Pollination Strategies Research Articles

Schottarum (Schismatoglottideae: Araceae) substantiated based on combined nuclear and plastid DNA sequences

Recent studies on Schismatoglottideae have resulted in the recognition of four new monophyletic genera, the resurrection of two additional genera following clarification of their monophyly, and the publication of many taxonomically novel species. However, generic boundaries among some parts of Schismatoglottideae remain unclear owing to several reasons: (1) more taxa are being revealed through our on-going fieldwork, now expanded to previously unsampled localities on Borneo; (2) established occurrence of a high level of homoplasies among the morphological characteristics hitherto used to delimitate genera; and (3) gene regions used in previous studies contradicted some of current taxonomic placements. Among the unsolved groups from previous studies a clade comprising Schismatoglottis sarikeensis and S. josefii needs further investigation. Therefore, phylogenetic analyses were carried out to investigate the position of these two species using the nuclear region, internal transcribed spacer and combined plastid regions: trnL intron and trnL-F intergenic spacer, coding matK+partial 3′ trnK, intergenic spacer trnH-psbA. A total of 23 accessions representing 16 taxa of Schismatoglottideae and Philonotieae were included in the study. Phylogenetic analyses of a total 4,658 bp combined dataset using parsimony, maximum likelihood, and Bayesian methods revealed that S. sarikeensis and S. josefii do not belong to Schismatoglottis, and therefore are transferred to Schottarum (≡Hottarum sarikeense ≡ Schismatoglottis sarikeense). Flowering mechanism, pollination strategy, and fruitset of S. sarikeense are also presented in the paper.

Self-Deception: A Teleofunctional Approach

This paper aims to offer an alternative to the existing philosophical theories of self-deception. It describes and motivates a teleofunctional theory that models self-deception on the subintentional deceptions perpetrated by non-human organisms. Existing theories of self-deception generate paradoxes, are empirically implausible, or fail to account for the distinction between self-deception and other kinds of motivated irrationality. Deception is not a uniquely human phenomenon: biologists have found that many non-human organisms deceive and are deceived. A close analysis of the pollination strategy of the mirror orchid (Ophrys speculum) provides the basis for an analysis of non-human deception as subintentionally purposive. This teleofunctional analysis is then used as the basis for a theory of self-deception that accounts for its normative and purposive features without defaulting to intentionalism. The teleofunctional theory accounts for distinction between self-deception and phenomena such as wishful thinking in an empirically plausible manner. Three objections to the theory are considered and rejected.

Pollen limitation and fruiting failure related to canopy closure inCalypso bulbosa(Orchidaceae), a northern food-deceptive orchid with a single flower

Natural fruit set is constrained by pollen limitation and fruiting failure, and pollen limitation is expected to be especially severe in deceptive orchids. We performed hand cross-pollinations in ten populations of a food-deceptive orchid, Calypso bulbosa, under sparse and dense canopies in three non-consecutive years. We explored the relationships between natural fruit set, pollen limitation and fruiting failure. Mean natural fruit set over the years was 60%, which is exceptionally high for a deceptive orchid. On average, hand cross-pollination increased fruit set by 23%. Among open-pollinated plants that did not set a fruit, 55.5% were estimated to be pollen limited and 44.5% to be limited by fruiting failure, i.e. inability to set a fruit after pollination. In species with high natural fruit set, hand cross-pollination experiments may not always detect statistically significant pollen limitation. In our case, pollen limitation tended to become significant when the natural fruit set dropped below 60%. Canopy cover had a significant effect on fruiting failure, which was more severe under a dense canopy. Although our results demonstrate pollen limitation in many cases, they also highlight the fact that food deception can be a very effective pollination strategy.

