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Examples of some bat-pollinated bromeliads: (a) Alcantarea imperialis, (b) Billbergia horrida, (c) Encholirium spectabile, (d) Pitcairnia albiflos, (e) Pitcairnia recurvata, (f) Tillandsia heterophylla, (g) Puya ferruginea, (h) Vriesea longiscapa, (i), Werauhia cf. sanguinolenta visited by Lonchophylla robusta. Note the pale-colored corolla in all species, and the predominance of the cup-like floral morphology in the members of the Tillandsioideae subfamily (e, f, h, i). Photos: A, C, D, H: Elton M. C. Leme. B: Ana Paula G. Faria. E, F, I, J: Pedro A. Aguilar-Rodríguez. G: Thorsten Krömer. I: Marco Tschapka.
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Background: Chiropterophily encompasses the floral traits by which bats are attracted as the main pollinators. Among the chiropterophilous flowering plants of the New World, Bromeliaceae is one of the most ecologically important families; however, information about the chiropterophilous interaction in this family is still scarce. Aims: We present a...
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... number of bromelia species confirmed or suggested to be bat-pollinated (i.e. with chiropterophilous floral traits) in the literature. Figure 1), and most (Table S2) are reported to possess a 'typical' chiropterophilous flower (short, wide and large flowers with pale petals; Figure 1) and are visited mainly by bats of the genus Anoura, especially at elevations of 1000 m a.s.l. or higher in tropical montane forests. ...
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... number of bromelia species confirmed or suggested to be bat-pollinated (i.e. with chiropterophilous floral traits) in the literature. Figure 1), and most (Table S2) are reported to possess a 'typical' chiropterophilous flower (short, wide and large flowers with pale petals; Figure 1) and are visited mainly by bats of the genus Anoura, especially at elevations of 1000 m a.s.l. or higher in tropical montane forests. ...
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... bat-pollinated bromeliads have pale yellow, green, creamy or white petals, occasionally with a reddish tint on the calyx, for example some species of Alcantarea and Vriesea (Martinelli 1994(Martinelli , 1997Sazima et al. 1999;Moura and Costa 2014;Versieux and Wanderley 2015). At least four distinct flower shapes can be distinguished among such bromeliads: i) the zygomorphic tube-type of some Billbergia, Vriesea, Pitcairnia and Puya species ( Sazima et al. 1999;Kessler and Krömer 2000;Schmid H 2000;Kowalski and Tardivo 2015;Macías-Rodríguez et al. 2007; Scultori Da Silva 2009; Figure 1(b,c,g)); ii) the zygomorphic bell-shape type present in Vriesea and Werauhia and even Tillandsia (Utley 1983;Grant 1995;Leme 1995; Tschapka and von Helversen 2007; Figure 1(f,h,i)); iii) the heliciform actinomorphic flower with strap-like petals and protruding stamens in Alcantarea and Pseudalcantarea (Martinelli 1994;Krömer et al. 2012;Versieux et al. 2012;AguilarRodríguez et al. 2014; Figure 1(a)); and iv) the brush- type flower of Encholirium ( Sazima et al. 1989;Christianini et al. 2013;Queiroz et al. 2016;Gomes et al. 2018; Figure 1(c); but see Encholirium horridum in Hmeljevski et al. 2017). ...
