Article

Pollination of Aristolochia pallida Willd. (Aristolochiaceae) in the Mediterranean

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Abstract

A first study of the pollination biology of a Mediterranean Aristolochia species in its natural habitat is presented. In all, 183 flowers of Aristolochia pallida Willd. were investigated, which in total contained 73 arthropods, dominated by two groups of Diptera, black fungus gnats (Sciaridae representing 37%) and scuttle flies (Phoridae representing 19%), respectively. However, only Phoridae are regarded as potential pollinators, since pollen has been found exclusively on the body of these insects. All Phoridae belong to the genus Megaselia and are recognised as three morpho-species. The measurements of flower and insect dimensions suggest that size is an important constraint for successful pollination: (a) the insects must have a definitive size for being able to enter the flower and (b) must be able to get in touch with the pollen. Only very few insect groups found in A. pallida fulfil these size requirements. However, size alone is not a sufficient filter as too many fly species of the same size might be trapped but not function as pollinators. Instead, specific attraction is required as otherwise pollen is lost. Since all trapped Phoridae are males, a chemical attraction (pheromones) is proposed as an additional constraint. Since the flowers are protogynous, the record of Megaselia loaded with pollen found in a flower during its female stage proves that this insect must have had visited at least one different flower during its male stage before. Further on, this observation provides strong evidence that the flowers are cross-pollinated. All these factors indicate a highly specialised pollination of A. pallida by Megaselia species.

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... So far, however, studies experimentally testing the deceptive strategies and determining the attractive signals are restricted to a single species, the Mediterranean A. rotunda, where a novel pollination strategy exploiting kleptoparasitic chloropid flies (kleptomyiophly) was discovered (Oelschlägel et al., 2015). Some weakly scented (to the human nose) Aristolochia species with strong male sex-bias in pollinators were suggested to mimic female sex pheromones of flies (Hall and Brown, 1993;Rulik et al., 2008). Other species, such as A. baetica L., A. fimbriata, A. macrophylla Lam., and A. maxima Jacq. ...
... As morphological flower traits (i.e. tube diameter and distance between utricle wall to stamens and stigma) define the size of potential pollinators in Aristolochia (Brantjes, 1980;Rulik et al., 2008), the generally smaller phorids are probably less effective pollinators than the larger drosophilids in A. baetica. ...
... The utricles of collected flowers were opened, the flower phase identified, the trapped arthropods collected and checked for pollen loads under a stereo microscope. Following the most conservative approach, only flower visitors that carried Aristolochia pollen in female-phase flowers were treated as pollinators (Oelschlägel et al., 2015;Rulik et al., 2008;Rupp et al., 2021). The so-called 'interphase' (Berjano et al., 2009) was considered as male phase, since the pollen is already released, although the trapping trichomes are still intact. ...
... Mohrig, 1998 sp. 14 (9) 54 (27) In pairwise comparisons with Ceropegia species, the average dissimilarity in floral scent between R. torulosa and any Ceropegia species was 98% (ANOSIM: R = 0.943, p < 0.001); a total of 10 compounds were also present in the floral scent of Ceropegia (see Table 2; see also [19]). In the NMDS analysis, R. torulosa grouped in the vicinity of C. haygarthii C, carnosa, and C. ampliata ( Figure 2). ...
... It is not known whether Ditassa flowers are deceptive. Representatives of deceptive and pollinator-trapping plant species in other families, such as Aristolochiaceae (Aristolochia; [25][26][27]), Araceae (Arisaema; [28][29][30][31][32]), and orchids (e.g., Pleurothallis [33], Trichosalpinx [34], Lepanthes [35], Pterostylis [36,37]) are likewise known to be visited/pollinated by Phoridae (Megaselia), Sciaridae (Bradysia, Corynoptera, Pseudolycoriella), Chloropidae (Oscinimorpha), and Ceratopogonidae (Forcipomyia). Furthermore, various economically important crops (e.g., Cacao; [38]) depend on the same dipteran families and genera for pollination. ...
... It is not known whether Ditassa flowers are deceptive. Representatives of deceptive and pollinatortrapping plant species in other families, such as Aristolochiaceae (Aristolochia; [25][26][27]), Araceae (Arisaema; [28][29][30][31][32]), and orchids (e.g., Pleurothallis [33], Trichosalpinx [34], Lepanthes [35], Pterostylis [36,37]) are likewise known to be visited/pollinated by Phoridae (Megaselia), Sciaridae (Bradysia, Corynoptera, Pseudolycoriella), Chloropidae (Oscinimorpha), and Ceratopogonidae (Forcipomyia). Furthermore, various economically important crops (e.g., Cacao; [38]) depend on the same dipteran families and genera for pollination. ...
Article
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Elaborated kettle trap flowers to temporarily detain pollinators evolved independently in several angiosperm lineages. Intensive research on species of Aristolochia and Ceropegia recently illuminated how these specialized trap flowers attract particular pollinators through chemical deception. Morphologically similar trap flowers evolved in Riocreuxia; however, no data about floral rewards, pollinators, and chemical ecology were available for this plant group. Here we provide data on pollination ecology and floral chemistry of R. torulosa. Specifically, we determined flower visitors and pollinators, assessed pollen transfer efficiency, and analysed floral scent chemistry. R. torulosa flowers are myiophilous and predominantly pollinated by Nematocera. Pollinating Diptera included, in order of decreasing abundance, male and female Sciaridae, Ceratopogonidae, Scatopsidae, Chloropidae, and Phoridae. Approximately 16% of pollen removed from flowers was successfully exported to conspecific stigmas. The flowers emitted mainly ubiquitous terpenoids, most abundantly linalool, furanoid (Z)-linalool oxide, and (E)-β-ocimene—compounds typical of rewarding flowers and fruits. R. torulosa can be considered to use generalized food (and possibly also brood-site) deception to lure small nematocerous Diptera into their flowers. These results suggest that R. torulosa has a less specific pollination system than previously reported for other kettle trap flowers but is nevertheless specialized at the level of Diptera suborder Nematocera.
... The different species are visited by a wide range of dipteran families, but often information on the actual pollinators is lacking (reviewed in Berjano et al., 2009). However, there is evidence that each Aristolochia species is specialized in just one or few pollinator families (e.g., Phoridae, Drosophilidae, and Chloropidae), and in some species fly attraction is sex-specific (Hime and Costa, 1985;Wolda and Sabrosky, 1986;Hall and Brown, 1993;Rulik et al., 2008;Berjano et al., 2009). Aristolochia species are long known for their spectacular, highly derived trap flowers (Knoll, 1929). ...
... Due to their often obvious and strong scents during the female phase, many authors suggested that Aristolochia flowers generally attract their pollinators by floral scent (Vogel, 1978;Hall and Brown, 1993;Bänziger and Disney, 2006;Trujillo and Séric, 2006;Rulik et al., 2008;Martin et al., 2017), which indeed was substantiated by behavioral assays in a few species (Cammerloher, 1923;Daumann, 1971;Oelschlägel et al., 2015). Based on the type of scent released, it is believed that the flowers generally mimic brood-sites of their respective pollinators, such as carrion, feces, decaying plants, or fungi, by chemical deception (Cammerloher, 1923;Vogel, 1978;Proctor et al., 1996;Martin et al., 2017). ...
... For each flower, we recorded the flowering phase (female or male), and the number of trapped arthropods with and without pollen. Applying the most conservative approach, only arthropods collected from female phase flowers that carried Aristolochia pollen were treated as pollinators (Rulik et al., 2008;Oelschlägel et al., 2015). The inaperturate exine is characteristic of Aristolochia-pollen (unpublished data), and since no other Aristolochia species were co-flowering at the study sites, we assumed that all Aristolochia pollen belonged to A. microstoma. ...
Article
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Deceptive flowers decoy pollinators by advertising a reward, which finally is not provided. Numerous deceptive plants are pollinated by Diptera, but the attractive cues and deceptive strategies are only identified in a few cases. A typical fly-deceptive plant genus is Aristolochia, which evolved sophisticated trap flowers to temporarily capture pollinators. Though rarely demonstrated by experimental approaches, Aristolochia species are believed to chemically mimic brood sites, food sources for adult flies, or utilize sexual deception. Indeed, for most species, studies on scent composition and attractive signals are lacking. In this study, we focused on Aristolochia microstoma, a peculiar Greek endemic with flowers that are presented at ground level in the leaf litter or between rocks and are characterized by a unique morphology. We analyzed flower visitor and pollinator spectra and identified the floral scent composition using dynamic headspace and gas chromatography coupled to mass spectrometry (GC/MS). Female and male phorid flies (Phoridae) are the exclusive pollinators, although the flowers are also frequently visited by Sciaridae, as well as typical ground-dwelling arthropods, such as Collembola and arachnids. The carrion-like floral scent mainly consists of the oligosulphide dimethyldisulfide and the nitrogen-bearing compound 2,5-dimethylpyrazine. These compounds together are known to be released from decomposing insects, and thus, we conclude that pollinators are likely deceived by chemical imitation of invertebrate carrion, a deceptive strategy not described from another plant species so far.
... The floral features of sapromyophily can be identified as follows: the brightly coloured inner side of perianth and the discrete (or dim) colouring of the outer side (dark brown, purple, or green); a perianth with a window area; the presence of osmophores (odorous glands); the missing nectar paths and nectar; etc. (Faegri & van der Pijl 1979;Vogel 1990;Proctor et al. 1996;Burgess et al. 2004). Aristolochia species have peculiar, protogynous flowers which can temporarily trap their pollinators, small dipteran insects from different families (Wolda & Sabrosky 1986;Razzak et al. 1992;Sakai 2002;Burgess et al. 2004;Murugan et al. 2006;Trujillo & Sérsic 2006;Valdivia & Niemeyer 2007;Rulik et al. 2008;Berjano et al. 2009;Hipólito et al. 2012;Stotz & Gianoli 2013;Oelschlägel et al. 2015Oelschlägel et al. , 2016Aliscioni et al. 2017;Martin et al. 2017). These flowers attract flies primarily by their specific scent (Vogel 1990) and by mimicking sex-specific pheromones (Wolda & Sabrosky 1986) or the same scent components that insects (chloropids) use to find their food sources (Oelschlägel et al. 2015). ...
... The absence of eggs belonging to Drosophilidae flies on the inner and outer surfaces of the perianth, and on the surface of stigma and style can be explained by the fact that these flies can lay eggs within floral tissues (Sakai 2002). According to data published in literature, flies of the family Phoridae pupate in flowers of Aristolochiaceae (Disney & Sakai 2001;Rulik et al. 2008). For example, Megaselia metropolitanoensis Disney, 2001 and Puliciphora pygmaea (Borgmeier, 1960) were reared from shed A. pallida Willd. ...
... For example, Megaselia metropolitanoensis Disney, 2001 and Puliciphora pygmaea (Borgmeier, 1960) were reared from shed A. pallida Willd. flowers collected from the forest floor (Rulik et al. 2008). Female M. sakaiae Disney, 2001 lay eggs inside flowers of A. inflata Kunth and A. maxima Jacq. ...