Morphological, ecological and genetic aspects associated with endemism in the Fly Orchid group

The European genus Ophrys (Orchidaceae) is famous for its insect-like floral morphology, an adaptation for a pseudocopulatory pollination strategy involving Hymenoptera males. A large number of endemic Ophrys species have recently been described, especially within the Mediterranean Basin, which is one of the major species diversity hotspots. Subtle morphological variation and specific pollinator dependence are the two main perceptible criteria for describing numerous endemic taxa. However, the degree to which endemics differ genetically remains a challenging question. Additionally, knowledge regarding the factors underlying the emergence of such endemic entities is limited. To achieve new insights regarding speciation processes in Ophrys, we have investigated species boundaries in the Fly Orchid group (Ophrys insectifera sensu lato) by examining morphological, ecological and genetic evidence. Classically, authors have recognized one widespread taxon (O. insectifera) and two endemics (O. aymoninii from France and O. subinsectifera from Spain). Our research has identified clear morphological and ecological factors segregating among these taxa; however, genetic differences were more ambiguous. Insights from cpDNA sequencing and amplified fragment length polymorphisms genotyping indicated a recent diversification in the three extant Fly Orchid species, which may have been further obscured by active migration and admixture across the European continent. Our genetic results still indicate weak but noticeable phylogeographic clustering that partially correlates with the described species. Particularly, we report several isolated haplotypes and genetic clusters in central and southeastern Europe. With regard to the morphological, ecological and genetic aspects, we discuss the endemism status within the Fly Orchid group from evolutionary, taxonomical and conservation perspectives.

Pre-adaptations and the evolution of pollination by sexual deception: Cope's rule of specialization revisited.

Pollination by sexual deception is arguably one of the most unusual liaisons linking plants and insects, and perhaps the most illustrative example of extreme floral specialization in angiosperms. While considerable progress has been made in understanding the floral traits involved in sexual deception, less is known about how this remarkable mimicry system might have arisen, the role of pre-adaptations in promoting its evolution and its extent as a pollination mechanism outside the few groups of plants (primarily orchids) where it has been described to date. In the Euro-Mediterranean region, pollination by sexual deception is traditionally considered to be the hallmark of the orchid genus Ophrys. Here, we introduce two new cases outside of Ophrys, in plant groups dominated by generalized, shelter-mimicking species. On the basis of phylogenetic reconstructions of ancestral pollination strategies, we provide evidence for independent and bidirectional evolutionary transitions between generalized (shelter mimicry) and specialized (sexual deception) pollination strategies in three groups of flowering plants, and suggest that pseudocopulation has evolved from pre-adaptations (floral colours, shapes and odour bouquets) that selectively attract male pollinators through shelter mimicry. These findings, along with comparative analyses of floral traits (colours and scents), shed light on particular phenotypic changes that might have fuelled the parallel evolution of these extraordinary pollination strategies. Collectively, our results provide the first substantive insights into how pollination sexual deception might have evolved in the Euro-Mediterranean region, and demonstrate that even the most extreme cases of pollinator specialization can reverse to more generalized interactions, breaking 'Cope's rule of specialization'.

The cowl does not make the monk: scarab beetle pollination of the Neotropical aroidTaccarum ulei(Araceae: Spathicarpeae)

Taccarum ulei (Araceae, Spathicarpeae) is a seasonal geophytic aroid, native to north-eastern Brazil, that flowers during two months of the rainy season. Patterns of floral thermogenesis, pollination biology, and floral traits associated with pollination syndromes were studied and compared with those of other Araceae. Two species of cyclocephaline scarabs (Scarabaeidae, Cyclocephalini) were recognized as effective pollinators: Cyclocephala celata and Cyclocephala cearae. Larvae of an unidentified species of fruit fly (Melanoloma spp., Richardiidae, Diptera) were also frequently observed in inflorescences at various maturation stages, feeding on the connectives of male florets and fruits, and thus lowering the reproductive success of individual plants. Beetles were attracted by odoriferous inflorescences in the early evening of the first day of anthesis, during the female phase. The emission of attractive volatiles was coupled with intense thermogenic activity in the entire spadix, unlike other aroids in which only certain zones of the spadix heat up. Pollen release, which marks the beginning of the male phase on the subsequent evening, was not related to floral thermogenesis. Comparative multivariate analysis of the floral traits of T. ulei points to a beetle-pollinated aroid, although some of the observed traits of the species are not common to other taxa sharing this pollination strategy. Such incongruence might be explained by the evolutionary history of the tribe Spathicarpeae and potential pollinator shifts. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ••, ••–••.

