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Ecological Aspects of the Interaction Between Chamaedorea Tepejilote, A

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Ecological Aspects of the Interaction Between Chamaedorea Tepejilote, A B6 PRINCIPES [Vor. 35 Principes,35(2), 1991, pp. 86 93 EcologicalAspects of the Interaction between Chamaedoreatepejilote, a DioeciousPalm and Calyptocephalamarginipennis, a Herbivorous Beetle, in a Mexican Rain Forest K. Oveue.* ANDR. DIRZo Centro d.eEcol.osia, WAM, Apartado Postal 7O-275, M6xico, 04510, D.F. Assrnecr ers have made studies regarding intraspe- cific differencesin herbivory in the context Liebm. ex Mart., an Chamaedorea tepejilote of polymorphic plant systemsin temperate abundant dioeciouspalm in the lowland rain forest of Los Tuxtlas (SoutheastMexico), typically bears evi- zones(e.g., Jones 1966, Cates1975, Dirzo dence of leaf damage by Calyptocephala margini- and Harper I982a). Dioecy constitutes pennis Bohem. (Chrysomelidae).The beetles mark- one of the most common polymorphic sys- edly prefer this palm in comparison to other plants tems, particularly in some Neotropical on this site, including other sympatric speciesin the same genus. Additionally, beetlesshowed some mar- communities(Croat 1979, Bawa and Opler ginal preference for the foliage of female plants under 1975, Bawa 1980, Flores and Schemske experimental and field conditions. Sex-related phy- 1984, Bawa et al. 1985). Suggestionin tochemical attributes of the palm, such as secondary the literature of sex-baseddifferences in and nutritional characteristicsof the foli- comoounds herbivory include male inflorescencessuf- age, did not differ markedly between sexes and did not seemto be the proximal causesof the inter-sexual fering higher levels of predation than female differences in herbivory. Thus, the hypothesis that inflorescences(Fryxell and Lulcefahr1967, differencesin resourceallocation to defenseand nutri Bawa and Opler 1978) and casesin which tional quality, resulting from sex-related differential herbivores preferentially feed either on allocation to reproduction, is not supported by this study. This investigation suggests that plant-herbi- leavesof female (Danell et al. 1985, Lovett vore interactionsin palms may be as complex as those Doust and Lovett Doust 1985) or male of other flowering plants. (Agren 1987) plants. However, folivory is extremely poorly known for dioecioustrop- Studieson the interactionsbetween her- ical plants, particularly palms. bivores and plants in tropical communities In the lowland rain forest of Los Tuxtlaso have been increasing markedly in recent SoutheastMexico, Charnaedoreatepej ilo- years (Coley 1983, Dirzo 1984, Janzen fe, a dioeciousunderstory palm, normally and Waterman 1984, Marquis 1984, bearsextremely variable levelsof leaf dam- among others). Also, much empirical evi- age and we explored the possibility that dencehas been obtainedwhich can be used this variation might explained, to some for the development of theories of plant- degreeat least, by inter-sexualdifferences. herbivore interactions (seeCrawley 1983, The plants on this site are normally eaten Rosenthaland Janzen 1979, Strong et al. by CaLyptocephala marginipennis, a l9B4). In this expandingfield, somework- chrysomelid beetle which appears to be largely monophagous.We measured the selectivity of this insect for the leaves of * Present address: Department of Botany, Faculty each sex in the field and laboratory. Also, of Science, Kyoto University, Kyoto, 606, Japan. we attempted to correlate the effect of L99ll OYAMA AND DIRZO: HERBIVORYCHAMAEDOREA herbivory with differences in contents of the leaves of male and female plants in the secondary compounds and nutritional field using two methods: a) an instanta- characteristicsof the sexes. neous measurement based on broad cat- egories of damage, and b) estimates of leaf area loss during a 3-month period. For the Materials and Methods first method, leaf damage was scored on an A to D scale (A < 25%; B > 25 < Study Species. The palm genus Cha- 5O%; C > 50 < 75%;D > 75%) n all maedorea includes approximately one the leaves of 48 male and 45 female plants. hundred speciesdistributed through Neo- The leaves of each plant were separated tropical forests from Brazil and Bolivia to into four age categories (I-IV). Contin- Guatemalaand the Central part of Mexico gency analyses (after a heterogeneity test (Standley and Steyermark 1958, Uhl and for the leaves of different age) were carried Dransfield 1987). All the species in the out to assess the distribution of categories genus are dioecious (H. Quero, pers. of damage of the leaves of both sexes. b) comm.). In Los Tuxtlas, Mexico, C. tepe- For the second method, leaf damage was jilote is a common palm that forms dense also scored with a grid of 893 points on a patches in the forest understory where transparent plastic sheet (I0 cm x 5 cm). slight natural disturbanceshave occurred. This grid was placed on the adaxial surface The tallest individuals reach up to 5.0 m of the pinnae in a way that we could count in height. Plants