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Systematics and Phytogeography |
Rancho Santa Ana Botanic Garden, Claremont, California 91711 USA
Received for publication December 15, 2003. Accepted for publication July 2, 2004.
| ABSTRACT |
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Key Words: archaethnobotany Bayesian analysis crop origins Opuntia Opuntia ficus-indica
| INTRODUCTION |
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Early European botanists (often referring to Pliny or Theophrastus) called this cactus Ficus indica (Donkin, 1977
), although some found this to be an unsuitable name, as the plant did not resemble the Indian fig (possibly Ficus benghalensis L.) already known (Anderson, 2001
). Linnaeus published Cactus opuntia and C. ficus-indica in Species Plantarum. Miller combined these into Opuntia ficus-indica in 1768. In the recorded history of the Old World, O. ficus-indica was certainly known at the beginning of the 16th century (Donkin, 1977
; Casas and Barbera, 2002
), and it is believed that this species accompanied Columbus in his first return to Lisbon in 1493 (Russell and Felker, 1987
; Anderson, 2001
), placing O. ficus-indica in the Caribbean by at least the late 1400s, although whether it is native there is unknown. The plants are also recorded in cultivation in Tlaxcala, Mexico, in 1519 (Diaz del Castillo, 1632 in Idell, 1957
). Opuntia ficus-indica fruits and shoots were also reportedly consumed by the Maya of southeastern Mexico (Coe, 1994
). There is also some evidence for the use of O. ficus-indica by the Nazca of Peru, placing these plants in South America at a very early date (Sejuro, 1990
). Other workers maintain that this taxon was unknown in pre-Columbian South America (Towle, 1961
; Baker, 2002
). The succulent, ever-fresh cladodes were certainly a novelty to late 15th century and later Europeans (Donkin, 1977
) and were widely included in ships' stores as insurance against scurvy (Kiesling, 1998
). This practice is thought to have contributed greatly to the present naturalized range of Opuntia ficus-indica throughout arid and semiarid habitats of the world (Anderson, 2001
; Casas and Barbera, 2002
; Sáenz-Hernandez et al., 2002
). This widespread propagation (intended and unintended) throughout the Mediterranean obscured the geographic origins of this species; many early European botanists regarded this cactus to be native (Donkin, 1977
; Barbera et al., 1992
), as reflected in Cactus opuntia L. (i.e., spiny plant from near Opus, Greece; Anderson, 2001
). This Mediterranean naturalization may now be conceived as complete, as the Israelis of the mid-20th century often adopted the (believed-indigenous) sabras as a symbol of their struggle (and humanity) in adverse desert conditions (Uris, 1959
).
Although most recent authors concede that this species is not native to the Old World, the geographic and evolutionary origins of Opuntia ficus-indica remain clouded in obscurity (Britton and Rose, 1919
; Bravo-Hollis, 1978
; Benson, 1982
; Anderson, 2001
). This mystery arises partly from the widespread cultivated distribution of the plants, both within and away from the New World (Anderson, 2001
), but also from the phenotypic variability and artificial selection of the plants (Benson, 1982
). The specific epithet (from P. Miller, 1768) may reflect the 16th-century European opinion that the plants are native to the West Indies (Barbera et al., 1992
; Casas and Barbera, 2002
). The name "tuna" is Caribbean in origin (Bravo-Hollis, 1978
; Kiesling, 1998
). Other early work proposed that O. ficus-indica was a spineless cultivar derived from O. megacantha, a species of central Mexico (Griffiths, 1914
). A recent, important paper explored this relationship with amplified fragment length polymorphism (AFLP) data and reported that O. ficus-indica had the closest affinity with O. megacantha (Labra et al., 2003
), corroborating the Griffiths (1914)
hypothesis. Other authors, however, consider O. megacantha also to be a cultivated taxon (Benson, 1982
) or a name applied to multiple ruderal reversions to spininess from escaped, cultivated O. ficus-indica (Kiesling, 1998
) and treat O. megacantha as a later synonym (Benson, 1982
; Kiesling, 1998
). Uphof (1968)
and Zeven and Zhukovsky (1975
, p. 164) place the origin of this cactus within "Mexico" or the "Central American and Mexican centre" of domestication, but without reference to a specific region. Some recent cytological work proposes that octoploid, cultivated O. ficus-indica is derived from a diploid Mexican (but unnamed) progenitor (Kiesling, 1998
). Benson (1982)
states that O. ficus-indica is native to mild tropical upland habitat (not deserts), but does not state where. Given the conflicting, sometimes confusing, and often unclear hypotheses as to the origins of this cactus crop, a wide molecular phylogenetic analysis has potential to elucidate the biogeography of this species. The current study addresses the biogeographic and evolutionary origins of O. ficus-indica through modern molecular means. To this end, I have gathered and analyzed DNA sequence data from a number of accessions of O. ficus-indica representing plantings throughout the world.