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... bat-pollinated bromeliads have pale yellow, green, creamy or white petals, occasionally with a reddish tint on the calyx, for example some species of Alcantarea and Vriesea (Martinelli 1994(Martinelli , 1997Sazima et al. 1999;Moura and Costa 2014;Versieux and Wanderley 2015). At least four distinct flower shapes can be distinguished among such bromeliads: i) the zygomorphic tube-type of some Billbergia, Vriesea, Pitcairnia and Puya species ( Sazima et al. 1999;Kessler and Krömer 2000;Schmid H 2000;Kowalski and Tardivo 2015;Macías-Rodríguez et al. 2007; Scultori Da Silva 2009; Figure 1(b,c,g)); ii) the zygomorphic bell-shape type present in Vriesea and Werauhia and even Tillandsia (Utley 1983;Grant 1995;Leme 1995; Tschapka and von Helversen 2007; Figure 1(f,h,i)); iii) the heliciform actinomorphic flower with strap-like petals and protruding stamens in Alcantarea and Pseudalcantarea (Martinelli 1994;Krömer et al. 2012;Versieux et al. 2012;AguilarRodríguez et al. 2014; Figure 1(a)); and iv) the brush- type flower of Encholirium ( Sazima et al. 1989;Christianini et al. 2013;Queiroz et al. 2016;Gomes et al. 2018; Figure 1(c); but see Encholirium horridum in Hmeljevski et al. 2017). ...
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... bat-pollinated bromeliads have pale yellow, green, creamy or white petals, occasionally with a reddish tint on the calyx, for example some species of Alcantarea and Vriesea (Martinelli 1994(Martinelli , 1997Sazima et al. 1999;Moura and Costa 2014;Versieux and Wanderley 2015). At least four distinct flower shapes can be distinguished among such bromeliads: i) the zygomorphic tube-type of some Billbergia, Vriesea, Pitcairnia and Puya species ( Sazima et al. 1999;Kessler and Krömer 2000;Schmid H 2000;Kowalski and Tardivo 2015;Macías-Rodríguez et al. 2007; Scultori Da Silva 2009; Figure 1(b,c,g)); ii) the zygomorphic bell-shape type present in Vriesea and Werauhia and even Tillandsia (Utley 1983;Grant 1995;Leme 1995; Tschapka and von Helversen 2007; Figure 1(f,h,i)); iii) the heliciform actinomorphic flower with strap-like petals and protruding stamens in Alcantarea and Pseudalcantarea (Martinelli 1994;Krömer et al. 2012;Versieux et al. 2012;AguilarRodríguez et al. 2014; Figure 1(a)); and iv) the brush- type flower of Encholirium ( Sazima et al. 1989;Christianini et al. 2013;Queiroz et al. 2016;Gomes et al. 2018; Figure 1(c); but see Encholirium horridum in Hmeljevski et al. 2017). ...
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... bat-pollinated bromeliads have pale yellow, green, creamy or white petals, occasionally with a reddish tint on the calyx, for example some species of Alcantarea and Vriesea (Martinelli 1994(Martinelli , 1997Sazima et al. 1999;Moura and Costa 2014;Versieux and Wanderley 2015). At least four distinct flower shapes can be distinguished among such bromeliads: i) the zygomorphic tube-type of some Billbergia, Vriesea, Pitcairnia and Puya species ( Sazima et al. 1999;Kessler and Krömer 2000;Schmid H 2000;Kowalski and Tardivo 2015;Macías-Rodríguez et al. 2007; Scultori Da Silva 2009; Figure 1(b,c,g)); ii) the zygomorphic bell-shape type present in Vriesea and Werauhia and even Tillandsia (Utley 1983;Grant 1995;Leme 1995; Tschapka and von Helversen 2007; Figure 1(f,h,i)); iii) the heliciform actinomorphic flower with strap-like petals and protruding stamens in Alcantarea and Pseudalcantarea (Martinelli 1994;Krömer et al. 2012;Versieux et al. 2012;AguilarRodríguez et al. 2014; Figure 1(a)); and iv) the brush- type flower of Encholirium ( Sazima et al. 1989;Christianini et al. 2013;Queiroz et al. 2016;Gomes et al. 2018; Figure 1(c); but see Encholirium horridum in Hmeljevski et al. 2017). ...
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... the latter species, the petals form an open corolla, in which only a bat can contact both the exerted anthers and the stigma simultaneously during a visit to the flower, while hummingbirds fail to do this due to their visitation behaviour and because of changes to floral parts following anthesis, for example the stigma pointing downwards due to turgor loss in the style, so the hummingbird does not come into direct contact with it (Aguilar- ). Pollen of bell-shaped chiropterophilous bromeliad flowers, for example in Werauhia and Vriesea, is placed on the head of the bat (Figure 1(k)), while in actinomorphic helicoiform flowers, such as in Pseudalcantarea macropetala, it is deposited on the ventral side of the wings (Aguilar- ). ...
