PRODUCTIVITY AND SUSTAINABILITY OF A VEGETABLE PALM (PHYTELEPHASAEQUATORIALIS, ) UNDER THeE MANAGEMENT REGIMES IN NORTHWESTERN 1

J. VELASQUEZ RUNK

Vehlsquez Runk, J. (Duke University, School of the Environment, Box 90328, Durham, NC 27708, USA; Present address: Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Reptiblica de Panamd). PRODUCTIVITYAND SUSTAINABILITYOF A VEGETABLE|VORY PALM (PHv- TELEPHAS AEQUATORIALIS, ARECACEAE)UNDER THREE MANAGEMENTREGIMENS IN NORTHWESTERN ECUADOR. Economic Botany 52(2):168-182. 1998. This study examines the productivi~, sus- tainabili~, and management of tagua nuts from the palm aequatorialis under three management regimes in northwestern Ecuador. Tagua nuts are used internationally as an ivor3.' substitute and the palm fronds are used locally as roof thatch. Plots were established in a control site and in three local management regimes: pure tagua groves; stands with tagua and cacao in an agroforestry system; and tagua groves where the undergrowth is cleared and old fronds removed. Data were collected for one year and local extractors were interviewed about their tagua management. Palm demography indicates heavy management with few subadult individuals and many more adult females than males. Productivity analyses do not demonstrate the influence of any one environmental variable on or infructescence productivi~.'. The pure tagua grove is the management regime most conducive to tagua sustainabili~.. The sus- tainabili~ results are underscored by the extractive methodology, where local extractors max- imize tagua collected while minimizing collection time. These results are incorporated into management recommendations to fi)ster tagua extraction while ensuring its sustainability and future use.

Este estudio se examina la productividad, sustentabilidad, y el manejo de nueces de tagua de la palma Phytelephas aequatorialis bajo tres regimenes de manejo en el noroccidente del Ec- uador Nueces de tagua estdn usados internacionalmente como un sustituto para mar[il y las frondas de la palms se usa Iocalmente para techos. Se estableci6 cuadrantes en un sitio de testigo yen trds regimenes de manejo local: arboledas de tagua pura; arboledas con tagua y cacao en un sistema agroforestal; arboledas de tagua donde se limpia el suelo y remueve las malezas y las frondas viejas. Se recolectaron datos por un afio y se entrevist6 extractores locales sobre su manejo de tagua. La demografia de las palmas indic6 que el manejo es intenso con pocos individuos de subadultos y mucho mds hembras adultas que varones. La anah'sis de la productividad no demostr6 la influencia de un solo variable en la productividad de hojas o infructescencias. La arboleda de tagua pura es el regimen de manejo rods conducible a sus- tentabilidad de tagua. Los resultados de sustentabilidad estdn subrayados por la metodologfa de extracci6n, en que extractores locales maximizan la recolecci6nn de tagua mientras min- imizan el tiempo de recolecci6n. Estos resultados estdn incorporados en unas recomendaciones a fomentar la extracci6n de tagua mientras asegura su sustentabilidad y uso futuro. Key Words: Phytelephas aequatorialis; ; tagua; Arccaceac; Ecuador; agro- forestry.

Non-timber forest products (NTFPs) have had while providing economic benefits to local res- substantial cultural and economic importance for idents (Peters, Gentry, and Mendelsohn 1989). local peoples. Over the past decade conserva- These non-wood biological resources, such as tionists have increasingly considered the use of , fruits, , and seeds, are removed from NTFPs as a means to maintain forest ecosystems natural forests to be sold locally, nationally, and/ or internationally. Among the recent crop of products for the environmentally conscious or t Received 17 November 1996; accepted 29 Novem- "green" consumer market are tagua nuts from ber 1997.

Economic Botany 52(2) pp. 168-182. 1998 91998 by The New York Botanical Garden, Bronx, NY 10458 U.S.A. 1998] VELASQUEZ RUNK: VEGETABLE IVORY PALM 169

Fig. 2. Phytelephas aequatorialis infructescence morphology. From left to right; individual fruit with fleshy, inner mesocarp; "woody" exocarp from one fruit; seeds with inner mesocarp from the infructes- cence; infructescence peduncle with individual fruit scars.

and seeds. The leaves are dried and used as roof thatch. The immature and palm heart are used for human consumption, the inner mesocarp for bait in fish and traps, the for birth control, and the trunks of the Fig. 1. Adult Phytelephas aequatorialis. Notice male for firewood (Coles-Ritchie 1993; Vehis- the large infructescences on the female palm, left. quez Runk 1995). In addition, dried tagua fronds have reportedly been used for brooms (Barfod the palm Phytelephas aequatorialis. The nuts 1991) and staminate flowers may be used as fod- are a type of vegetable ivory used for buttons der for cattle in the upper elevations of the and carvings. palm's range (Kozoil and Borgtoft Pedersen Phytelephas aequatorialis [syn. Palandra ae- 1993). The residual tagua powder from button quatorialis (Spruce) O.E Cook] is a dioecious processing is supplemented with protein and palm endemic to western Ecuador. It grows from used as a livestock fodder (Barfod, Bergman, sea level to 1500 meters on Andean slopes and Borgtoft Pedersen 1990) and the non-gov- (Borgtoft Pedersen 1993). Large trees may grow ernmental organization Conservation Interna- to 10 m in height (Fig. 1). The infructescence is tional is marketing the powder as an industrial very large, 30-40 cm in diameter with a abrasive (Frank 1995). "woody" surface. A single infrnctescence may Tagua has been traded in international mar- have from seven to twenty-two fruits, each con- kets since about 1865 (Acosta Solis 1948). It taining five to seven seeds (Fig. 2). Borgtoft was used primarily for buttons as well as for Pedersen (1993) estimated fruit development to some toys and figurines. In 1887, the export of take 3 years in lowland sites. Seeds are collected tagua from the port city of Esmeraldas account- after the infrnctescence matures and falls to the ed for 76.2% of that port's earnings (Barfod ground, breaking open and exposing the seeds. 1989). The demand for tagua declined dramati- Although tagua seeds are known to have a cally during World War II, with the advent of prolonged dormancy, information is scant on plastics (Barfod 1989). In 1987-1988, the inter- seed germination rates (Barfod 1991). Neither is national market for tagua increased sharply it known how many years juveniles take to de- (Borgtoft Pedersen 1993) and continues to grow velop a stem, that is, become a subadult, nor with rising consumer interest in "green" prod- how many years subadults require to reach ma- ucts (Macray pers. comm.). turity. Eighty-six percent of present-day tagua ex- Local people have a number of uses for Phy- tractors in Ecuador's Comuna Rio Santiago- telephas aequatorialis, most of which involve its Cayapas work exclusively in agriculture (Coles- 170 ECONOMIC BOTANY [VOL. 52

