Article

Caladium bicolor (Araceae) and Cyclocephala celata (Coleoptera, Dynastinae): A Well-Established Pollination System in the Northern Atlantic Rainforest of Pernambuco, Brazil

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Abstract

Flowering, pollination ecology, and floral thermogenesis of Caladium bicolor were studied in the Atlantic Rainforest of Pernambuco, NE Brazil. Inflorescences of this species are adapted to the characteristic pollination syndrome performed by Cyclocephalini beetles. They bear nutritious rewards inside well-developed floral chambers and exhibit a thermogenic cycle which is synchronized to the activity period of visiting beetles. Heating intervals of the spadix were observed during consecutive evenings corresponding to the beginning of the female and male phases of anthesis. Highest temperatures were recorded during the longer-lasting female phase. An intense sweet odour was volatized on both evenings. Beetles of a single species, Cyclocephala celata, were attracted to odoriferous inflorescences of C. bicolor and are reported for the first time as Araceae visitors. All the inflorescences visited by C. celata developed into infructescences, whereas unvisited inflorescences showed no fruit development. Findings of previous studies in the Amazon basin of Surinam indicated that Cyclocephala rustica is a likely pollinator of C. bicolor. This leads to the assumption that locally abundant Cyclocephalini species are involved in the pollination of this species.

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... Cyclocephala celata was until now associated only with species of Araceae, as a pollinator of Caladium bicolor L., Tacaccarum ulei Engl. and K. Krause and Philodendron acutatum Schott, all in Pernambuco state (Maia & Schlindwein 2006, Maia et al 2010. On the other hand, C. vestita had already been registered as pollinator of soursop in Pernambuco and Bahia states in Brazil, and recommend as an important pollinator of this culture (Cavalcante 2000, Maia et al 2010. ...
... A poorly known species, C. celata, (Fig 2e, f) was successfully reared (Souza et al 2014), although with a low emergence rate, probably because the authors frequently moved the larvae to make measurements, most likely stressing and killing them. The species was originally described from Paraguay and was reported for the first time for Brazil by Maia & Schlindwein (2006) and since then frequently collected in northeastern Brazil (Souza et al 2014, this study). We also studied two specimens from Bolivia that constitute a new country record for this species (CERPE). ...
... The anterior tibiae are bidentate in males with a proximal notch, suggesting a third tooth (Ratcliffe 2003), but tridentate in females, with the proximal tooth removed from the distal ones. It was reported on soursop plant by Vilalta (1988), but according to most of the literature (e.g., Ratcliffe 2003;Maia & Schlindwein 2006;Moore & Jameson 2013), it is most frequently found in Arecaeae species, making the records in Annonaceae dubious, and also probably a result of species misidentification. ...
Article
The soursop (Annona muricata L., Annonaceae) is an important fruit crop in several countries of South America, including Brazil, and the presence of the scarab beetles in this orchads can reduce pollination deficits and increase the productivity. For this reason, we report Cyclocephala celata Dechambre, 1980 as a flower visitor and potential pollinator of the soursop. Additionally, this work presents an updated list of Cyclocephala species found on A. muricata, corrects some misidentifications, and provides comments and an identification key for all taxa cited as floral visitors of this crop.
... Nocturnal or crepuscular habits are associated with anthophilia, especially for Cyclocephalini (Endrödi, 1985;Gottsberger, 1999Gottsberger, , 1986Maia and Schlindwein, 2006;Maia et al., 2012Maia et al., , 2010. The pollination of many tropical fruit trees that have night anthesis, such as Annona spp. ...
... The pollination of many tropical fruit trees that have night anthesis, such as Annona spp. (Annonaceae), palms (Arecaceae) and aroids (Araceae), is dependent on pollinating beetles, especially species of the genera Cyclocephala Latreille and Erioscelis Burmeister, which ensure the reproductive success of these trees (Endrödi, 1985;Gottsberger, 1986Gottsberger, , 1999Silberbauer-Gottsberger et al., 2003;García-Robledo et al., 2004;Croat, 2004;Maia and Schlindwein, 2006;Ratcliffe, 2008Ratcliffe, , 2003Maia et al., 2012Maia et al., , 2010. ...
... & K. Krause (Araceae), as the species has been identified as a pollinator of this plant species (Maia et al., 2013). Such nocturnal habits, and period of activity, may be associated with the cantharophilous pollination of species of Araceae and Annonaceae, which provide special chambers and exudates for feeding and mating (Maia and Schlindwein, 2006;Cavalcante et al., 2009;Maia et al., 2010Maia et al., , 2013. Due to this relationship, some groups have evolved flight periods that are similar to the flowering time of their preferential host plants (Maia and Schlindwein, 2006;Maia et al., 2010). ...
Article
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Dynastinae is one of the most representative subfamilies of Melolonthidae (Scarabaeoidea) and has considerable ecological importance due mainly to interactions with plants of the families Araceae and Annonaceae. This relationship has led to the evolution of nocturnal activity patterns, which are influenced by environmental conditions. In the present study, abiotic factors were investigated to comprehend the influence on the flight patterns and identify the sex ratio of beetles from this subfamily. A study was conducted the Campo de Instrução Marechal Newton Cavalcante in northeastern Brazil between December 2010 and November 2011. Thirteen species of Dynastinae were identified, most of which were from the genus Cyclocephala. Abundance and richness were greater in the dry season. Six species exhibited peak flight activity at specific periods of the night. More females than males were recorded for Cyclocephala distincta and C. paraguayensis. The present findings suggest that that rainfall reduces the flight activity of these beetles and different time schedules may be related to mating behavior, foraging behavior and the avoidance of interspecific resource competition.
... Floral associations between cyclocephaline beetles (Scarabaeidae: Dynastinae) and Araceae species have been widely documented in the Neotropics (Gottsberger and Amaral 1984;Gottsberger 1986;Young 1986Young , 1988Young , 1990Gottsberger and Silberbauer-Gottsberger 1991;Beath 1999;Gibernau et al. 1999;Gibernau and Barabé 2002;Gibernau 2003;Gibernau et al. 2003;Maia and Schlindwein 2006;Maia et al. 2010;Gottsberger et al. 2013). Neotropical cyclocephaline species have been recorded from the inflorescences of ten Araceae genera native to that biogeographic region (Moore and Jameson 2013). ...
... Two species of Cyclocephala Dejean have been reported in the inflorescenes of Caladium bicolor (Aiton) Vent.: Cyclocephala celata Dechambre in Pernambuco, Brazil (Maia and Schlindwein 2006) and Cyclocephala rustica (Olivier) in Brownsberg, Suriname and French Guiana (Pellmyr 1985;Moore and Jameson 2013). Cyclocephala atricapilla Mannerheim was observed in the inflorescences of an unidentified Caladium species from Maranhão, Brazil (Gottsberger 1986). ...
... These heated floral chambers are energy savers for the beetles, and they are used as shelters and mating sites (Seymour et al. 2003). Aroid floral visitation patterns observed in this study are consistent with many angiosperms (Gottsberger and Amaral 1984;Pellmyr 1985;Gibernau et al. 1999;Gibernau and Barabé 2002;Maia and Schlindwein 2006;Maia et al. 2010). Early developmental stages of flowers in C. bicolor and Philodendron species are similar (Barabé and Lacroix 2002), suggesting similar conditions. ...
... 9 All Caladieae species studied so far, 22 including Caladium, the sister genus of Syngonium, are pollinated by beetles ( Figure S3) (Coleoptera: mainly Cyclocephalini scarab beetles, Dynastinae) that feed on floral food bodies (enlarged sterile male flowers) and pollen. [37][38][39][40][41] These plant species are mostly co-visited by high numbers of other Neella plant bug species that do not enter the pollination chamber and thus cannot act as pollinators. 26,27,42 Similar communities of floral visitors, including scarab beetles as pollinators and plant bugs as abundant florivores, are found in other, more distantly related Araceae genera including Dieffenbachia and Philodendron. ...
... 24,[43][44][45][46] Anthesis and pollination biology of S. hastiferum differ distinctly in various properties from scarab beetle-pollinated Neotropical Araceae ( Figure 6). Food bodies, which are characteristic of scarab beetle-pollinated species (e.g., Syngonium schottianum; species of Caladium, Dieffenbachia, and Philodendron 26,41,44,[46][47][48], are absent in S. hastiferum. Heat and scent production occur in the morning instead of evening hours, 24,26,47-52 and a novel, previously unknown scent compound, (Z)-3-isopropylpent-3-en-1-ol (gambanol), is responsible for pollinator attraction. ...
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Angiosperm flowers and their animal visitors have co-evolved for at least 140 Ma, and early flowers were likely used mainly as mating and feeding sites by several groups of insects, including beetles, flies, true bugs, and thrips. Earlier studies suggested that shifts from such neutral or antagonistic relationships toward mutualistic pollination interactions between flowers and insects occurred repeatedly during angiosperm evolution. However, the evolutionary mechanisms and adaptations, which accompanied shifts toward effective pollination, are barely understood, and evidence for such scenarios has been lacking. Here, we show that Syngonium hastiferum (Araceae), a Neotropical representative of an otherwise beetle-pollinated clade, is pollinated by plant bugs (Miridae; Heteroptera), which are florivores of Syngonium schottianum and other Araceae species. We found that S. hastiferum differs in several floral traits from its beetle-pollinated relatives. Scent emission and thermogenesis occur in the morning instead of the evening hours, and its pollen surface is spiny instead of smooth. Furthermore, the floral scent of S. hastiferum includes a previously unknown natural product, (Z)-3-isopropylpent-3-en-1-ol, which we show to have a function in specifically attracting the plant bug pollinators. This is the first known case of a specialized plant bug pollination system and provides clear evidence for the hypothesis that the adoption of antagonistic florivores as pollinators can drive flower diversification. Video abstract Video Camera Download : Download video (48MB)
... Among the angiosperm families with beetle-pollinated species (cantharophilous), Araceae represents one of the most diverse in the Neotropics with an estimated 800 species associated with cyclocephaline scarabs (Melolonthidae, Dynastinae, Cyclocephalini) alone. Although these highly specialized scent-oriented pollinators have been documented in association with the inflorescences of Caladium Ventenat (Maia and Schlindwein 2006), Dieffenbachia Schott (Valerio 1984;Young 1986Young , 1990Cuartas-Hernandez 2006;Gibernau 2015aGibernau , 2015b, Gearum N.E. Brown (Gonçalves and Maia 2006), Montrichardia H. Crüger (Gibernau et al. 2003), Syngonium Schott (Beath 1998), Taccarum Brongniart ex Schott (Maia et al. 2013), and Xanthosoma Schott (Garcia-Robledo et al. 2004;Gottsberger et al. 2020), it is the megadiverse genus Philodendron Schott with ca. ...