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Background and aims – Interactions of insects with trap flowers of Aristolochia manshuriensis, a relic woody liana with fragmented natural populations from south-eastern Russia, were studied. Pollination experiments were conducted to identify the causes of the poor fruit set in this plant.Material and methods – The study was carried out at two ex situ sites within the natural range of A. manshuriensis in the suburban zone of the city of Vladivostok (Russia). The floral morphology was examined to verify how it may affect the process of pollination in this species. To test for a probability of self-pollination, randomly selected flowers at the female phase of anthesis (day 1 of limb opening) were hand-pollinated with pollen from the same plant. The daily insect visitation was studied. The pollen limitation coefficient and the number of visitors to the flowers were determined. To identify insects that lay eggs on the flowers, the insects were reared from eggs collected from fallen flowers. Both caught and reared insects were identified.Key results – The floral morphology and the colour pattern of A. manshuriensis are adapted to temporarily trap insects of a certain size. The hand-pollination experiment showed that flowers of this plant are capable of self-pollination by geitonogamy and require a pollinator for successful pollination. The positive value (2.64) for the pollen limitation coefficient indicates a higher fruit set after hand-pollination compared to the control without pollination. The number of visitors to the flowers was low (0.17 visitors per flower per day). Insects from three orders were observed on the flowers: Diptera (up to 90.9%), Coleoptera (8.3%), and Hymenoptera (0.8%). Four species of flies (Scaptomyza pallida, Drosophila transversa (Drosophilidae), Botanophila fugax, and Botanophila sp. 1 (Anthomyiidae)) are capable of transferring up to 2500–4000 pollen grains on their bodies and can be considered as pollinators of A. manshuriensis. Data of the rearing experiment indicate that flies of the families Drosophilidae (S. pallida, D. transversa), Chloropidae (Elachiptera tuberculifera, E. sibirica, and Conioscinella divitis), and Anthomyiidae (B. fugax, B. sp. 1) use A. manshuriensis flowers to lay eggs. Beetles were also collected from the flowers, but they were probably not involved in pollination, because no pollen grains were observed on them during our study.Conclusions – Pollinators of A. manshuriensis include mainly Diptera that lay eggs on the flowers. The poor fruit set (2%) in A. manshuriensis is associated with pollen limitation due to the lack of pollinators, as the number of visitors to flowers was extremely low. This may be due to the fact that the flowers of this species are highly specialized on insects of a certain size for pollination.
... The pollinators of Aristolochia are species of diverse families of Diptera, including Anthomyiidae, Calliphoridae, Ceratopogonidae, Chloropidae, Cypselidae, Drosophilidae, Heleomyzidae, Milichiidae, Muscidae, Phoridae, Sarcophagidae, Sepsidae, Syrphidae and Ulidiidae (Cammerloher 1923;Petch 1924;Brues 1928;Lindner 1928;Vogel 1978;Brantjes 1980;Costa & Hime 1981, 1983Wolda & Sabrosky 1986;Razzak et al. 1992;Hall & Brown 1993;Sakai 2002;Burgess et al. 2004;Trujillo & Sérsic 2006;Rulik et al. 2008;Oelschlägel et al. 2015). ...
... Several studies were published on the reproductive biology of different species of Aristolochia (Branjtes 1980;Hall & Brown 1993;Endress 1998;Sakai 2002;Bänziger & Disney 2006;Trujillo & Sérsic 2006;Nakonechnaya et al. 2008;Rulik et al. 2008;Stotz & Gianoli 2013;Oelschlägel et al. 2015); however, anatomical studies are more limited (Costa & Hime 1983;Vogel 1990;Erbar 2014;Erbar et al. 2017). ...
... The flowers of Aristolochia show a pronounced protogyny and attract flies by odour only during the female phase (Oelschlägel et al. 2009). The presence of insects with pollen loads inside the flowers in this phase proves that these insects must have visited at least one different flower during its male stage beforehand (Rulik et al. 2008). A study in A. bracteolata Lam. ...
Article
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The genus Aristolochia L. has a specialised pollination system based on flowers that represent traps for insects. The floral anatomy and micromorphology of the perianth were studied to detect structural characters and secretory cells that are essential for the pollination and reproduction in A. esperanzae, A. fimbriata and A. triangularis. Additionally, we identified the insects collected inside the flowers in natural populations of the north of Argentina. Our observations indicate the presence of osmophores in the limbs of A. esperanzae and A. triangularis, and in the tube of A. fimbriata, because in this last species the fimbriae probably play the first optical attraction role. The papillose cells and epicuticular waxes in the limbs and tube entrances, particularly in A. esperanzae and A. fimbriata, ensure that insects slip into the tube, and the non-secretory conical hairs of the tube, when turgid, guide the insects to the utricle. Idioblasts on the abaxial surfaces of the tubes and utricles were observed in A. esperanzae and A. fimbrita, and probably represent ethereal oil cells that protect flowers from predation. Secretion of sugar was confirmed on all internal surfaces of the utricle in the three species, and was more abundant at the entrance. The insects were diverse species of Diptera (Sarcophagidae, Muscidae, Ulidiidae, Drosophilidae and Anthomyiidae) and Coleoptera (Staphylinidae), although pollen loads on the body were only detected on specimens of Ulidiidae and Sarcophagidae.
... In Aristolochia gigantea, in which in same races a lemon scent overlaps a carrion-like one (Costa and Hime, 1982;Leins and Erbar, 2010), the corresponding chemical compounds are found (dimethyl disulphide, linalool, citral, betacitronellol; Raguso, 2006;Hipólito et al., 2012). In two Aristolochia species, namely Aristolochia littoralis (Hall and Brown, 1993) and Aristolochia pallida (Rulik et al., 2008), either a great part of or all flies found in the flowers are male. The authors discuss an attractive olfactory cue like pheromones. ...
... However, males seeking for mating opportunities might also be attracted and act as pollinators, potentially being rewarded with sexually mature females (Renner, 2006). In the cases of high number of males found in the kettles of Aristolochia littoralis (Hall and Brown, 1993) and A. pallida (Rulik et al., 2008), the authors propose a chemical attraction by pheromones. Most authors have not recorded the sex of the trapped flies, however some studies report a pronounced bias in favour of females (Wolda and Sabrosky, 1986;Carr, 1924;Hime and Costa, 1985;Trujillo and Sérsic, 2006). ...
... In the Aristolochia species stud- ied so far, flies of (more than 20) different families have been reported as pollinators, including Calliphoridae, Ceratopogonidae, Chloropidae, Drosophilidae, Muscidae, Mycetophilidae, Otitidae, Phoridae, Sarcophagidae, and Sciaridae (e.g. Cammerloher, 1923;Daumann, 1971;Razzak et al., 1992;Vogel, 1978;Costa and Hime, 1983;Hilje, 1984;Hall and Brown, 1993;Disney and Sakai, 2001;Sakai, 2002;Burgess et al., 2004;Bello et al., 2006;Trujillo and Sérsic, 2006;Rulik et al., 2008;Berjano et al., 2009;Hipólito et al., 2012;Oelschlägel et al., 2015). Both, in the Tropics and in the Mediterranean region, Phoridae are the most frequently recorded pollinators. ...
Article
The flowers of the eight . Aristolochia species studied in this paper (A. arborea, . A. clematitis, . A. gigantea, . A. grandiflora, . A. rotunda, . A. salvadorensis, . A. sempervirens, . A. tricaudata) represent three different pollinator deceit cases in this genus: brood site mimic in carrion-fly and fungus-gnat blossoms and food source mimic. In any case, all of them have nectaries on the inner wall of the kettle which provide a small quantity of nectar. As regards the histology, two different types of nectaries can be discerned: the trichomatous type and the so-called nectarioles (termed by Vogel, 1998). The latter type, characterized by clusters of independent and isolated glandular units, is reported for the first time in the kettles of . Aristolochia, namely in . A. arborea, . A. salvadorensis and . A. tricaudata.The nectar production in the kettles of the investigated species is regarded as food ensuring survival of the imprisoned pollinators during captivity. Additionally, the location of the nectaries on the inner kettle wall can be interpreted as an adaptation directing the trapped flies to the exit after pollination and fresh pollen loading.
... Aristolochia flowers have been postulated to mimic oviposition sites for flies and generally exhibit a sapromyiophilous or micromyiophilous pollination system mediated through flower scent (Vogel, 1978;Faegri & van der Pijl, 1979;Johnson & J€ urgens, 2010). Other studies discuss mimicry of sex pheromones, as trapped flies were either males (Hall & Brown, 1993;Rulik et al., 2008) or females (Trujillo & S ersic, 2006). The flowers of several tropical species emit unpleasant, carrion-like odors (e.g. A. grandiflora; Burgess et al., 2004). ...
... Female-stage flowers were collected and examined for trapped insects during annual field trips from 2009 to 2012. As A. rotunda flowers are proterogynous, all collected insects from female-stage flowers that carried Aristolochia pollen must have visited an Aristolochia flower in the male stage before and are regarded as pollinators (Rulik et al., 2008; the most conservative approach). All pollinators were identified to family level (Oosterbroek, 2006) and within Chloropidae individual species were identified (Collin, 1946;Narchuk et al., 1970Dely-Draskovits, 1981;Beshovski, 1985;Nartshuk & Andersson, 2013). ...
... Aristolochia pollen was found on all inspected individuals. As only A. rotunda occurs at the study sites and other Aristolochia species that might potentially be present are pollinated by Phoridae (Rulik et al., 2008), the pollen must belong to A. rotunda. The collected fly specimens were deposited at Senckenberg Naturhistorische Sammlungen, Museum f€ ur Tierkunde Dresden (MTD). ...
Article
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Pollination of several angiosperms is based on deceit. In such systems, the flowers advertise a reward that ultimately is not provided. We report on a previously unknown pollination/mimicry system discovered in deceptive Aristolochia rotunda (Aristolochiaceae). Pollinators were collected in the natural habitat and identified. Flower scent and the volatiles of insects (models) potentially mimicked were analyzed by chemical analytical techniques. Electrophysiological and behavioral tests on the pollinators identified the components that mediate the plant-pollinator interaction and revealed the model of the mimicry system. The main pollinators of A. rotunda were female Chloropidae. They are food thieves that feed on secretions of true bugs (Miridae) while these are eaten by arthropod predators. Freshly killed mirids and Aristolochia flowers released the same scent components that chloropids use to find their food sources. Aristolochia exploits these components to deceive their chloropid pollinators. Aristolochia and other trap flowers were believed to lure saprophilous flies and mimic brood sites of pollinators. We demonstrate for A. rotunda, and hypothesize for other deceptive angiosperms, the evolution of a different, kleptomyiophilous pollination strategy. It involves scent mimicry and the exploitation of kleptoparasitic flies as pollinators. Our findings suggest a reconsideration of plants assumed to show sapromyiophilous pollination. © 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.
... The representatives of the genus Aristolochia are well known because of their peculiar flowers, which attract pollinators by odour (Cammerloher, 1923(Cammerloher, , 1933Daumann, 1971;Vogel, 1978;Burgess et al., 2004;Bänziger & Disney, 2006;Trujillo & Sérsic, 2006). Most of the insects attracted to the flowers belong to various families of Diptera, among others Calliphoridae, Ceratopogonidae, Cypselidae, Drosophilidae, Heleomyzidae, Muscidae, Phoridae, Sepsidae and Ulidiinae (Lindner, 1928;Vogel, 1978;Razzak et al., 1992;Sakai, 2002;Burgess et al., 2004;Trujillo & Sérsic, 2006;Rulik et al., 2008). ...
... Once captured, insects are fed by nectar produced by glands located on the utricle wall (Daumann, 1959;Trujillo & Sérsic, 2006). According to Brantjes (1980) and Rulik et al. (2008) pollinators must fulfill the following size requirements: they have to be small enough to pass through the smallest part of the tube but also big enough to be able to interact with the gynostemium to unload and upload pollen grains. Proximally around the gynostemium (Fig. 3b,d) the utricle wall shows a pale translucent circle ('light window';McCann, 1943). ...
... The anthers 'burst explosively' and thus the whole utricle is dusted with pollen grains (Razzak et al., 1992). Insects usually carry most pollen grains on their thorax (Razzak et al., 1992;Burgess et al., 2004;Trujillo & Sérsic, 2006;Rulik et al., 2008). The release of insects takes place 24-48 h after the beginning of anthesis (Razzak et al., 1992). ...