whole, exported whole, or taken to a local fac- tory for button processing. The whole seeds are used for carvings and figurines. Tagua nuts are a non-timber with current and historical importance for the cultures and economy of local communities in northwestern Ecuador. Like many NTFPs the economics of tagua extraction has been studied (Barfod 1989; Barfod et. al 1990; Coles-Ritchie 1993; Hardner 1995), but the ecology has re- ceived less attention. In general, the enthusiasm for extractive systems has been tempered by the lack of ecological information surrounding NTFP extraction (Godoy and Bawa 1993; Hall and Bawa 1993; Peters 1994; Peters 1996; Pan- ayotou and Ashton 1992; Nepstad 1992; Red- ford and Padoch 1992). Conservation and de- velopment specialists often assume that the re- moval of such products does not impact ecosys- Fig. 3. Boys cutting off inner mesocarp from Phy- tem structure and function and/or is sustainable telephas aequatorialis seeds. for the exploited population. This research joins several other recent studies in addressing this void by considering impacts of non-timber Ritchie 1993) (Fig. 3). The same percentage forest product extraction, specifically, the eco- manage tagua in agroforestry systems (Coles- logical and socio-ecological impacts of tagua Ritchie 1993) usually involving cacao trees extraction from Phytelephas aequatorialis in (Theobroma cacao) cultivated in the shade of northwestern Ecuador. tagua palms. The remaining 14% of extractors tend pure tagua stands, where tagua palms occur STUDY AREA without other crops (Coles-Ritchie 1993). Most The study area is the Comuna Rio Santiago- tagua extractors conduct limited management of Cayapas in Esmeraldas Province, Ecuador. The their tagua groves. However, twenty-eight per- area is classified as moist tropical forest and cent of extractors "clear" their tagua groves very moist tropical forest in the Holdridge sys- (Coles-Ritchie 1993), which involves cutting tem (OAS 1979), and the nearby town of Bor- vegetation from around the base of palms and bon receives about 2.3 m of rain annually, with removing old fronds and lianas. This clearing the rainier months occurring from December to facilitates the collection of tagua nuts because June. The area is considered a biological "hot- they are more easily seen. Some farmers also spot" for its high endemism (Myers 1990; My- believe that clearing increases the production of ers 1988) and borders the western side of the nuts. Cotacachi-Cayapas Ecological Reserve. An unpublished 1995 study showed that ex- The 63 000 hectare Commune is inhabited by tractors received between US $4.35 and $6.25 8000-10 000 Afro-Ecuadorians, who are subsis- for 100 pounds of tagua (Hardner pers. comm.). tence cultivators, relying mainly on tagua, plan- Tagua extraction represented about 40% of tains, cacao, and timber managed in forest gar- monthly income for the extractors. The market dens. The Commune is occupied almost entirely for tagua thatch is informal and supplied on an by secondary forest, with old-growth forest re- as-needed basis. stricted to upland areas (Foster 1990) and small Once tagua nuts are extracted, they are taken patches on individual farms (Calero pers. to local collection centers. From these centers, comm.). Estimates from overflights and site vis- they are either taken to nearby tagua drying pat- its indicate that tagua palms occupy 2.1%-2.5% ios or shipped to the city of Manta where they of the Commune. Tagua often grows in relative- are dried for about 3 months (Macray pers. ly small (0.75 ha), monotypic stands (taguales). comm.). Once dried, they are shelled and used It is not clear whether residents have enriched 1998] VELASQUEZ RUNK: VEGETABLE IVORY PALM 171 these stands over time, although local extractors to measure accurately, height was not measured believe these taguales occur naturally. among younger size classes. Light exposure was estimated visually on a scale of 0 to 5, in incre- METHODS ments of 0.5, with a measure of 0 indicating to- This study examined whether the extraction of tal shade and 5 indicating total exposure. Foliage tagua nuts has an impact on the growth, repro- cover was measured using the standard method duction, and sustainability of tagua palm popu- from the base of the adult palm to the tip of the lations under three different management re- longest leaf. The absence of lianas, vines, and/ gimes: pure tagua groves; tagua and cacao in an or termites was noted as a measure of the overall agroforestry system; and "cleared" tagua health of a palm. groves. Under all three management regimes, To track productivity, and in- extractors collect tagua nuts. fructescences were counted and painted with Palms were sampled using 20 x 30 m plots colors that alternated with each field visit. Fol- in each management regime (Pinard and Putz lowing Borgtoft Pedersen's method (1993), only 1992; Pifiero, Martinez-Ramos, and Sarukh~in infructescences greater than or equal to 20 cm 1984). Plot sites were chosen to minimize eco- in diameter were marked. To gauge leaf produc- logical differences, such as habitat, palm densi- tivity of adults and subadults, a loose nylon cord ty, degree of human disturbance, and soil type. was placed around the spear leaf and the three Each management type was represented by one closest extended leaves were painted as a back- plot that was less seasonally inundated and one up procedure in the event the cord was lost. For plot that was more seasonally inundated. the juveniles, leaves were painted to track pro- Throughout the rest of this paper "dry" is used ductivity. Juvenile palm leaf length, number of to indicate the less inundated plots and "wet" leaves, leaf productivity, light exposure, and to indicate the more inundated plots. The wet density were also measured. sites were periodically inundated with slightly Each data collection period coincided with a brackish, sediment-laden river water during the period of nut extraction. Seeds in each plot were six-month rainy season. Water may stagnate at marked and recaptured in order to obtain seed- such sites. A seventh unmanaged, "control" plot crop estimates. The viable, non-viable, and ger- was located in an area of old growth forest on minating seeds that remained on the ground a hill: although habitat and soil type differed for post-collection were counted. Heavier seeds this plot, it was the only area located where tag- without holes were tallied as viable, while light, ua remained unexploited. The control plot was punctured, or infested seeds were tallied as non- within several kilometers of the other sites at viable. All seeds were painted with colors that about 300 m elevation. Soil at this site was a alternated with each field visit, and then turned clayey tropothent whereas the other sites were so that the color was not visible to the extractors, silty fluventic dystropepts or aquic eutropepts. who continued their typical extraction regimes All individuals were mapped and classified throughout the course of the study. This method into one of three growth stage classes: juveniles, assumes that the movement of seeds into the plot subadults, and adults. Juveniles are those indi- is negligible. Questionnaires were used to obtain viduals ranging in size from seedlings to un- data on tagua management from the extractors stemmed with large fronds. Subadults are involved in the studies. In addition, participant immature, caulescent palms and adults are ma- observation of extractors at each site was used ture (reproductive), caulescent palms. Data were to obtain data on tagua collection methods. collected every three months for one year. For The sustainability analysis was done using an each plot, a number of variables were recorded: individual-based model, POMIB (Philippi, Peet, adult palm density, distribution, sex, stem and Christensen n.d.). The POMIB simulation height, light exposure, foliage cover, number of begins by comparing the relative growth rate leaves, number of cut leaves, number of dead and annual growth increment of each juvenile to leaves, and presence of leaves, inflorescences, determine the best estimator of growth. The data and infructescences. Stem height was measured for all juveniles were pooled and relative growth from the ground to the margin of the leaf sheath rate was used to estimate size-dependent survi- surrounding the spear leaf of adult and subadult vorship probabilities and growth rates for each palms. Because juvenile palm height is difficult height. Growth rates and survival probabilities 172 ECONOMIC BOTANY [VOL. 52