... High visitation rates were also reported for other cyclocephaline scarab-pollinated, such as P. solimoesense (100%, Gibernau et al. 1999) and P. acutatum Schott (>91.5%, Maia et al. 2010), Caladium bicolor (Aiton) Ventenat (100%, Maia and Schlindwein 2006), and Dieffenbachia nitidipetiolata Croat & Grayum (reported as D. longispatha) (70%, Young 1986). When pollinators are less abundant and/or the habitat is highly fragmented/disrupted, visitation rates can be lower as in the cases of Taccarum ulei Engler & K. Krause (60%, Maia et al. 2013), Montrichardia linifera (Arruda) Schott (58%, Gibernau et al. 2003), or Xanthosoma daguense Engler (46%, Garcia-Robledo et al. 2004). ...
Article
In French Guiana, Philodendron fragrantissimum (Hook.) G.Don was specifically pollinated by a single species of night-active cyclocephaline scarab, Cyclocephala simulatrix Höhne. Its inflorescences exhibited short-lasting anthesis (~30 h) and characteristic floral traits such as floral thermogenesis, edible/nutritious floral tissues, and profuse floral scent emission. Our insect exclusion experiments indicate that entomophilous cross-pollination is obligatory for P. fragrantissimum, with olfactory signalling playing a pivotal role in pollinator attraction. Three volatile organic compounds – methyl benzoate, (Z)-jasmone, and dehydrojasmone – dominate the floral scent of pistillate phase inflorescences (84–99%), but their relative proportions were different according to the headspace sampling method used, dynamic or static. Local pollination services provided by both female and male C. simulatrix are effective, as evidenced by a high visitation rate (>70%) that resulted in 55% of the inflorescences maturing into infructescences. Moreover, pollination was highly efficient, since 2–3 beetles sufficed to pollinate an average 94% of the pistillate flowers.
... Observou-se uma intensificação do odor atrativo durante os picos de temperatura. Contudo, foi impossível avaliar a importância do aquecimento floral em A. muricata, uma vez que a variação de temperatura foi pequena, em comparação com outros estudos realizados em plantas cantarófilas (Maia & Schlindwein 2005, 2006). ...
... Não existem registros de indivíduos de C. vestita coletados em flores de outras espécies de plantas, mas a ampla distribuição da espécie no Nordeste brasileiro evidencia a utilização de diferentes recursos alimentares pelos adultos de espécies nativas de Annonaceae (Barbosa et al. 1996, Silberbauer-Gottsberger et al. 2003 e Araceae (Barbosa et al. 1996, Maia & Schlindwein 2006. Cavalcante (2000) observou que a presença e abundância de C. vestita em pomares de A. muricata estão relacionadas à proximidade de áreas com cobertura vegetal nativa. ...
... Pistillate and sterile staminate flowers are distributed within the tube. Among several genera of the Caladieae, including Xanthosoma, either nitidulids (Nitidulidae) or cyclocephaline scarab (Melolonthidae, Cyclocephalini) beetles act as key pollinators (Maia & Schlindwein 2006;Chartier et al. 2014;Gibernau 2016;Milet-Pinheiro et al. 2017). ...
... In spite of the existence of several studies on pollination and scent emission in tropical Araceae (Maia & Schlindwein 2006;Maia et al. 2012;Dötterl et al. 2012;Maia et al. 2013a;b;Gottsberger et al. 2013;Pereira et al. 2014;Gibernau 2016;Milet-Pinheiro et al. 2017), we know little about the constancy of pollinating beetle species in different regions and whether floral scent emissions are similar among distant populations or not. To this end, the present paper presents data on pollinators and floral biology of the widely distributed species Xanthosoma striatipes from sites in the Brazilian states of São Paulo, Minas Gerais, and Maranhão. ...
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We report on the reproductive biology, pollinators, and thermogenesis of the widely distributed Araceae species, Xanthosoma striatipes. We analyzed the floral scent in one population and compared it to a previous study. Xanthosoma striatipes was studied at three sites in Brazil, São Paulo, Minas Gerais and Maranhão. The results showed constancy of the reproductive biology and main pollinator species across sites. The pollinators at all three sites sites were two species of cyclocephaline scarabs, Cyclocephala atricapilla and C. ohausiana. The scent composition of the strongly scented inflorescences of plants of a site in São Paulo differed from plants recently studied in Central Brazil. Of the 39 and 23 compounds detected, in the present and the previous study, respectively, only seven compounds, were in common. We discuss that the two main constituents identified in samples at both sites, the aromatic compound methyl salicylate and the terpenoid (E)-4,8-dimethyl-1,3,7-nonatriene, alone seem to be sufficient for the attraction of the beetles. However, it is also possible that different compounds are involved in attracting the same beetle species to plants of the different populations.
... The average and maximum MVT emission rates reported for C. bicolor are 790 and 2250 μg h −1 per inflorescence [78], whichtaking inflorescence fresh weight to be ∼2 g [80] would be four orders of magnitude above THZ synthesis rates in leaves (Table 1). If the MVT was derived from THZ produced by the normal suicidal action of THI4, these massive emission rates would require THI4 synthesis rates of 234 and 666 mg h −1 per inflorescence, respectively. ...
... However, these organisms are hyperthermophilic anaerobes that grow in sulfide-rich environments and contain millimolar internal levels of sulfide [84]. It is therefore uncertain whether their 'catalytic instead of suicidal THI4' strategy is possible in arum lily flowers, which operate at moderate temperatures [80], fairly high oxygen tensions [85], and at internal sulfide levels that are probably in the 10-100 μM range [86,87]. Alternatively, C. bicolor flowers could have a novel biosynthetic pathway to MVT, although it is not obvious what the initial substrate(s) and intermediates in such a pathway might be. ...
... This might indicate that a particular floral scent compound can exhibit different biological attractiveness towards different cyclocephaline species, even if this compound is predominant or co-dominant in the floral bouquet of the host plants. Scented traps containing (S)-2-hydroxy-5-methyl-3hexanone, in contrast, successfully lured beetles of Cyclocephala cearae Höhne, 1923, which are pollinators of T. ulei (Maia et al., 2013a), and Cyclocephala celata Dechambre, 1980, which are pollinators of Caladium bicolor, P. acutatum, and T. ulei (Maia & Schlindwein, 2006, Maia et al., 2010, 2013a: results apparently demonstrating that the same compound is capable of attracting more than one species. Our study plant is overall most similar to P. acutatum (Fig. 4), but does not share a pollinator with this species. ...
... observ.). This 'dangerous' local dependence on one or sometimes two highly effective, specialized pollinator species seems to be the rule for all cyclocephaline scarab-pollinated Araceae (Gottsberger, 1999;Gibernau et al., 1999Gibernau et al., , 2000Gibernau et al., , 2003Maia & Schlindwein, 2006;Maia et al., 2010). In contrast, the animal counterpart of the association appears to be less specific and not entirely dependent on a single host plant. ...
Article
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Cyclocephline scarabs and their host plants are documented as highly specialized plant–pollinator associations, with various fine-tuned adaptations. We studied the association between Philodendron adamantinum, a species endemic to the Espinhaço Range in Minas Gerais, South-East Brazil, and its exclusive pollinators. We focused on the pollination mechanism and reproductive success of P. adamantinum, analysed its floral scent composition, and performed field bioassays to verify the scent-mediated attraction of pollinators. The reproductive success of P. adamantinum depends on the presence of Erioscelis emarginata (Scarabaeidae, Cyclocephalini), its sole pollinator. At dusk, the inflorescences heat up to 18 °C above the surrounding ambient air temperature and give off a strong sweet odour, from which 32 volatile compounds were isolated. Dihydro-β-ionone, the major constituent in the floral scent bouquet, lures individuals of E. emarginata when applied to scented artificial decoys, either alone or blended with methyl jasmonate. We attribute the low fruit set of P. adamantinum at our study sites to pollinator limitation of small and isolated populations and geitonogamic pollen flow of vegetatively generated clonal plant groups. The interaction between P. adamantinum and E. emarginata shows common traits typical of the known plant–pollinator associations involving cyclocephaline scarabs: the asymmetrical dependence of plants on their pollinators, and the scent-mediated interaction between flowers and beetles. In addition to updating the current catalogue of active compounds of cantharophilous pollination systems, further experimental studies should elucidate the role of the specific chemical compounds that attract pollinators along different time and biogeographic scales. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, ●●, ●●–●●.
... Over nearly 150 years of selection and breeding, more than 2000 varieties have been cultivated [16][17][18]. At present, there are more than 90 varieties on the market, among which more than 50 varieties are grown for large-scale flower production [19]. Since the beginning of this century, China has successively introduced some varieties of colorful Caladium as ornamental plants, which are greatly appreciated by retailers and young people [20]. ...
Article
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Caladiums are promising colorful foliage plants due to their dazzling colors of the leaves, veins, stripes, and patches, which are often cultivated in pots or gardens as decorations. Four wild species, including C. bicolor, C. humboldtii, C. praetermissum, and C. lindenii, were employed in this study, where their chloroplast (cp) genomes were sequenced, assembled, and annotated via high-throughput sequencing. The whole cp genome size ranged from 162,776 bp to 168,888 bp, and the GC contents ranged from 35.09% to 35.91%. Compared with the single large copy (LSC) and single small copy (SSC) regions, more conserved sequences were identified in the inverted repeat regions (IR). We further analyzed the different region borders of nine species of Araceae and found the expansion or contraction of IR/SSC regions might account for the cp genome size variation. Totally, 131 genes were annotated in the cp genomes, including 86 protein-coding genes (PCGs), 37 tRNAs, and eight rRNAs. The effective number of codons (ENC) values and neutrality plot analyses provided the foundation that the natural selection pressure could greatly affect the codon preference. The GC3 content was significantly lower than that of GC1 and GC2, and codons ending with A/U had higher usage preferences. Finally, we conducted phylogenetic relationship analysis based on the chloroplast genomes of twelve species of Araceae, in which C. bicolor and C. humboldtii were grouped together, and C. lindenii was furthest from the other three Caladium species occupying a separate branch. These results will provide a basis for the identification, development, and utilization of Caladium germplasm.
