A literature review of 34 families of flowering plants containing at least one species pollinated primarily by beetles is presented. While the majority of species are represented by magnoliids and basal monocotyledons specialized, beetle-pollinated systems have evolved independently in 14 families of eudicotyldons and six families of petaloid monocots. Four, overlapping modes of floral presentation in plants pollinated exclusively by beetles (Bilabiate, Brush, Chamber Blossom and Painted Bowl) are described. Chamber Blossoms and Painted Bowls are the two most common modes. Chamber Blossoms, found in magnoliids, primitive monocotyledons and in some families of woody eudicots, exploit the greatest diversity of beetle pollinators. Painted Bowls are restricted to petaloid monocots and a few families of eudicots dependent primarily on hairy species of Scarabaeidae as pollen vectors. In contrast, generalist flowers pollinated by a combination of beetles and other animals are recorded in 22 families. Generalist systems are more likely to secrete nectar and exploit four beetle families absent in specialist flowers. Centers of diversity for species with specialized, beetle-pollinated systems are distributed through the wet tropics (centers for Brush and Chamber Blossoms) to warm temperate-Mediterranean zones (centers for Painted Bowls and a few Bilabiate flowers). It is unlikely that beetles were the first pollinators of angiosperms but specialized, beetle-pollinated flowers must have evolved by the midlate Cretaceous to join pre-existing guilds of beetle-pollinated gymnosperms. The floras of Australia and western North America suggest that mutualistic interactions between beetles and flowers has been a continuous and labile trend in angiosperms with novel interactions evolving through the Tertiary.

Convergent evolution and adaptive radiation of beetle-pollinated angiosperms

The fi rst three branches of the angiosperm phylogenetic tree consist of eight families with ~201 species of plants (the ANITA grade). The oldest fl ower fossil for the group is dated to the Early Cretaceous (115 - 125 Mya) and identifi ed to the Nymphaeales. The fl owers of extant plants in the ANITA grade are small, and pollen is the edible reward (rarely nectar or starch bodies). Unlike many gymnosperms that secrete " pollination drops, " ANITA-grade members examined thus far have a dry-type stigma. Copious secretions of stigmatic fl uid are restricted to the Nymphaeales, but this is not nectar. Floral odors, fl oral thermogenesis (a re- source), and colored tepals attract insects in deceit-based pollination syndromes throughout the fi rst three branches of the phylo- genetic tree. Self-incompatibility and an extragynoecial compitum occur in some species in the Austrobaileyales. Flies are primary pollinators in six families (10 genera). Beetles are pollinators in fi families varying in importance as primary (exclusive) to secondary vectors of pollen. Bees are major pollinators only in the Nymphaeaceae. It is hypothesized that large fl owers in Nympha- eaceae are the result of the interaction of heat, fl oral odors, and colored tepals to trap insects to increase fi tness.

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Pollination biology of basal angiosperms (ANITA Grade)

Dieffenbachia is an monoecious understory herb of tropical rain forests that exhibits a complex and specialized relationship with its beetle pollinators. The erect protogynous inflorescence has the spadix divided, with the female flowers in the basal half and male flowers in the upper half. Dieffenbachia longispatha Engler & Krause is pollinated by scarab beetles in the genera Cyclocephala and Erioscelis. The enveloping spathe of the inflorescence opens in the evening, but no flowers are sexually functional until the stigmas become receptive about 24 hr later. Beetles fly to the inflorescence in darkness, suggesting that floral odors play a role as an attractant. Beetles remain in the inflorescence for 24 hr, eating protein-rich staminodia that surround the stigmas. On the evening of the third day the anthers dehisce and beetles become covered with pollen as they crawl up the spadix in the process of leaving. Beetles fly an average of 80 m between inflorescences, usually to the nearest female inflorescence, although distances of 4001,000 m have been observed. Minimal estimate of genetic neighborhood sizes are large for D. longispatha (750 to 8,900 plants) and neighborhood areas encompass 41,000 to 67,000 M2. Experiments demonstrate that the species is self-compatible and that fruit production is pol-

https://www.jstor.org/stable/pdfplus/2444306.pdf

Beetle pollination of Dieffenbachia longispatha (Araceae)

Dynastid scarab beetles are the main or exclusive pollinators ofAnnona spp. with large flowers and wide floral chambers. These nocturnal beetles are attracted by the characteristic odours which are caused by measurable temperature elevation of the flowers up to 10°C or even 15°C above the ambient air temperature. TheAnnona spp. investigated showed different floral rhythms varying from one to three days. The elaborate and wellcoordinated flowering processes, along with floral heating and olfactorial attraction of the night-activeCyclocephala spp., result in a very precise and effective pollination. The floral chamber provides alimentation for the beetles in the form of nutritious food-tissues and pollen and offers a mating place and a well developed shelter against predation and environmental changes. A staggered flowering period and an alternative attraction of different beetle species seems to be more a device for diminishing competition between the cooccurringAnnona spp. than a hybridization barrier.

Beetle pollination and flowering rhythm of Annona spp. (Annonaceae) in Brazil

Previously published data on flowering phenology and pollination of neotropical families are reviewed and compared with new observations, in order to evaluate some trends of pollination strategies in neotropical savannas and forests. Central American forestBignoniaceae apparently diminish competition between sympatric species with identical pollination strategies by alternating flowering. On the other hand, in theBignoniaceae of Brazilian cerrado savannas coincident flowering is more common, suggesting different pollination strategies. Bees specialized in nectar robbery are associated withBignoniaceae, particularly mass-flowering species. These nectar robbers stimulate pollinator movements between flowers and thus increase pollen-flow. The flowering phenology of theMalpighiaceae from forests and savannas apparently follows the phenology of theAnthophorinae (mainlyCentridini)-bees, on which this plant group almost exclusively depends for pollination in the Neotropics. As the flowers of different genera and species are very uniform, the bees quite indiscriminately pollinate coexistingMalpighiaceae taxa. In theMalvaceae, the more primitive tree or shrub habit and an allogamous reproduction system is quite significantly associated with an ornithophilous pollination mode. Hummingbirds in the Neotropics have probably influenced the origin of the primitive bird-pollinatedMalvaceae flower. In conjunction with the diversification of theMalvaceae and their migration into more open regions, pollination by bees has apparently become more prominent. ChiropterophilousMalvaceae seem to have developed more recently from ornithophilous ancestors within the neotropical forest regions. As a result of this ecological radiation, parallel modifications of the basic ornithophilous flower principle can be traced in different groups of theMalvaceae.
 
Dynastinae-beetles are pollinators of South and Central American species ofAnnona, Cymbopetalum, Caladium, Talauma, Philodendron, Victoria, Nymphaea, Cyclanthus, Bactris and others. In many cases the relationship between flowers and theirDynastinae visitors is not very specific, principally reflecting differences in the faunistic spectrum of diverse regions and vegetation types. The attraction of these beetles as flower visitors in several unrelated groups of Angiosperms must have occurred repeatedly. It is suggested that the flower visiting species of this beetle group may have acquired their preference for certain pungent or aromatic flower odours by their feeding habits during the larval stage.

Some pollination strategies in neotropical savannas and forests

Comparaison de la biologie florale de 2 especes de Philodendron dans leur habitat naturel a Sao Paulo

Gerhard Gottsberger

Papers

Pollination strategies in brazilian Philodendron species