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Based on the conceptual framework of pollination syndromes, pollination networks should be composed of well‐delimited subgroups formed by plants that diverge in floral phenotypes and are visited by taxonomically different pollinators. Nevertheless, floral traits are not always accurate in predicting floral visitors. For instance, flowers adapted to...
Citations
... They contribute significantly to the floristic diversity of vascular epiphytic floras in the Neotropics (Cascante-Marín and Nivia-Ruíz, 2013). Bromeliads possess specialized pollination systems that involve vertebrate pollinators (hummingbirds and nectarivorous bats) and insects to a lesser degree, mainly bees (Benzing, 2000;Kessler and Krömer, 2000;Aguilar-Rodríguez et al., 2019a;Kessler et al., 2020). Even though most bromeliads exhibit adaptations for cross-pollination, nearly two-thirds of the species investigated for their reproductive systems are capable of selfing. ...
... updated) of epiphytic life-form and distributed mainly on the mountains of southern Central America (Costa Rica and Panama) (Grant, 1995;Morales, 2003). Previous studies in Werauhia indicate the presence of specialized pollination systems involving nocturnal nectarivorous bats (Aguilar-Rodríguez et al., 2019a) and hummingbirds (Lasso and Ackerman, 2004), as well as high selfing ability in W. gladioliflora (Cascante-Marín et al., 2005;Tschapka and von Helversen, 2007), W. nutans and W. noctiflorens (Aguilar-Rodríguez et al., 2019b), and W. sintenisii (Lasso and Ackerman, 2004). ...
... Within the Bromeliaceae family, chiroterophily is present in subfamily Pitcairnioideae (Pitcairnia) but is better represented in Tillandsioideae, mainly in Pseudoalcantarea and Vriesea, and Werauhia is thought to be the genus with the greatest specialization in bat pollination (reviewed by Aguilar-Rodríguez et al., 2019a). Fenster and Martén-Rodríguez (2007) suggested that specialized pollination is frequently associated with floral mechanisms to self-pollinate; however, several examples indicate that for bat pollination such association is weak. ...
Plants with specialized pollination systems frequently exhibit adaptations for self-pollination, and this contradictory situation has been explained in terms of the reproductive assurance function of selfing. In the Neotropics, several plant lineages rely on specialized vertebrate pollinators for sexual reproduction, including the highly diverse Bromeliaceae family, which also displays a propensity for selfing. Thus far, the scarce evidence on the role of selfing in bromeliads and in other neotropical plant groups is inconclusive. To provide insights into the evolution and persistence of self-fertilization in the breeding systems of Bromeliaceae, we studied four sympatric epiphytic species from the genus Werauhia (Tillandsioideae) in Costa Rica. We documented their floral biology, pollination ecology, and breeding systems. We estimated the contribution of selfing by comparing the reproductive success between emasculated flowers requiring pollinator visits and unmanipulated flowers capable of selfing and exposed to open pollination across two flowering seasons. The studied species displayed specialized pollination by nectar-feeding bats as well as a high selfing ability (autofertility index values > 0.53), which was attained by a delayed selfing mechanism. Fruit set from natural cross-pollination was low (<26% in both years) and suggested limited pollinator visitation. In line with this, we found a very low bat visitation to flowers using video-camera recording, from 0 to 0.24 visits per plant per night. On the contrary, the contribution of selfing was comparatively significant since 54-80% of the fruit set from unmanipulated flowers can be attributed to autonomous self-pollination. We concluded that inadequate cross-pollination services diminished the reproductive success of the studied Werauhia, which was compensated for by a delayed selfing mechanism. The low negative effects of inbreeding on seed set and germination likely reinforce the persistence of selfing in this bromeliad group. These results suggest that selfing in bat-pollinated bromeliads may have evolved as a response to pollinator limitation.