... The floral scent chemistries of two other cyclocephaline scarab beetle-pollinated aroid genera have been studied: Montrichardia and Caladium (Gibernau et al. 2003;Maia and Schlindwein 2006;Maia et al. 2012). Floral VOCs identified in samples of Montrichardia arborescens (Gibernau et al. 2003) are common among other species associated with cyclocephaline scarab beetles, namely (Z)-jasmone (37%), 1,3,5-trimethoxybenzene (26%), methyl salicylate (7%), and methyl benzoate (8%). ...
Article
We provide a characterization of the floral scent chemistry of nine species of Philodendron present in French Guiana. The scent samples were obtained by dynamic headspace, in situ in French Guiana, or from plants in the living aroid collection at the Botanical Garden of the University of Brasilia. Overall, we identified 43 volatile organic compounds (VOCs) belonging to the compound classes of monoterpenes, sesquiterpenes, irregular terpenes, benzenoids, aliphatics, jasmone derivatives, and nitrogen-containing compounds. Five of these floral VOCs are new reports for Philodendron. Interestingly, the floral blends in the investigated species were comprised of 3–22 VOCs; but when considering only major VOCs (≥5%), each fragrant blend could be summarized into one up to four constituents. Except for P. acutatum and P. billietiae, whose floral scents were largely dominated (≥93%) by 4-vinylanisole, fragrances of the investigated species were dominated (≥62%) each by a different VOC (i.e. dehydrojasmone, dihydro-β-ionone, (E)-4,8-dimethylnona-1,3,7-trien-5-yl acetate, methyl benzoate, 1,3,5-trimethoxybenzene, dihydrobovolide, or (Z)-jasmone). Consequently, all floral scent compositions significantly differed from one other. We compare our results to published data to provide an insight into the floral scent chemistry within the genus Philodendron. We further discuss the diversity of VOCs in cyclocephaline scarab-pollinated aroids.
... Within the private Atlantic Forest we actively searched inside inflorescences of Philodendron acutatum Schott, Caladium bicolor (Aiton) Vent., and Taccarum ulei Engl. & K. Krause (Araceae), known hosts of flower-visiting C. celata and C. cearae (Maia and Schlindwein, 2006;Maia et al., 2010). Since most species of cyclocephaline scarabs are attracted to light sources at night (Ratcliffe and Cave, 2009;Albuquerque et al., 2016), light traps were also installed during early evening hours, from 17h30 to 21h00. ...
Article
Different species of Cyclocephala scarab beetles (Scarabaeidae, Dynastinae) perform key functional roles in both natural and agricultural systems, such as the cycling of organic matter and pollination, while also being known as destructive pests both as immatures and adults. Therefore, the identification of biological parameters is crucial for defining strategies for their conservation and efficient pest management. In a forest fragment within the Brazilian Atlantic Forest biodiversity hotspot, we field-captured adult individuals of Cyclocephala cearae , C. celata , and C. paraguayensis then reared and bred them under controlled temperature and humidity conditions. On a daily basis, we individually weighted eggs of all three species, from oviposition until hatching, and monitored egg development parameters (i.e., incubation duration, viability, and egg weight increase). Our findings provide novel empirical evidence showing (i) a positive correlation between egg weight and incubation duration, (ii) idiosyncratic characteristics on egg development, and (iii) a negative (involuntary) effect of manipulation on egg development and viability. Thus, the successful breeding and rearing of Cyclocephala spp. is correlated with egg integrity and the targeted species. Our analyses present a quantitative understanding of the egg phase and can assist in refining strategies for ovicidal activity and pest management of Cyclocephala spp. in agriculture systems. Moreover, they can provide a basis for new studies related to captivity breeding, pollinator management, and developmental biology for biodiversity conservation.
... Scarab beetles (Coleoptera: Scarabaeidae) are a diverse group with a wide range of adult feeding habits, including phytophagous species that consume leaves, flowers, and fruits of diverse plant species; some species cause damage to cultivated plants (Solís 2004, Maia & Schlindwein 2006, Shaughney & Ratcliffe 2015, Rodrigues et al. 2016, Ferreira et al. 2018. Moreover, larvae of some phytophagous scarabs feed on roots and also may cause economic damage to crops (Santos & Ávila 2009, Cherman et al. 2011, Coutinho et al. 2011. ...
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Adults of the beetle Cyclocephala literata Burmeister, 1847 are important pollinators to some Magnoliaceae. Is known that insects could find host plants by detecting volatiles through antennal sensilla. Cyclocephala has its three distal antennomeres lamellate, and the surface of each lamella has sensilla trichodea, chaetica, placodea, coeloconica, basiconica and ampullacea. Three kinds of sensilla placodea were found (type I, II and III), and two kinds of sensilla coeloconica were observed (type I and II). Females have on average 10,776 sensilla, of which 10,214 are sensilla placodea, 536 are sensilla coeloconica, and 26 are sensilla basiconica. Males have on average 10,386 sensilla, of which 9,873 are sensilla placodea, 464 are sensilla coeloconica, and 49 are sensilla basiconica. Males and females have similar quantities of sensilla, and sensilla placodea are predominant. The differences observed in the number of sensilla of males and females were found in other beetles and were attributed to the detection of cospecific sexual pheromones by one of the sexes, or to the detection of plant volatiles. The antennal sensilla of C. literata is described and quantified in present study, and some perspectives about the differences kind of chemical communication, pollination, and antennae dimorphism is discussed.
... Studies have shown that adults of Scarabaeidae use different plant species as sources of food and they use this sites for mating (e.g., Maia and Schlindwein, 2006;Maia et al., 2013;Martínez et al., 2013;Rodrigues et al., 2014bRodrigues et al., , 2017. The adult aggregation is intermediated by the detection of valatines by the antennae of these insects (Leal, 1995(Leal, , 1998. ...
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ABSTRACT Species of the scarab beetle genus Liogenys are potential pests to several crops in Brazil. This study aimed to describe the antennal sensilla of Liogenys suturalis (Blanchard, 1851). Adults were collected in a pasture area in Bálsamo, São Paulo state, Brazil, using a light trap. The antennae were dissected and images of the antennal sensilla were obtained using a scanning electron microscope. Sensilla ampulacea (pores), s. auricilica, s. basiconica, s. placodea, and s. trichodea are present in the lamellae. The antenna of females have 4399 sensilla, of which 3671 (83.5%) are s. placodea, 422 (9.5%) s. coeloconica, and 306 (6.9%) s. auricilica. The antennae of males have 4039 sensilla, of which 3117 (77.1%) are s. placodea, 353 (8.7%) s. coeloconica, and 569 (14.1%) s. auricilica. The antennal sensilla of the genus Liogenys have been described for the first time.
... These, in turn, can be sent off for analysis by gas chromatography coupled to mass spectrometry (GC-MS). This has been a major pursuit of some researchers, particularly Artur Maia at the Federal University of Paraiba in Brazil, who has used this technique on a variety of different plants, including palms, water lilies and aroids (Maia & Schlindwein, 2006;Maia et al., 2010Maia et al., , 2013. Florian Etl at the University of Vienna in Austria has shown for example that not only is there a difference in the scent produced, but has shown that bees can be induced to visit inflorescences only at specific times, so that it is a combination of the scent produced and the exact timing of that scent production that enable species to be in flower at seemingly the same time without being crosspollinated. ...
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This paper discusses the family Araceae, emphasizing its worldwide distribution and the diversity of morphological and ecological characteristics of the family that have enabled it to reach such a prominent position with diverse habitats. Few families of its size have come to inhabit such a diverse spectrum of habits and biomes. The family has important habit forms and growth structures that have enabled such distributional patterns. The very broad spectrum of life forms it demonstrates is one of the main characteristics of the family, namely broad habitat diversity. This coupled with high species diversity; high rates of endemism and the presence of large numbers of unknown species (probably the highest of any family percentage-wise) constitute principal characteristics of the family.
... Caladium bicolor (Aiton) Vent is a member of Araceae family widely believed to originate from South and Central America but globally used as an ornamental plant. Due to its horticultural value, it is commonly cultivated in and around homes in different regions and climate of the world [1]. C. bicolor has diverse nomenclature in different parts of the world. ...
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To assess nephropathic potential of Caladium bicolor methanolic extract within the renal parenchyma of experimental Wistar rats. Twenty four albino Wistar rats (weighing between 180-195 g) were divided into four groups which include Group I treated with distilled water (2 ml/kg b.w.), Group II treated with methanolic extract of C. bicolor (100 ml/kg b.w.), Group III treated with methanolic extract of C. bicolor (200 ml/kg b.w.), Group IV treated with methanolic extract of C. bicolor (300 ml/kg b.w.). All administrations were done orally and once daily for a period of thirty five days. Body weight of animals was recorded during days 0, 7, 14, 21, 28 and 35 of study. After the study period, kidney tissue of study animals was harvested, weighed and processed for histopatho logical study. Staining of renal tissue sections was done using H & E technique, examined under microscope for observable histopathological changes that were scored using image-J software. Results of this study showed that sub-chronic exposure to methanolic extract of C. bicolor caused reduction in body and renal tissue weight. Moreover, exposure to the extract increases significantly (p < 0.05) renal histopatho logical changes including inflammation, necrosis, glomerular congestion and tubular dilatation within the renal parenchyma of study animals. Therefore, methanol extract of C. bicolor exhibited dose-independent nephropathic effect on the renal parenchyma following a sub-chronic exposure in experimental Wistar rats.
... Regarding adults of Cyclocephala, several data on biology, life cycle, association with plants, and sexual behavior have been provided (e.g. Maia and Schlindwein, 2006;Maia et al., 2013;Mondino et al., 1997;Moore and Jameson, 2013;Nogueira et al., 2013;Oliveira and Ávila, 2011;Rodrigues et al., 2010Rodrigues et al., , 2018. ...
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Some species of Cyclocephala Dejean, 1821 (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini) were reported as crop or pasture pests. Within this genus, Cyclocephla tucumana Bréthes, 1904 was noticed in cultivated areas in South Brazil, 2009 − 2010. Other study found larvae of the species associated withpastures in Aquidauana, Mato Grasso do Sul State (MS), 2009. In the present study, larvae of C. tucumanawere collected in pasture areas from August 2015 to July 2016, in Cassilândia, MS. The material was reared in laboratory for the description of immatures, and the regional population was studied to clarify its dynamics. Immatures were abundant throughout the years and are described for the first time. Taxonomic discussion, updated key to larvae and pupae, and a teratological note are also added.
... Phytophagous scarab beetles (Coleoptera: Scarabaeidae) in Neotropical region usually swarm during the hot and humid period of the year, when they can be observed feeding on leaves, flowers and fruit of several plant species (Oliveira & Ávila 2011, Gottsberger et al. 2012, Martínez et al. 2013, Moore & Jameson 2013, Ferreira et al. 2016). The different plant species used for feeding are important breeding sites for Scarabaeidae (Morón 1996, Maia & Schlindwein 2006, Rodrigues et al. 2017. ...