Article
*Catching insects to ensure pollination is one of the most elaborate and specialized mechanisms of insect-plant interactions. Phylogenetically, Aristolochiaceae represent the first angiosperm lineage that developed trap flowers. Here we report the structure and function of specific trichomes contributing to the highly specialized trapping devices. *Investigations were carried out on six Mediterranean Aristolochia species. The morphology and arrangement of the trapping trichomes were investigated by scanning electron microscopy (SEM) and cryo-SEM. To demonstrate frictional anisotropy of the trapping trichome array, a microtribological approach was used. *The results of our experiments support a hypothesis long proposed, but never tested, regarding the trapping mechanism in proterogynous Aristolochia flowers: that an array of highly specialized trichomes arranged eccentrically to the underlying surface is responsible for the easy entrance of insects into flowers but impedes their escape. As they enter the male stage of anthesis, flowers significantly modify their inner surface characteristics, allowing insects to leave. *We have demonstrated the substantial contribution of trapping trichomes to the capture, retention and release of pollinators, an important prerequisite for making cross-pollination possible in most Aristolochia species. Finally, we compare trapping trichomes of Aristolochia with similar structures found in other trapping flowers as well as in pitchers of carnivorous plants not optimized for insect release.
... Some studies (e.g. those on A. littoralis, A. maxima, A. inflata or A. pallida) have revealed that only one or a few dipteran species are pollinators, which points to a high level of ecological specialisation (Hall & Brown 1993;Sakai 2002;Rulik et al. 2008). Nevertheless, when a single species is extensively studied, a higher level of generalisation can be uncovered (e.g. A. grandiflora ;Hart 1897;Cammerloher 1923;Petch 1924;Hilje 1984;Burgess et al. 2004). ...
... Hilje 1984;Hall & Brown 1993;Sakai 2002). Little is known, however, about the pollination of Mediterranean Aristolochia species (Rulik et al. 2008). In Mediterranean ecosystems, environmental conditions are unpredictable (Cowling et al. 1996;Rodó & Comín 2001) and pollinator availability Knuth (1909), Cammerloher (1923, Carr (1924), Petch (1924), Brues (1928), Daumann (1971), Stebbins (1971), Brantjes (1980), Costa & Hime (1981, 1983, Crosswhite & Crosswhite (1984), Hilje (1984), Havelka (1982), Hime & Costa (1985), Wolda & Strabrosky (1986), Siqueira (1988), Razzak et al. (1992), Blanco (2002), Sakai (2002), Burgess et al. (2004), Nakonechnaya et al. (2005), Bä nziger & Disney (2006), Murugan et al. (2006), Trujillo & Sé rsic (2006), Valdivia & Niemeyer (2007) and Rulik et al. (2008) (see bibliography for complete references). ...
... Little is known, however, about the pollination of Mediterranean Aristolochia species (Rulik et al. 2008). In Mediterranean ecosystems, environmental conditions are unpredictable (Cowling et al. 1996;Rodó & Comín 2001) and pollinator availability Knuth (1909), Cammerloher (1923, Carr (1924), Petch (1924), Brues (1928), Daumann (1971), Stebbins (1971), Brantjes (1980), Costa & Hime (1981, 1983, Crosswhite & Crosswhite (1984), Hilje (1984), Havelka (1982), Hime & Costa (1985), Wolda & Strabrosky (1986), Siqueira (1988), Razzak et al. (1992), Blanco (2002), Sakai (2002), Burgess et al. (2004), Nakonechnaya et al. (2005), Bä nziger & Disney (2006), Murugan et al. (2006), Trujillo & Sé rsic (2006), Valdivia & Niemeyer (2007) and Rulik et al. (2008) (see bibliography for complete references). *Data provided by Ollerton (unpublished): Calliphoridae found in A. flos-avis carried pollen loads that were not identified. ...
Article
The pollination of Aristolochia involves the temporary confinement of visitors inside the flower. A literature review has shown that some species are visited by one or a few dipteran families, while others are visited by a wider variety of dipterans, but only some of these are effective pollinators. We observed flowering phenology and temporal patterns of pollinator attendance in diverse populations of Aristolochia baetica and A. paucinervis, two species that grow in SW Spain, frequently in mixed populations. The two species had overlapping floral phenologies, extended flowering periods and long-lived flowers. A. baetica attracted a higher number of visitors than A. paucinervis. Drosophilids and, to a lesser extent, phorids, were the main pollinators of A. baetica, whereas in A. paucinervis, phorids were the only pollinators. Attendance to A. paucinervis flowers by phorids in mixed populations was markedly lower than in pure populations. This effect was more evident in years with lower pollinator density. Our results suggest that A. baetica and A. paucinervis may compete for pollinators in mixed populations.
... One significant factor is the intricate timing and coordination required for successful pollination, with protogyny and specific interactions with insect pollinators playing a crucial role (Razzak et al., 1992;Oelschlägel et al., 2016). Furthermore, the availability and effectiveness of pollinators, particularly small flies, can influence the pollination process and lead to reduced fruit set (Rulik et al., 2008;Aliscioni et al., 2017). Recent studies examining Aristolochia spp. in various ecological settings have contributed to our understanding of pollination mechanisms and adaptations within the Aristolochiaceae family. ...
... For pollination, the pollinator must move freely from the outside to the inside of the flower, and vice versa. There are significant relationships between pollinator size, in terms of either length or width, and the smallest diameter of the perianth tube (Burgess et al., 2004;Trujillo and Seŕsic, 2006;Rulik et al., 2008). Furthermore, A. contorta exhibits several adaptive traits that make it suitable for pollination by Ceratopogonidae spp. ...
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Insects are vital pollinators for angiosperms, playing a crucial role in their reproductive success and fruit production. Aristolochia contorta is a perennial herbaceous vine that occurs in fragmented populations across East Asia. One notable feature of this plant is its trap flower, which employs a unique mechanism to attract, trap, retain, and release insects, ensuring effective pollination. The presence of this trap flower significantly influences the pollination system of A. contorta. Field surveys and pollination experiments were conducted to understand the processes and effectiveness of its pollination mechanism. It was allogamous and was pollinated by the species from Ceratopogonidae. During the insect attraction stage, 11.57% of the flowers contained insects, primarily Ceratopogonidae spp. Most Ceratopogonidae spp. concentrated in few flowers, indicating that although overall attraction might be modest, specific flowers acted as significant focal points for gathering. Trichomes effectively trapped Ceratopogonidae spp. inside flower tubes. In the retention stage, 26.16% of Ceratopogonidae spp. were loaded with pollen grains, but only 7.91% of those exited the flowers in the release stage. The sticky texture of the perianth’s internal cavity posed challenges during this release, leading to adhesion and clogging of the narrow perianth tube. Consequently, a significant portion of Ceratopogonidae spp. became trapped on the perianth wall and perished. This highlights that despite the significant energy and resources invested in flower development, the perianth contributes to the low pollination effectiveness. This study revealed additive factors with negative effects on pollination, including the densely clustered distribution of its pollinators within only a few flowers, insufficient pollen loading onto pollinators, hindered release of entrapped pollinators due to the perianth adhesive surface, and a high rate of defective pollen grains in A. contorta. These factors account for the observed phenomenon of low fruit set (7.7%) and contribute to the diminished rate of sexual reproduction in A. contorta populations. This might lead the species to heavily rely on asexual reproduction, which could potentially lead to gene erosion within populations. The implications of these findings extend to the ecological and conservation aspects, emphasizing the need to understand and conserve the unique pollination system of A. contorta.
... INTRODUCTION Rulik et al. (2008) recorded 73 arthropods trapped in 183 flowers of Aristolochia pallida Willd. at various sites in Italy in 2005. ...
... Phoridae appear to be important pollinators of species of Aristolochia across the world, from North (Hall & Brown, 1993) and South America (Disney & Sakai, 2001;Trujillo & Sérsic, 2006) to Thailand (Disney & Bänziger, 2009). The observations reported above reinforce the report of Rulik et al. (2008) with regard to their similar role in Europe. ...
... The three most predominant components The most abundant insect visitors to N. lutea were phorid fl ies. Phorids are typically recorded in fl owers with moist decaying matter and are noted for feeding on decaying organic matter, but were also recorded as important pollinators of the Araceae where spathes serve as sites for larval development ( Sakai, 2002 ;Rulik et al., 2008 ). In studies on Herrania (Sterculiaceae), phorids transported pollen on the notal and head areas ( Young, 1984 ). ...
... In this study, phorids carried pollen primarily on the proximal areas of pro-and mesothoracic legs. Previous studies showed that phorids are capable of transporting pollen and affecting pollination in the Orchidaceae and Aristolochiaceae ( Borba and Semir, 2001 ;Rulik et al., 2008 ;respectively). For N. lutea , phorids would likely be ineffectual in transferring pollen from bisexual-to female-phase fl owers (cross-pollination), because most (92%) were observed in bisexual-phase fl owers ( Table 2 ) and hence, not moving frequently to femalephase fl owers ( Tables 5 & 6 ). ...
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Floral thermogenesis is an unusual floral trait with a well-documented physiological process, and yet, there is limited understanding of how this trait influences plant reproduction. The current study was undertaken to gain a better understanding of how floral thermogenesis in Nelumbo lutea impacts pollinator attraction and consequent plant reproduction. We conducted field studies on floral thermogenesis and thermoregulation, flower sexual development, floral visitation patterns, breeding system, pollen transfer dynamics, and floral scent production. The most abundant visitors to the thermoregulatory flowers included the Phoridae (Diptera), Chrysomelidae (Coleoptera), and Hymenoptera. Chrysomelid beetles, particularly Diabrotica, were frequent visitors to both first-day female- and second-day bisexual-phase flowers, while phorid flies were most common in bisexual-phase flowers. Pollen transfer experiments indicated that Diabrotica was equally effective in depositing pollen on stigmas, as were the less frequent, but pollen-loaded halictid bees. Flowers received a taxonomically wide assemblage of floral visitors and appear adapted to attract beetles, primarily Chrysomelidae and medium-sized bees. This study is the first to provide strong support that beetles can comprise the dominant portion of floral visitors and are as effective in pollen transfer as bees. Thermogenesis aids in dispersing the main floral scent component-1,4-dimethoxybenzene-attracting both chrysomelids and bees, while thermoregulation causes chrysomelid beetles to actively seek out new flowers for evening residence. This search behavior likely results in chrysomelids affecting cross-pollination.
... Most species in the family produce an odour similar to rotting meat (Endress 1994; Proctor et al. 1996; Burgess et al. 2004; Trujillo and Sérsic 2006). There is a strong association between these kinds of flowers and flies (Diptera), which tend to seek places to lay eggs or to find a mate; thus, flies are attracted to flowers of several species of Aristolochia that deceive their visitors (Sakai 2002; Rulik et al. 2008; Nakonechnaya et al. 2008). Most dipteran families are Phoridae in Aristolochia maxima Jacq., Aristolochia inflata H. B. & K. (Sakai 2002 ), Aristolochia argentina Griseb. ...
... (Nakonechnaya et al. 2008) and Sciaridae in Aristolochia pallida Willd. (Rulik et al. 2008). The association between Aristolochia spp. ...
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The genus Aristolochia presents conserved features in its basic structural plan of trap flowers and in its pollination syndrome. Visitors, usually flies, are attracted to the rotting-meat scented flowers and remain trapped until the second day, when the protogynous flowers release them. Aristolochia gigantea Mart. and Zucc showed many of these floral traits but display a citronella-like odour, giving the opportunity to study the reproductive ecology in this atypical species. Characteristics and behavior of pollinators, coupled with the distances between individual plants and populations, are important factors in patterns of genetic diversity. Thus, our objectives were to study the flower visitors, reproductive success, and gene flow in A. gigantea in the Chapada Diamantina, Bahia, Brazil. Results suggest that pollination by flies remains as a conserved interaction within the genus Aristolochia. The main pollinators appeared to be Megaselia spp. (Phoridae), evidenced primarily by the frequency of their occurrence within flowers and the transference of compatible pollen. On the other hand, the moderate to low levels of intrapopulation genetic variability found in A. gigantea, combined with low effective pollination and its reproductive dependence on biotic vectors, indicate that this species may be suffering gene erosion in the remaining populations located at Chapada Diamantina.