... Puya flowers are generally long, tubular, and wide, which allows for many hummingbird species to access the nectar (Smith 1969, Krömer et al. 2006, Gonzalez & Loiselle 2016. Many researchers report that hummingbirds are the main floral visitors and pollinators of several Puya species (García-Meneses & Ramsay 2012, Hornung-Leoni et al. 2013, Restrepo-Chica & Bonilla-Gómez 2017, Gonzalez et al. 2019, Kessler et al. 2020, Velásquez-Noriega et al. 2020; however, a few species of bats and moths also consume their nectar and might act as pollinators (Hornung-Leoni & Sosa 2005, Krömer et al. 2006, Aguilar-Rodríguez et al. 2019. Passerine birds were observed using the plant's inflorescences as well, mainly as perches, but they also chew the corollas, often destroying the flowers (Rees & Roe 1980, Salinas et al. 2007, Hornung-Leoni et al. 2013, Velásquez-Noriega et al. 2020. ...
... Regarding nocturnal floral visitors, none have been reported, although the characteristics of the flowers and its nectar, such as the greenish-yellow petals, the nectar composition, and the concentration of sugar, would appear to be adaptations to attract nocturnal visitors. However, only 7 % of all bromeliads in Bolivia are known to be pollinated by bats (Kessler & Krömer 2000), suggesting that there might be many more undetected cases (Aguilar-Rodríguez et al. 2019). ...
Background:
The terrestrial bromeliad Puya ctenorhyncha is a near-threatened endemic species that grows between 2,500-4,050 m asl.
Hypotheses:
Hummingbirds are the most important visitors of this plant given its morphological and floral traits.
Study site:
The study was carried out along the “Death Road”, located in the Yungas montane cloud forest, La Paz, Bolivia.
Methods:
We monitored an average of 216 individuals monthly for two years (2018-2019). Besides we conducted direct observations of focal plants to quantify the visits of animals.
Results:
The reproductive season lasted from April to August in the first year and from May to June in the second. The mean nectar volume was 8.09 ± 1.59 μL per flower and its composition included fructose, glucose, and 86 % of water. With 341 hours of observations, we determined that Coeligena torquata (Trochilidae) is the main visitor and putative pollinator, followed by C. violifer. Males of both species were more frequent visitors than females. Coeligena torquata showed the highest activity during the afternoon while C. violifer was more active in the morning.
Conclusions:
Floral phenology seems to be annual and unimodal. Although the main pollinators seem to be hummingbirds, the nectar concentration and composition is more similar to bromeliad species of other genera which are pollinated by bats or passerine birds. It is important to evaluate, in future studies, the dependence of both hummingbirds and their behavioural response towards the plant.
... Bromeliads fascinate with their spectacular inflorescences, colorful bracts, and beautiful flowers, and increase their charm even more when we observe that they are visited by a hummingbird, bat, or an insect (Aguilar-Rodríguez et al., 2019;Kessler et al., 2020). However, there is a genus of the subfamily Bromelioideae that is somewhat shy because it is hardly possible to observe its hidden flowers with the naked eye as their inflorescences are not present on a conspicuous peduncle but located at the base of the leaves. ...
Bromeliads fascinate with their spectacular inflorescences, colorful bracts, and beautiful flowers, and increase their charm even more when we observe that they are visited by a hummingbird, bat, or an insect. However, there is a genus of the subfamily Bromelioideae that is somewhat shy because it is hardly possible to observe its hidden flowers with the naked eye as their inflorescences are not present on a conspicuous peduncle but located at the base of the leaves. The identity of its pollinators is also still a mystery and basically nothing is known about the sexual reproductive biology of this genus. We refer to Greigia Regel, whose large members have leaves arranged in a rosette pattern, more or less caulescent, of terrestrial and/or rupicolous habit, leaves with marginal spines, the flowers tubular, purple or white and borne in flattened inflorescences that are deeply arranged in the axils of the leaves. The inflorescences are usually covered by accumulated organic matter (e.g., leaf litter), so their presence is hardly visible. This is why its 36 unspectacular species, distributed mainly in humid montane forests from Mexico to Chile have gone largely unnoticed by botanists, resulting in few collections and a low number of herbarium specimens.