of new juveniles were simulated with 100000 iterations until individuals either died or became subadults. A random number generator was used to select an individual from the population prob- ability density function, and simulate whether it lived or died based on its size-specific survival probability. If the individual survived, the model simulated how much it grew based on the size- specific relative growth rate. These last two steps were repeated until each individual died or became an adult. The result is an estimate of the proportion of individuals that will survive to adulthood, and an estimate of the time necessary for them to do so. Because this model is indi- vidual-based it is more advantageous than anal- ysis via a Leftkovitch stage-based matrix, for which a number of assumptions are required, such as guessing the length of seed dormancy and approximating the duration of stage classes.

RESULTS AND DISCUSSION PALM DEMOGRAPHY The number of palms per each 600 m 2 site varied substantially (Fig. 4). The plots in the wetter sites contained many fewer juvenile palms than did drier sites. This is likely due to more persistent flooding in the wetter plots, which may impede seedling establishment. Overall, the number of subadults was unex- pectedly low, a result of extractors' cutting ju- veniles and preventing them from reaching older stage classes. When extractors enter a tagual, they typically cut some of the competing vege- tation, including tagua seedlings, to see seeds better. Although the sample size is small, the absence of subadults in the cacao and tagua plots Fig. 4. Palm Life Stage Distribution. could be due to additional shading from cacao trees or, more likely, greater cutting to manage both in those sites. Extractors at the ca- neck in managed stands, limiting the number of cao sites did not note any different management adults. The number of adults is fairly constant, techniques from the other sites. The unmanaged with the exception of the unmanaged plot, as a plot had a ratio of 1:1.5 subadults to adults, with result of extractors' management for what they noticeably more subadults per palm than in the believe is an appropriate density. That is, they other plots. Bernal (in preparation) noted ap- may maintain the tagual at a preferred density proximately equal numbers of older juveniles and remove subadults and/or adults above that and adults in his work on P. seemannii. Many density. palm demographic studies show a large propor- The female:male ratio differs noticeably be- tion of older juvenile stage classes (Enright and tween the unmanaged and managed sites. In the Watson 1992; Olmstead and Alvarez-Buylla unmanaged plot, the ratio of females to males is 1995; Oyama 1990; Sullivan, Konstant, and 2:1, whereas that of the managed plots is an av- Cunningham 1995). The lack of juveniles to the erage of 7:1. This ratio is an artifact of manage- subadult stage appears to be a population bottle- ment, as extractors select against male palms be- 1998] VELASQUEZ RUNK: VEGETABLE IVORY PALM 173

Fig. 5. Palm Germination with Annual Rainfall. cause they do not, bear fruit. Borgtoft Pedersen's (1993) research with this same species showed a 1:1 female : male ratio in upland agroforestry plots where palms are not managed for seed col- lection and Bernal's (n.d.) work with P. seeman- nii also demonstrated a 1:1 female to male palm ratio. In addition, a 1:1 female to male palm ra- tio was found in several studies of other palm species (Oyama 1990; Pifiero, Martinez-Ramos, Fig. 6. Annual Leaf Productivity per Quarter. Each and Sarukh~in 1984; Sarukh~in, Martinez-Ramos, circle indicates a single adult palm (male or female). and Pifiero 1984). The relatively low male palm density did not seem to reduce pollination rates or the ability of the females to set fruit. found a germination peak in Chamaedora bart- During the year no individuals in any plot lingiana after the highest annual precipitation. grew into the adult or subadult size classes. Fig- In addition, the wet sites demonstrate fewer ure 5 illustrates seed germination over the year individuals recruited into the seedling popula- and consequent recruitment into the juvenile size tion than do the dry sites. Perhaps the increased class. The values in Fig. 5 indicate a seasonal flooding of these site prohibits greater seedling trend, with increased germination during the dry establishment by promoting a negligible change season (July-November). A temporal analysis of of water potential. This idea may be supported seed availability (that is, seed production minus by the unmanaged site's lack of a seasonal ger- extraction) demonstrated that more seeds are mination trend: its position on a steep slope may available during the dry season. However, be- prevent change of water potential, since water cause of an extended dormancy period, perhaps moves through the soil profile much more quick- greater than two years (Barfod 1991), it is un- ly at the site. likely that germination rates reflect seasonal availability. Higher germination rates in the dry PRODUCTIVITY season may be triggered by the change of water Neither leaf nor infructescence productivity potential following the wet season. Soaking differed per season (Fig. 6 and Fig. 7). DeSteven palm seeds in standing or running water was et. al. (1987) found both seasonal productivity found to be a successful germination treatment of leaves and inflorescences among a palm as- in a number of ornamental palms (Odetola semblage in Panam~i. Additional years of data 1987). As well, Ataroff and Schwarzkopf (1992) might serve to verify such a trend. In all sites, 174 ECONOMIC BOTANY [VOL. 52

