Size Variations of Flowering Characters in Arum Maculatum (Araceae)
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M. CHARTIER AND M. GIBERNAU, 2009 153 Size Variations of Flowering Characters in Arum maculatum (Araceae) Marion Chartier and Marc Gibernau* Universite Paul Sabatier ~boratoire d'Evolution & Diversite Biologique (UMR 5174) Bat.4R3·B2 31062 Toulouse cedex 9, France ABSTRACT KEYWORDS For entomophilous species, the size Size-advantage model, resources alloca advantage model predicts a bias to the tion, flower number, floral sex-ratio. female function with resource increase. This model has been shown to be ex INfRODUCTION pressed on Arum italicum Mill. (Araceae), Proposed by Ghiselin 0969, quoted in whose bigger or latest inflorescences are Lloyd & Bawa, 1984), the size-advantage female-biased. Here, we characterise the reproductive-trait variations of Arum ma model postulates a sex change when an increase in body size is related to differen culatum 1., a closely related species that is tial abilities to produce or sire offspring. For smaller and produces a single inflorescence per year. entomophilous plant species, such as Aroids, the size-advantage model predicts By measuring the different floral parts a female-biased gender expression with and counting the flower number of differ size increase (Klinkhamer et al., 1997). This ently-sized inflorescences, we showed that is expressed by a proportionally bigger (1) for A. maculatum, 34% of spadix size is increase of the female function than the allocated to the appendix, 12% for female male function in bigger individuals. This flowers and 7% for male flowers. Due to its topic has been mainly studied in Araceae size range, A. maculatum could in a on Arisaema species presenting an extreme manner be considered as a "small" A. reproductive variation with a gender mod italicum. (2) As for A. italicum, the ification according to size (Gusman & appendix size is the longest part of the Gusman, 2003). Apart from Arisaema, this inflorescence, and increases proportionally question has been mainly studied on Arum more than the male and female parts size italicum. It has been shown in this species with plant vigour. This is not surprising, as that there is a disproportionate allocation to the appendix is an essential attracting the appendix versus the floral parts, with a organ for pollinators. (3) There is no length representing 44% of the spadix size correlation in A. maculatum between the (Mendez, 1998; Gibernau & Albre, 2008), flowers number and the other floral char and a mass representing 29% of the total acters, and no bias for female part size in inflorescence weight (Mendez, 2001). Such bigger inflorescences. Thus, contrary to A. results are not surprising as the appendix is italicum, there is no expression of the size the inflorescence's attractive organ, pro advantage model for A. maculatum, whose ducing heat and volatile organic com range of sizes might be too small. Another pounds (Bermadinger-Stabentheiner & Sta explanation is selection for a high number bentheiner, 1995; Kite et al., 1998; Diaz & of male gametes that may counterbalance Kite, 2002). In general, fertile parts and the potential bias for female flowers. appendix weights increase with plant vigour, which is representative of an * e-mail: [email protected] increase of the reproductive effort when 154 AROIDEANA, Vol. 32 resources become more abundant (Men one diptera species, the midge Psychoda dez, 1998; Gibernau & Albre, 2008). In phalaenoides whereas A. italicum is polli Spanish A. italicum, Mendez (2001) nated by several diptera species (Lack & showed that the mass allocated to male Diaz, 1991; Diaz & Kite, 2002; Albre et al., flowers increases proportionally more than 2003). In a way, we can consider from a for female flowers with increase in the pollination point of view that A. macula inflorescence's weight, which is in contra tum may be a "specialist" and A. italicum diction with the size-advantage model rather an "opportunist" species. Another prediction. However, the same author difference is that mature plants of Arum (1998) found no significant difference maculatum generally produce only one between the proportional increase of male inflorescence whereas mature Arum itali and female flower number with spadix size cum produce an average of two inflores increase. On the contrary, in the South of cences (Albre & Gibernau, 2008). France, Gibernau & Albre (2008) found that The aim of this paper is to document the female flower number increased pro the reproductive characters of Arum portionally more than male flowers with maculatum in three different French spadix size increase in A. italicum, show populations, and to characterize its allo ing that a bias to female parts can exist, cation strategy. Does A. maculatum, which is in accordance to the