... We coded pollination syndromes based on flower and floral bract traits, following Neves et al. (2020a), who validated the utility of pollination syndromes in Vriesea based on information on confirmed pollinators from the literature. There are some Vriesea species known to be visited and/or pollinated by both hummingbirds and bats (Sazima et al. 1995, Aguilar-Rodríguez et al. 2019. These studies show that bat-pollinated flowers can be visited by hummingbirds at dawn, but they feed on the small amount of nectar left by the bats in withered flowers. ...
... With this analysis we aim to detect if plant species from hummingbird and bat pollination syndromes are spatially segregated or not. Because distribution patterns are likely correlated to preferences of habitat, physiology, and other ecological aspects of their specific pollinator species (Aguilar-Rodríguez et al. 2019, Kessler et al. 2020, we used data from the literature on documented Vriesea pollinator interactions to build a plant-pollinator network. Taken together, this approach allowed a better visualization and an integrated discussion of Vriesea spatial distribution and pollinator-specific interactions. ...
... Recent studies identified repeated evolution of pollination syndromes occurring broadly in Bromeliaceae (Givnish et al. 2014, Aguilar-Rodríguez et al. 2019 and in other diverse tropical groups such as the Gesneriaceae and Campanulaceae (Lagomarsino et al. 2017, Serrano-Serrano et al. 2017), Acanthaceae (Tripp and Manos 2008), and Passifloraceae (Abrahamczyk et al. 2014). We found two shifts from bat to hummingbird pollination (Fig. 3A), whereas the reverse pattern (from hummingbird to bat pollination) is more frequently documented across angiosperms. ...
Mutualistic plant-pollinator interactions play a critical role in the diversification of flowering plants. The spatiotemporal correlates of such interactions can be understood in a phylogenetic context. Here, we generate ddRAD-seq data for the highly diverse Vriesea-Stigmatodon lineage to test for correlated trait evolution among pollination syndromes and life form, habitat type, and altitude. Our results show that pollination syndromes are correlated with changes in life form and habitat type. The ancestor of the Vriesea-Stigmatodon lineage was likely bat pollinated, rock dwelling and inhabited open, mid-elevation forests. Transitions from bat to hummingbird pollination are correlated with transitions to the epiphytic life form in shaded habitats, whereas bat pollination is correlated with the rock-dwelling life form and open habitats. Our dated phylogenetic tree reveals independent origins of hummingbird pollination, occurring twice in Vriesea at c. 5.8 and 5.4 Mya. The timing for the shifts in pollination syndrome coincides with geological and environmental transformations across the Serra do Mar Mountain Chain, which increased habitat heterogeneity where Vriesea and their mutualists diversified. The phylogenetic tree reinforces the non-monophyly of taxonomic sections within the genus Vriesea previously defined by flower morphology, indicating that some lineages should be treated as species complexes. This study identifies synergetic drivers of speciation in a tropical biodiversity hotspot.
... rosea show pale to yellow petals. These features are shared with most members of section Xiphion and are typically associated with chiropterophilous syndrome (Moura, 2011;Sazima et al., 1995;Aguilar-Rodríguez et al., 2019;Neves et al., 2020). In fact, bat pollination has been demonstrated in V. bituminosa and reported to V. fenestralis (Aguilar-Rodríguez et al., 2019;Neves et al., 2020). ...
... These features are shared with most members of section Xiphion and are typically associated with chiropterophilous syndrome (Moura, 2011;Sazima et al., 1995;Aguilar-Rodríguez et al., 2019;Neves et al., 2020). In fact, bat pollination has been demonstrated in V. bituminosa and reported to V. fenestralis (Aguilar-Rodríguez et al., 2019;Neves et al., 2020). There is no data on the pollination biology of V. platynema var. ...