| METHODS AND MATERIALS |
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| RESULTS AND DISCUSSION |
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Natural hybridization
Polyploidy and hybridization are well documented in the opuntioid cacti (Benson and Walkington, 1965
; Grant and Grant, 1971b
, 1979
; Baker and Pinkava, 1987
, 1999
; Griffith, 2001a
, b
, 2003
; Pinkava, 2002
). Opuntia ficus-indica is octaploid (Pinkava, 2002
), possibly through ancient allopolyploidy. Natural biological events may have led to the pattern of relationships in O. ficus-indica recovered for the ITS data set.
Human manipulation
If O. ficus-indica is of hybrid origin, it is conceivable that the hybridization was human facilitated. This may have occurred indirectly, by bringing allopatric ancestors into cultivated sympatry. One intriguing possibility is that hybridization of ancestral O. ficus-indica stock was instead performed directly by Mesoamerican agriculturalists. There is evidence for the intensive selective breeding of fruit crops by pre-Columbian peoples (Bergh, 1995
); it is possible that Mesoamerican knowledge of plant reproduction included pollination technology. Descriptions and depictions of Mesopotamian date palm (Phoenix spp.) cultivation dating to 1750 BC or earlier are interpreted as the earliest documentation of human use of artificial pollination in fruit crop selection (Paley, 1976
; Roth, 2000
; Janick, 2005
). Although there is no direct evidence of pollination for crop manipulation of O. ficus-indica in contemporary Mesoamerica, the early knowledge of its biological function is suggested by the depiction of hummingbird pollination in the Primeros Memoriales of Sahagún (ca. 1558; Fig. 7).
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Lineage sorting
Another possible explanation for this pattern of genetic relationships is lineage sorting (Pamilo and Nei, 1988
) of multiple ITS copies in a widespread ancestral population of Opuntia that gave rise to O. ficus-indica and its close relatives.
These four hypotheses should be tested through other independent molecular investigations. Other DNA sequence-based phylogenetic estimations may be of use, but the low level of variation in the ITS region for the genus Opuntia may indicate that a genome-wide approach may yield more useful variation. Reticulate evolution in Opuntia has been investigated via random amplified polymorphic DNA (RAPD) data (Mayer et al., 2000
; Griffith, 2003
), and some modern clones of O. ficus-indica have been characterized via this method (Wang et al., 1998
) as being either for fruit or vegetable production. Other data that may elucidate these relationships might include AFLP (as in Labra et al., 2003
) and microsatellite data. This population genetic approach may elucidate potential differences in heterozygosity between O. ficus-indica populations from different localities within central Mexico and abroad, which could further pinpoint the location of the wild progenitors of this species. Ongoing investigations will involve these data types interpreted in a historical and archaeological context and may illuminate the ancient selection of these plants by the oldest inhabitants of Mexico.
| FOOTNOTES |
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2 E-mail: michael.patrick.griffith{at}cgu.edu ![]()
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