size-advan which produces a single inflorescence tage model. Thus, A. italicum floral sex per year, express the size-advantage ratio, which is male-biased (mean = .74), model for big inflorescences? Does its decreases with spadix size increase (Giber higher pollinator-specificity result in a nau & Albre, 2008). Furthermore, the sex different resource allocation in terms of ratio from the last inflorescences (which are the relative appendix size or the numbers smaller) tends to be female-biased (Men of fertile flowers? dez, 1998) which is also in accordance with the sized-advantage model, as the more MATERIALS AND METHODS vigorous plants are the ones able to produce up to three inflorescences per 34 inflorescences of Arum maculatum L. year. Then, when considering the floral were collected during the spring of 2008 in parts size and mass variations, the resourc 3 populations situated in Bagneres-de es allocation in A. italicum seems to be Bigorre (Midi-Pyrenees, France), La Louba different for female parts, for which an tiere (Languedoc-RoussilIon, France) and increase in size would rather be due to an Smarves (Poitou-Charentes, France). The increase in flower size, than it is for male inflorescences were collected before the parts, for which an increase in size may release of pollen and preserved in 70% likely be due to an increase of flower alcohol. number (Mendez, 2001; Gibernau & Albre, The lengths of the different parts of the 2008). spadix, i.e. fertile male and female zones Here, we study the floral variations and and appendix, were measured with a the expression of the size-advantage model digital calliper. Moreover, male and female of Arum maculatum (L.). Its flowering flowers per inflorescence were counted on cycle is protogynous; female flowers being 13 inflorescences. receptive before stamens release the pol Data analyses consisting of linear regres len. Arum maculatum is pollinated by sions, Shapiro tests, Bartlett tests, ANCOVA deceived diptera attracted by the floral (for differences between regression odour mimicking the odour of their ovi slopes), as well as a Kruskal-Wallis test positing site (sapromyophily) (Gibernau et followed by a multiple comparison test (for al., 2004). The deceiving odour is pro inter populations differences of floral part duced by a sterile organ situated at the apex sizes) were performed using R 2.5.1 (2007) of the inflorescence called the appendix. software. Codes are available from M. Arum maculatum is mainly pollinated by Chartier. M. CHARTIER AND M. GIBERNAU, 2009 155 RESULTS '3~-;'C; g ..... rv") ..... Morphology ~~t ~C\j""'; The mean sizes of the different parts of a ~ tI tI +1 ~=d-1 ,-... C"cc the inflorescences are shown in Table 1. ~C\j<V)" r <'1') ..... The appendix appears to be the most ~ ~ II S',u~z important part of the inflorescence, rep I'-' fIl resenting 33.8 :!: 5.4% of the total spadix ~ i21 length. The fertile parts are shorter than =U \0 '<I'· ('(').. 00 '<I' the appendix representing 11.9 :!: 1.9% of 5 ~1 ~ .-t I'.-t . rIJ 1 U I'-' the total spadix length for the female ,-,i-sa: U e tI +1 tI tI ('(')'<1'\0\0 zone and 7.2 :!: .9% for the male zone rl '§ "d 0 "NOO...r (Table 1). Moreover, the female zone 13 ; = \0 N appears to be 1.6 times longer than the 1 d.!3U~ U =~ male zone. The inflorescences from the ~ ~ 2 population of Bagneres-de-Bigorre ap . 00 1rI.......... " . pear to be longer than in the two other "d~ a1Q Q~ populations (Table 1). Zone lengths are ; j ~ : ~ +1 +1 tI significantly different. between popula 8oq~~ S' 5.,s ;: ..... 1rIrv") ..... " tions only for the male part (X22• 4= 7.07, 3 rIJ 8. ~ ~ df = 2, P = .03), Bagnere-de-Bigorre +1 ~ fIl .-t .d Q having significantly longer male zone ~ ; ~ !! '-' length than Smarves (P < .05). U~ U 0 ~ ..........· N'<I'.-t The mean numbers of male and female ,g ~ froB .-t\O.-t.-t 1'-' fIlrf')U"d flowers are shown in Table 2. Here again, +1 +1 +1 tI in the population of Bagneres-de-Bigorre, ~.s-s ; I' N \0 If\ v\"oov\ inflorescences seems to have more flowers I'N ~fIl: than in the two other populations. But f~· 1 ~tfll inflorescence numbers per population are too low to test for any difference between .s]5i C'o.!C populations. ~C\j~ ~ ~!~ ,-... ~ +1 +1 tI Intra-plant relationships C'<;t<CC'o.! ~2;,s ~cC\j" ~ rv") ..... The three measured floral zones sizes eQl~ '-' were positively correlated with the spadix ~~=ar ,~ size, which is logical due to the autocorre i!Jl I i 1). 00 I' If\ lation of these traits (Fig. The appendix Q!,sd i • .. '<I' length increases proportionally more ~ ~ 1I' If\.-t • t.s Q 3 '-' tI +1 tI +1 (slope = .5) than the male (slope = .06) a U ~ ~ ~ Q, .!3 \O.-tNo\ and female zone lengths (slope= .07). is "d. oo"",;oo...r fIl ; rl \0 N Surprisingly, male and female zone size i is U ~ 1 slopes are not Significantly different (F1•32 f ts 'i = .11, P= .75) i.e.