Orchis × colemanii hybridization: Molecular and morphological evidence, seed set success, and evolutionary importance

Despite highly specialized pollination strategies, hybridization is a common phenomenon among Mediterranean deceptive orchids. Food-deceptive species sire a progeny of F1 unfertile plants, which work as a late post-zygotic barrier. Conversely, when pre-zygotic barriers of sexually deceptive (Ophrys) species are absent, the hybrids are fertile and an extensive introgression may occur. Here, we have performed molecular analysis and hand pollination treatments to characterize a hybrid zone of two food-deceptive species, O. mascula and O pauciflora. Hybrids (called O.×colemanii) have shown different amounts of parental nrDNA, strongly supporting that they are F2 and/or successive hybrid generations. Comparable high levels of reproductive success have been detected in natural conditions and in experimental crosses suggesting the absence of effective reproductive barriers either between hybrids, either between hybrids and parental species. In light of ecological and distributional features of O.×colemanii across its distribution range, we hypothesize that these populations have originated by secondary contact in the periglacial belt of Apennines. Moreover, the rare and localized O. pauciflora could benefit a genetic enrichment by hybridizing with a widespread related species. O.×colemanii is not a dead end population, but may have a role as potential reserve of adaptive variability and is an unusual stage along the speciation process.

Comparative analysis of floral scents in four sympatric species of Serapias L. (Orchidaceae): clues on their pollination strategies

Orchid species of Mediterranean genus Serapias often live in sympatry, exhibit similar floral morphology, bloom in the same period and share the same pollinators. Previous studies on Serapias species have ascertained that reproductive isolation is based on pre-pollination barriers, that secretory cells and trichomes are typically distributed on the floral labellum and that flowers produce aliphatic compounds. In this study we compare the floral scent composition of four widespread, co-occurring Serapias species, namely Serapias lingua, Serapias parviflora, Serapias vomeracea and Serapias cordigera. Our goals are to assess if differences in floral scent may act as interspecific pre-pollination barriers and if these olfactory signals may be involved in the pollination strategy of Serapias. We find that all the selected species produce C20–C29 alkanes and alkenes and, in addition, have detected the presence in S. cordigera of large amounts of oleate and stearate ethyl ester. Our findings help to clarify that the sympatric Serapias species have slightly different floral scent signatures that may account for their relevant role as pre-pollination barriers. Therefore, the pollination strategy of Serapias relies not only on the tubular shape of their floral corolla but also on the production of olfactory signals that may lure potential pollinators and even assure a sufficient degree of pollinator fidelity.

Divergence on floral traits and vertebrate pollinators of two endemicEncholiriumbromeliads

Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self-compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E.vogelii. Pollination treatments demonstrated that E.heloisae and E.vogelii were partially self-compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E.vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.

Distribution of calcium oxalate crystals in floral organs of Araceae in relation to pollination strategy

Many flowers are pollinated by potentially hungry insects, yet flowers also contain gametes and embryos which must be protected from predation. Microscopic calcium oxalate crystals in plant tissues have been proposed to protect against herbivory. Aroids, which have an unusual diversity of such crystals, also exhibit diverse pollination strategies. Many species have pollinators that do not feed while visiting the flowers, while other species, especially those pollinated by beetles, offer sterile staminodia as food rewards. We examined flowers of 21 aroid species with various pollination strategies to test the hypothesis that crystals protect vital gametes and embryos while allowing consumption of food bribes. Aroid inflorescences collected from the field or from greenhouse material were sectioned, cleared, and examined by bright field and polarization microscopy. All species examined, regardless of pollination strategy, arrayed crystals around unshed pollen and ovules. Less vital tissues, such as odoriferous appendages, had few crystals. Staminodia offered as food to beetle pollinators, however, differed greatly between species in their crystal contents. Some had minimal crystals; some had crystals in patterns suggesting they limit beetle feeding; still others had abundant crystals in no obvious pattern. The results are consistent with crystals protecting against insect predation of gametes and embryos. However, the role of crystals in food-bribe staminodia is unclear. They may limit and direct feeding by beetles in some species, while in others they might have no protective role.