... Although, this observation would not be in accordance with the finding of no double peaks in the sequences or shifts due to indels in the two alleles of the single copy gene. Taking the genetic distance of the hybrid parents into account, one would expect at least double peaks due to allelic variation.Hybridization also highlights other factors which then need to be involved in the building of a hybrid swarm, such as both species need to have the same fly pollinator which would be in contrast toRulik et al. (2007) who postulated species specific pollinators. Another possibility for interspecific pollen transfer would be that both species share individual pollinators from individual pollinator guilds of each species.This would be in accordance withBerjano et al (2009b) who showed that two not closely related Aristolochia species (A. ...
... In addition, hybridization can only act when pollen can be transferred from one parent to the other and cross pollination is not inhibited trough genetic barriers(Thompson 2005). The transfer of pollen is simpler when populations of different species are occurring syntopic especially when pollen is transferred by small flies that might not be able to conquer large distances as is the case in Mediterranean Aristolochia(Rulik et al. 2007, Berjano et al. 2009b. Possible patterns of biogeographical shared areas by populations of different species recovered in the cpDNA tree might indicate that hybridization is acting if the nuclear gene tree is recovering different groupings.Our phylogenetic results, although the statistical support is not always to our satisfaction, clearly demonstrate reticulation through signals pointing into different directions and support biogeographical orientation of potential parents when hybridization was supposed traditionally (see A. maurorum case below) but also reveal many more cases where hybridization might have influenced evolution in the East Mediterranean Aristolochia species. ...
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The Aristolochiaceae are one of the largest angiosperm families, the family has been divided into two subfamilies: Asaroideae, which include Asarum and Saruma, and Aristolochioideae, which includes Thottea sensu lato and Aristolochia sensu lato (Kelly and Gonzales, 2003). Aristolochia sensu lato comprise between 450 and 600 species, distributed throughout the world with centers of diversities in the tropical and subtropical regions (Neinhuis et al., 2005, Wanke et al., 2006a, 2007). However, the extended Mediterranean region including Turkey, the Caucasus and the Near East is likely to be the only diversity hotspot of the genus Aristolochia in the northern hemisphere were up to 60 species and subspecies could be observed (Wanke 2007). Most important contributions to the knowledge of these species were published by Nardi (1984, 1988, 1991, 1993) and Davis & Khan (1961, 1964, 1982), all of these studies were based on morphological characters only. In recent years, with the progress of molecular techniques and in light of the systematic chaos, a detailed study was needed to unravel the evolutionary history prior to a taxonomic revision of this group. The first chapter of my thesis should be regarded as the starting point for more detailed investigation on population level. Preliminary molecular phylogenitic analysis recovered the Mediterranean Aristolochia species as monophyletic (de Groot et al 2006). However, only very few members were included in that study. The latest phylogenetic study by Wanke (2007) dealed with west Mediterranean Aristolochia species and sampled also few members belonging to the east Mediterranean and Caucasian species (3 from Greece, 2 from Georgia and 1 from Turkey). This study reported the Mediterranean Aristolochia species as two molecular and morphologically well supported clades, which were sister to each other. Furthermore, the two closely related species A. sempervirens and A. baetica which have an east west vicariance and are known as Aristolochia sempervirens complex has been recovered as sister group to the remaining west Mediterranean species. A detailed investigation of the evolutionary history of this group is the topic of the second chapter of my thesis (Chapter 2). The Aristolochia sempervirens complex is characterized by an unusual growth form and has a circum Mediterranean distribution. The investigation of these species complex seem to be of great importance to understand speciation and colonization of the Mediterranean by the genus Aristolochia and might shade light in historical evolutionary processes of other plant lineages in the Mediterranean. Furthermore, I test applicability and phylogenetic power of a nuclear single copy gene (nSCG) region to reconstruct well resolved and highly supported gene genealogies as a prerequisite to study evolutionary biology questions in general. Furthermore, a comprehensive overview of leaf epicuticular waxes, hairs and trichomes of 54 species from the old and new world taxa of the genus Aristolochia were investigated using scanning electron microscopy (SEM) to clarify taxonomic status of theses species in contrast to their molecular position. Also this study which is the third chapter of this thesis (Chapter 3), has a strong focus on Mediterranean Aristolochia and tries to provide additional support for molecular findings based on epicuticular waxes and to test them as synapomorphies. Each chapter has its own introduction and abstract resulting in a short general introduction here.
... The genus Aristolochia Linnaeus (1753: 960) is the largest in the family Aristolochiaceae, comprising about 600 species widely distributed across tropical, subtropical, and temperate regions (Huber 1993;Neinhuis et al. 2005;. Due to its remarkable species diversity, unique floral morphology, and complex ecological interactions with pollinators and parasitic insects (Rulik et al. 2008;Zhang et al. 2019;Allio et al. 2021;Alpuente et al. 2023), Aristolochia has been a focal point for botanical and ecological research, leading to numerous recent discoveries. The taxonomic, nomenclatural and systematic studies of this genus have garnered significant scholarly attention (González 1999b;Wanke et al. 2006;Ohi-Toma & Murata 2016;Zhu et al. 2019a). ...
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Aristolochia pulvinata Yi-Fan Wang & Zhi-Jian Yin, a new species of Aristolochia subgenus Siphisia (Aristolochiaceae) from Yunnan Province, southwest China, is described and illustrated. Aristolochia pulvinata is distinguished by its unique characters: an elevated and thickened pulvinate structure in the inferior calyx limbs of its perianth. Morphologically, it resembles A. ovatifolia, A. melanocephala, A. utriformis, and A. luudamcui due to the similar thickening on the calyx limb region, but it differs in terms of degree, surface texture, color, and angle, as well as leaf morphology. This species, discovered and known only from one region, is assessed as Critically Endangered (CR) according to IUCN categories and criteria. This publication includes detailed morphological illustrations of A. pulvinata, with comparative photo plates and a table that highlights distinguishing features relative to similar species.
... Questa pianta appartiene alle specie del genere Aristolochia a fiori solitari (come confronto, A. clematitis presenta fiori a fascetti), con portamento erbaceo non lianoso e con foglie pubescenti (puberule) su entrambe le pagine fogliari. Si tratta di una geofita rizomatosa, caratterizzata da una grossa radice irregolare e tuberiforme (caulorizza), con fusti aerei annui erbacei generalmente quadrangolari (raramente esangolari) (Rulik et al., 2008;Pignatti, 2017). É una specie termofila, tipica delle fasce collinari e montane in boschi caducifogli e loro margini in ambienti a clima submediterraneo (Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959, Aeschimann et al., 2004Mucina et al., 2016), indicata per tutta l'Italia continentale ad eccezione delle regioni Trentino Alto-Adige, Puglia e Basilicata (Portale Flora Italiana, 2023). ...
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The present study updates the distribution of Zerynthia polyxena and its host plant Aristolochia pallida in the alpine region of Aosta Valley, NW Italy. Unpublished data and confirmations of historically known sites are provided and updated to the years 2022-2023. We searched for the two species within 212 cells of one square kilometer, and we confirmed both species to be more widespread in the region than known, with reproductive confirmations for Z. polyxena within 73 cells in an extent of occurrence of 221 km 2 and A. pallida within in 82 cells in an extent of occurrence of 239 km 2. Regarding the altitudinal distribution, rather high observations are reported for both species, with confirmation of reproduction for Z. polyxena up to 1.570 m, the highest altitude ever reported in Italy and Europe, and the presence of A. pallida up to 1.630 m. Z. polyxena was observed on wing between 25th of March and 11th of June, while the earliest data of observations for A. pallida was recorded on the 25th of February. For both species, we report the semi-quantitative results carried out in a sample area that returned density values of considerable interest: 24 adults of Z. polyxena along a transect of 400 m and 50 preimagine instar within a 25 m2 sample area. Finally, some management and conservation aspects concerning Z. polyxena, a protected species by “Habitat” Directive 92/43/EEC, are considered, confirming its presence in Aosta Valley exclusively outside of protected areas of the Natura 2000 network.
... They are easily recognizable as specimens, due to their simple but characteristic wing venation (Mohrig et al., 2012). They play an important role in natural ecosystems (Menzel & Mohrig, 2000;Menzel & Schulz, 2007;Menzel et al., 2020) where larvae are involved in the decomposition of forest litter (Hövemeyer, 1989;Deleporte & Rouland, 1991;Deleporte & Charrier, 1996;Menzel et al. 2020) whereas adults are implicated in the transmission of fungal basidiospores (Schmidt, 1979;Menzel et al., 2020) and in plant pollination (Vogel & Martens, 2000;Rulik et al., 2008;Menzel et al., 2020). However, these small insects are not always beneficial for some ecosystems, because they are the most common greenhouse pests in the order Diptera, damaging edible mushrooms and vegetables. ...
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The present work deals with the spatial distribution of the sciarid species (black fungus gnats) recorded from Morocco throughout the major biogeographical regions: Rif, Eastern Morocco, Atlantic Plain, Middle Atlas, High Atlas, and Anti-Atlas, providing for the first time an atlas of the distribution of Moroccan sciarid fauna. The analysis of the species distribution showed differences between the regions, revealing that the High Atlas and the Rif hosted the greatest specific richness. Of the surveyed sites, forests, crop fields, and aquatic habitats seem to be the most favourable for supporting many sciarid species. Ecological preferences for each species are discussed, indicating a clear preference for medium altitudes ranging from 500 to 1000 m. Alongside the study on the distribution of species, a review of the species recorded in Morocco and gathered from the literature has enabled us to update the checklist of sciarid species, which comprises 65 so far in Morocco. Particular emphasis focused on black fungus gnats considered potentially harmful, with the aim of assessing their distribution in the country, has revealed that Bradysia transitata, B. trivittata, B. xenoreflexa, Lycoriella sativae, Scatopsciara subarmata were collected from strawberry greenhouses (Rosaceae: strawberries), Bradysia placida, B. santorina, B. scabricornis, B. tilicola, B. trivittata and Camptochaeta jeskei were collected from fruit trees (Rosaceae: almond trees), Bradysia santorina, Corynoptera saccata and C. semipedestris were found on palms (Arecaceae: date palms), Scatopsciara atomaria and S. curvilinea were captured in crop fields (Poaceae, Fabaceae), Bradysia scabricornis, Bradysiopsis vittata were collected from Liliaceae, Scatopsciara atomaria was also found on Asteraceae and Solanaceae, and Corynoptera praeparvula was exclusively reported from Solanaceae.
... The phylogenetic relationships argue for a high morphological variation within geographic subclades (Figure 4). The diverse flower morphology may be linked to the specialized pollination syndromes in this group (Rulik et al., 2008). Comparable results were found for the Turkish A. hirta group, indicating that morphological variation could have been conserved in refugial areas or evolved recently, also as result of hybridization (Mahfoud, 2010). ...
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The taxonomy of the Mediterranean Aristolochia pallida complex has been under debate since several decades with the following species currently recognized: A. pallida, A. lutea, A. nardiana, A. microstoma, A. merxmuelleri, A. croatica, and A. castellana. These taxa are distributed from Iberia to Turkey. To reconstruct phylogenetic and biogeographic patterns, we employed cpDNA sequence variation using both noncoding (intron and spacer) and protein-coding regions (i.e., trnK intron, matK gene, and trnK-psbA spacer). Our results show that the morphology-based traditional taxonomy was not corroborated by our phylogenetic analyses. Aristolochia pallida, A. lutea, A. nardiana, and A. microstoma were not monophyletic. Instead, strong geographic signals were detected. Two major clades, one exclusively occurring in Greece and a second one of pan-Mediterranean distribution, were found. Several subclades distributed in Greece, NW Turkey, Italy, as well as amphi-Adriatic subclades, and a subgroup of southern France and Spain, were revealed. The distribution areas of these groups are in close vicinity to hypothesized glacial refugia areas in the Mediterranean. According to molecular clock analyses the diversification of this complex started around 3-3.3 my, before the onset of glaciation cycles, and the further evolution of and within major lineages falls into the Pleistocene. Based on these data, we conclude that the Aristolochia pallida alliance survived in different Mediterranean refugia rarely with low, but often with a high potential for range extension, and a high degree of morphological diversity.