of both sexes bear from the number of visible points coinciding with 3 to 5 leaves.The plants flower from Octo- ruptures of the lamina (as a product of ber through December. Both sexes have herbivore damage). We took care to check yellow green flowers; male flowers tend to all types of damage and differentiated them be slightly smaller (1.5 to 2.5 mm) than from the specific damage produced by C. females(2.5 to 3.5 mm). Single female marginipennis; this was, by far, the most plantscan bear from 5 to 400 fruits (Oyama abundant type of damage (see Oyama 1984). Mortality of adult plants is due l9B4). We then calculated an index of mostly to branches or other debris falling leaf area removed as the quotient of the from the canopy. Tree falls frequently number of points registered (ruptured) to break apical meristems and kill the palms the total number of points of the grid (i.e., (Oyama1987, 1990). 893). Following this second method, we Eight speciesof Calyptocephala have recorded the cumulative damage on the been reported in Neotropical forests from leaves by marking a permanent area of Mexico to Brazil, paralleling the geo- observation (of equal size to the grid) on graphic distribution of the Chamaedorea the last two (youngest) leaves produced by species.The beetles are small (ca. 6 mm the plant. We recorded the damage long) with pale brown elitrae and a dark monthly for three consecutive months. For line at the margin. This insect feeds on this analysis, we chose randomly 15 indi- leavesof C. tepejilote making specificlon- viduals of each sex in three categories of gitudinal ruptures along the leaf veins, height (0 to 2.0 m; 2.0 to 3.5 m; > 3.5 which are easily recognizedin the field. m). Two-way analyses of variance were Study Site. This study was conducted done in function of sex and height (age) in a 600 m2 permanent plot in a lowland of the plants, after arcsin transformation rain forest in the Gulf of Mexico (95"14'- of the data. 95o09'W, lBo34'-18o36'N),in the bio- In a second line of investigation, intact logical reserve Estaci6n de Biologia Trop- pinnae of male and female plants of the ical Los Tuxtlas at 150-650 m altitude. same size and age were bagged together Methods. We recorded herbivory on and exposed to 6 beetles previously starved B8 PRINCIPES [Vor-.35 of m'ale Table 1. Contingency analysis for the number of leaaes of dffirent age _(M) in each category do'ftiage. Values and.female g1 ptanis o/Ciramaedorea tepejilote -of in pJrenthese. b"io* each pair of values are the absolutedeviations from those expected if beetlesdamaged leaves regardless of sex; the sign of the deviation is shownin parentheses after each ,r"G. Chi-tqnare analysesare shown for leaf age, the pooled data of leaves .d@ Category of beetle damage Leaf age Sex . Total X2 I M 90576 6 924 6.96 5 n.s. F 1,045 l5 I 6 r,067 .ll II M 827 46 13 897 t.29 3 n.s. F 929 60 II 14 1,014 il1 M 7rr (+) 72(-) 22(+) 19(-) 424 26.88 <0.001 F 690(-) 147(+) 2r (-) 34 (+) 892 (38) (33) (l) (6) M 470(+) r22 (-) 40 (-) 28 (-) 660 36.76 <0.001 F 314(-) I53(+) 43(+) 56 (+) s66 (48) (26) (5) (t 7) Total 71.89 I2 <0.001 Pooled M 2,e13(-) 247(-) Bl (+) 64 (-) 3,30s 31.07 3 F 2,978(+) 375(+) 76(-) 1r0 (+) 3,700 (6s) (r28) (s) (46) Heterogeneity 40.42 T2 <0.001 for 24 hours. The experiments liverecar- Results ried out in a laboratory arranged for this purpose. We recorded the leaf area con- In the survey of leaf damagescored on iumed for each type (sex) of plant 12 and a visual scale, the heterogeneity test 24 hours after. Each experiment consisted between leaves of different age was sig- : oI six replicates.Statistical analyses (t-tests) nifrcant (X" 40.82:.P < 0.001) and the were done to compare the mean leaf area resultsof eachleaf agewere examinedinde- eaten of male and female plants. All the pendently (Table l). The heterogeneity statisticalanalyses were done following Zar appears to be due to the fact that leaves (1974). in categories I and II did not show sex- A third line of study consistedof a phy- related differenceswhile categoriesIII and tochemical screening of secondary com- IV were the only caseswhere sexesdiffer pounds (alkaloids, tannins, flavonoids, sa- in the levels of damage. Moreover, the ponins and cyanogenic compounds) and leavesof the youngestcategory were mostly nutritional characteristics (protein, fiber, undamaged (905/924 for female and sugar, mineralsand fat contents)as Pos- I,045/L,067 for maleplants) and damage sible factors affecting the intensity of her- gradually increases with leaf age. Thus, bivory. Young and mature leaves of each sex-related differences could only be sex were collected, dried and analyzed in deteetedwhen damagewas high in general. the laboratory (see Dominguez(1979) for For leaves of age categories III and IV, a description of techniquesfor analysesof clearly females were over-representedin secondarycompounds, and Flores(I977) the heaviest category of damage (D) and for a descriptionof techniquesfor assessing in category B, while males were under- nutritional characteristics).
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