Labellum transcriptome reveals alkene biosynthetic genes involved in orchid sexual deception and pollination-induced senescence

One of the most remarkable pollination strategy in orchids biology is pollination by sexual deception, in which the modified petal labellum lures pollinators by mimicking the chemical (e.g. sex pheromones), visual (e.g. colour and shape/size) and tactile (e.g. labellum trichomes) cues of the receptive female insect species. The present study aimed to characterize the transcriptional changes occurring after pollination in the labellum of a sexually deceptive orchid (Ophrys fusca Link) in order to identify genes involved on signals responsible for pollinator attraction, the major goal of floral tissues. Novel information on alterations in the orchid petal labellum gene expression occurring after pollination demonstrates a reduction in the expression of alkene biosynthetic genes using O. fusca Link as the species under study. Petal labellum transcriptional analysis revealed downregulation of transcripts involved in both pigment machinery and scent compounds, acting as visual and olfactory cues, respectively, important in sexual mimicry. Regulation of petal labellum senescence was revealed by transcripts related to macromolecules breakdown, protein synthesis and remobilization of nutrients.

Innate Adjustment of Visitation Behavior to Rewarding and Reward‐Minimized Petunia axillaris (Solanacea) Plants by Hawkmoth Manduca sexta (Sphingidae)

AbstractPlant–pollinator interactions offer an excellent system to study the stability of mutualistic interactions. While nectar production requires resources and a reduction could in principle benefit plant fitness, only few angiosperms lack nectar, and thus cheat from a pollinator's perspective. Cheating behavior may be scarce because of pollinator foraging behaviors that select for nectariferous plants. Shorter inspection duration, interaction with fewer flowers, or even complete avoidance of plants with low/no nectar may reduce the fitness of cheating plants. The effectiveness of pollinator strategies may depend on how they are implemented. Innate strategies would invariably decrease the fitness of a cheating plant, while learned responses allow cheaters to exploit naïve pollinators. Here, we studied the foraging strategies of the hawkmoth Manduca sexta during interactions with nectariferous and reward‐minimized Petunia axillaris. We found that neither naïve nor experienced hawkmoths discriminated a priori between rewarding and nectar‐less plants. However, naïve hawkmoths displayed reduced probing time per flower and number of flowers visited on reward‐minimized plants during the first trial, without showing further improved discrimination with experience. In conclusion, the foraging decision rules of hawkmoths that may reduce the fitness of reward‐minimized plants appear to be innate, with little scope for additional learning.

Floral and vegetative cues in oil-secreting and non-oil-secreting Lysimachia species.

Unrelated plants pollinated by the same group or guild of animals typically evolve similar floral cues due to pollinator-mediated selection. Related plant species, however, may possess similar cues either as a result of pollinator-mediated selection or as a result of sharing a common ancestor that possessed the same cues or traits. In this study, visual and olfactory floral cues in Lysimachia species exhibiting different pollination strategies were analysed and compared, and the importance of pollinators and phylogeny on the evolution of these floral cues was determined. For comparison, cues of vegetative material were examined where pollinator selection would not be expected. Floral and vegetative scents and colours in floral oil- and non-floral oil-secreting Lysimachia species were studied by chemical and spectrophotometric analyses, respectively, compared between oil- and non-oil-secreting species, and analysed by phylogenetically controlled methods. Vegetative and floral scent was species specific, and variability in floral but not vegetative scent was lower in oil compared with non-oil species. Overall, oil species did not differ in their floral or vegetative scent from non-oil species. However, a correlation was found between oil secretion and six floral scent constituents specific to oil species, whereas the presence of four other floral compounds can be explained by phylogeny. Four of the five analysed oil species had bee-green flowers and the pattern of occurrence of this colour correlated with oil secretion. Non-oil species had different floral colours. The colour of leaves was similar among all species studied. Evidence was found for correlated evolution between secretion of floral oils and floral but not vegetative visual and olfactory cues. The cues correlating with oil secretion were probably selected by Macropis bees, the specialized pollinators of oil-secreting Lysimachia species, and may have evolved in order to attract these bees.