dation variables is indicative of each variable's impact on annual leaf productivity. To further examine the impact of management technique on annual leaf productivity, a separate analysis was conducted with each management type vari- able (Table 2). When these data are sorted per management technique it becomes clear that the modeling de- scribed in Table 1 does not hold true for each management technique, but instead, aggregates the impact of each variable in all three manage- ment techniques. For example, in the tagua and cacao plots, age is much more important for leaf productivity (P = 0.0133) than in the tagua grove or cleared tagua grove sites (P = 0.3336 and P = 0.2561, respectively). In the overall re- gression equation, the greater importance of age in the tagua and cacao management regime el- evates the importance of that variable across all management techniques, where it is very signif- icant (P = 0.0055). At the same time, the degree of seasonal inundation also affects the leaf pro- ductivity across all management regimes. In ad- dition, the R 2 value for pure tagua grove man- agement is noticeably higher than the R 2 values for tagua and cacao and "cleared" tagua man- agement regimes (Table 2), although all R 2 val- ues are significant (F --- .0001). A high positive correlation between palm Fig. 7. Annual Infructescence Productivity Per Quarter. Each circle indicates a single female, adult growth and light is well established (Oyama palm. 1990; Pifiero and Sarukhfin 1982; Pinard and Putz 1992). Borgtoft Pedersen's 1993 study in- dicates a significant correlation between growth productivity of both leaves and infructescences and light in tagua palms from lowland agrofor- appeared to be negatively affected by inundation estry systems (P = 0.0001), but not for those on in the wetter plots. To further investigate the im- the Andean slopes. These data did not support pact of several variables on leaf and infructes- that same analysis. In addition, leaves grown in cence productivity, general linear models were the shade are thicker and more suitable for roof- used. ing thatch, indicating that there is value in hav- The results from the general linear model ing palms that are shaded (Borgtoft Pedersen (GLM) for leaf productivity are presented in Ta- 1993). Male palms are used more often for roof- ble 1. GLM results for annual leaf productivity ing because the local residents recognize that the indicate a robust model using five variables: removal of fronds from the female palms inhib- management, degree of seasonal inundation, an its infructescence productivity. Borgtoft Peder- interaction of management and seasonal inun- sen (1993) also found that male palms produce dation, age (using height as a proxy variable), more leaves than female palms, however the and light level, with an R 2 = 0.55983 (Table 1). small number of male palms in this study pre- The inclusion of the management and inun- vented assessment of difference between female dation interaction term in the model demon- and male leaf productivity. Given that leaves strates that variability exists between plots, re- grown in the shade are better for roofing thatch gardless of the management technique or degree and that males fronds are more frequently used of inundation. Yet, the significance level of both for roofing, it is suggested to leave male palms the management and degree of seasonal inun- in areas which receive indirect sun, so that pro- 1998] VELfitSQUEZ RUNK: VEGETABLE IVORY PALM 175

TABLE 1. GENERAL LINEAR MODEL PARAMETER ESTIMATES FOR ANNUAL LEAF PRODUCTIVITY.

Parameter Parameter estimate Significance probability Standard error Management Tagua Grove 3.0805 0.0001 0.7065 Tagua and Cacao 6.3963 0.0001 0.8917 "Cleared" Tagua 4.4651 0.0001 0.8801 Degree of Seasonal Inundation Dry 1.8091 0.0051 0.6273 Wet 0.0000 -- -- Management and Degree of Seasonal Inundation Tagua Grove, Dry 1.7328 0.0972 1.0320 Tagua Grove, Wet 0.0000 -- -- Tagua and Cacao, Dry -2.5203 0.0124 0.9846 Tagua and Cacao, Wet 0.0000 -- -- "Cleared" Tagua, Dry 0.0000 -- -- "Cleared" Tagua, Wet 0.0000 -- -- Age (using Height as a proxy) 0.3902 0.0055 O. 1367 Light Level 0.2901 0.2949 0.2751

General linear model parameter estimate table for annual leaf productivity. The equation is fit basing comparisons against the cleared tagua and wet variables, therefore, when the interaction term of management and degree of seasonal inundation include either the cleared tagua or wet variable, the mean estimate does not differ from that variable alone. The R 2 value for the full model fit is 0.55983. Statistically significant parameter estimates are italicized (a 0.01). ductivity and high quality thatching can be toft Pedersen (1993) found a strong correlation maintained. A positive correlation of palm age, between light exposure and production of in- using height as a proxy variable, with leaf pro- fructescences in a lowland site (OL = 0.001). His ductivity is a relationship that was anticipated study was limited to one lowland site, so the by tagua extractors (field notes 1993). However, correlation may exist only on that site, parallel- these data did not demonstrate that relationship. ing the differences of the importance of light The general linear model for infructescence intensity in these data. productivity yielded results similar to that of leaf productivity. The GLM for infructescence pro- SUSTAINABILITY ductivity used five variables: management, de- Evaluation of ecological sustainability of tag- gree of seasonal inundation, an interaction of ua management was based on a POMIB com- management and seasonal inundation, age, leaf puter simulation (Philippi, Peet, and Christensen productivity, and height with an R: = 0.5860 at in preparation). A 5 m frond length was taken a P = 0.0001 (Table 3). Because infructescence as the minimum size of adult or subadult palms productivity was found to be highly correlated and thus the final point in the simulations, where with leaf productivity, 0.6536 (a = 0.0001), the the palm had survived and grown to subadult- latter was removed from the model. hood. In all plots, juveniles with frond lengths The model above is the fit for infructescence from 2 to 5 m were rare or absent, due to inten- productivity, yet light level is the only variable sive management. The survivorship of those in- that borders on statistical significance (a = dividuals was easily interpolated, as both indi- 0.01). Thus, although this might be the best fit viduals with >2 m fronds and subadults had ex- model, these data rely upon the other variables tremely high survivorship (>95%). Once palms to be significant. When these data were divided develop stems (i.e., subadults and adults) they per management technique, the probability sig- generally have a very low mortality (Kahn and nificances were further muddled (Table 4). de Granville 1992; Sarukh~in, Martinez-Ramos, This table demonstrates that among all the and Pifiero 1984). Of the few individuals in the variables and within all the sites, only light was 2-5 m frond range, the majority had negative statistically significant, at the tagua and cacao growth rates: the longest frond measured after site, using a lower alpha value (a = 0.05). Borg- one year was shorter than the longest frond mea- 176 ECONOMIC BOTANY [VOL. 52

TABLE 2. GENERAL LINEAR MODEL PARAMETER ESTIMATES FOR ANNUAL LEAF PRODUCTIVITY SORTED BY MANAGEMENT TYPE.