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When suitable, adults of Scarabaeidae usually form swarms to find food and breeding sites. The steps of mating behavior can be mediated by chemical communication, and antennal sensilla are released volatiles detection structures, as sexual pheromones. In present work the mating behavior and the antennal sensilla of Anomala inconstans Burmeister, 1844 are described. The study was conducted at the Universidade Estadual de Mato Grosso do Sul, Cassilândia, Brazil from March 2015 to December 2017. Adults were collected through a light trap and taken to the laboratory for studies. Field and laboratory observations provided data for the description of the steps of mating behavior. Adults swarms were registered from September to November 2015 at 05:30 pm to 00:00 am. Females display a calling behavior from 05:25 pm to 08:00 pm, in which they rub their posterior legs against their abdomen, and after a few minutes males are able to locate them. In laboratory, the mating process lasted 20.4 minutes on average, and the possibility of chemical communication between adults was here discussed. The antennae of the species have trichoid, chaetica, placoid types I, II and III, and coeloconic types I and II sensilla. Placoid sensilla are the most abundant and females have more sensilla than males.
... Although deception is the most common pollination syndrome among the Araceae, some plants have a mutualistic relationship with their pollinator(s) (Chartier et al. 2014). Plants pollinated exclusively by beetles (cantharophily) (Maia and Schlindwein 2006;Gottsberger et al. 2013) provide specific cues that attract these beetles (Maia et al. 2012(Maia et al. , 2013Pereira et al. 2014). For example, the cantharophilous Philodendron selloum (K. ...
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Western skunk cabbage, Lysichiton americanus (Araceae), is pollinated mainly by the rove beetle Pelecomalium testaceum (Staphylinidae). Our objective was to determine the floral semiochemical(s) of L. americanus that attract(s) P. testaceum. Porapak Q headspace volatile extracts of L. americanus inflorescences were analyzed by gas chromatographic–electroantennographic detection (GC–EAD) and GC–mass spectrometry. In GC–EAD analyses, three floral odorants [(E)-4 nonene, (E)-5-undecene, indole] elicited consistent responses from the antennae of female P. testaceum. In field experiments, traps baited with a blend of these three components (“3-CB”) captured significantly more P. testaceum than unbaited control traps. Traps baited with the 3-CB, the two hydrocarbons, or indole, each captured significantly more beetles than unbaited control traps, indicating redundancy in the semiochemical blend. Moreover, traps baited with indole captured significantly more beetles than traps baited with either the 3-CB, or the hydrocarbons, indicating that indole is a key floral attractant for P. testaceum. Many necrophilous and coprophilous insects respond to indole in search of carrion or feces, but P. testaceum has never been associated with these types of resources. Both electrophysiological and behavioral responses of P. testaceum to two hydrocarbon semiochemicals, which are not signature odorants of carrion or feces, may indicate that the beetles recognize the odor of L. americanus as an honest signal, seek and pollinate its inflorescences, and get rewarded with pollen and on-plant mating opportunities.
... Thermogenesis in plant reproductive structures is critical in the pollination systems of basal angiosperms, some monocots and cycads, where it functions as an attractant and/or reward for insects (Seymour & Schultze-Motel 1997;Ervik & Barfod 1999;Thien et al. 2000Thien et al. , 2009. Both odour and thermogenesis are believed to represent adaptive mechanisms that regulate insect pollinator behaviour and composition (Kumano & Yamaoka 2006;Maia & Schlindwein 2006;Theis et al. 2007). Variations in these traits can therefore reflect shifts in pollinators that may ultimately lead to speciation in plants (Xu et al. 2011). ...
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• Heat and odour production can have profound effects on pollination in cycads. It is therefore expected that these traits would co‐vary geographically with pollinator assemblages. Such intraspecific variation, may lead to the evolution of pollination ecotypes, which can be an early stage of pollinator‐mediated speciation. • We measured cone temperatures using miniature temperature data loggers and examined the composition of cone volatile odours using headspace sampling and analysis with gas chromatography‐mass spectrometry in four populations spanning the range of the African cycad Encephalartos ghellinckii. Pollinator assemblages were also investigated in three populations. • Male and female cones were thermogenic at pollen shed and receptive stages, respectively, but patterns of thermogenesis did not vary among populations. Scent emissions from cones in populations in the Drakensberg Mountains were characterised by cis‐β‐ocimene, β‐myrcene and (3E)‐1,3‐octadiene, while camphene and α‐pinene were characteristic of scent emissions from cones in populations closer to the coast. These differences in volatile blends corresponded with differences in insect assemblages. • These results confirm intraspecific variation in volatile emissions of E. ghellinckii and support the predictions that intraspecific variation in volatile emissions will be associated with shifts in pollinator assemblages. While further work needs to be done to test for local adaptation in this system, this preliminary evidence is consistent with the formation of pollination ecotypes in the E. ghellinckii species complex.
... The family Scarabaeidae (Coleoptera) comprises a large group of beetles, many of which feature larvae that feed on decaying wood in forested areas, thereby playing an important role in recycling organic material (Garcia et al. 2013). Some species of this group are pollinators in the adult stage (Maia and Schlindwein 2006;Cavalcante et al. 2009;Maia et al. 2013), while others may be pests of several crops (Oliveira et al. 2007;Santos and Avila 2009;Coutinho et al. 2011;Rodrigues et al. 2011). ...
... The family Scarabaeidae (Coleoptera) comprises a large group of beetles, many of which feature larvae that feed on decaying wood in forested areas, thereby playing an important role in recycling organic material (Garcia et al. 2013). Some species of this group are pollinators in the adult stage (Maia and Schlindwein 2006;Cavalcante et al. 2009;Maia et al. 2013), while others may be pests of several crops (Oliveira et al. 2007;Santos and Avila 2009;Coutinho et al. 2011;Rodrigues et al. 2011). ...
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The third instar and pupa of Geniates borelli Camerano, 1894 and Anomala testaceipennis Blanchard, 1856 are described and illustrated. Adults were collected with light traps from September to December in 2009 and 2010 at the experimental farm of the Universidade Estadual de Mato Grosso do Sul, muncipality of Aquidauana, Mato Grosso do Sul state, Brazil. Laboratory rearing of eggs from these adults was conducted to obtain immature insects. An identification key to the known third instars of Rutelinae of the New Word is also provided.
... Within the Caladieae, Caladium is the only other genus adequately investigated in regards to aspects of reproductive biology (Maia and Schlindwein, 2006) and floral scent composition (Maia et al., 2012;Maia unp. data). ...
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The Neotropical aroid genus Xanthosoma (Caladieae, Araceae) embraces about 140 species of tuberous herbs whose night-blooming inflorescences emit strong, pleasant floral scents, presumably involved in the atttaction of pollinators. The number of studies on the floral scent chemistry of aroids, as well as on their role in pollinator attraction, has undergone a considerable increase in the last decade. In Xanthosoma, however, neither have never been properly assessed. In the present study, we provide an overview on the reproductive and pollination biology of Xanthosoma species and add new reports on the pollinators of some species. Furthermore, we investigate the floral scent chemistry of 13 species through dynamic headspace sample collection and analysis by gas chromatography coupled to mass spectrometry (GC-MS). A thorough literature survey and novel data from our own personal findings revealed that floral visitors are known for only 12 species of Xanthosoma, which represent less than 10% of its currently documented species diversity. Cyclocephaline scarabs (Melolonthidae, Cyclocephalini) were reported in inflorescences of 11 of the 12 species and are assumed to be the main pollinators associated with them. A total of 76 VOCs, mainly terpenes and benzenoids, were detected. Floral scent bouquets diverge considerably among analyzed species of Xanthosoma mainly due to the presence of unique prominent compounds that ensure originality of the blends rather than because of their quantitative complexity (number of constituents), as analysed samples were typically dominated by few (2–4) VOCs that together accounted for >80% of the total relative scent discharge. A number of these dominant VOCs had already been isolated in other species of aroids and reported as effective attractants of cyclocephaline scarabs in field biossays, namely (Z)-jasmone, p-vinylanisole and dihydro-β-ionone and we speculate that along with other dominant compounds they are directly involved in pollinator selection in Xanthosoma.
... Such a long flowering period is rare in the subfamily Aroideae (species with unisexual flowers) and only known to occur in Arisarum (Herrera 1988 ) and Ambrosina (Killian 1929). Contrary to Arisaema, most of the genera of the Aroideae, such as Arum, Alocasia, Amorphophallus, Caladium, Colocasia, Helicodiceros, Peltandra, and (Yafuso 1993; Patt et al. 1995; Kite et al. 1998; Seymour et al. 2003; Ivancic et al. 2004; Sivadasan and Kavalan 2005; Maia and Schlindwein 2006). In Araceae, a long flowering period is generally characteristic of genera with bisexual flowers such as Anaphyllopsis (Chouteau et al. 2006 ), Anthurium (Croat 1980), or Symplocarpus (Wada and Uemura 2000). ...
... In this case, pollinators meet in the inflorescence to mate and eat floral parts (e.g. Maia and Schlindwein 2006, Seymour et al. 2009, Maia et al. 2010. The third pollination type is a nursery pollination mutualism and is associated with Diptera. ...
... Of the several species observed in Homalomena (Homalomeneae: Araceae) on Borneo, these species, which are pollinated by scarab beetles, also flowered before dawn (Kumano & Yamaoka 2006;Tung et al. 2010;Hoe et al. 2011). Elsewhere in the related tribe Philodendreae, which are pollinated by scarab beetles, the major events occur at dusk (Gibernau et al. 1999;Maia & Schlindwein 2006). It seems that Schismatoglottideae and Homalomeneae tend to flower at dawn in the Old World tropics but related taxa in the Neotropics flower at dusk. ...