... In addition to acoustic cues, floral scents comprise an important sensory signal for the location of flowers by bats (Gonzalez-Terrazas et al., 2016). Currently, reported floral scents associated with chiropterophily in bromeliads are the garlic-like fragrance of opened flowers (Aguilar-Rodríguez et al., 2019;Neves et al., 2020). To our knowledge, no connection has been drawn, experimentally or not, regarding bat pollination and the scent emanated by the floral bract secretions in bromeliads. ...
... The floral features of S. enigmaticus, i.e., red bracts and yellow perianth, exserted stamens and stigma, and diurnal anthesis, are unique in the genus Stigmatodon but common in Vriesea (Costa et al. 2014, Couto et al. 2022. These characteristics, associated with hummingbird pollination syndrome (ornithophily), seem to be the ancestral state among bromeliads, while bat pollination (chiropterophily) originated multiple times in the family as a whole (Aguillar-Rodríguez et al. 2019), as well as in Vriesea (Kessler et al. 2020;Neves et al. 2020), and is supported as an ancestral state in Stigmatodon species (Couto et al. 2022). All the 33 previously known species of Stigmatodon bear chiropterophilous flowers, therefore the floral features of S. enigmaticus could result from the retention of the ancestral state or a reversion to it, a hypothesis that needs to be tested with molecular phylogeny. ...
Stigmatodon enigmaticus is described and illustrated as a new lithophytic taxon from the quartzitic Campos Rupestres within the biodiversity hotspot of the Atlantic Forest, in Minas Gerais state, Southeastern Brazil. This puzzling new species presents floral features that are unique in the genus as they are associated with ornithophily, as in most Vriesea, contrasting with the chiropterous features of all previously known Stigmatodon species. It is assigned to the “S. limae group”, being most similar to S. vexatus, with which it is compared. Stigmatodon enigmaticus is a microendemic species and is here assessed as Critically Endangered due to its restricted habitat, reduced population size, and threats from invasive species. This finding adds to several others that reinforce the relevance of the currently unprotected Serra do Padre Ângelo as a priority for conservation.
... The suborder Megachiroptera has one family, Pteropodidae, containing P. alecto (Nowak 1994). Bats have a vital function in regulating the balance of ecosystems as agents of fruit pollination (Stewart and Dudash 2016;Aziz et al. 2017;Thavry et al. 2017;Lim et al. 2018;Rodriguez et al. 2019;Sheherazade et al. 2019;Ng et al. 2020;Baqi et al. 2021) and seed dispersal, which plays a vital role in forest regeneration and maintenance (Sarmento et al. 2014;Oleksy et al. 2017;Ashwin and Jayakumar 2019;Shah et al. 2021). However, bat meat is also used as a source of food by some people (Mildenstein et al. 2016;Suwannarong and Schuler 2016;Ransaleleh et al. 2020) Wild animal meat may be eaten as halal food by the Minahasa people of North Sulawesi, Indonesia, particularly Christians. ...