Parameter Significance Standard Parameter estimate probability error Tagua Grove R 2 = 0.7505 Dry 5.9158 0.0087 2.0619 Wet 3.3390 0.0035 1.0267 Age (using Height as a proxy) 0.2049 0.3336 0.2075 Light Level 0.8646 0.1723 0.6138 Tagua and Cacao R 2 = 0.2737 Dry 5.1405 0.0034 1.6085 Wet 6.3786 0.0001 1.1341 Age (using Height as a proxy) 0.5949 0.0133 0.2250 Light Level 0.0721 0.8443 0.3635 "Cleared" Tagua R 2 = 0.2535 Dry 6.6483 0.0013 1.8109 Wet 4.8420 0.0139 1.8173 Age (using Height as a proxy) 0.3693 0.2561 0.3171 Light Level 0.1842 0.7804 0.6530 Statistically significantparameter estimates are italicized(a = 0.01). sured at the initial census, perhaps due to cutting assumption should have little effect on the frac- of fronds and/or measurement errors. Therefore, tion becoming adults. Given the number of palm a rough initial estimate assumed that juveniles studies that indicate that juveniles are more pro- with frond lengths greater than 2 m grow at the ductive than seedlings (Ataroff and Schwarz- same rate as juveniles with 2 m frond lengths. kopf 1992; DeSteven et al. 1987; Olmstead and Because once established, such large, juvenile Alvarez-Buylla 1995; Pinard 1993), this as- palms demonstrate high survivorship (Bernal sumption is a conservative underestimation of n.d.; Olmstead and Alvarez-Buylla 1995), this juvenile growth rates.

TABLE 3. GENERAL LINEAR MODEL PARAMETER ESTIMATES FOR ANNUAL INFRUCTESCENCE PRODUCTIVITY.

Parameter Parameter estimate Significanceprobability Standard error Management Tagua Grove - 1.0886 0.6621 2.4771 Tagua and Cacao -0.3846 0.8720 2.3756 "Cleared" Tagua -2.2245 0.3872 2.5521 Degree of Seasonal Inundation Dry 2.9342 0.1171 1.8429

Wet 0.0000 -- -- Management and Degree of Seasonal Inundation Tagua Grove, Dry -0.1339 0.9628 2.8539 Tagua Grove, Wet 0.0000 -- -- Tagua and Cacao, Dry 0.6330 0.8162 2.7096 Tagua and Cacao, Wet 0.0000 -- -- "Cleared" Tagua, Dry 0.0000 -- -- "Cleared" Tagua, Wet 0.0000 -- -- Age (using Height as a proxy) 0.2065 0.5904 0.3814 Light Level 1.7930 0.0148 0.7122 General linear model parameter estimate table for annual infructescenceproductivity. The equation is fit basing comparisonsagainst the cleared tagua and wet variables, therefore,when the interactionterm of managementand degree of seasonal inundation include either the cleared tagua or wet variable, the mean estimatedoes not differ from that variable alone. The R2 value for the full model fit is 0.5860. Statisticallysignificant parameter estimates are italicized (ct = 0.01). 1998] VELASQUEZ RUNK: VEGETABLE IVORY PALM 177

TABLE 4. GENERAL LINEAR MODEL PARAMETER ESTIMATES FOR ANNUAL INFRUCTESCENCE PRODUCTIVITY SORTED BY MANAGEMENT TYPE.

Parameter Significance Standard Parameter estimate probability error

Tagua Grove R2 = 0.4693 Dry 6.2127 0.4020 7.2159 Wet 5.0919 0.3322 5.0914 Age (using Height as a proxy) -1.0768 0.1942 0.7945 Light Level 2.8296 0.1490 1.8665 Tagua and Cacao R 2 = 0.6478 Dry 0.6074 0.8464 3.0983 Wet - 1.5598 0.4847 2.1945 Age (using Height as a proxy) 0.7359 0.1012 0.4302 Light Level 1.5316 0.0320 0.6688 "Cleared" Tagua R 2 = 0.2227 Dry 2.3318 0.6240 4.6440 Wet - 1.2206 0.8100 4.9754 Age (using Height as a proxy) 1.0359 0.2510 0.8622 Light Level 0.0938 0.9620 1.9340

Statistically significant parameter estimates are italicized (a - 0.01).

The most striking outcome of this analysis is and as a result, more seeds are available for ger- the pure tagua sites exhibit a higher percentage mination and to survive to subadulthood. of individuals surviving and growing to subadult The ages of individuals surviving to the sub- height (Table 5). The percentage of individuals adult class are similar to informal estimates of reaching subadult stage is, on average, an order age at first flowering (Acosta Solis 1994; Borg- of magnitude lower for the tagua and cacao toft Pedersen 1993). There is no apparent trend plots, cleared plots, and the old-growth forest in the number of years to become a subadult plot. Also, the dry tagua grove site had the high- between the wet or dry sites. The time required est percent survival to subadult stage, but also to grow from the subadult class to adult class had zero existing subadults, indicating stand or could not be estimated due to the small sample management changes. In the past, the dense size of the subadults. Interestingly, the time to adult palm canopy may have limited light avail- become a subadult in the pure tagua sites is lon- ability in the understory so that fewer subadults ger than in half of the sites. The distinction be- were established on site. Alternatively, current tween subadults and adults is not very clear as seed extraction may be less than in past years adults often had stems only 0.25-1.0 m taller than subadults. It is suggested that one to five years are necessary to grow from subadult stage

TABLE 5. PALM GROWTH SIMULATION RESULTS. to adulthood based on these data and Borgtoft Pedersen's (1993) data on stem height increment Percentage Mean per year. Further growth information is needed that survive number of and grow years to for larger juveniles, subadults, and in replicate Site to subadult subadult sites to increase the accuracy of these estimates. Tagua Grove, Dry 1.122 10.4 These results demonstrate the need to alter Tagua Grove, Wet 0.538 10.3 current management practices to promote juve- Tagua and Cacao, Dry 0.121 10.5 nile establishment. A management alternative to Tagua and Cacao, Wet 0.014 6.4 a poor seed crop is to cut fewer juveniles. To Cleared Tagua, Dry 0.150 7.6 Cleared Tagua, Wet 0.005 12.8 determine the percent of individuals that will Unmanaged, Dry 0.041 9.6 survive to subadulthood, the fraction of individ- uals that survive from juvenile to subadult must Values for each site are the percentage of individuals surviving and growing from newly germinated seedlings to subadults and the mean be multiplied by the number of palms recruited yeats or age of individuals attaining subadult size. into the juvenile stage class via germination. For 178 ECONOMIC BOTANY [VOL. 52

extraction to be sustainable, the annual number of palms that grow to subadulthood must be greater than annual adult mortality, which equals the fraction of adult mortality multiplied by the number of adults (Table 6). For example, in the dry tagua-dominated site, 159 individuals ger- minated during the one-year course of this study. Multiplying 159 by the percentage which will grow to subadulthood (1.122%) yields 1.78. That is, less than two individuals out of 159 will become subadults. Given adult palm mortality during the course of this study, a high value of annual adult palm mortality is estimated at 3% and a low value at 1% using a 95% confidence interval. Therefore, if 1.78 individuals survive and grow to subadults and 2 adults die on an Z annual basis, the tagua population will remain stable or possibly decrease. II II II II II Iq II E