Article
Homoplastic evolution of 'unique' morphological characteristics in the Schismatoglottideae - many previously used to define genera - prompted this study to compare morphology and function in connection with pollination biology for Aridarum nicolsonii, Phymatarum borneense and Schottarum sarikeense. Aridarum nicolsonii and P. borneense extrude pollen through a paired of horned thecae while S. sarikeense sheds pollen through a pair of pores on the thecae. Floral traits of spathe constriction, presence and movement of sterile structures on the spadix, the comparable role of horned thecae and thecae pores, the presence of stamen-associated calcium oxalate packages, and the timing of odour emission are discussed in the context of their roles of pollinator management. Pollinators for all investigated species are determined to be species of Colocasiomyia (Diptera: Drosophilidae). This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
... On the other hand, the eight tropical species (six Philodendron species, one Caladium and one Dieffenbachia) are all pollinated by mutualistic crepuscular/ night scarab beetles (Cyclocephala spp.). These beetles feed and copulate within the inflorescences (Young 1986;Gibernau et al. 1999;Maia and Schlindwein 2006;Seymour and Gibernau 2008;Maia et al. 2010). It should be noted that the observed pollination similarity in these tropical species is an evolutionary convergence since the three genera belong to three independent tribes (Chartier et al. 2014). ...
Article
Staining and histochemistry of Volatile Organic Compounds (VOCs) were performed at different inflorescence developmental stages on nine aroid species; one temperate, Arum italicum and eight tropical from the genera Caladium, Dieffenbachia and Philodendron. Moreover, a qualitative and quantitative analysis of VOCs constituting the scent of A. italicum, depending on the stage of development of inflorescences was also conducted. In all nine species, vesicles were observed in the conical cells of either the appendix or the stamens and the staminodes. VOCs were localised in intracellular vesicles from the early stages of inflorescence development until their release during receptivity of gynoecium. This localisation was observed by the increase of both number and diameter of the vesicles during one week before receptivity. Afterward, vesicles were fewer and smaller but rarely absent. In A. italicum, staining and gas chromatography analyses confirmed that the vesicles contained terpenes. The quantitatively most important ones were the sesquiterpenes but monoterpenes were not negligible. Indeed, the quantities of terpenes matched the vesicles’ size evolution during one week. Furthermore, VOCs from different biosynthetic pathways (sesquiterpenes and alkanes) were at their maximum quantity two days before gynoecium receptivity (sesquiterpenes and alkanes) or during receptivity (isobutylamine, monoterpenes, skatole and p-cresol). VOCs seemed to be emitted during gynoecium receptivity and/or during thermogenesis. FADs, accumulated after pollination in the spadix. These complex dynamics of the different VOCs could indicate specialisation of some VOCs and cell machinery to attract pollinators on the one hand and to repulse/protect against phytophagous organisms and pathogens after pollination on the other hand.
... and Taccarum ulei Engl. and K. Krause (Maia and Schlindwein 2006;Maia et al. 2010Maia et al. , 2013. The life cycle of C. celata, however, is unknown, and the main focus of the present study is to provide detailed structural descriptions of its immature stages and to better understand the life cycle of the species in captivity, so contributing to the formulation of accurate management and conservation plans for native pollinators. ...
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We collected 76 specimens of the Cyclocephala celata Dechambre in Igarassu, Pernambuco, Brazil, in March 2008 for captive breeding and rearing to document its life cycle and to describe the immatures. A total of 98 eggs was obtained from captive-reared insects, each individually enclosed in an egg chamber assembled with the surrounding substrate. Viability was highest in the egg phase (92.8%). Pupae were enclosed in pupal cells. The duration of the life cycle was 164 days (n = 2), and only 2% of the eggs developed into adults. Third instar characters documented for the first time in the genus are: epitorma on the epipharynx, three dorsolongitudinal striae on the right mandible and one fringe of setae on the hypopharynx. Our data support the possibility of captive rearing and breeding of C. celata, contributing to the formulation of accurate management and conservation plans for native pollinators.
... Cyclocephala celata was identified as the only effective pollinator of native populations of Philodendron acutatum in the northern coastal region of the state of Pernambuco, northeastern Brazil. In the same region, Maia and Schlindwein (2006) have reported this cyclocephaline scarab in exclusive association with sympatric and coflowering populations of another aroid, Caladium bicolor (Aiton) Vent. Cyclocephaline scarabs are commonly acknowledged as indiscriminate toward floral host choice (Schatz 1990). ...
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Philodendron acutatum (Araceae) is a hemiepiphyte common to the Atlantic Forest of northeastern Brazil. In two localities, we studied the species' breeding system and associations with flower-visiting insects, along with an analysis of its floral scent composition. The fruit set of self-incompatible P. acutatum was high, more than 90%, and inflorescences were exclusively pollinated by one species of scarab beetle, Cyclocephala celata (Scarabaeidae, Dynastinae). Pollinators are drawn toward the inflorescences at dusk by strong floral fragrances given off during the female phase of anthesis, along with endogenous heating of the spadix, whose temperatures were recorded at more than 11 degrees C above ambient air. Two other species of flower-visiting Cyclocephala were also consistently recovered in blacklight trappings during the flowering period of P. acutatum. The fact that only C. celata was found in association with P. acutatum suggests a local reproductive dependence of the plant to this scarab beetle species. Dihydro-beta-ionone and 2-hydroxy-5-methyl-3-hexanone, a rare volatile molecule so far unreported as a floral compound, together accounted for more than 97% of the unique scent composition of P. acutatum and might be involved in specific attraction of C. celata.
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The Order Coleoptera provides good examples of morphological specializations in the reproductive apparatus, gonadic maturation, and allometry differing between the sexes. The female and male reproductive apparatus has been modified to ensure reproduction between individuals of the same species. The genus Cyclocephala has more than 500 species distributed in America, and Cyclocephala barrerai Martínez is an economically important species in the central part of Mexico. The objective of this work was to study the reproductive system, gonadic maturation, and allometry of C. barrerai. We used light, scanning electron, and laser scanning confocal microscopy to describe the reproductive apparatus and gonadic maturation of females and males. The relationship between adult weight and different parts of the body was established by linear regression. Regardless, the reproductive apparatuses of C. barrerai are like those of other Melolonthidae: the genital chamber, the type II accessory glands, and the ventral plaques of the female and the ejaculator bulb and genital capsule of the males are specific to C. barrerai. The gonads are fully developed when 18 d old. The weight of adult C. barrerai has a positive linear relationship with distinct parts of its body, while the antennae of males are larger than those of the females.
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Floral thermogenesis is a well-documented physiological process. However, floret (or inflorescence) development, thermogenesis and pollinator activity is rarely studied in association. In this study, the relationship between blooming, floral temperature and insect activity in Alocasia odora were observed. Based on the phenotypic characteristics of spathes and development of florets, four stages of blooming were categorized in A. odora: stage I (spathe closing), stage II (spathe unfolding, female florets receptive), stage III (male florets releasing pollen), and stage IV (spathe withering). Temperature analyses revealed that the appendix and male zone were the main thermogenic structures, but with different temporal patterns. The male zone showed a 3-day continuous episode of thermogenesis with three waves, to maintain an internal higher temperature, whereas the appendix showed a circadian rhythm of six cycles in 6 days with a duration of 6–7 h every day. Fruit flies were observed to mate, feed on the inflorescence, and enter the female zone to lay eggs when the spathe just opened at stage II. At the same time, the pollen that they carried from previous visits to another inflorescence was deposited on the stigma of female florets. The patterns of blooming and heating were closely tied to the activities of fruit flies. In particular, thermogenesis and behavior of flower-visiting insects were well synchronized and correlated during inflorescence development. Our results therefore suggest that floral thermogenesis plays an important role in successful reproduction.
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Eu estudei sistemas de polinização altamente especializados envolvendo aráceas Neotropicais e besouros escarabeídeos da tribo Cyclocephalini (Scarabaeidae, Dynastinae), e também elucidei aspectos da ontogenia e desenvolvimento embrionário de espécies do gênero Cyclocephala mantidas em cativeiro. Machos e fêmeas antófilos de Cyclocephala celata e C. latericia apresentam resposta atrativa direta a sinais químico-olfatórios simples, presentes nos odores florais de Caladium bicolor, Philodendron acutatum e Taccarum ulei. Padrões puros de 4-metil-5-viniltiazol e (S)-2-hidroxi-5-metil-3-hexanona, dois compostos raros e biosintenticamente diversos isolados em amostras de fragrâncias florais das três espécies vegetais estudadas, foram utilizados com sucesso na atração seletiva de espécies polinizadoras de Cyclocephala. Quatro espécies do gênero nativas da Floresta Atlântica do Nordeste brasileiro, C. celata, C. distincta, C. latericia e C. paraguayensis, foram mantidas em cativeiro durante todo o ciclo biológico através da aplicação de um protocolo simples e econômico. Foram obtidos ovos viáveis de todas as espécies, e imagos de três destas se desenvolveram a partir de uma dieta larval exclusivamente saprófaga. Perspectivas para a reprodução controlada e em maior escala para espécies polinizadoras de Cyclocephala num futuro próximo oferecem oportunidades interessantes para a gestão ambiental e manejo em sistemas biológicos
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Mating behavior and description of immature stages of Cyclocephala melanocephala (Fabricius, 1775) (Coleoptera: Scarabaeidae: Dynastinae), identification key and remarks on known immatures of Cyclocephalini species. Some species of Cyclocephala Dejean, 1821 are regarded as rhizophagous crop pests and others as beneficial species. The objective of this work was to report the mating behavior and to describe the immature stages of C. melanocephala. The studies were developed at the Universidade Estadual de Mato Grosso do Sul in Cassilândia, Mato Grosso do Sul state, Brazil. Adults were collected with a light trap from September to December 2014 and 2015 to carry out studies of mating behavior, breeding, and descriptions of immature stages. Copulation lasted 10.4 ± 4.3 min and took place from 19:00 to 24:00 h. Some females refused males for mating and moved away from them. Regarding flight period, adults were collected in larger quantities from 20:00 to 23:00 h. Identification keys to immatures of three genera of Cyclocephalini, including several Cyclocephala species are presented.
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Cyclocephaline scarab beetles represent the second largest tribe of the subfamily Dynastinae, and the group includes the most speciose genus of dynastines, Cyclocephala . The period following publication of Sebő Endrődi’s The Dynastinae of the World has seen a huge increase in research interest on cyclocephalines, and much of this research has not been synthesized. The objective of this catalog and bibliography is to compile an exhaustive list of taxa in Cyclocephalini. This paper provides an updated foundation for understanding the taxonomy and classification of 14 genera and over 500 species in the tribe. It discusses the history of cataloging dynastine species, clarifies issues surrounding the neotype designations in Endrődi’s revision of Cyclocephalini, synthesizes all published distribution data for cyclocephaline species, and increases accessibility to the voluminous literature on the group by providing an easily searchable bibliography for each species. We propose the nomen novum Cyclocephalarogerpauli , new replacement name , for C.nigra Dechambre.