... lertid ikke Sprengel klart å avdekke alle hemmelighetene, for han omtaler også denne arten som en narresaftplante. Seinere undersøkelser har vist at blomstene i pipeurtslekta har nektar (Rulik et al. 2008). Nyere undersøkelser har også avdekka en annen spissfindighet ved pipeurtblomsten som Sprengel overså, nemlig at det i bunnen av blomsten er en gjennomskinnelig del som fører til at det kommer inn lys slik at besøkerne tolker lyset som veien ut og dermed i mindre grad søker tilbake mot den egentlige utgangen. ...
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Grindeland, J.M. 2020. Christian Konrad Sprengel - the father of pollination ecology. Blyttia 78: 165-181. The theologian and teacher Christian Konrad Sprengel is considered by many as the father of pollination ecology. His classic book from 1793, The discovered secret of nature in the form and fertilisation of flowers, was published after a few intense summers collecting data through observations in nature. Sprengel was headmaster of a primary school in Spandau, near Berlin. At the behest of his physician he spent much time in nature and developed a keen interest for botany which probably led him to neglecting his teaching duties and finally, leaving the position as headmaster in 1793. In 1787 Sprengel noticed the tiny hairs at the basis of the petals of Geranium sylvaticum straight above the nectaries. Sprengel thought that «the wise Creator» would not have created anything unnecessary and thus he concluded that the nectar must have an important function for the plants. He soon deduced that nectar is systematically sought out by insects which in the process of collecting nectar will pollinate the flowers. Sprengel went on to conclude that colour and odour of flowers help insects locate the flowers and that many flowers have stripes and dots-honey guides-to further aid the insect in finding the nectar and operating the flower in a useful way from the plant's perspective. With his groundbreaking agenda Sprengel went on to discover many other morphological traits of plants that seem to be aimed at the crosspollination of plants, such as dichogamy, herkogamy, heterostyly, deceit pollination and trap-flowers. These and some additional aspects of pollination ecology that Sprengel details in his book are presented together with a brief discussion of the (lack of) impact of the book and Charles Darwin's rediscovery and advocacy some fifty years after Sprengels death.
... The Sciaridae is one of the largest families of Diptera, rich in both species and individuals, and plays a significant role in natural ecosystems (summarised in Menzel and Schulz 2007). For example, the larvae are important for the litter decomposition in forests (Hövemeyer 1989;Deleporte and Rouland 1991;Deleporte and Charrier 1996), and the adults for the transmission of basidiospores of fungi (Schmidt 1979) and the pollination of plants (Vogel and Martens 2000;Rulik et al. 2008). Sciarids are also well-known as pests in mushroom farms and greenhouses, or as common inhabitants of pot plants in houses (e.g., . ...
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Black Fungus Gnats (Sciaridae) are a megadiverse, cosmopoliltan family of bibionomorph Diptera. Even in Europe, the continent with the longest tradition in sciarid taxonomy, numerous taxonomic issues remain unresolved and countless species await discovery and description. The fauna of Norway is in these respects no exception. Recognising considerable knowledge gaps, the Norwegian Biodiversity Information Centre provided substantial funding for a detailed inventory of the Sciaridae species occurring in Norway, which was realised in 2014–2018. The results of this project will be published in a series of papers, of which the first is presented here, summarising available data on the taxonomy, faunistics, and autecology of Norwegian Sciaridae beginning with Zetterstedt’s pioneering work in 1838 and ending with 31 December 2019 as the cut-off date. All published records from that period were analysed. The result is a list of 143 species and four unplaced names. Following a consistent scheme, verified locality details are provide including alternative spellings, habitats, and flight times of adults in Norway, literature citations for the faunistic records, and general taxonomic references for classification or identification. A checklist of the sciarid fauna of Norway and a complete list of the relevant literature are also presented.
... As espécies são caracterizadas principalmente pela morfologia floral, incluindo perianto monoclamídeo, zigomorfo, com sépalas petaloides fusionadas em uma estrutura tubular, na qual destacam-se o tubo inferior (geralmente denominado de utrículo, bojo ou colo), e o tubo superior (geralmente denominado de tubo), que termina nos lábios que margeiam a fauce (também denominado de limbos), androceu e gineceu fusionados em uma estrutura chamada ginostêmio e ovário ínfero (Hoehne 1927(Hoehne , 1942Abreu 2015). As espécies do gênero apresentam adaptações relacionadas aos mecanismos de polinização por diversos grupos de Dípteros (Sakai 2002;Rulik et al. 2008). ...
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We present the floristic treatment for the species of Aristolochiaceae from the canga of the Serra dos Carajás, state of Pará, Brazil. One genus and three species are recorded: Aristolochia mossii with Amazonian and periamazonic distribution, A. rugosa with distribution in the Cerrado and Amazonia, and Aristolochia aff. disticha with uncertain identification. An identification key, descriptions, illustrations, photos, and comments are also provided. © 2018 Instituto de Pesquisas Jardim Botanico do Rio de Janeiro. All rights reserved.
... Many published studies have focussed on groups of long-tongued insects or birds, describing the positive relationship between the length of corolla tubes and the lengths of insect proboscises (Stang et al. 2009) or bird bills (Cronk and Ojeda 2008). Fewer studies have evaluated the relationship between corolla tube size and size of the whole body of small pollinators in limiting pollinator access to floral rewards (Rulik et al. 2008). More recent studies have reported the mechanisms by which fly pollinators are filtered according to head size in a stapeliad (Shuttleworth et al. 2017) and to body size in an orchid (Martos et al. 2015). ...
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Although Ceropegia species are well known for their complex pitfall flowers that temporarily imprison their pollinators, various aspects of their pollination ecology are still unknown. This study investigated flowering phenology, functional floral traits, and insect visitation in a natural population of a rare endemic lithophyte, Ceropegia thaithongiae Kidyoo. Flowering of C. thaithongiae was not synchronous but staggered. Anthesis lasted mostly 1–2 days, but its duration was shorter in flowers with a pollinium inserted into the stigmatic chamber. Several different insects visited flowers, but only chloropid and milichiid flies were effective pollinators. At anthesis, the epithelial osmophores on the corolla lobes emitted a floral scent that was simple in composition. Nectar of high viscosity was exuded from the nectaries hidden behind the guide rails. When transported by an insect, the pollinarium was attached to bristles on the mouthparts. Size and shape of the thin pellucid margin of the pollinium enable it to fit into the stigmatic chamber in a lock-and-key arrangement. The pollen transfer efficiency was 6.8%. The plant’s staggered flowering and pollination-induced ending of anthesis are advantageous in decreasing competition for pollinators when flower-visiting insects are scarce.
... Although hard to quantify in economic terms, Sciaridae play an important role in the maintenance of ecosystem functions, e.g., as decomposers of plant material (compost, fallen leaves, pine needles, tree bark and deadwood) or animal wastes (cow and horse manure), in food chains and webs (as prey of nematodes, spiders, predatory insects and amphibians) and even as pollinators (Rulik et al. 2008;Vogel & Martens 2000). ...
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Considering taxonomic decisions and other results (published descriptions, synonymies and records) into account, there are currently 101 species of Sciaridae known to occur in the Afrotropics. Of these, 78 species in 23 genera remain to be properly classified and the remaining 23 species have not yet been studied and are therefore, regarded as Sciaridae incertae sedis for the time being. Kirk-Spriggs & Stuckenberg (2009) estimated the Afrotropical fauna to contain 2,000+ species of Sciaridae.
... The vast majority of species of the family Sphaeroceridae are associated with decaying vegetation (Rohá cek et al. 2001). Spaerocerid flies were recognized as pollinators of Araceae, Arecaceae, Aristolochiaceae, etc. (Bernal & Ervik 1996;Rulik et al. 2008;Urru et al. 2011). They were noted as visitors and pollinators of Aspidistra for the first time. ...
Article
The genus Aspidistra comprises about 160 species of herbaceous plants. In previous investigations, different authors suggested that Aspidistra is pollinated by flies, fungus gnats, slugs, amphipods and even collembolans. The vast majority of data was based on only one species, A. elatior, and was not confirmed by direct observations. Recent field observations of anthetic plants confirm the idea of pollination by flies for two other species of the genus. The present study summarizes direct observations of flowering phenology and pollination of five previously unstudied species of Aspidistra. Flowering biology was studied using visual observations as well as video and photo monitoring. Flowers and captured pollinators were investigated using scanning electron microscopy. As a result, knowledge of the flowering biology of Aspidistra has been increased. The absence of heat production and an ultraviolet pattern were verified. The present data show that flowers of Aspidistra can serve as a food resource for animals. The investigated species of Aspidistra are pollinated by dipteran insects: A. formosa, A. marasmioides and A. subrotata are pollinated by fungus gnats; A. multiflora and A. oviflora are pollinated by flies. The occurrence of fungus-gnat pollination was directly demonstrated for the first time in the genus Aspidistra. The present study confirmed the idea of myiophily in the genus Aspidistra. Apparently, myiophily is common and diverse in Aspidistra. More likely, pollinators are attracted by scent, which is undetectable by humans, but the attraction mechanism is still not completely clear.
... In contrast, males accounted for 334 of 349 visitors (mostly Megaselia scalaris, M. aurea and M. perdita) to 32 flowers of the Brazilian species A. littoralis (= elegans) naturalized in Florida, USA (Hall and Brown, 1993). Similarly, all individuals of three pollen-carrying Megaselia morpho-species visiting A. pallida in Italy were male (Rulik et al., 2008). Clearly, further insights on the floral evolution of many Neotropical Aristolochia species will require a deeper understanding of the chemical and reproductive ecology of Megaselia flies, whose 1500 described species represent a small fraction of actual phorid diversity (Disney, 1994). ...
... Insects that carried pollen and were trapped in a female stage flower must have previously been trapped in another flower. Only these insects have proven to repeatedly visit Aristolochia flowers and thus are potential pollinators (Rulik et al. 2008). Therefore, only arthropods found in the utricle of female stage flowers were subject to subsequent analysis. ...
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Pollination success of highly specialised flowers is susceptible to fluctuations of the pollinator fauna. Mediterranean Aristolochia rotunda has deceptive trap flowers exhibiting a highly specialised pollination system. The sole pollinators are kleptoparasitic flies in search of food. This study investigates these pollinators on a spatio‐temporal scale and the impact of weather conditions on their availability. Two potential strategies of the plants to cope with pollinator limitation, i.e . autonomous selfing and an increased floral life span, were tested. A total of 6156 flowers were investigated for entrapped pollinators in 10 Croatian populations. Availability of the main pollinator was correlated to meteorological data. Artificial pollination experiments were conducted and the floral life span was recorded in two populations according to pollinator availability. Trachysiphonella ruficeps (Chloropidae) was identified as dominant pollinator, along with less abundant species of Chloropidae, Ceratopogonidae and Milichiidae. Pollinator compositions varied among populations. Weather conditions 15–30 days before pollination had a significant effect on availability of the main pollinator. Flowers were not autonomously selfing, and the floral life span exhibited considerable plasticity depending on pollinator availability. A. rotunda flowers rely on insect pollen vectors. Plants are specialised on a guild of kleptoparasitic flies, rather than on a single species. Pollinator variability may result in differing selection pressures among populations. The availability/abundance of pollinators depends on weather conditions during their larval development. Flowers show a prolonged trapping flower stage that likely increases outcrossing success during periods of pollinator limitation.