In this simulation, the percentage of juveniles < which survive and grow to subadult includes Z both female and male juvenile palms; the eco- < nomically valuable female palms are thought to represent only one-half of the percentage of ju- _o veniles that survive and grow to subadulthood. < (Until the palms flower, there is no way to de- AI VI VI Vl AI VI VI termine the sex of an individual, so the female: )- ...1 male ratio is estimated at 1:1.) If the sustaina- o bility assessment considers only the female < z palms, then the values are not sustainable. < Several factors may account for the variation o

in sustainability levels. The small number of in- -1 dividuals surviving to the subadult stage likely < II II II II II II II .= Z results from the practice of clearing undergrowth Z with machetes. In tagua and cacao agroforestry < N plots, the increased mortality and stymied ,d growth of juveniles may result from additional _oo8=8o .~ ,..1 0~0~0~0 ~ shading by cacao trees. A much higher percent- < age of juveniles survive and grow in the tagua- ..= ._ dominated groves than in the unmanaged plot, possibly because light and moisture availability are lower in the old-growth forest. In the wet sites, sustainability levels are noticeably less than in the dry sites. This decreased recruitment into the juvenile size class, may be due to the lower establishment of germinated palms result- ing from greater seasonal flooding (Fig. 5). Giv- en the low sustainability levels, tagua extraction levels should not increase within the Commune. At each site the viable seeds that were uncol-

lected by local extractors were counted. These o data indicate that more seeds are left during the dry season than during the wet season. More seeds may be available during the dry season for E 1998] VELASQUEZ RUNK: VEGETABLE IVORY PALM 179 at least three reasons: 1) most leaf litter falls imals which extractors indicated eat the meso- from the canopy at this time, obscuring the carp are spiny rats, squirrels, opossums, and do- seeds; 2) more infructescences fall; 3) unequal mesticated . Tagua seeds with their me- effort by extractors. Residual seed availability socarp are currently being used as feed for spiny appears to be primarily an artifact of extraction rats (Proechimys spp.) in a local development effort because it does not coincide with the num- project that breeds the rats as a source of food ber of infructescences fallen per site. protein. Also, the tagua mesocarp is used in an- imal and fish traps. In addition to previously- MANAGEMENT reported uses of tagua, extractors indicate that Towards the end of the study, interviews were the male palm inflorescences are burned to keep conducted with the six extractors who harvest at away mosquitoes (Vel~squez Runk 1995). the research sites (the seventh site is unmanaged Four separate observations of tagua collection and therefore unharvested). All extractors indi- were completed within a two week period. Each cated that they did not plant the tagua, but rather, observation was carried out within the confines it grew there naturally. One extractor explained of the respective study plot, and therefore rep- that he protected the tagua trees by culling out resent equal area. Typically, the extractor en- other species. Most extractors said that they tered the plot and walked directly to a palm that cleared undergrowth from the plots twice a year, had fallen fruit at its base. This technique might taking two to twenty-two working days each help explain why a variable number of palms time. The range in time required for clearing is were visited, given that each plot had a different due to a common technique of clearing under- number of recently fallen infructescences. None growth in a 1-2 m radius circle around the base of the collectors visited all the palms in the plot of each palm rather than clearing the entire area. and seeds were not collected from all the palms One extractor also cleared the dead leaves from visited. Once the extractor left the plot, double each palm twice a year. The extractors do not the time was spent looking for any uncollected burn the taguales to kill the undergrowth but in- seeds (Table 7). stead clear it with a machete. All six extractors Perhaps most evident is that the number of claimed to use no other management methods. seeds collected by each extractor varies substan- Many indicated they are aware that it is better tially among sites (Table 7). The site from which to cut fronds for thatch from male palms rather no seeds were collected is a relatively unpro- than female palms, so that infructescence pro- ductive site, so the lack of seeds collected is not ductivity is not decreased. surprising. The weights of the seeds collected Contrary to information presented by Foster did not vary tremendously even if the number (1990), extractors indicated that they knew of no of seeds did. places where tagua went uncollected. This also Once the extractor had finished collecting, ad- was the researcher's impression based on trying ditional seeds were found at the site in all cases, to find such a site for a control when establish- even at a site where an extractor collected no ing the study. The one exception mentioned was seeds. In three instances the number of seeds an uncollected tagual in another town, but an found by doubling the collection time was great- extractor indicated that it was too far away and er than the original number of seeds collected too small to be of consequence. The lack of by the extractor. Moreover, the number of seeds places where tagua is uncollected may reflect the collected by doubling the collection time was increased importance of tagua for local income much lower than the numbers found by system- since Foster's 1990 report. atically inventorying the plot for residual, viable Extractors also noted that several worms and seeds (which were uncollected by extractors) beetles prey upon tagua seeds. Observation in- without time constraints during the ecological dicates that termites may be an additional seed data gathering. The observations of collectors predator, as they sometimes infest entire infruc- combined with data on the uncollected seeds and tescences that are still maturing on the tree (Ve- seedling germination, indicate that seeds remain l~isquez Runk 1995). Animals eat the fleshy me- on the ground post-collection and are not har- socarp of the seed, but not the seed itself. vested during future collection periods. In ad- and may sequester seeds in their burrows dition, although much wildlife has been elimi- (Barfod 1991; Vel~isquez Runk 1995). Other an- nated in the Commune, some animals still are 180 ECONOMIC BOTANY [VOL. 52

TABLE 7. RESULTS OF TAGUA COLLECTOR OBSERVATIONS.