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The cyclocephaline scarabs (Scarabaeidae: Dynastinae: Cyclocephalini) are a speciose tribe of beetles that include species that are ecologically and economically important as pollinators and pests of agriculture and turf. We provide an overview and synopsis of the 14 genera of Cyclocephalini that includes information on: 1) the taxonomic and nomenclatural history of the group; 2) diagnosis and identification of immature life-stages; 3) economic importance in agroecosystems; 4) natural enemies of these beetles; 5) use as food by humans; 6) the importance of adults as pollination mutualists; 7) fossil cyclocephalines and the evolution of the group; 8) generic-level identification of adults. We provide an expanded identification key to genera of world Cyclocephalini and diagnoses for each genus. Character illustrations and generic-level distribution maps are provided along with discussions on the relationships of the tribe’s genera.
Article
Receiver bias in plant-animal interactions is here defined as "selection mediated by behavioral responses of animals, where those responses have evolved in a context outside the interactions." As a consequence, the responses are not necessarily linked to fitness gains in interacting animals. Thus, receiver bias can help explain seemingly maladaptive patterns of behavior in interacting animals and the evolution of plant traits that trigger such behavior. In this review, I discuss principles of receiver bias, show its overlap with mimicry and how it differs from mimicry, and outline examples in different plant-animal interactions. The most numerous and best documented examples of receiver bias occur within plant-pollinator interactions. I elaborate on the ability of some plants to heat up their flowers (i.e., floral thermogenesis) and argue that this trait likely evolved under receiver bias, especially in pollination systems with oviposition mimicry. Further examples include signals in insect-mediated seed dispersal and plant defense through repellence of aphids. These examples show that receiver bias is widespread in different plant-animal interactions. For a broader understanding of the role of receiver bias in those interactions, we need more data on how animals respond to plant signals, the context and evolutionary history of those behaviors, and the evolutionary patterns of plant signals.
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A publicação do livro Artrópodes do Semiárido: Biodiversidade e Conservação1 no ano de 2014 (Bravo & Calor 2014) foi um marco para o conhecimento da diversidade alfa dos artrópodes desse Bioma. Abordar todos ou um grande número de grupos de artrópodes é tarefa impossível para um pesquisador ou um grupo de pesquisadores, como é o caso do grupo que integrou o PPBio Semiárido Invertebrados. Os 21 capítulos do livro de 2014 aborda alguns táxons e nos brinda informação valiosa sobre a presença e distribuição de espécies em diferentes áreas do Semiárido. Este livro é uma atualização sobre a informação da diversidade alfa de artrópodes, ou seja, a riqueza de espécies de alguns táxons, além de dois capítulos com informação inédita. São oito capítulos deste livro, seis deles com atualizações sobre a riqueza de espécies no Semiárido: Collembola, Ephemeroptera, Plecoptera, Cerambycidae (Coleoptera), Trichoptera, Psychodidae. Os dois capítulos com informação inédita são os de: 1) Opiliões Laniatores; 2) Dynastinae e Melolonthinae (Coleoptera: Scarabaeoidea, Melolonthidae).
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What are the evolutionary mechanisms and ecological implications behind a pollinator choosing its favourite flower? Sixty-five million years of evolution has created the complex and integrated system which we see today and understanding the interactions involved is key to environmental sustainability. Examining pollination relationships from an evolutionary perspective, this book covers both botanical and zoological aspects. It addresses the puzzling question of co-speciation and co-evolution and the complexity of the relationships between plant and pollinator, the development of which is examined through the fossil record. Additional chapters are dedicated to the evolution of floral displays and signalling, as well as their role in pollination syndromes and the building of pollination networks. Wide-ranging in its coverage, it outlines current knowledge and complex emerging topics, demonstrating how advances in research methods are applied to pollination biology.
Book
Allee effects are broadly defined as a decline in individual fitness at low population size or density, that can result in critical population thresholds below which populations crash to extinction. As such, they are very relevant to many conservation programmes, where scientists and managers are often working with populations that have been reduced to low densities or small numbers. There are a variety of mechanisms that can create Allee effects, including mating systems, predation, environmental modification, and social interactions among others. The abrupt and unpredicted collapses of many exploited populations is just one illustration of the need to bring Allee effects to the forefront of conservation and management strategies. This book provides an overview of the topic, collating and integrating a widely dispersed literature from various fields: marine and terrestrial, plant and animal, theoretical and empirical, academic and applied. © F. Courchamp, L. Berec, and J. Gascoigne 2008. All rights reserved.
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Abstract The scarab beetle tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) is the second largest tribe of rhinoceros beetles, with nearly 500 described species. This diverse group is most closely associated with early diverging angiosperm groups (the family Nymphaeaceae, magnoliid clade, and monocots), where they feed, mate, and receive the benefit of thermal rewards from the host plant. Cyclocephaline floral association data have never been synthesized, and a comprehensive review of this ecological interaction was necessary to promote research by updating nomenclature, identifying inconsistencies in the data, and reporting previously unpublished data. Based on the most specific data, at least 97 cyclocephaline beetle species have been reported from the flowers of 58 plant genera representing 17 families and 15 orders. Thirteen new cyclocephaline floral associations are reported herein. Six cyclocephaline and 25 plant synonyms were reported in the literature and on beetle voucher specimen labels, and these were updated to reflect current nomenclature. The valid names of three unavailable plant host names were identified. We review the cyclocephaline floral associations with respect to inferred relationships of angiosperm orders. Ten genera of cyclocephaline beetles have been recorded from flowers of early diverging angiosperm groups. In contrast, only one genus, Cyclocephala, has been recorded from dicot flowers. Cyclocephaline visitation of dicot flowers is limited to the New World, and it is unknown whether this is evolutionary meaningful or the result of sampling bias and incomplete data. The most important areas for future research include: 1) elucidating the factors that attract cyclocephalines to flowers including floral scent chemistry and thermogenesis, 2) determining whether cyclocephaline dicot visitation is truly limited to the New World, and 3) inferring evolutionary relationships within the Cyclocephalini to rigorously test vicarance hypotheses, host plant shifts, and mutualisms with angiosperms.
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Observations and experiments show that Erioscelis emarginata (Dynastinae, Scarabaeidae), the sole pollinator of Philodendron selloum (Araceae), is guided at dusk to the inflorescences in their female stage by a combination of olfactory and visual stimuli. At close range, visual stimuli are induced by olfactory stimuli and the beetles orient themselves by the light inner side of the spathe. Light models are more attractive than dark ones. Nonfragrant spathes or spathe models, which are put closest to the odor sources, are most visited. Odoriferous inflorescences in the female stage emit stimuli for Erioscelis that are present to remain, while in male-stage inflorescences such stimuli are absent. Light inhibits beetles from leaving male-stage inflorescences.
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Examples of beetle pollination in the South American tropics are given. The case histories refer to representatives of the families Cyclanthaceae, Annonaceae, and Araceae. In most cases the cantharophilous syndrome does not seem to be the result of coevolution but of a one-sided adaptation of flowers to the behavior, necessities, and sense-capacities of beetle species. It is emphasized that cantharophily may not necessarily be an indication of ancientness of a flower/insect relationship. Besides primitive angiosperms, there are derived and modern groups which, in their pollination, are specialized for beetles.Carludovica palmata (Cyclanthaceae) is pollinated by small curculionids which are attracted by odors in the early morning hours. Annona coriacea (Annonaceae) and Philodendron selloum (Araceae) are pollinated by large dynastid scarab beetles in the evening hours. In all three species thermogenesis occurs. Although belonging to widely separate systematic groups, Annona flowers and Philodendron inflorescences have a similar pollination biology, which indicates that their adaptations to beetles are the result of convergent evolutionary developments.
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Data on the pollination biology of neotropical aroids are still scarce and concern only a few species (Beath 1999, Croat 1997, Gibernau et al . 1999, 2000; Mayo et al . 1997 and references cited therein). It appears from these studies that Anthurium , Monstera and Spathiphyllum are on the whole pollinated mainly by bees (but see Kraemer & Schmitt 1999), whereas Dieffenbachia , Homalomena , Syngonium , Philodendron and Xanthosoma are generally beetle-pollinated. Although araceous inflorescences may be visited by several insect taxa (Madison 1979), only a few are the legitimate pollinators for each species (Seres & Ramirez 1995, Valerio 1984, Young 1986).
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Inflorescences of the neotropical arum lily, Philodendron selloum, are strongly thermogenic for 2 d during anthesis. Continuous measurements of spadix temperature (Ts) and rate of oxygen consumption (VO2) were made outdoors in whole inflorescences attached to the plants. Some inflorescences were exposed to uncontrolled ambient temperature (Ta) while others were enclosed in clear water-jackets that produced nearly constant ambient conditions. A repeatable, diphasic pattern of heat production appeared, most clearly in water-jacketed inflorescences, and it comprised a short peak phase at sunset followed by a plateau phase that lasted until the following sunset. Regulation of Ts occurred in both phases, but at different levels. Peak phase Ts was regulated in the region of 38–42°C, but plateau phase Ts was usually in the range of 25–36°C. Both VO2 and total heat produced throughout anthesis increased at lower Ta. The data imply that the short peak phase is related to the enhancement of odour production that attracts a single species of large scarabaeid beetle in its native Brazil, and regulation of maximum Ts may prevent overheating. Thermoregulation in the long plateau phase produces equable temperatures inside the inflorescence that may facilitate the resident beetles' activities as a direct energetic reward.
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The Annonaceae show a broader flower biological radiation than originally thought, with flowers being pollinated not only by beetles, but also by thrips, flies and even bees. The majority of species have hermaphroditic protogynous flowers. Species with white or yellowish-white, small, delicate, day-active flowers, may be pollinated either solely by thrips, or by thrips and small beetles (e.g., species of Bocageopsis, Oxandra, Xylopia). Several of these thrips-pollinated species have stamens with an elongated, tongue-like connective. Pollination by flies is not well documented for American species, notwithstanding it appears to be more common in Old World species, e.g., in the genus Pseuduvaria. The mitriform flowers exhale an unpleasant smell, produce nectar in purple-colored petal glands and have a sapromyiophilous syndrome. Flies enter the flower center through large openings between the inner petals. Beetle-pollinated Annonaceae have flowers with comparatively thick and, often, fleshy petals, which, during anthesis frequently form a pollination chamber with the petals inclining over the flower center. The stamens usually have peltate connective shields, probably a device for protection against voracious beetles. Some cantharophilous species have flowers which are day-active while others are night-active. When they are in their pistillate phase, the beetles are attracted by characteristic odor components. They enter the pollination chamber and usually remain in the interior of the flower until the flower has changed to its staminate phase, when pollen is shed and afterwards petals and stamens detach. Two lines of cantharophilous Annonaceae are recognizable on the basis of present knowledge. Species with smaller and more delicate flowers are pollinated by small beetles (Nitidulidae, Curculionidae, Chrysomelidae and Staphylinidae), whereas species with large, more robust flowers in the Neotropics are pollinated by large beetles of the family Scarabaeidae, subfamily Dynastinae. Some species of the cantharophilous Annonaceae, especially the large-flowered ones, but also some species with smaller flowers, produce beat during anthesis (thermogenesis). Food bodies, developed on the adaxial surfaces or sides of the petals, provide unique nourishment possibilities for beetles when they stay inside the flowers during the pistillate phase. In the staminate stage of the flower, after pollen is shed, beetles also feed on pollen. Apparently, no dynastid-flower relationship has evolved in Asia and Australia. Pollination by bees was discovered recently in Unonopsis guatterioides in Amazonia and Uvaria concava in North Queensland: the first by scent-collecting euglossine males and the second by pollen-collecting Meliponinae. The general trends in morphological/functional floral characteristics in the family are discussed in a presumptive phylogenetic context.