Did Pollination Shifts Drive Diversification in Southern African Gladiolus? Evaluating the Model of Pollinator-Driven Speciation

The pollinator-driven ecological speciation model has frequently been invoked to explain plant richness in biodiversity hotspots. Here, by focusing on Gladiolus (260 species), a flagship example of a clade with diverse pollination biology, we test the hypothesis that high species diversity in southern Africa, one of the world's most floristically rich regions, has primarily been driven by ecological shifts in pollination systems. We use phylogenetic methods to estimate rates of transition between the seven highly specialized pollination strategies in Gladiolus. We find that pollination systems have evolved multiple times and that some pollination strategies arose by a variety of evolutionary pathways. Pollination shifts account for up to one-third of all lineage splitting events in the genus, providing partial support for the pollinator-driven speciation model. Transitions from the ancestral pollination mode to derived systems have also resulted in increased rates of diversification, suggesting that certain pollination systems may speed up speciation processes, independently of pollination shifts per se. This study suggests that frequent pollination shifts have played a role in driving high phenotypic and species diversity but indicates that additional factors need to be invoked to account for the spectacular diversification in southern African Gladiolus.

Floral scent in Annonaceae

Many species of Annonaceae are known for their distinctive, penetrating floral aromas. Numerous pollination studies have documented floral scents which probably play a key role in specialized pollination strategies. In particular, floral scents appear to play crucial roles in deceptive pollination strategies, contributing to floral mimicry of ripe or decaying fruits, fungi and, potentially, carrion or faeces. Occasionally, floral scent may advertise genuine floral rewards, as is the case for two species of Unonopsis pollinated by male euglossine bees. To date, ten studies have chemically characterized floral scent for 24 species representing 11 genera of Annonaceae. In this review, I discuss the chemical composition and diversity of the analysed floral scents in Annonaceae. I also summarize and discuss a wide range of (human) perceptual descriptions of floral scent found throughout the literature on Annonaceae. I have framed discussions of floral scent in Annonaceae in ecological and evolutionary contexts whenever possible. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169, 262–279.

An orchid colonization credit in restored calcareous grasslands

Habitat restoration comprises the re-creation of suitable environmental conditions with the intention of recolonization by certain target species. In previously abandoned calcareous grasslands, however, many characteristic plant species have been reported missing even decades after the reinstatement of traditional mowing or grazing management. Such grasslands are said to exhibit a colonization credit. This may be particularly true for orchid species, which often rely on highly specialized pollination strategies and mycorrhizal associations for completion of their life-cycle. In this study, we investigated whether restored calcareous grasslands exhibited an orchid colonization credit, whether this credit was associated with the degree of grassland fragmentation, and with particular species' life history traits. Applying the Beals index as a quantitative method to identify suitable habitats, several orchid species were indeed found missing from grasslands deemed suitable. There was no relation, however, between the extent of the colonization credit and the spatial isolation or size of the grasslands. Of all life history traits examined, only a high degree of pollinator specialization could be related to delayed colonization. This may suggest that restoring the pollinator community is an important bottleneck in calcareous grassland restoration.

Ecological features of the flowers of Aesculus hippocastanum L. and characteristics of Aesculus L. pollen seasons under the conditions of central-eastern Poland

The content of <em>Aesculus </em>pollen in aeroplankton of Lublin, located in central-eastern Poland, was studied during the period 2001–2012. The volumetric method was used for the study. The following parameters were determined: start, end, and duration of the pollen season, days of maximum concentration, and total annual pollen counts. Correlation coefficients were calculated between the start of the pollen season and meteorological parameters. The study showed that during the period 2007–2012 the onset of the <em>Aesculus </em>pollen season occurred on average 4 days earlier than in the years 2001–2006. The total annual pollen counts were in the range of 53–271, with an average number of 117. The average pollen counts did not differ significantly in the first and second year of a 6-year period of the study. The analysis of the correlation coefficients showed that there were significant positive correlations only between the season start date and minimum temperature of the first decade of May. The highest <em>Aesculus </em>pollen concentration occurred between 6 and 22 May. The flowers of <em>A. hippocastanum </em>demonstrate an ambophilous pollination strategy due to partial insect and wind pollination.