... Sex-biased attraction of Aristolochia pollinators is unclear, as case studies have shown strong preferences of either male or female flies visiting the flowers of different species. Male flies of Megaselia (Phoridae) are the exclusive or predominant pollinators of the Mediterranean A. pallida (Rulik et al., 2008) and the neotropical A. elegans (Hall & Brown, 1993; however, this study should be taken with caution as A. elegans is likely not native from the study site in the United States). Conversely, female Chloropidae, Drosophilidae, Milichiidae and Phoridae flies are the exclusive or predominant pollinators of the neotropical A. labiata (Hime & Costa, 1985), A. maxima (Sakai, 2002), and A. pilosa (Wolda & Sabrosky, 1986). ...
... Phoridae adults often live near decaying vegetation (Richards and Davies 1984), and species of Megaselia have been reported as pollinators of Aristolochia spp. (Hall and Brown 1993, Sakai 2001, Rulik et al. 2008, Berjano et al. 2009), Ceropegia spp. (Ollerton et al. 2009) and some orchids (Borba andSemir 2001, Albores-Ortiz andSosa 2006). ...
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Apium bermejoi is a critically endangered species that is native to Menorca, an island of 702 km2 located in the western Mediterranean basin. We investigated the reproductive biology of this species and evaluated flowering/fruiting phenology and reproductive output in wild subpopulations over four years (2007–2010). We used ex situ cultivated plants to study floral biology and performed bagging experiments to elucidate the breeding system. We addressed the role of ants in seed dispersal/predation in wild subpopulations by analysing the presence of A. bermejoi seeds in the waste accumulated on the refuse piles of all anthills. Our observations indicated that flowers are protandrous but that there is no dichogamy between flowers within and among umbels of the same individual. Moreover, individuals are mixed in the field because the close germination of several seeds and the plant’s caespitose habit promote cross-pollination. Bagging experiments demonstrated that the species is autogamous, and seeds from selfing (bagged with muslin) showed high final germination (>80%). However, the fruit set of selfed umbels (0.3–0.4 per 1) was lower than that of open-pollinated umbels (0.6–0.8 per 1), and the high pollen/ovule (P/O) ratio (3641±133, n= 30) was also indicative of xenogamy. Plants bagged with muslin, bagged with tulle or barred from contact from ants exhibited similar fruit sets, lower than those observed in isolated and control groups, suggesting that anemophily and pollination by flying insects were less effective than ant-based pollination. We identified eight species of floral visitors, and we noted that ants were the most frequent visitors. Although we found some A. bermejoi seeds in the refuse piles of Messor bouvieri anthills, most of the seeds were not damaged (99.1%), and the number of seeds found represents only a small proportion of the overall fruit production. Because sexual reproduction is the main reproductive mechanism of these plants, despite their high potential for space colonisation and vegetative propagation, the reproductive output for each year determines the subpopulation configuration in the following year.
... Insects are kept imprisoned inside the flower by downwardoriented trichomes located at the inner surface of the flower. During the male phase, insect pollinators are released via the loss of turgor in the trichomes (Fuller 1924; Burgess et al. 2004; Trujillo and Sérsic 2006; Rulik et al. 2008; Berjano et al. 2009; Oelschlägel et al. 2009 ). Being protogynous is a successful strategy to avoid self-pollination in self-incompatible species, which is generally the case for Aristolochia species (Fuller 1924; Sakai 2002; Burgess et al. 2004; Trujillo and Sérsic 2006; Valdivia and Niemeyer 2007; Nakonechnaya et al. 2008 ). ...
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... Dipterans commonly serve as pollinators in Aristolochia [42,8384858687, sometimes having specialized, mutualistic relationships involving egg deposition in the flowers88899091 . Dipteran pollinators are thought to be attracted to secondary metabolites mimicking the aroma of a food source [42,91] , or acting as pheromones to attract species-specific , sex-specific pollinators [92], or stimulating oviposition [89]. Aristolochia species are also important host plants for the larval stages of swallowtail butterflies (Papilionidae, Lepidoptera)939495969798. ...
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Background Previous studies in basal angiosperms have provided insight into the diversity within the angiosperm lineage and helped to polarize analyses of flowering plant evolution. However, there is still not an experimental system for genetic studies among basal angiosperms to facilitate comparative studies and functional investigation. It would be desirable to identify a basal angiosperm experimental system that possesses many of the features found in existing plant model systems (e.g., Arabidopsis and Oryza). Results We have considered all basal angiosperm families for general characteristics important for experimental systems, including availability to the scientific community, growth habit, and membership in a large basal angiosperm group that displays a wide spectrum of phenotypic diversity. Most basal angiosperms are woody or aquatic, thus are not well-suited for large scale cultivation, and were excluded. We further investigated members of Aristolochiaceae for ease of culture, life cycle, genome size, and chromosome number. We demonstrated self-compatibility for Aristolochia elegans and A. fimbriata, and transformation with a GFP reporter construct for Saruma henryi and A. fimbriata. Furthermore, A. fimbriata was easily cultivated with a life cycle of just three months, could be regenerated in a tissue culture system, and had one of the smallest genomes among basal angiosperms. An extensive multi-tissue EST dataset was produced for A. fimbriata that includes over 3.8 million 454 sequence reads. Conclusions Aristolochia fimbriata has numerous features that facilitate genetic studies and is suggested as a potential model system for use with a wide variety of technologies. Emerging genetic and genomic tools for A. fimbriata and closely related species can aid the investigation of floral biology, developmental genetics, biochemical pathways important in plant-insect interactions as well as human health, and various other features present in early angiosperms.
... The genus Pseudolycoriella Menzel and Mohrig, 1998 may be closely associated with living plants, as the probably very closely related Eugnoriste Coquillett, 1896 species feed on flowers (Steffan, 1966). According to Rulik et al. (2008), some species of Pseudolycoriella are phytosaprophagous and their adults are Enderlein (1911); (B) Cladogram after Lengersdorf (1928Lengersdorf ( -1930; (C) Cladogram after Frey (1942); (D) Cladogram after Röschmann and Mohrig (1995); (E) Cladogram after Menzel and Mohrig (2000) and frequently found in Aristolochia pallida Willd. flowers (Aristolochiaceae). ...
... Small arthropods are known to be important pollinators of a range of other plant species. These include the Dipteran families Cecidomyiidae (Yuan et al. 2007), Ephydridae (Nagasaki 2007), Drosophilidae (Miyake & Yafuso 2005), Sciaridae and Mycetophilidae (Mesler et al. 1980), Phoridae (Rulik et al. 2008), Chironomidae (Murugan et al. 2006), Psychodidae (Albre et al. 2003), small Coleoptera (Krantz 2004;Blanche et al. 2006), small Hymenoptera (Herre et al. 2008) and Thysanoptera (Kirk 1997). Moreover, the propensity for small arthropods, which may carry pollen, to be carried in air currents over many kilometres suggests a possible route for inter-crop and crop-weed pollen movement and may elevate the importance of these arthropods in the contamination of seed crops. ...
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Flower-visiting small arthropods (body length <3 mm) that may vector pollen can be carried via air currents over long distances. Therefore, assessing their diversity, abundance and distribution within crops is an important step in determining their role in the transfer of pollen between crops and weeds leading to crop contamination. A window trap survey of eight flowering pak choi field trials in the North and South Islands of New Zealand demonstrated that small arthropods, particularly flies and thrips, were highly abundant within all fields, greatly dominating arthropod counts. However, the relative abundance of different taxa could vary substantially. For example, Drosophilidae individuals represented 65.5% of total arthropod individuals counted in a Pukekohe field and only 7.0% in a Hastings field. The large representation of small arthropods within flowering fields across New Zealand warrants further investigation of their contribution to pollination and pollen flow.
... However, the insects that interact with A. fimbriata are small brachyceran flies, e.g. those of the genus Megaselia (Phoridae) [24], whose mass m = 0.3 mg [25] is lower than that of the syrphid fly Plactycheirus angustatus considered in the above calculations. Since during walking, the weight is passed only through a half of the legs, the load on one foot of the Megaselia fly is ...
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It is known that attachment ability of insects depends among others on the texture of the substrate. On plant surfaces, covered with microscopic epicuticular wax crystals, insect attachment was found to be highly reduced for many insect species. In some plants, this effect may be explained by the contamination of insect adhesive organs (pads) by wax crystals. In the present study, mechanics of the wax crystals fracture during contact formation between insect adhesive pads and plant surface is examined, in order to explain the observed contamination of pads by wax. It is shown that mechanisms of the wax crystal fracture may be rather different, depending on the slenderness ratio of the crystals. Crystals with high values of the ratio may buckle elastically or in an elastic-plastic way. For five plant species under consideration, the critical value of the slenderness ratio for elastic buckling is 26.5, while for elastic-plastic buckling it is 18.7. If the values of the slenderness ratio are lower than the critical values, one has to consider bending of the crystals under the weight of insects. Although this study considered only crystals of a tubular shape, the general approach is valid also for crystals of other shapes.
... The fruit set of some non-autogamous orchids is low as a consequence of a deficient pollen transference between plants (Ackerman and Montalvo, 1990;Calvo, 1990;Janzen et al., 1980;Schemske, 1980), although other features, as inflorescence size, population density, flowering synchronization and flower duration, also can exert strong influence on the reproductive success of animal-pollinated plants (e.g., Augspurger, 1983;Flores-Palacios and Garcı´a-Franco, 2003;House, 1992;Human˜a et al., 2008;Kunin, 1997;Marquis, 1988;Pansarin and Amaral, 2008;Schmitt et al., 1987). Furthermore, fruit production may depend on several reproductive characteristics as flower size, shape, color, reward production and breeding systems (Rulik et al., 2008;Zimmerman and Pyke, 1988). ...
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Reproductive biology and pollination of Stanhopea lietzei and Stanhopea insignis were studied in a semi-deciduous mesophytic forest in the Serra do Japi (SJ), and in the coastal plain of Picinguaba, both in the State of São Paulo, Brazil. Floral morphology, pollination, breeding system and fruit set of both species were investigated. S. lietzei and S. insignis are pollinator-specific, being pollinated by male bees of Eufriesea (Apidae, Euglossini), which collect the fragrance produced by pluricellular osmophores at the base of the saccate hypochile. S. lietzei and S. insignis were pollinated by Eufriesea pulchra and Eufriesea purpurata, respectively. Observations using substances present in the floral fragrance of both studied species as chemical baits were also performed. E. purpurata was attracted by benzyl alcohol, the major compound of the perfume of S. insignis, while E. pulchra was attracted by none of the compounds used. Both studied Stanhopea are self-compatible but pollinator dependent. Self-pollination, however, tends to be avoided by floral mechanisms. In experimental self- and cross-pollinations the proportion of fruit abortion was high and related to resource limitation. The reproductive success of S. lietzei and S. insignis was low as a consequence of deficient pollen transference while pollinator scarcity was the main factor.
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Most Aristolochiaceae species studied so far are from temperate regions, bearing self-compatible protogynous trap flowers. Although self-incompatibility has been suggested for tropical species, the causes of self-sterility in this family remain unknown. To fill this gap, we studied the pollination of the tropical Aristolochia esperanzae, including the physical and physiological anti-selfing mechanisms. Floral visitors trapped inside flowers were collected to determine the pollinators. Protogyny was characterized by observing the temporal expression of sexual phases and stigmatic receptivity tests. The breeding system was investigated using hand-pollination treatments. Pollen tube growth was observed using epifluorescence to identify the self-incompatibility mechanism. Flies were the most frequent visitors found inside A. esperanzae trap flowers, with individuals from the family Ulidiidae being potential pollinators since they carried pollen. The characteristic flower odour and presence of larvae indicate that A. esperanzae deceives flies through oviposition-site mimicry. Although this species showed incomplete protogyny, stigmatic receptivity decreased during the male phase, avoiding self-pollination. Fruits developed only after cross- and open pollination, indicating that the population is non-autonomous, non-apomictic, and self-sterile. This occurred through a delay in the growth of geitonogamous pollen tubes to the ovary and lower ovule penetration, indicating a late-acting self-incompatibility mechanism. Our findings expand the number of families in which late-acting self-incompatibility has been reported, demonstrating that it is more widespread than previously thought, especially when considering less-studied tropical species among the basal angiosperms.