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This study describes a pollination system in a species of Araceae that involves three species of beetle, one of which is also a fruit predator. In a tropical cloud forest in Colombia, inflorescences of Xanthosoma daguense opened at dusk, releasing a sweet scent and raising their temperature 1–3 °C. Soon after, two species of Scarabaeidae (Dynastinae; Cyclocephala gregaria and C. amblyopsis) and one species of Nitidulidae (Macrostola costulata) arrived with pollen. Cyclocephala beetles remained inside the inflorescence for 24 h. The next night, Cyclocephala beetles left the inflorescence after picking up the freshly shed pollen, almost always moving to the nearest inflorescence available. The probability of inflorescence abortion and number of fruits set after the visit of one individual was equivalent for both Cyclocephala species. However, C. gregaria was much more abundant than C. amblyopsis, so it was the most important pollinator. There was a positive relationship between the number of dynastine visits and the number of fruits produced. Besides carrying pollen to the inflorescences, nitidulid beetles had a negative effect on female reproductive success through fruit predation. Nitidulid larvae developed inside the infructescence and preyed on up to 64% of the fruits. However, 8% of inflorescences not visited by dynastines were probably pollinated by nitidulids, because hand-pollination experiments showed that self-pollination was unlikely. Inflorescences potentially pollinated by nitidulids comprised 25% of the fruit crop in the year of our study. This interaction with a fruit predator that is also a potential pollinator resembles brood-site pollination systems in which pollinators prey on part of the fruit set (e.g. Ficus, senita cacti, Yucca), making this system substantially more complex than previously described dynastine-pollinated systems in aroids.
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Night-flowering water lilies (Nymphaeaceae) in South America are pollinated by Cyclocephala scarab beetles (Scarabaeidae: Cyclocephalini) in a specialized relationship involving synchronized flowering movements, strong floral scent, food tissues, and heat-producing flowers. We report that a similar and closely related association exists in West Africa between Nymphaea lotus L. and Ruteloryctes morio Fabricius (Cyclocephalini). This finding strongly supports a late Early Cretaceous origin of a symbiosis between the night-flowering water lilies and pollinating Cyclocephalini beetles. We believe that this is the first unambiguous evidence of a plant-pollinator relationship of this age. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 539–543.
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The most prominent pollination mode in neotropical Annonaceae is pollination by beetles. Flowers are protogynous and have fruity, spicy or unpleasant odors. The floral chambers, formed by the petals closing over the flower center, emit these specific odors which entice the beetles to enter the flowers. There, the beetles are sheltered from daylight and predators and encounter food (tissues and pollen) as well as opportunities for mating. The amount of food offered, the thickness of the petal tissue and the dimension of the flowers increases with size, number and voraciousness of the attracted beetles. Three main groups of beetle-pollinated Annonaceae can be distinguished. Two of them have relatively small floral chambers, exhibit diurnal and/or nocturnal anthesis and are visited and pollinated by relatively small beetles (Nitidulidae, Curculionidae, Chrysomelidae). Large flowers with a large pollination chamber and very thick petals are associated with nocturnally active, large and voracious dynastid scarab beetles, which are attracted by strong odors promoted principally by thermogenetic processes of the flowers during the time of the beetles’ main activity. The dynastid beetle-pollinated species appear to have the most adapted and most specialized flower characteristics in the Annonaceae. Some small-flowered Annonaceae are pollinated by thrips and, for a few species, pollination by flies is suspected. Some genera, such as Guatteria, are uniform with respect to flower biology and seem to have adapted principally to nitidulids as pollinators. In contrast, the genus Annona, basically a group pollinated by dynastid beetles, is diversified with respect to flower morphology and pollination. The neotropical Annonaceae as a whole may have started as a group pollinated by sucking and pollen-eating Thysanoptera and non-destructive beetle groups. Species pollinated by such Thysanoptera and/or rove beetles (Staphylinidae) still show the laminar, plesiomorphic stamen type of Annonaceae with a tongue-shaped connective prolongation. The disc-like sclerified connective shield of the majority of Annonaceae is apparently a secondary and modified structure, especially prominent in the beetle-pollinated species. The densely aggregated stamens with their connective shield appear to be a kind of antipredator structure in Annonaceae adapted to pollination by beetles.
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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, beetlepollinated flowers must have evolved by the midlate Cretaceous to join pre-existing guilds of beetlepollinated 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.
Article
Caladium bicolor (Aiton) Ventenat (Araceae) is reported here for the first time from Bangladesh. Recent nomenclature, important synonyms, illustration, flowering and fruiting time, ecology, geographical distribution and occurrence within Bangladesh are provided.
Chapter
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.
Article
Current knowledge about the diversity and biogeography of aroid taxa from central Brazil is here presented and discussed. For practical reasons, the study area was limited to the Cerrado phytogeographic province. In all, 18 genera and 64 species of Araceae were found naturally occurring in the studied area. The genera found were: Philodendron (23 species), Anthurium (11), Xanthosoma (7), Dracontium (3), Monstera (3), Caladium (2), Dieffenbachia (2), Spathicarpa (2), Taccarum (2), as well as Asterostigma, Gearum, Heteropsis, Pistia, Rhodospatha, Scaphispatha, Spathiphyllum, Urospatha, and Zomicarpa (1 species each). The Jaccard similarity index between aroid flora of the Cerrado province and the two main contiguous forested provinces (Mala Atlântica and Amazonia) was calculated for both genera and species. The values obtained show a closer similarity in species composition between the Cerrado and Amazonia (0.127) than between the Cerrado and Mata Atlântica (0.088). However, the generic similarity wilh Mata Atlântica (0.591) is higher than with Amazonia (0.414). Approximalely 32% of the aroid species of central Brazil can be considered endemic to the Cerrado province; the only genus known to be endemic to the Cerrado is Gearum.
Article
This volume provides a comprehensive overview of tropical flower diversity, thereby forming an indication of evolutionary trends. An introductory chapter provides an evolutionary context and introduces tropical flowers. Six chapters then deal with general structural and biological features of flowers and illustrate facets of their diversity: floral organization (structural units and floral symmetry, perianth, androecium, gynoecium, and floral phyllotaxis); floral construction/architecture; adaptation to different pollinators; differentiations associated with pollinator attraction (for example - nectaries, resin glands/flowers, optical displays, and tactile guides); differentiations associated with breeding systems (for example - sex expression, dichogamy, herkogamy, heterostyly, and agamospermy); and the process of anthesis. Chapter eight then outlines floral diversity and evolution of selected tropical systematic groups: Magnoliales (Magnoliidae); Laurales (Magnoliidae); Aristolochiales (Magnoliidae); Lecythidales (Dilleniidae); Violales (Dilleniidae); Fabales (Rosidae); Gentianales (Asteridae); Scrophulariales (Asteridae); Zingiberales (Zingiberidae); and Orchidales (Liliidae). Next, the salient aspects of flower evolution are reviewed. To conclude, the author underlines the need for research synthesis at all levels. -S.R.Harris
Article
Several reproductive strategies have evolved within the genus Nymphaea. Sexual reproduction is mostly protogynous; flowers are open two to several days (depending on the species), with pollen release usually commencing on the second day. The five subgenera of Nymphaea exhibit differences in floral biology. There is considerable variation in the timing of flower opening, floral odor, flower color, and the form and function of various flower parts. These differences may contribute to genetic isolation between species both through temporal separation of flowering and attraction of different pollinators. While sexual reproduction in many species is dependent on xenogamy or geitonogamy, other species have adaptations to promote autogamy. No species of Nymphaea is known to be agamospermous, but several other modes of asexual reproduction are exhibited: detachable tubers, stolon formation, and proliferations of floral and foliar tissue. Wide-ranging tropical species all avoid being totally reliant on pollinators by employing one or more reproductive alternatives to outcrossing; clearly some obligate outcrossers are limited in their colonizing by pollinator availability. The most important of these reproductive alternatives is autogamy. Wide-ranging temperate species, which mostly rely exclusively on outcrossing for sexual reproduction, probably avoid pollinator limitations by utilizing a broader range of pollinators.
Article
Despite a recent taxonomic treatment of Styrax from North America and the Caribbean, the taxonomy of this genus has remained poorly understood. In an effort to clarify the taxonomy of Styrax from western Texas through Mesoamerica, morphological characteristics of over 2500 specimens from 29 herbaria were examined. The previous treatment emphasized vegetative characters for circumscription, whereas the present revision includes heretofore poorly characterized features of the androecium. This revision comprises 19 species and 24 taxa in the area considered, nearly three times as many species as recognized previously. It includes six new species (S. gentryi, S. incarnatus, S. nicaraguensis, S. radians, S. steyermarkii, S. tuxtlensis), two new subspecies (S. platanifolius subsp. mollis, S. nicaraguensis subsp. ellipsoidalis), three new combinations (S. platanifolius subsp. stellatus, S. platanifolius subsp. texanus, S. platanifolius subsp. youngiae), and two new species names (S. austromexicanus, S. lanceolatus). Two names (S. glabrescens var. pilosus and S. jaliscanus) are lectotypified. Nearly 70% of the species recognized in the revision are narrow endemics. This work has provided the basis for a taxonomic revision of all neotropical Styrax that is currently under way.
