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Biology of the Bamboo <Emphasis Type="Italic">Chusquea Culeou </Emphasis> (Poaceae: Bambusoideae) in Sout

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Biology of the Bamboo <Emphasis Type= Vegetatio 111: 93-126, 1994. © 1994 Kluwer Academic Publishers. Printed in Belgium. 93 Biology of the bamboo Chusquea culeou (Poaceae: Bambusoideae) in southern Argentina Anita K. Pearson 1, Oliver P. Pearson i & Isabel A. Gomez 2 l Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA 2 Grupo de Andlisis de Sistemas Ecol6gicas, Bariloche, Argentina Accepted 3.2.1993 Keywords: Biomass, Chusquea culeou, Nothofagus forests, Patagonia, Productivity, Bamboo Abstract Over a period of 7 years the biology and phenotypic variability of Chusquea culeou were studied at 5 locations in cool temperate forests of southern Argentina. Excavated rhizomes had an average of 1.1 successful rhizome buds, and an average of 2.1 years elapsed between successive generations of rhi- zomes. Rhizome buds usually develop within the first four years after a rhizome forms. Height, volume and weight of a culm can be calculated from its diameter 1 m above the ground. Culm size, length of foliage leaf blades, and pattern of secondary branching differed among study sites. Dead culms were numerous and commonly remained erect for more than 7 years after dying. New culm shoots appear in spring and reach full size within a few months. Shoots can grow more than 9 cm/day. Less than half of the shoots survived a year; most were killed by moth larvae. Multiple primary branch buds emerge through the culm leaf sheaths in the second spring. The mean number of branch buds at mid-culm nodes varied between 34.8 and 81.5, and the mean number of primary branches was between 22.8 and 40.8. Number and length of branches, and number and length of foliage leaf blades at each node is related to the position of the node on a culm. Most branches grow about 3 cm and produce 1 to 3 foliage leaves annually. Foliage leaf blades generally live 2 years or more; few survive 6 years. Relative lengths of fo- liage leaf blades and their spacing along a branch permit recognition of annual cohorts. Both gregarious and sporadic flowering have been reported, and every year a few isolated plants flower and die. Length of the life cycle is unknown. Seedlings require up to 15 years to produce culms of mature size. Foliage branches may live more than 23 years, and culms may survive 33 years. Extensive loss of new shoots to predation suggests that gregarious flowering may be driven by a need to escape parasit- ism. C. culeou clumps expand slowly. Average annual rate of increase of the number of live culms in a clump was 4.6?o. Methods of seed dispersal are undocumented. A dense stand of Chusquea culeou had an estimated phytomass of 179 tons/hectare (dry weight), 28 ?o of which was underground. Net annual production was about 16 t/ha dry weight. Nomenclature: Voucher specimens from bamboo plants in this study were deposited in the following herbaria: Inst. de Bot~nica Darwinion, San Isidro, Argentina (SI); Iowa State University, Ames, Io- wa (ISC); Smithsonian Institution, Washington D.C. (US); and University of California, Berkeley (UC). Specimens, photographs and drawings of the monitored clumps were examined by Dr. Lynn Clark of Iowa State University, who identified all as Chusquea culeou E. Desvaux in Gay (1853), Hist. Chile, Bot. 6: 450. 94 Introduction 47°S in the southern cone of South America (Parodi 1945)• In Argentina, Chusquea culeou oc- Bamboos of the genus Chusquea are widely dis- cupies a narrow band along the eastern Andean tributed in Central and South America (Clark slopes of the Provinces of Neuquen, Rio Negro, 1989). However, unlike Asiatic bamboos that and northern Chubut; Parque Nacional Nahuel have been extensively utilized by native popula- Huapi lies near the center of its range, at latitude tions and have been cultivated and studied for 41°S (Fig. 1). Here in the park's glacier-carved hundreds of years (Janzen 1976), little use is made valleys are great native forests dominated by of Chusquea. It is not cultivated, and few studies beech trees of the genus Nothofagus, with bamboo have been published on any aspect of its biology as a major understory component• Although man (see Veblen 1982). has had relatively little impact on these forests, Our observations on Chusquea were begun in glaciers, avalanches, mudslides, volcanic ashfalls 1978 incidental to a study of small mammals in and fires have all influenced the distribution of the bamboo habitats in Parque Nacional Nahuel beech trees and the bamboo• Precipitation also Huapi, southern Argentina (Pearson & Pearson has had an effect: bamboo is more abundant in 1982; Pearson 1983). One goal was to document interactions between small mammal populations and the flowering cycle of the bamboo, but the lack of well documented information about the natural history and flowering behavior of the Chusquea, and uncertainties about its taxonomy encouraged us to focus on this plant. The abun- dance of Chusquea in many plant communities within the park (Mermoz & Martin 1987) offered an unusual opportunity to study the bamboo in its natural, relatively undisturbed environment. 