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• Deceptive pollination has been reported in the genus Aristolochia, but the floral biology and pollination strategy of Aristolochia bianorii, an endemic species to the Balearic Islands, have not been studied yet. Here, we aimed to investigate the floral anthesis, the mating system, the pollinators and the volatile organic compounds (VOCs) emitted by its flowers. • Flower buds were marked and monitored daily to define floral stages and their duration. Experimental bagging and hand pollination treatments were performed to test for autonomous self-pollination, induced self-pollination, and cross-pollination. Flowers were collected to analyse the presence of entrapped pollinators. VOCs emitted by flowers were evaluated by means of a solid phase microextraction followed by immediate gas chromatography-mass spectrometry. • Anthesis lasted between 63 and 96 hours, and the species exhibited autonomous self-pollination with a moderate inbreeding depression. Pollinators were mainly females of Oscinomorpha longirostris (Diptera; Chloropidae), and the number of pollinators inside flowers was affected by the floral stage and the time of flowering. The most common VOCs were alkanes, oximes, esters, alkenes, cyclic unsaturated hydrocarbons, isocyanates, amides and carboxylic acids. • Aristolochia bianorii can set seeds by autonomous self-pollination, in contrast to other Aristolochia species in which both protogyny and herkogamy prevent from autonomous self-pollination. However, the species may encourage cross-pollination by attracting female chloropid flies though the emission of floral scents that may mimic an oviposition site and possibly, freshly killed true bugs (i.e., Heteroptera). In conclusion, A. bianorii promotes cross-pollination but delayed autonomous self-pollination assures the reproductive success in putative absence of pollinators.
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Ontogeny and several aspects of the reproductive biology of Aristolochia elegans Mast., cultivated in a greenhouse of the Botanical Garden-Institute, Far Eastern Branch of RAS (BGI FEB RAS, Vladivostok, Russuia) were studied. A. elegans is ornamental and medicinal plant, woody liana and the native species of South and Central America flora. Plants were cultivated from reproduction seeds of Tallinn Botanical Garden. Three periods and seven age states from seed to mature generative individual were identified in the plants ontogeny. Nearly 85 % of individuals successfully change their age status. The plants bloom in 10–12 months after sowing. Blossoming and fruitage are every year. Flowers are protogynous and adaptive to pollination by insects. Life duration of individual flower is 2–3 days. The presence of self-compatibility was confirmed for A. elegans, but pollinators are required for successful pollination. Members of Drosophilidae family (order Diptera) are pollinators in ex situ. Fruit is six ocular dry capsule 30.3±0.95 mm. length, 13.2±0.35 mm wide. The seed length with wing is 5.52±0.08 mm and width 4.39±0.06 mm; length of seeds without wings is 3.18±0.05 mm and width 3.02±0.04 mm. Seeds obtained in the BGI FEB RAS, are smaller than seeds of Tallinn Botanical Garden. However, germinating ability 84±2% as well as high index of ovules formation per fruit 374±16, viable seeds per fruit 207±16 and seed set 55±4% were shown. Regular flowering and high reproductive ability of Aristolochia elegans ex situ confirmed that this species adapts to ecological and geographical conditions of the new area and its potential success in the introduction in Primorsky krai of Russia.
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Aristolochia genus, which has about 450 species all ower the world flora attracts attention especially with its unique generative properties besides it's highly variable vegetative properties. In this study we tried to explain the structure of flower, fruit and seed and pollination and breeding properties of Aristolochia genus.
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A cladistic analysis of the subfamily Aristolochioideae (Aristolochiaceae) based on morphological characters is presented. The analysis includes 65 taxa within the ingroup, which represent all the tribes, subtribes, genera, and infrageneric taxa formally described within the subfamily Aristolochioideae sensu Schmidt. The analysis shows that Aristolochia s.l. is paraphyletic and that Euglypha and Holostylis are not different lineages from Aristolochia. Two of the three subgenera recognized within Aristolochia (Siphisia and Pararistolochia) are shown to be monophyletic. The third subgenus (Aristolochia) is paraphyletic. Most taxa at lower rank levels (sections, subsections, series, and subseries) are shown to be either polyphyletic or paraphyletic. The characters traditionally used for the recognition of these taxa are evaluated and new characters are introduced. Finally, a revised system of classification of the Aristolochioideae based on monophyletic groupings and consisting of two tribes, two subtribes, and five genera is proposed here.
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The present work contains a revision of the Black Fungus Gnats (Insecta, Diptera, Nematocera: Sciaridae) of the Palaearctic Region. The opening section contains a historical review in which important workers in the field of sciarid research are discussed and their taxonomic oeuvre evaluated. Other sections contain fundamental information on the occurrence, distribution, biology and individual development. Our discussion of this family also devotes much space to describing successful methods of rearing, the role of sciarids in nature, their economic importance as pests of greenhouse plant cultures, and the biological control of the major pest species. A separate section is devoted to the migratory behaviour of sciarid larvae, the "army worm" phenomenon. These introductory sections are completed with a detailed characterisation of the family and a discussion of its systematic position within the Diptera. The results of our revision of the sciarid fauna of the Palaearctic region have led not only to great changes in taxonomy and nomenclature but also to a considerable increase in the number of species. At the species level, it has been necessary to make 375 lectotype designations, 143 new combinations and 3 changes of status. In addition, 4 preoccupied names were replaced and 199 new synonyms discovered. At the present time the list of described Palaearctic Sciaridae stands at 836 valid species, 412 synonyms, 24 species incertae and 41 nomina dubia. So far as supra-specific taxa are concerned, 91 names have been recorded from the Palaearctic region of which 33 are synonyms according to present knowledge. A hypothetical phylogenetic tree for the Palaearctic sciarid genera could be developed. The weak points in this tree [convergence, non-monophyla] or alternative hypotheses of relationship are discussed at the relevant points. From this analysis, we recognise in this revision a classification of the Palaearctic Sciaridae composed of 28 genera and 30 subgenera. In order to facilitate identification of the species, 61 species groups have been defined and under each genus a list is given of the included species. To resolve various taxonomic-nomenclatural, phylogenetic or zoogeographic problems we also partially revised selections of material from the Afrotropical, Australian, Nearctic, Neotropical and Oriental regions. The results from this study were of considerable significance for the present revision because they contributed to almost every one of the topics covered, substantially enhanced the discussion of Palaearctic faunistic elements, and consequently they run through the entire work. Some of the non-Palaearctic types that we revised are of direct relevance to this Palaearctic revision. For this reason, we have included an appendix in which redescriptions are given of certain type-species and of relevant species from other faunal regions. The present work has very much the character of a Handbook, in which everything of value and interest about the Black Fungus Gnats of the Palaearctic Region has been included. The bibliography claims to be complete at least as far as the descriptive literature on the recent Palaearctic fauna is concerned. The results of our revision are illustrated by 612 figures, 4 diagrams and 3 tables. As a work of reference, it contains the most recent research results on the taxonomy and classification, a comprehensive list of sources, and an index of sciarid names. Moreover, the updated check list, which is based on a revised nomenclature and a stabilised classification, subsumes current "state-of-the-art" knowledge of the species of the Palaearctic region into a species inventory. The 18 identification keys provide the user with a much improved tool for identification, which includes for the first time all the revised species of the Palaearctic region. With these keys species can be assigned to genera, subgenera and species groups. In addition, the non-specialist, confronted with the task of identifying sciarids, is provided with the possibility of narrowing down considerably the number of species that he has to take into consideration. [in German]
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The flowers of Aristolochia grandiflora are sapromyiophilous trap blossoms that deceive their visitors with optical and olfactory promise of hidden protein-rich substrate. The most effective pollinators are large Diptera, mostly Calliphoridae, which become trapped in the protogynous flowers on the first day of anthesis. Although the flowers are protogynous and subsequently release pollen, a variety of floral changes occur that discourage further insect visitation after pollination and allow the pollinators to escape on the second day of anthesis. On the first day of anthesis the flowers' strong carrion odor and color gradients draw pollinators toward the receptive gynostemium deep within the flower. Constricting floral tubes with trichomes oriented toward the gynostemium aid in capturing and holding the insects. On the next day, the flowers change to male phase and pollen is deposited on the pollinator. Flower structure and function then change to release the pollen-dusted pollinator. To aid in pollinator release, the floral odor disappears, color cues change, hairs relax, and the constricting areas of the tube are opened. Pollination appears to be a two-day process for any given flower with floral senescence by the third day. Floral visitors do oviposit in the flowers, but we suggest that this is not relevant to pollination. Comparisons are made with other Aristolochiaceae.
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González: Aristolochia grandiflora Sw. (Aristolochiaceae): Desarrollo y morfología de la flor más larga del mundo. Rev. Acad. Colomb. Cienc. 30(115): 181-194. 2006. ISSN 0370-3908. Aristolochia grandiflora Sw. posee las flores más largas del mundo. La distribución de esta especie incluye México, Centro América, Cuba, Jamaica, Colombia, Trinidad y Ecuador, con algu-nos registros posiblemente procedentes de poblaciones cultivadas o naturalizadas en St. Thomas, Guadalupe y Martinica. Aunque es una especie bien conocida en cultivo, debido a la morfología floral tan extraña y conspicua, es poco conocida en su estado nativo. Además de las dimensiones extremas de la flor, la morfología del perianto es atípica con respecto a las demás especies neotropicales del subgénero Aristolochia, en cuanto a la zigomorfía lateral (i.e. la que modifica el perfil del perianto) y frontal (i. e. la que afecta únicamente al limbo floral), y el desarrollo de dos de las estructuras internas más especializadas, la siringe y el anillo. Se describe el desarrollo floral, con énfasis en el perianto, y se documenta el que sin duda, es el ejemplar con la mayor longitud del perianto hasta ahora conocido en estado silvestre. Finalmente, se discuten las posibles relaciones filogenéticas de la especie.
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A combined phylogenetic analysis of the Aristolochioideae was conducted based on 72 morphological characters and molecular data sets (matK gene, trnK intron, trnL intron, trnL-trnF spacer). The analysis sampled 33 species as the ingroup, including two species of Thottea, 30 species of Aristolochia, and the monotypic genus Euglypha, which represent all the infrageneric taxa formally described; Saruma henryi and Asarum caudatum were used as the outgroup. The results corroborate a sister-group relationship between Thottea and Aristolochia and the paraphyly of Aristolochia with respect to Euglypha, which consequently should be included in Aristolochia. Two of the three subgenera within Aristolochia (Isotrema and Pararistolochia) are shown to be monophyletic, whereas the signal obtained from the different data sets about the relationships within subgenus Aristolochia is low and conflicting, resulting in collapsed or unsupported branches. The relationship between the New World and the Old World species of subgenus Aristolochia is conflictive because morphological data support these two groups as monophyletic, whereas molecular data show the monophyletic Old World species of Aristolochia nested within the New World species. A sister-group relationship is proposed between Aristolochia lindneri and pentandrous species, which suggests that a group of five species from central and southern South America (including A. lindneri) could be monophyletic and sister to Aristolochia subsect. Pentandrae, a monophyletic taxon consisting of ca. 35 species from the southern United States, Mesoamerica, and the West Indies.