Ransaleleh TA, Wahyuni I, Kawatu M, Laatung S, Lambey L, Rembet UNWJ. 2022. Growth of Pteropus alecto bats (Chiroptera: Pteropodidae) in cages. Biodiversitas 23: 6313-6319. Black Flying Fox (Pteropus alecto), known as Paniki yaki, is widely consumed by the residents of Minahasa, North Sulawesi, Indonesia. It is imported from outside the provinces of Gorontalo, Central Sulawesi, and South Sulawesi. The peak of selling bats in Sulawesi is during Thanksgiving Day and religious holidays such as Easter, Christmas, and New Year. Even though the conservation status of this species, according to the International Union for Conservation of Nature (IUCN), is of Least Concern, this species will one day become extinct due to continuous exploration without control. Conservation or cultivation is one of the efforts to preserve this type of bat. Cultivation will be successful when the growth of this mammal is outside the habitat. Therefore, research has been conducted on breeding this animal in cages using five bats. The bats had been taken from the captivity in Wanea Sub-district, Manado city, 58-533 days old. The fruits given as food are papaya, banana, and mango, and the variables measured were the amount of consumption, body weight gain, and morphometry. Furthermore, the data obtained were tabulated and narrated descriptively. The results showed that consumption of P. alecto 1,2,3,4 and 5 fruits were 110.64, 147.86, 192.61, 249.18, and 331.61 g/day, with body weight gain of 0.50, 0.53, 0.96, 1, 17, 1, and 50 g/day. The morphometric growth of P. alecto 1 was total body length 0.10, forearm 0.06, tibia 0.03, ear 0.01 and wingspan 0.08 mm/day. Sequentially, the morphometric growth of P. alecto 2 was 0.11, 0.07, 0.05, 0.01 and 0.12 mm/day, while P. alecto 3 was 0.29, 0.13, 0.06, 0.03, and 0.89 mm/day. The growth of P. alecto 4 was 0.19, 0.02, 0.02.0.01 and 0.89 mm/day, while P. alecto 5 was 0.20, 0.01, 0.01, 0.01 and 0.85 mm/day. In conclusion, the 1st and 2nd P. alecto were in early bone growth, while the 3rd P. alecto was in bone and meat growth, meanwhile, the 4th and 5th P. alecto were in meat and slow bone growth.
... However, the relative abundance of the photosynthesis pathway is very different within the genera; e.g., almost all analyzed species of the genera Aechmea, Billbergia, or Quesnelia show CAM photosynthesis, whereas the species of the genera Alcantarea, Pitcairnia, or Werauhia show only C3 photosynthesis (Crayn et al., 2015). Most bromeliads are pollinated by vertebrates, either hummingbirds or bats (Benzing, 2000;Krömer et al., 2006;Aguilar-Rodríguez et al., 2019). Bat-pollinated (chiropterophilous) species are usually species with C3 photosynthesis, whereas hummingbirdpollinated (trochilophilous) bromeliads show either C3 or CAM photosynthesis (Pierce et al., 2002). ...
Floral nectar contains mainly sugars as well as smaller amounts of amino acids and further compounds. The nectar composition varies between different plant species and it is related to the pollination type of the plant. In addition to this, other factors can influence the composition. Nectar is produced in and secreted from nectaries. A few models exist to explain the origin of nectar for dicotyl plant species, a complete elucidation of the processes, however, has not yet been achieved. This is particularly true for monocots or plant species with CAM photosynthesis. To get closer to such an elucidation, nectar, nectaries, and leaves of 36 bromeliad species were analyzed for sugars, starch, amino acids, and inorganic ions. The species studied include different photosynthesis types (CAM/C3), different pollination types (trochilophilous/chiropterophilous), or different live forms. The main sugars in nectar and nectaries were glucose, fructose, and sucrose, the total sugar concentration was about twofold higher in nectar than in nectaries, which suggests that sugars are actively transported from the nectaries into the nectar. The composition of amino acids in nectar is already determined in the nectaries, but the concentration is much lower in nectar than in nectaries, which suggests selective retention of amino acids during nectar secretion. The same applies to inorganic ions. Statistical analyses showed that the photosynthesis type and the pollination type can explain more data variation in nectar than in nectaries and leaves. Furthermore, the pollinator type has a stronger influence on the nectar or nectary composition than the photosynthesis type. Trochilophilous C3 plants showed significant correlations between the nitrate concentration in leaves and the amino acid concentration in nectaries and nectar. It can be assumed that the more nitrate is taken up, the more amino acids are synthesized in leaves and transported to the nectaries and nectar. However, chiropterophilous C3 plants show no such correlation, which means that the secretion of amino acids into the nectar is regulated by further factors. The results help understand the physiological properties that influence nectaries and nectar as well as the manner of metabolite and ion secretion from nectaries to nectar.