Collector observations Measure 1 2 3 4 No. of Seeds Collected by Extractor 35 0 63 34 Weight of Seeds Collected (in pounds) 5 0 5.5 6 Time Spent Collecting (in minutes) 8 5 4 11 No. of Palms Visited 10 2 2 13 No. of Palms Visited and Collected From 3 0 2 3 Additional Seeds Collected with Double the Time and Systematic Collection by Researcher 40 19 24 43 Additional Seeds Collected: Seeds Collected 40:35 19:0 24:63 43:34 Seeds Collected During Systematic Plot Inventory 69 55 39 192

Observations were completed on four collectors within a two-week period. Observation 1 and 2 is of same collector who is responsible for two sites. The final row are values from the systematic plot inventory of seeds during palm data collection.

present to hide tagua nuts among roots and un- out the possibility of negatively affecting female der logs. It is not uncommon to see a seed with palms' fruit production. A fourth strategy is to tooth marks on it. These factors, combined with maintain other tree species in tagua groves and water dispersion of tagua seeds (along the river local farms in the event of tagua market insta- bottoms), likely maintain a source of new tagua bility. These management strategies are being seedlings. presented to local extractors using an illustrated, These data demonstrate that the current ex- educational brochure. Inherent to these strategies tractor methodology is not an exhaustive one. are the local communities' needs to improve Although tagua has become a significant source their quality of life by maximizing tagua extrac- of income since the late 1980s, it is not being tion while minimizing ecological impacts, keep- harvested completely. Extractors do not scour ing control over their land and resources, mini- the ground looking for tagua, but rather go to mizing expenses, and maintaining their culture. trees with fallen infructescences, in an optimal These management recommendations are based foraging-type strategy. That is, the tagua extract- upon conservative interpretations of data from ors make a rational decision to optimize tagua one year. Based upon these results, additional acquisition while minimizing their time and risk studies have been planned for older juvenile and constraints, leading to their net gain (Smith subadult life stages, male palms, seed germina- 1991). It also is possible that tagua extractors' tion, and palm distribution. The results of these decisions, years ago, to maintain tagua palms further studies are crucial for statistically sound and other useful species while culling out other results from which to make sound management species is an extension of optimal foraging, as decisions. resources are optimized while minimizing time Although non-timber forest products remain a and risk. These results suggest several management potentially sustainable means of conserving for- strategies that might better foster sustainable est ecosystems while providing local income, tagua extraction. The first strategy is to maintain greater emphasis must be given to the analysis some seeds and seedlings in sunlit areas where of extraction effects on productivity and sustain- the extractors would like new adult palms, es- ability of the species in question. By employing pecially in areas that flood. The intent is to foster ecological studies, local residents and research- recruitment, particularly in those flooded areas ers can better ensure that both development and where seedlings are difficult to establish. The conservation objectives are being met in non- second strategy is to protect some of the sub- timber forest product extraction efforts. In this adult palms from clearing, fostering the produc- case study of tagua extraction, applying these tive, adult population into the future. The third ecological data to management strategies strategy is to leave at least several male palms strengthens the culture, forest conservation, and in each tagual, so that fruit sets. It also helps standard of living of the residents of the Comuna assure that fronds are available for thatch with- Rio Santiago-Cayapas. 1998] VEL,~SQUEZ RUNK: VEGETABLE IVORY PALM 181

ACKNOWLEDGMENTS October 1994. Unpublished report for Conservation Many thanksto the people of Comuna Rio Santiago-Cayapasfor their International, Washington, DC, USA. assistance and cooperation, especially Antonio Arroyo and Andr6s Arce Godoy, R. A., and K. S. Bawa. 1993. The economic for their work in the field. I appreciated the field support of the Ecua- value and sustainable harvest of plants and animals dorian Foundationfor Socio-EnvironmentalTraining, Research, and De- form the tropical forest: assumptions, hypotheses, velopment (CIDESA) and ConservationInternational. Grateful thanksto John Terborgh, Ram Oren, Lynn Maguire, Tom Philippi, Lisa Moore, and methods. Economic Botany 47:215-219. Henrik Borgtoft Pedersen, and John Tuxill for technical and statistical Hall, P., and K. Bawa. 1993. Methods to assess the advise. Financial support was provided by the SustainableUse of Bio- impact of extraction of non-timber tropical forest logical Resources consortium,The Tinker Foundation, and the Student products on plant populations. Economic Botany 47: International DiscussionGroup of Duke University'sSchool of the En- vironment. 234-247. Hardner, J. 1995. Unpublished draft report on house- hold economics of tagua for Conservation Interna- LITERATURE CITED tional, Washington, DC, USA. Acosta Solis, M. 1948. Tagua or vegetable ivory--a Kahn, F., and J. de Granville. 1992. Palms in forest forest product of Ecuador. Economic Botany 1:46- ecosystems of Amazonia. Springer-Verlag Berlin 57. Heidelberg, New York, NY, USA. 91944. La tagua. Publicaciones Cientfficas, Qui- Koziol, M. J., and H. Borgtoft Pedersen. 1993. Phy- to, Ecuador. telephas aequatorialis in human and animal nutri- Ataroff, M., and T. Schwarzkopf. 1992. Leaf produc- tion. Economic Botany 47:401-407. tion, reproductive patterns, field germination, and Myers, N. 1990. The biodiversity challenge: expanded seedling survival in Charnaedora bartlingiana, a di- hot-spot analysis. The Environmentalist 10:243-256. oecious understory palm. Oecologia 92:250-256. --. 1988. Threatened biotas: "hot spots" in tropi- Barfod, A. S. 1991. A monographic study of the sub- cal forests. The Environmentalist 8:187-208. family Phytelephantoideae (Arecaceae). Opera Bo- Nepstad, D. C. 1992. Conclusions and recommenda- tanica 105:1-73. tions: the challenge of non-timber forest product ex- 91989. The rise and fall of vegetable ivory. Prin- traction. Pages 143-146 in D. C. Nepstad and S. cipes 33:181-190. Schwartzman, eds., Non-timber products from trop- , B. Bergman, and H. Borgtoft Pedersen. ical forests: evaluation of a conservation and devel- 1990. The vegetable ivory industry: surviving and opment strategy. New York Botanical Gardens, doing well in Ecuador. Economic Botany 44:293- Bronx, NY, USA. 300. OAS (Organization of American States). 1979. Mapa Bernal, R. n.d. Demography and impact of seed har- de zonas de vida. Organization of American States, vesting on populations of the vegetable ivory palm Washington, DC, USA. Phytelephas seemannii in . In preparation. Odetola, J. A. 1987. Studies on seed dormancy, viabil- Borgtoft Pedersen, H. 1993. Ivory nuts, fruits, and ity, and germination in ornamental palms. Principes thatch: Use and management of Phytelephas aequa- 31:24-30. torialis (Palmae) in Ecuador. Part 3d in Extractivism Olmstead, I., and E. R. Alvarez-Buylla. 1995. Sus- in Ecuador with special emphasis on management tainability harvesting of tropical trees: demography and economic exploitation of native palms. PhD dis- and matrix models of two palm species in . sertation, Institute of Biology, University of Aarhus, Ecological Application 5:484-500. Denmark. Oyama, K. 1990. Variation in growth and reproduction Coles-Ritchie, M. 1993. Preliminary report on tagua in the neotropical dioecious palm Chamaedorea te- industry surveys, report for Fundaci6n IDEA, Quito, pefilote. Journal of Ecology 78:648-663. Ecuador. Panayotou, T., and P. S. Ashton. 1992. Not by timber DeSteven, D., D. M. Windsor, F. E. Putz, and B. de alone: economics and ecology for sustaining tropical Le6n. 1987. Vegetative and reproductive phenolo- forests. Island Press, Washington, DC, USA. gies of a palm assemblage in . Biotropica 19: Peters, C. M. 1996. The ecology and management of 342-356. non-timber forest resources. The World Bank, Wash- Enright, N. J., and A. D. Watson. 1992. Population ington, DC, USA. dynamics of the nikau palm, Rhopalostylis sapida --. 1994. Sustainable harvest of non-timber plant (Wendl. et Drude), in a temperate forest remnant resources in tropical moist forest: an ecological near Aukland, New Zealand. New Zealand Journal primer. The Biodiversity Support Program, Washing- of Botany 30:29-43. ton, DC, USA. Foster, R. B. 1990. Report: the forest and Phytelephas --, A. H. Gentry, and R. Mendelsohn. 1989. Val- populations in Comuna Rio Santiago, Esmeraldas, uation of an Amazonian ralnforest. Nature 339:655- Ecuador. Unpublished report for Conservation Inter- 656. national, Washington, DC, USA. Philippi, T., R. K. Peet, and N. Christensen. Age and Frank, R. 1995. Keidanren Grant Report, April 1993- size specific demography of tree seedlings in a Pied- 182 ECONOMIC BOTANY [VOL. 52