Generic status of <i>Quisqualis</i> (Combretaceae), with notes on the taxonomy and distribution of <i>Q. parviflora</i>

The taxonomic status of Quisqualis L., a genus closely related to and sometimes considered congeneric with Combretum Loefl., is discussed. The genus Quisqualis is maintained to accommodate selected African, Indian, Asian and Malesian species (including Q. indica L. and Q. parviflora Sond.). Diagnostic characters for Quisqualis include: leaves with abundant subepidermal crystalliferous idioblasts, each containing a large druse of calcium oxalate; leaves, stems and flowers with stalked glands but no scales; persistent petiole bases that develop into curved spines; petals well developed; hypanthium tubular or cylindrical, usually longer than 20 mm; stamens and style not or scarcely exserted; style adnate to hypanthium for more than half its length. Quisqualis shares a number of morphological similarities with Combretum Loefl. subgen. Cacoucia (Aubl.) Exell Stace sect. Poivrea (Comm. ex DC.) G.Don. Some species of Combretaceae from West, West Central and East Africa have the style adnate to the upper hypanthium and display features reminiscent of both Quisqualis and Combretum. These species also have characters of their own and in the past were placed in different sections under Combretum. It is suggested that at least some of these species may be best classified in genera distinct from Combretum and Quisqualis, one of which is Campylogyne Hemsley. Combretum s.str. is defined on the basis of a combination of characters and includes species of which the upper hypanthium is variable in shape, but when tubular or cylindrical, then always shorter than 20 mm. Other diagnostic characters include: stamens exserted well beyond petals; style exserted and free, but when shortly adnate to upper hypanthium (only at the base or for a short distance), then stamens long-exserted. It is suggested that different pollination strategies have developed independently in the Combretum–Quisqualis clade, resulting in convergent morphological trends in floral morphology. These homoplasious similarities in floral morphology are at the root of the difficulties experienced to demarcate genera. An alternative classification is provided for those preferring to include the southern African Quisqualis parviflora under Combretum s. l. For this purpose, a new combination and name, Combretum sylvicola O.Maurin is provided. Quisqualis parviflora is confined to the Eastern Cape and KwaZulu-Natal coastal regions and does not extend beyond this area as has been claimed by some. A comparative table to differentiate among four groups in Quisqualis and Combretum in Africa, as well as a photo of a herbarium specimen and a distribution map of Quisqualis parviflora, are provided.

723. GORTERIA DIFFUSA

The beetle daisy, Gorteria diffusa Thunb., is a South African and Namibian spring-flowering annual with a very interesting pollination strategy involving insect-mimicking petal spots, details of which are given here, in addition to notes on its history, habitat and cultivation, accompanied by a watercolour painting of the subsp. diffusa and a habitat photograph.

Investigating the relationship between pollination strategies and the size-advantage model in zoophilous plants using the reproductive biology of Arum cylindraceum and other European Arum species as case studies

The size-advantage model (SAM) explains the temporal variation of energetic investment on reproductive structures (i.e. male and female gametes and reproductive organs) in long-lived hermaphroditic plants and animals. It proposes that an increase in the resources available to an organism induces a higher relative investment on the most energetically costly sexual structures. In plants, pollination interactions are known to play an important role in the evolution of floral features. Because the SAM directly concerns flower characters, pollinators are expected to have a strong influence on the application of the model. This hypothesis, however, has never been tested. Here, we investigate whether the identity and diversity of pollinators can be used as a proxy to predict the application of the SAM in exclusive zoophilous plants. We present a new approach to unravel the dynamics of the model and test it on several widespread Arum (Araceae) species. By identifying the species composition, abundance and spatial variation of arthropods trapped in inflorescences, we show that some species (i.e. A. cylindraceum and A. italicum) display a generalist reproductive strategy, relying on the exploitation of a low number of dipterans, in contrast to the pattern seen in the specialist A. maculatum (pollinated specifically by two fly species only). Based on the model presented here, the application of the SAM is predicted for the first two and not expected in the latter species, those predictions being further confirmed by allometric measures. We here demonstrate that while an increase in the female zone occurs in larger inflorescences of generalist species, this does not happen in species demonstrating specific pollinators. This is the first time that this theory is both proposed and empirically tested in zoophilous plants. Its overall biological importance is discussed through its application in other non-Arum systems.