... Ramírez and Hokche (2019) found that the high frequency of protandry recorded in herbaceous-shrubby communities is associated with the relationship between herbaceous species, dichogamy, and the C 4 pathway, which concurs with that registered for the SDF. The high frequency of succulent-CAM-protandrous species matches, to a large extent, with herbaceous-epiphytes that employ specialized pollination systems and are dispersed by wind, as recorded for the main epiphytic families, Orchidaceae and Bromeliaceae (Gravendeel et al. 2004;Nunes et al. 2018;Aguilar-Rodrígueza et al. 2019). In contrast, protandrous-herbaceous-C 4 species are correlated with generalist pollination and dispersal systems. ...
Plant sexual and breeding systems, temporal variations in sex expression, and herkogamy were investigated. Of the 294 plant species surveyed, 73.8% were hermaphroditic, 18.7% monoecious, and 7.5% dioecious. The frequency of adichogamy (71.7%) was higher than that of dichogamy (28.3%) in hermaphrodite and monoecious species, and protandry was more common than protogyny. There was a higher proportion of herkogamous species (80.6%) to non-herkogamous species. Dioecious species were associated with trees, frugivory, and late seral stages, and monoecy was associated with herbaceous life forms, the C4 metabolism, and disturbed habitats. Outbreeding systems were the most abundant in a subsample of 84 plant species: no agamospermy (98.4%), non-spontaneous self-pollination (64.3%), xenogamy (65.4%), and self-incompatibility (39.3%), followed by mixed breeding systems. Non-spontaneous self-pollinated and xenogamous plants were mostly woody species from undisturbed areas dispersed either by frugivores or abiotically. Self-incompatibility was also mainly a strategy of woody species that flowered during the dry period. Mixed breeding strategies: partial spontaneous self-pollination, partial xenogamy, and partial self-incompatibility were more frequent in disturbed areas. Thus, outbreeding was the main form of reproduction in the undisturbed forest. We recommend that these forest relics be maintained as reservoirs in order to protect them in their original state.
... In addition to hummingbirds, other nectar feeding animals with a diurnal behaviour, such as butterflies and bees, also benefit from the sequential flowering and floral resources offered by bromeliad species (Varassin & Sazima 2000;Siqueira Filho & Machado 2001;Machado & Semir 2006). Bats form the second main group of vertebrates acting like bromeliad pollinators, these animals being attracted by floral scent and abundant nectar (Sazima et al. 1989;Benzing et al. 2000;Aguilar-Rodríguez et al. 2019). ...
Background and aims – The Bromeliaceae family has great importance in the maintenance of neotropical communities. In the Brazilian Atlantic Forest, bromeliads are among the major groups responsible for maintaining the local flora and fauna and participate in important ecological interactions with insects, anurans, and hummingbirds. This work reports on aspects of the reproductive biology and the interactions between two endemic bromeliad species from the Atlantic Forest (Aechmea bruggeri and Quesnelia indecora) and their floral visitors to assess the impact of these relationships on the reproductive success and conservation of these plants.
Material and methods – Reproductive phenology, floral biology, pollination experiments, and the reproductive success of both species were investigated. To determine the floral visitors, we made direct observations on flowers and collected floral visitors that could not be identified in the field.
Key results – Aechmea bruggeri and Quesnelia indecora presented the individual and population flowering phenological pattern classified as annual with intermediate duration. The species are partially and totally self-incompatible, respectively. Both species presented a varied visitation guild, and although Q. indecora presented flowers with ornitofilous characteristics, no hummingbirds were recorded for this species. The hummingbird Thalurania glaucopis was the main visitor for Aechmea bruggeri and the bee Trigona cf. braueri was the main visitor for Quesnelia indecora. Nectar thieving by lepidopterans was observed for both species. Pollen robbing by beetles and nectar robbing by bees were registered for Aechmea bruggeri and Quesnelia indecora, respectively. Fruit and seed set of both species were highly affected by herbivory, which may negatively affect their reproductive success.
Conclusion – Our work highlights the important role of bromeliads in neotropical communities, showing how floral visitors and plants interact by participating in maintaining biological diversity in the studied forest remnant.
