant in the soil under H. rigida compared with under bare ground, the potassium level was substantially higher, and the sodium level was substantially lower. All four elements varied greatly with depth, N and K decreasing and P and Na increasing. Based on the known growth responses of A. deserti and F. acanthodes to these four elements, growth was predicted to be higher for plants in soil from the shallower layers, most of the differences being due to nitrogen.
Scarification in sulfuric acid consistently increased germination of Opuntia edwardsii sp. nov, O. discata Griffiths, and O. lindheimeri Engelm. ♯ OPULI seeds over that of untreated seeds. Optimum constant temperatures for germination were generally 25 to 35 C and germination was not enhanced by alternating temperatures. There was a trend for increased germination following leaching in water for 12 h which suggested the presence of chemical germination inhibitors. Seeds passed through the digestive tracts of cattle exhibited average germination percentages that were 1.5 times greater than seeds removed from ripe fruits.
This chapter discusses different structural aspects of coat-imposed dormancy. Dormancy in seeds provides a means by which germination is delayed until favorable conditions for growth and establishment in the field are met. The water-impermeable coat protects the embryo from adverse storage and environmental conditions and actively promotes seed longevity. However, it is still unknown whether impermeability is due to mechanical processes (for example, shrinkage and closer packing of the cells in the seed coat as the seed matures) or to chemical effects (for example, impregnation of the cell walls with hydrophobic substances). A combination of both may exist and the operative mechanisms may differ among various species. The strength and other mechanical properties of the coat need to be measured for many more species to determine if mechanical constraints to embryo expansion or radicle extension prevent germination. From a small number of studies, it is evident that the lens in mimosoid seeds and an area near the chalazal discontinuity in malvaceous seeds are underlain by a single or double layer of thin-walled, sub-palisade cells, whose walls break readily under stress. In papilionoid seeds, the hilum functions as a hygroscopically activated, one-way valve that permits drying out of the seed during maturation. The mechanism(s) of natural breakdown of the seed coat and the effects of passage through birds and animals are two virtually unexplored areas. The available reports of the effects of artificial treatments on the seed coat are sometimes vague and frequently contradictory. There is a need for co-operative research between workers in various fields to understand these problems and to minimize unnecessary proliferation of speculation and hypotheses.