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CACTACEAE) ROOT GORELICK Department of Biology Carleton University 1125 Raven Road Ottawa, Ontario K1S 5B6 Canada Email: [email protected]

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CACTACEAE) ROOT GORELICK Department of Biology Carleton University 1125 Raven Road Ottawa, Ontario K1S 5B6 Canada Email: Root.Gorelick@Carleton.Ca Haseltonia 22: 18–21. 2016 18 SUBDERMAL WOODY THORNS WITH SECONDARY GROWTH MAY SUPPORT SHOOTS OF SENITA, LOPHOCEREUS SCHOTTII (CACTACEAE) ROOT GORELICK Department of Biology Carleton University 1125 Raven Road Ottawa, Ontario K1S 5B6 Canada email: [email protected] Abstract: Lophocereus (Pachycereus) schottii has lignified thorns with secondary growth connecting areoles to the vascular cylinder, which may help support their massive highly parenchymatous shoots. INTRODUCTION biomechanical case study that could shed light on Lophocereus schottii (Engelm.) Britton & Rose the ability to produce tall and stiff stems capable of (synonym Pachycereus schottii (Engelm.) D.R. Hunt), growing to substantial heights, yet without benefit senita, is a columnar cactus with a candelabra of the large amounts of wood found in many other growth habit. It has fairly massive shoots that are dicot arborescent species.” The same authors noted mostly composed of cortex plus wide pith, with a that, “cacti also provide numerous opportunities relatively small vascular cylinder, i.e. without much to identify the extent to which anatomical features lignification. Almost all new shoots arise as branches correlate with tissue stiffness and strength” (Niklas, from the base of existing shoots and each grows fair- Molina-Freaner, and Tinoco-Ojanguren 1999: 774). ly vertically. Therefore much of the weight on shoots Here I hypothesize one possible unexplored mecha- is supported by the ground and does not have to be nism by which L. schottii, may stiffen its shoots. supported by other shoots. Yet still there is need for Each Lophocereus schottii shoot starts off growing stiffness of shoots, especially when buffeted by wind. slowly. Until one meter tall, on average, shoots grow With regards to Lophocereus schottii, Molina- 6 cm per year, with growth rates decreasing as shoots Freanar et al. (1998: 1083) noted that “the stems approach one meter in height (Parker 1988). How- of columnar cactus species present an interesting ever, once shoots exceed one meter in height, aver- Figure 1. An unusual Lophocereus schottii with a pseudocephalium on each shoot. Pachycormus discolor in left background. Near Cataviña, Baja California. HASELTONIA VOL. 22. 2016 19 Figure 2. Decomposing Lophocereus schottii skeleton showing Figure 3. Lophocereus schottii skeleton with thin thorns and disarticulation of woody rods. Note unusual aerial branch at disarticulating woody rods. far left. This and remaining figures are of plant #95 at Organ Pipe Cactus National Monument. and Nobel, 1986). However, L. schottii differs from all other Pachycereeae skeletons that I have seen, in- age growth rates become 15 times faster than that cluding C. gigantea, S. thurberi, and Neobuxbaumia of juvenile shoots, increasing to a growth rate of 88 mezcalensis (Bravo) Backeb.; clavija), in that only L. cm per year (Parker 1988). This transition to much schottii have highly lignified protuberances, essen- faster growth occurs at roughly the same time shoots tially thorns, extending from the vascular cylinder reach sexual maturity, producing an apical pseudo- of the massive shoot out to the areoles (Figures 3–5). cephalium with many long flexible spines per areole Lignified thorns connecting the vascular cylinder to (Martorell, Vega, and Ezcurra 2006). Most shoots the areoles may provide extra support for the mas- remain shorter than one meter tall, with individual sive highly parenchymous shoots. These completely plants only having one or a few taller shoots with subdermal thorns are relatively narrow near the api- pseudocephalia (Martorell, Vega, and Ezcurra 2006), cal meristem of the massive shoot and relatively nar- although in a few specimens most or all shoots ap- row near the areole. Thorns become broader near pear to produce pseudocephalia (Figure 1). the base of the massive shoot and become broader Skeletons of Lophocereus schottii resemble those centripetally, i.e. become broader closer to the vas- of closely related cacti with overlapping ranges, such cular cylinder. It thus appears that Lophocereus schot- as saguaro (Carnegiea gigantea Britton & Rose) and tii has secondary growth of the long vascular traces organ pipe (Stenocereus thurberi (Engelm.) Buxb.; connecting areoles to the deeply embedded vascular pitaya dulce) in having a vascular cylinder that de- cylinder, with a hollow center to each thorn where composes into a ring of vertically oriented parallel the vascular traces originally were (Figure 5). There discrete lignified rods, with one woody internal rod is probably both a fascicular and interfascicular cam- for each external rib of the shoot (Figure 2). This is bium present in the secondary growth of subdermal a typical architecture for the Pachycereeae, in which thorns of L. schottii given the solid nature of these the woody rods can be spread farther apart as the woody thorns, vice the reticulate look of wood in shoot develops, allowing