mont forest. Botany Department, Duke University, Sarukhfin, J., M. Martinez-Ramos, and D. Pifiero. NC, USA. In preparation. 1984. The analysis of demographic variability at the Pinard, M. 1993. Impacts of stem harvesting on pop- individual level and its population consequences. ulations of 1. deltoideae (Palmae) in an extractive Pages 83-106 in R. Dirzo and J. Sarukh~, eds., Per- reserve in Acre, . Biotropica 25:2-14. spectives on plant population ecology. 1984. Sinauer Pinard, M. A., and F. E. Putz. 1992. Population matrix Associates, Inc., Sunderland, MA, USA. models and palm resource management. Bulletin de Smith, E. A. 1991. Inujjuamiut foraging strategies: l'Institut Franqais d'l~tudes Andines 21:637-649. evolutionary ecology of an Arctic econo- Pifiero, D., M. Martinez-Ramos, and J. Sarukhfin. my. Walter de Gruyter, Inc., NY, USA. 1984. A population model of Astrocaryum mexican- Sullivan, S., T. L. Konstant, and A. B. Cunningham. um and a sensitivity analysis of its finite rate of in- 1995. The impact of utilization of palm products crease. Journal of Ecology 72:977-991. on the population structure of the vegetable ivory Pifiero, D., and J. Sarukhfin. 1982. Reproductive be- palm (Hyphaene petersiana, Arecaceae) in north- haviour and its individual variability in a tropical central Namibia. Economic Botany 49:357-370. palm, Astrocaryum mexicanum. Journal of Ecology Vehisquez Runk, J. 1995. Integrating conservation 70:461-472. and development: ecological impacts of tagua nut Redford, K. H., and C. Padoeh. 1992. Conservation extraction in Comuna Rio Santiago-Cayapas, Ec- of neotropical rainforests. Columbia University uador. Thesis. School of the Environment, Duke Press, NY, USA. University, NC, USA.

BOOK REVIEW

Diversity and Classification of Flowering Plants. At- but rather 'one of the focal points of biology' (Huxley men Takhtajan. 1997. Columbia University Press, 1942), a synthetic interdisciplinary science that crowns 562 W. ll3th Street, New York, NY 10027. x + the whole edifice of biology." I wish this wide accep- 643 pp. (hardcover). $95.00. ISBN 0-231-10098-1. tance was more real than imagined. From this point he goes on to discuss his views of Takhtajan has given us an updated view of his in- important topics. Included, are monophyly, compara- terpretation of the evolutionary relationships of the tive character analysis, the significance of primitive flowering plants. This book, devoted "To the memory characters (he still uses that judgmental word, although of Arthur Cronquist (1919-1992)," is a fitting dedi- he equates it with plesiomorphic), convergence and cation that surely would have made Cronquist smile. parallelism, weighting of taxonomic characters and During the years that these two collaborated, it became heterobathmy (differences in grades of characters with- less clear who was the originator of some of their in the same taxon), the Linnean hierarchy, and a sum- ideas; their classification schemes evolved to resemble mary of the recent "successive approximation" each other more and more. (phrase from Hull 1967) of a natural system of clas- This book has an introduction, an outline of his sification. While his comments make one wish for a scheme, and then segments on the various parts of the complete book on these topics, they are merely a lucid Magnoliophyta that Takhtajan perceives. After intro- summary here. ducing the Phylum (sic) Magnoliophyta, that he Takhtajan says that many will accuse him of being equates with the formal name Division within the text, a "splitter." That is as true now as it was in 1942. he breaks out the Classes Magnoliopsida and Liliop- Still, he emphatically defends not splitting where there sida. Next he discusses the Subclasses within each is no good (at least to him) reason. For example, he Class. Each taxon is followed by a list of pertinent finds it absurd to split the Asteraceae, Fabaceae, Ro- references. Along with the index, this results in a mas- saceae, Poaceae and Orchidaceae. Many of us agree sive volume packed with literature, descriptions, and with those points even if we do not with other cases. interpretations. Everyone who specializes in a family will likely find This book is the culmination of studies that began points where they disagree with the author. However, before 1942--thus, it was over 55 years in its creation. he has again done an admirable and boundlessly useful In my view, the Introduction is one of the best parts job in compiling this opus and making his views avail- of this book. Takhtajan begins by slamming the nar- able. For the size of this book, it is comparatively row-minded, outdated, egocentric individuals (my de- cheap at today's prices. At least make sure your library scriptors, not his) in other disciplines with this delight- has a copy. You will find yourself using it often. ful, if too long, sentence. "'It is now widely accepted DANIEL E AUSTIN that systematics is neither just a "great catalogue," or DEPARTMENT OF BIOLOGICALSCIENCES an enumeration of names and descriptions, nor simply FLORIDA ATLANTICUNIVERSITY an identification service (which is desperately needed) BOCA RATON, FL 33431