Article
Dieffenbachia longispatha is pollinated by scarab beetles (Cyclocephala spp. and Erioscelis sp.) at the La Selva Biological Station, Costa Rica. The 3 most abundant beetle species differed in abundance, recapture rates, pollen loads carried, and behavior at inflorescences, yet the effect of a single visit on fruit-set of D. longispatha (the "pollinator effectiveness' of each species) did not vary. Use of common indices of pollinator contribution to seed set, such as pollinator abundance and the number of seeds produced as a result of a single visit, would lead to erroneous conclusions if applied to Dieffenbachia and its beetle pollinators, or to other pollination systems where there is cumulative floral damage with increasing visitation frequencies. -from Authors
Article
The geographical characteristics of a total of 1839 forest fragments surrounded by sugar cane fields were studied in the Brazilian Atlantic rain forest region of the northeastern state of Pernambuco. The size and shape of the fragments as well as inter-fragment distances and the effects of varying edge width were examined using a geographical information system. The analyses show that the fragments are relatively small and close to each other. Approximately 48% of the rain forest fragments are 100 hectares. Forest fragments are close to each other, as fragments located 50m or less apart formed groups that included ca. 50% of the total forest area. At 350m inter-fragment distance, 98% of the rain forest area was included in groups of fragments. Due to the small size and irregular shape of the fragments, the total area of edge zone exceeds that of the interior habitat when the edge width is ca. 60m. At an edge width of 300m ca. 94% of the total fragment area is edge zone. For conservation purposes, ways of establishing networks of forest fragments connected by corridors and stepping stone fragments are demonstrated using GIS. Simulations using these techniques show that reforestation of sugar cane fields between the forest fragments would considerably increase the area of interior forest habitat and connectivity between fragments.
Article
Spadix temperature was measured in three species of Philodendron: P. acutatum, P. pedatum and P. solimoesense. These species showed two different patterns of spadix temperature during their flowering cycle. In P. acutatum and P. pedatum (subgenus Philodendron), the spadix warmed up twice during the beginning of each flowering night with a temperature not significantly different from that of ambient air between the two peaks. In P. solimoesense (subgenus Meconostigma), the spadix temperature rose up to 14oC above that of ambient air during the first night, then it progressively cooled down but remained 3-6oC above ambient air temperature. We propose that the heat production and the spadix temperature patterns observed may reflect different physiological processes and have a taxonomic significance in the genus Philodendron.
Article
In French Guiana, inflorescences of Philodendron squamiferum Poepp. (Araceae) were regularly visited by the scarab beetle Cyclocephala simulatrix Hölne and (Scarabaeidae, Coleoptera) occasionally by Cyclocephala tylifera Hölne. The flowering cycle lasted 2 days, and the protogynous inflorescence exhibited features typical of beetle polli-nation (floral chamber, food rewards, flower heating). The spadix temperature was measured during the entire flowering cycle in French Guiana and at the Montreal Botanical Garden. Both sets of measurements gave a similar temperature pattern. The spadix warmed up twice at the beginning of each evening with no notable heat production between the two peaks. These two temperature peaks were well synchronized with the arrival and departure of beetles. Résumé : En Guyane française, les inflorescences de Philodendron squamiferum Poepp. (Araceae) sont régulièrement visitées par Cyclocephala simulatrix Hölne (Scarabé-, Coléoptère) et occasionnellement par Cyclocephala tylifera Hölne. Le cycle de floraison dure 2 jours avec une inflorescence protogyne qui possède des caractères typiquement liés à la pollinisation par des coléoptères (chambre florale, ressource alimentaire, production de chaleur). La température du spadice a été mesurée durant toute la durée du cycle floral en Guyane française et au Jardin botanique de Montréal. Les deux types de mesures ont donné des résultats similaires. Le spadice chauffe par deux fois en début des deux soi-rées du cycle floral avec aucune production de chaleur entre ces deux événements. Ces deux pics de chaleur sont bien synchronisés avec d'une part l'arrivée, et d'autre part le départ des coléoptères de l'inflorescence.
Article
The pollination ecology of Philodendron melinonii was studied at two locations in French Guiana. Inflorescences of P. melinonii were regularly visited by Cyclocephala colasi, a scarab beetle also found in inflorescences of P. solimoesense in the same area. The flowering cycle lasted two days and the inflorescence exhibited features typical for beetle pollination (e.g., floral chamber, food rewards, flower heating). The flowering process is well synchronized with the night activity of Cyclocephala visitors. The spadix temperature was measured during the entire flowering cycle in the field in French Guiana and in greenhouse conditions at the Montreal Botanical Garden. Both measurements gave a similar temperature pattern with a two hour delay at the Botanical Garden. The spadix warmed up twice during the flowering period and its temperature was not significantly different from that of ambient air between the two peaks. These two temperature peaks are also well synchronized with the arrival and departure of the beetles on two consecutive nights.
Article
Spadix temperature was measured in two species of Philodendron: P. melinonii (subgenus Philodendron) and P. solimoesense (subgenus Meconostigma). For each species, the temperature of the male zone, the sterile male zone and the female zone of the spadix were recorded. In both species, the temperature of the male zone warmed up at the beginning of each of the two flowering nights. In P. melinonii, the temperature of the male sterile zone increased the first day but remained not significantly different from that of the ambient air during the second day. The temperature of the male zone warmed up slightly on the second day. In P. melinonii, the temperature of the three zones was not significantly different from that of the ambient air between the two peaks. In P. solimoesense, the temperature of the male zone and sterile zone rose to above that of the ambient air during the first night and then progressively cooled down but remained 3–6°C above the ambient air temperature until the second peak. In both species the temperature of the female zones remained more or less constant during the entire flowering cycle, very close to the temperature of the ambient air. We suggest that the heat production and the spadix temperature patterns observed may reflect a general physiological process common to all species of Philodendron. The biphasic pattern present in the subgenus Meconostigma can be seen as a variant of the ‘two peaks’ pattern, occurring in the subgenus Philodendron, with a ‘plateau’ phase between them. The comparison of the different thermogenic cycles occurring in Philodendron, Arum and Dracunculus seems to indicate some clear evolutionary trends. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 139, 79–86.
Article
The pollination of Philodendron solimoesense (subgenus Meconostigma) was studied in four populations of French Guiana. Flowering is asynchronous within each population during July, and the flowering cycle is a 2-d process. Numerous insects visit Philodendron inflorescences, but the main pollinator seems to be Cyclocephala colasi (Scarabaeidae, Dynastinae). The pollination process displays aspects typical of beetle pollination: the production of heat and of a strong odor, the presence of a food reward (stigmatic secretion and sterile male flowers), and the presence of a copulation chamber. Flower heat production is important (ca. 11 degrees C above the ambient air) and may help to volatilize the fragrance. Attraction and choice-test experiments showed that C. colasi is not likely to depend on chemical information (such as pheromone) to localize conspecifics but may rely instead on stimuli produced by the inflorescences in order to meet mating partners.
Article
In neotropical forests, adults of many large scarab beetle species spend most of their time inside the floral chambers of heat-producing flowers, where they feed and mate throughout the night and rest during the following day, before briefly flying to another flower. Here we measure floral temperatures in Philodendron solimoesense (Araceae) in French Guiana and the respiration rates of Cyclocephala colasi beetles at floral and ambient temperatures, and show that the the beetles' extra energy requirements for activity are 2.0-4.8 times greater outside the flower than inside it. This finding indicates that heat produced by the flower constitutes an important energy reward to pollinators, allowing them to feed and mate at a fraction of the energy cost that would be required outside the flower.
Dynastine scarab beetle pollination in Dieffenbachia longispatha (Araceae) on Barro Colorado Island (Panama) compared with La Selva biological station (Costa Rica)
  • D N Beath
Beath, D. N. (1999) Dynastine scarab beetle pollination in Dieffenbachia longispatha (Araceae) on Barro Colorado Island (Panama) compared with La Selva biological station (Costa Rica). Aroideana 22, 63-71.
The Beetles of the United States (A Manual for Identification) Washington, DC: The Catholic University of Ameri-ca Press Zonal thermogenetic dynamics of two species of Philodendron from two different sub-genera (Araceae)
  • R H Arnett
Arnett, R. H. (1963) The Beetles of the United States (A Manual for Identification). Washington, DC: The Catholic University of Ameri-ca Press, p.1112. Barabé, D., Gibernau, M., and Forest, F. (2002) Zonal thermogenetic dynamics of two species of Philodendron from two different sub-genera (Araceae). Botanical Journal of the Linnean Society 139, 79 – 86.
BR 50670-420 Brazil E-mail: artur_campos_maia@yahoo.com
  • Recife
Recife, BR 50670-420 Brazil E-mail: artur_campos_maia@yahoo.com.br Editor: M. Ayasse Plant Biology 8 (2006) A. C. D. Maia and C. Schlindwein
Controlling White Grubs in Turfgrass
  • M F Potter
  • D A Potter
Potter, M. F. and Potter, D. A. (1999) Controlling White Grubs in Turfgrass. US, Kentucky: Cooperative Extension Service.
Beetles of the world. A key and information system for families and subfamilies. Version 1.0 for MS-Windows
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  • T A Paine
  • E J Zurcher
Lawrence, F. A., Hasting, A. M., Dallwitz, M. J., Paine, T. A., and Zurcher, E. J. (1999) Beetles of the world. A key and information system for families and subfamilies. Version 1.0 for MS-Windows.
Turfgrass Disorder: White Grubs
  • R C Williamson
  • P J Pellitteri
Williamson, R. C. and Pellitteri, P. J. (2004) Turfgrass Disorder: White Grubs. US, University of Wisconsin: Cooperative Extension Publications.
White Grub Pests on Turfgrass. US: The University of Georgia College of Agricultural and Environmental Sciences
  • W Hudson
  • B Sparks
Hudson, W. and Sparks, B. (2002) White Grub Pests on Turfgrass. US: The University of Georgia College of Agricultural and Environmental Sciences, Cooperative Extension Service.
Insects and Flowers: The Biology of a Partnership
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Barth, F. G. (1991) Insects and Flowers: The Biology of a Partnership. New Jersey: Princeton University Press, p. 408.
Controlling White Grubs on Lawns and Turf. US, Alabama: Cooperative Extension System
  • P P Cobb
Cobb, P. P. (1998) Controlling White Grubs on Lawns and Turf. US, Alabama: Cooperative Extension System.
  • Caladium Bicolor
  • Cyclocephala Celata
Caladium bicolor and Cyclocephala celata: Pollination System Plant Biology 8 (2006) 533
The genera of Araceae. Kew: The Trustees
  • S J Mayo
  • J Bogner
  • P C Boyce
Mayo, S. J., Bogner, J., and Boyce, P. C. (1997) The genera of Araceae. Kew: The Trustees, Royal Botanic Gardens, p. 370.