the pith to enlarge (Gibson shoots of Cylindropuntia (Engelm.) F.M. Knuth 20 GORELICK—LOPHOCEREUS WOODY THORNS Figure 4. Thick thorns on lower part of shoot. Note branch Figure 5. Thick thorns with hollow centres on lower part of at upper left of this thorny shoot. shoot. and vascular traces to flowers of Hylocereus undatus secondary growth connecting areoles to a vascular (Haw.) Britton & Rose (Weidlick 1974; Schwager, cylinder probably occurs in other individuals of L. Neinhuis, and Mauseth 2015). Vascular traces with schottii. While photographs of dead and dying speci- lignified secondary growth are known to provide mens are rare, especially of this relatively photogenic structural support to shoots (Tomlinson et al. 2005). pseudocephalium-bearing species, Arnaud Martin The exceptionally slow growth of the massive shoots (personal communication) took a photo of a plant of L. schottii, at least until one meter tall, may pro- from near Cataviña, Baja California with one tall vide time for the shoots to develop large lignified dead shoot that appears to show numerous thin sub- subdermal thorns to support the shoot, especially to dermal thorns about 1.0–1.5 meters above ground. support the fast-growing pseudocephalium. Lower portions of the dead shoot are obscured in his The attached photos of Lophocereus schottii skel- photo, which is low-resolution and backlit. Therefore etons were from a single plant at Organ Pipe Cactus further confirmation of the subdermal thorns in L. National Monument (Figure 6). This locale is at the schottii is needed. extreme northern range for this species, which seems Thick, woody, subdermal thorns make sense for to be cold limited. The photographed plant is tagged supporting shoots with massive cortex and pith, as as specimen #95, near the north terminus of Senita found in the Pachycereeae with their fastigiate rods Basin Road, less than ten kilometers from the Mex- of a vascular cylinder, especially in very slow growing ico/US border. There are only a few hundred plants juvenile shoots. What makes less sense is the rapid of senita native in the U.S. (Parker 1989). Some of shoot elongation once shoots reach sexual maturity these individuals may be clones of one another, with at approximately one-meter tall to form a pseudo- tall shoots eventually bending over after reaching cephalium. Adult shoots with pseudocephalia often 2.5–4.0 meters tall (Martorell, Vega, and Ezcurra lean, sometimes precipitously, because they do not 2006), and then possibly (although this has never have time to grow thick subdermal thorns. But why been thoroughly documented) striking adventitious the rapid shoot elongation? These plants do not need roots. This phenomenon of subdermal thorns with to produce many new areoles because of non-existent HASELTONIA VOL. 22. 2016 21 Figure 6. Overview of plant #95, with another senita at upper right, one Stenocereus thurberi between the two senitas, and many saguaros in the background on the opposite side of Senita Basin Road. constraints of determinate growth, as with Melocac- Cactus and Succulent Journal 77: 132–143. tus Link & Otto. L. schottii is able to produce viable Molina-Freaner F, Tinoco-Ojanguren C, Nik- flowers and fruits from old areoles of pseudocepha- las KJ. 1998. Stem biomechanics of three co- lia and are able to simultaneously produce multiple lumnar cacti from the Sonoran Desert. American flowers and fruits per areole (Gibson and Nobel Journal of Botany 85: 1082–1090. 1986)l. Could this rapid elongation of reproductive Niklas KJ, Molina-Freaner F, Tinoco-Ojan- shoots and subsequent breakage of overly tall shoots guren C. 1999. Biomechanics of the columnar be another maladaptive trait, such as with abscission cactus Pachycereus pringlei. American Journal of of pseudocephalia on the closely related Backebergia Botany 86: 767–775. militaris (Audot) Bravo ex Sánchez-Mej., tiponchi, Parker KC. 1988. Growth rates of Stenocereus thur- synonym Pachycereus militaris (Audot) D.R. Hunt beri and Lophocereus schottii in southern Arizona. (Mauseth et al. 2005)? Botanical Gazette 149: 335–346. Parker KC. 1989. Height sturcture and reproduc- ACKNOWLEDGMENTS tive charactersistics of senita, Lophocereus schottii (Cactaceae), in southern Arizona. Southwestern Thanks to Barbara Terkanian who braved camp- Naturalist 34: 392–401. ing overnight at +3°C and 5cm of rain without a Schwager H, Neinhuis C, Mauseth JD. 2015. tent or stove in order to visit these senitas with me. Secondary growth of the leaf and bud traces in Thanks to Arnaud Martin for sharing photos. Hylocereus undatus (Cactaceae) during the forma- tion of branches or flowers. International Journal REFERENCES of Plant Sciences 176: 762–769. Tomlinson PB, Fisher JB, Hallé F, Villalobos Gibson AC, Nobel PS. 1986. The cactus primer. R. 2005. Development of woody branch at- Harvard University Press, Cambridge. tachments in Schefflera (Araliaceae or Apiaceae). Martorell C, Vega E, Ezcurra E. 2006. Mor- American Journal of Botany 92: 1765–1773. phological consequences of the trade-off between Weidlick WH. 1974. The induction of excessive growth and reproduction in a columnar cactus amounts of secondary growth in the petioles of (Lophocereus schottii). Plant Ecology 183: 125– Norway maple (Acer platanoides).
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