1 From 1984 through 1991 we observed and mea- I \ sured selected Chusquea culeou plants in an effort I to document phenotypic variability and to quan- tify growth, productivity and survival. Similar de- tailed, long-term observations are not available for other American bamboos. We also wished to learn what we could about the flowering cycle. Like other woody bamboos, the bamboos of Parque Nacional Nahuel Huapi flower and die r., after a long vegetative period, but there are few precise records of the life cycle of any species of ,a Chusquea. P arodi ( 1941, 1945) recognized three species of Chusquea in southern Argentina (C. culeou Des- vaux, C. argentina Parodi and C. montana Phil- ,a Verana a ippi), and in addition described the subspecies I I I I I C. culeou longiramea. All four were said to occur 0 20 km in a small western portion of Parque Nacional Nahuel Huapi (Parodi 1941, 1945; Nicora 1978), Fig. 1. Location of Parque Nacional Nahuel Huapi in south- but only C. culeou, the species we have studied, is ern Argentina. The five areas where clumps of C. culeou were widely distributed, occurring between 35°S and studied are indicated. 95 Table 1. Characteristics of study areas with marked clumps. Locality Altitude Annual Exposure Light intensity index* Associates within 2 m precip. Periphery Periphery + 2 m Puerto Blest (71 ° 49' W; 41 ° 1' S Clump # 1 765 m 400 cm Dense shade 6.20; 6.3°o Nothofagus dombeyi Myrceugenia chrysocarpa Berberis linearifolia Clump #2 765 m 400 cm Dense shade 5.4°; 9.4°; N. dornbeyi Castafio Overo (71 ° 47' W; 41 ° 12' S) Clump # 1 870 m 200 cm Valley floor 32.40 95.1~; N. antarctica Some shade Clump #2 870 m 200 cm North slope 20.4°0 70.6°; N. dombeyi, N. pumilio, Blechnum sp., Berberis linearifolia, B. darwini, B. sp., herbs Llao Llao (71 ° 33' W; 41 ° 3' S) Clump # 1 780 m 240 * Dense shade 13.5 ° o 9.60o N. dombeyi Clump # 2 780 m 240 Dense shade 1.4°o 4.200 N. dombeyi La Veranada (71 ° 28' W; 41 ° 29' S) Clump # 1 1090 m ... Valley floor 34.0 o; 80.0 o5 N. antarctica, Berberis Some shade buxifolia, thick grass Clump # 2 1090 m ... Some shade 28.4?, o 68.4°0 N. antarctica, Ribes cucullatum, grass and herbs Clump # 3 1090 m ... Some shade 21.20o 74.4°0 N. antarctica, B. buxifolia, grass and herbs Cerro Otto (71 ° 20' W; 41 ° 8' S) Clump # 1 1145 m 130 * SE facing 29.5°0 100.0°0 Berberis sp., Ribes sp. slope; open Gaultheria sp., Ovidia pillo-pillo, grass and herbs Clump # 2 1160 m 130 SE facing 6.5) o 34.6°o N. pumilio slope; shade * Estimated as described in Methods. the wet, mild climate near the Chilean border Methods (over 4000 mm annual precipitation), and disap- pears toward the drier, colder eastern edge of the Over a period of 13 years we recorded our own park 60 km away (600 mm annual precipitation). observations and information gathered from local Chusquea grows and thrives near Lago Nahuel inhabitants concerning the vegetative character- Huapi (elevation 760 m), where most precipita- istics and flowering behavior of Chusquea culeou. tion falls as rain, but is also found up to 1450 m During six years, detailed measurements were along the ski slopes of Cerro Catedral. It occurs made on two clumps of bamboo at each of five both in pure stands in the open as well as beneath locations differing in climate, altitude and plant the dense canopy of Nothofagus forests. associations (Fig. 1 and Table 1). Although the 96 Table 2. Characteristics of monitored clumps. Clump Year Clump Culms over 1 meter in height marked area a (m2 ) N N/m 2 Live:N Mean diam b Max N Mean height c Mass culms Mass culms Mass clump -+ se nodes + se - foliage d + foliage ° FW/m 2 (mm) (m) (kg) (kg) (kg) Puerto Blest #1 1984 6.19 123 45.6 108(88~o) 20.25_+0.29 58 5.14_+0.07 ll2.1 138.2 22.3 Puerto Blest #2 1985 0.72 52 72.2 47(90%) 12.53+0.25 40 3.33_+0.06 15.7 20.4 28.4 Castafio Overo # 1 1984 6.42 121 18.8 107 (889o) 8.27 + 0.37 42 2.33_+ 0.09 17.2 25.6 4.0 Castafio Overo #2 1985 2.35 44 18.7 38 (869°) 5.73 + 0.40 36 1.74 + 0.09 4.0 8.2 3.5 LlaoLlao#1 1984 4.82 146 30.3 108(749o) 21.47_+0.69 44 5.43_+0.16 121.8 149.6 31.0 LlaoLlao #2 1984 7.36 269 36.6 203(75~o) 15.35_+0.26 39 3.99_+0.06 120.1 151.9 20.6 LaVeranada #1 1984 2.01 60 29.8 56(9390) 7.98_+0.34 33 2.27_+0.08 7.5 11.3 5.6 LaVeranada #2 1985 7.06 138 19.4 121(88%) 5.74_+0.20 32 1.74_+0.05 6.2 12.5 1.8 La Veranada #3 1988 10.52 189 17.2 180 (95~o) 8.12_+ 0.15 32 2.30 + 0.04 25.6 38.2 3.6 CerroOtto #1 1984 5.82 386 66.3 313(81~o) 18.49_+0.52 58 4.73_+0.12 259.4 322.0 55.3 CeroOtto #2 1984 5.48 234 42.7 195(83%) 17.07_+0.24 42 4.39+0.06 161.1 201.3 36.7 Calculated from measured circumference of clump 1 m above ground.
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