Article
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The floral biology of Aristolochia argentina (Aristolochiaceae) was studied in natural populations in Córdoba, Argentina. This native vine has flowers that attract mainly scuttle fly pollinators of the genus Megaselia (Phoridae). The trap-like perianth is formed by a limb, a tube, and a basal utricle. The limb produces an odor that recalls decaying plant tissues, which apparently mimics the natural oviposition substrate of the flies. The insects stay entrapped inside the utricle for approximately 24 h, making contact with the sexual organs of the flower. When released, they can become captured again in other flowers. The mechanisms of herkogamy and protogyny are efficient. Although self-compatibility exists, as demonstrated by the high percentage of fruits produced by geitonogamy, fruits were not produced under natural or artificial autogamous conditions. Natural pollination showed significantly lower fruit set than xenogamous and geitonogamous crosses.
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The Diptera are the second most important order among flower-visiting (anthophilous) and flower-pollinating insects worldwide. Their taxonomic diversity ranges from Nematocera to Brachycera, including most families within the suborders. Especially important are Syrphidae, Bombyliidae, and Muscoidea. Other families, especially of small flies, are less appreciated and often overlooked for their associations with flowers. We have compiled records of their flower visitations to show that they may be more prevalent than usually thought. Our knowledge of anthophilous Diptera needs to be enhanced by future research concerning (i) the significance of nocturnal Nematocera and acalypterate muscoids as pollinators, (ii) the extent to which the relatively ineffective pollen-carrying ability of some taxa can be compensated by the abundance of individuals, and (iii) the role of Diptera as pollinators of the first flowering plants (Angiospermae) by using phylogenetic and palaeontological evidence. Specializations in floral relationships involve the morphology of Diptera, especially of their mouthparts, nutritional requirements, and behaviour, as well as concomitant floral attributes. The South African flora has the most highly specialized relations with dipterous pollinators, but in arctic and alpine generalist fly–flower relations are important in pollination and fly nutrition.
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This study presents breeding and pollination systems of Aristolochia maxima and A. inflata in a seasonal tropical forest of Panama. Aristolochia is the most diverse genus of Aristolochiaceae, with ∼120 species distributed throughout the tropics and subtropics. All the Aristolochia species studied so far are pollinated by saprophagous flies of different families, which are presumably deceived by floral odor. Flowers of many species have trap-and-release mechanisms. The flowers attract and imprison pollinators during the female stage first day of flowering and release them after anther dehiscence. Pollination systems of A. maxima and A. inflata are different from those of other Aristolochia in lacking trap mechanisms. Furthermore, the pollinators oviposit in the flowers, and their larvae grow on the fallen, decaying flowers on the ground. Therefore, the plants have a mutualistic relationship with their pollinators. Self-compatible A. inflata is pollinated by Megaselia sakaiae (Phoridae, Diptera). The pollinator may be specialized to Aristolochia flowers, which is the only substrate for larval development. On the other hand, self-incompatible A. maxima is pollinated by Drosophila spp. (Drosophilidae, Diptera), which utilize Aristolochia flowers as a breeding site only occasionally. This pollination mutualism might have evolved from deceit pollination.
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Evolution of Piperales – matK gene and trnK intron sequence data reveal lineage specific resolution contrast. Piperales are one of the largest basal angiosperm orders with a nearly worldwide distribution. The order includes three species rich genera, Piper (ca. 1,000 species), Peperomia (ca. 1,500-1,700 species), and Aristolochia s. l. (ca. 500 species). Sequences of the matK gene and the non-coding trnK group II intron are analysed for a dense set of 105 taxa representing all families (except Hydnoraceae) and all generic segregates (except Euglypha within Aristolochiaceae) of Piperales. A large number of highly informative indels are found in the Piperales trnK/matK dataset. Within a narrow region approximately 500 nt downstream in the matK coding region (CDS), a length variable simple sequence repeat (SSR) expansion segment occurs, in which insertions and deletions have led to short frame-shifts. These are corrected shortly afterwards, resulting in a maximum of 6 amino acids being affected. Furthermore, additional non-functional matK copies were found in Zippelia begoniifolia, which can easily be discriminated from the functional open reading frame (ORF). The trnK/matK sequence data fully resolve relationships within Peperomia, whereas they are not effective within Piper. The resolution contrast is correlated with the rate heterogenity between those lineages. Parsimony, Bayesian and likelihood analyses result in virtually the same topology, and converge on the monophyly of Piperaceae and Saururaceae. Lactoris gains high support as sister to Aristolochiaceae subf. Aristolochioideae, but the different tree inference methods yield conflicting results with respect to the relationships of subfam. Asaroideae. In Piperaceae, a clade formed by the monotypic genus Zippelia and the small genus Manekia (=Sarcorhachis) is sister to the two large genera Piper and Peperomia. Systematics of pipevines – Combining morphological and fast-evolving molecular characters to investigate the relationships within subfamily Aristolochioideae (Aristolochiaceae) A combined phylogenetic analysis of the Aristolochioideae was conducted based on 72 morphological characters and molecular datasets (matK gene, trnK intron, trnL intron, trnL-trnF spacer). The analysis sampled 33 species as the ingroup, including two species of Thottea and 30 species of Aristolochia and the monotypic genus Euglypha, which represent all the infrageneric taxa formally described; Saruma henryi and Asarum caudatum were used as the outgroup. The results corroborate a sister-group relationship between Thottea and Aristolochia, and the paraphyly of Aristolochia with respect to Euglypha that consequently should be included into Aristolochia. Two of the three subgenera within Aristolochia (Isotrema and Pararistolochia) are shown to be monophyletic, whereas the signal obtained from the different datasets about the relationships within subg. Aristolochia is low and conflicting, resulting in collapsed or unsupported branches. The relationship between the New World and the Old World species of subgenus Aristolochia is conflictive because morphological data support these two groups as monophyletic, whereas molecular data show the monophyletic Old World species of Aristolochia nested within the New World species. A sister group relationship is proposed between A. lindneri and pentandrous species, which suggests that a group of five species from central and southern South America (including A. lindneri) could be monophyletic and sister to Aristolochia subsection Pentandrae, a monophyletic taxon consisting of about 35 species from southern USA, Mesoamerica, and the West Indies. Colonisation, phylogeography and evolution of endemism in Mediterranean Aristolochia (Aristolochiaceae). This study provides evidence for a multiple colonisation of the western Old World from Asian ancestors within Aristolochia section Diplolobus (subsection Aristolochia and Podanthemum). Within subsection Podanthemum it is assumed, that the colonisation of the African continent happened at least two times independently. In contrast, for subsection Aristolochia, a rapid morphological radiation in the Near East (or close to this area) with subsequent star like colonisation of the different current distribution areas, which is not paralleled on the molecular level, appears to be more likely. Phylogenetic tree reconstruction is unsupported for these clades, but most clades are highly supported as monophyletic. Interestingly the Mediterranean and temperate Eurasian species, which are morphologically distinct (A. pistolochia, A. clematitis) are not clustering within the main clades, but are independent lineages. Analogue, A. rigida a species from Somalia is well-supported sister to the subsection Aristolochia. Within subsection Podanthemum the colonisation event from an Asian ancestor is clearly traceable, whereas in subsection Aristolochia the path is not traceable, since the ancestors are extinct or not present in the connecting areas. Within the Mediterranean, Near East and Caucasian species of subsection Aristolochia two morphologically and biogeographically well supported groups can be identified: the Near East/Caucasian species and the West Mediterranean species. The previous groupings for the latter, based on morphological characters, could be substantiated only partly by our results. This study provides the first phylogeny of all West Mediterranean species. In addition an independent complex is established including some micro endemic species. The phylogenetic results are discussed with respect to biogeography, and morphology, to give a first insight into the radiation and colonisation of the genus Aristolochia in the Mediterranean region. Universal primers for a large cryptically simple cpDNA microsatellite region in Aristolochia. We provide a new and valuable marker to study species relationships and population genetics in order to trace evolutionary, ecological, and conservational aspects in the genus Aristolochia. Universal primers for amplification and subsequent sequencing of a chloroplast microsatellite locus inside the trnK intron are presented. Utility of the primers has been tested in 32 species representing all clades of Aristolochia, including population studies within the A. pallida complex, A. clusii and A. rotunda. The microsatellite region is characterized as a (AnTm)k repeat of 22–438 bp containing a combination of different repeats arranged as ‘cryptically simple’. Trapped! Pollination of Aristolochia pallida Willd. in the Mediterranean A first study of the pollination biology of a Mediterranean Aristolochia species in its natural habitat is presented. 183 flowers of Aristolochia pallida were investigated, which in total contained 73 arthropods, dominated by two groups of Diptera, Sciaridae (37%) and Phoridae (19%). However, only Phoridae are regarded as potential pollinators, since pollen has been found exclusively on the body surfaces of these insects. All Phoridae belong to the genus Megaselia and are recognised as four undescribed species. The measurements of flower and insect dimensions suggest that size is an important constrain for successful pollination: 1) the insects must have a definitive size for being able to enter the flower and 2) must be able to get in touch with the pollen. Only very few insect groups found in Aristolochia pallida fulfil these size requirements. However, size alone is not a sufficient constrain as too many fly species of the same size might be trapped but not function as pollinators. Instead, specific attraction is required as otherwise pollen is lost. Since all trapped Phoridae are males, a chemical attraction (pheromones) is proposed as an additional constrain. Since A. pallida flowers are protogynous, the record of Megaselia loaded with pollen found in a flower during its female stage proves that this insect must have been visited at least one different flower during its male stage before. Further on, this observation provides strong evidence that the flowers are cross-pollinated. All these factors indicate a highly specialised pollination of Aristolochia pallida by Megaselia species.
Article
Part I of this work provided a key to the 17 genera and 32 species of Phoridae other than those in the giant genus Megaselia. This part, starting from a list of four previously recorded species of Megaselia from Arabia, provides keys to the 84 species now known from this region. Revisionary papers leading up to this work described 15 new species, which describes and names a further 33 new species and characterises eight species currently only known in the female sex (which are not named until linked to their males). Megaselia tiagoensis Disney, 1991 is synonymised with M. sokotrana Beyer, 1965.
Article
With the previous reporting in the literature of only four species of scuttle fly from Arabia, an attempt has been made to fill this glaring gap in our knowledge of world Phoridae. Keys are provided for the 18 genera and 32 species of Phoridae, other than those in the giant genus Megaselia, now recorded from Arabia. Three new genera, Arabiphora, Miclepta and Zygtaxphora and 12 new species are described. Two species, including the type species of Zygtaxphora, Z. orientalis, and Z. sulawesae from Indonesia are also described, A female previously assigned to Razorfemora nussbaumi is now recognised as belonging to a different species. It is formally named R. zaragozae n. sp and distinguished from the true female of R. nussbaumi, which is described from a series of associated males and females from Yemen.
Article
A comparative study of the development and morphology of the perianth in 42 species of Aristolochia is presented. These species represent all the subgenera, sections, and subsections formally proposed within this genus. Additional observations on the perianth of Asarum, Saruma and Thottea are also included because perianth morphology has been crucial for the classification of the Aristolochiaceae. The results support the interpretation of the perianth of Aristolochia, Euglypha and Holostylis as a trimerous calyx. Five main types of perianth development were found in Aristolochia which differ in the degree of fusion between the perianth lobes, the direction of floral curvature, and the symmetry of the perianth limb. The interpretation of the perianth of Aristolochia as a calyx is supported in terms of position, morphology, development, and comparison to related taxa.
Chapter
The larvae of the Phoridae are very diverse. A preliminary review of this diversity is provided by Ferrar (1987). Schmitz (l938n) discussed larval morphology.
Article
Aristolochia dinghoui is described from southwestern Thailand. It is morphologically similar toA. harmandiana Pierre ex Lecomte, from Cambodia and Laos, and also bears some resemblance toA. philippinensis Warb. andA. sericea Blanco, both from the Philippines. A key to these four species is included, and some of their diagnostic features are listed.
Article
An extensive list of corrections to the Chironomidae part of the Catalogue of Palaearctic Diptera (ASHE and CRANSTON, 1990) is given.
Article
Piperales represent the largest basal angiosperm order with a nearly worldwide distribution. The order includes three species rich genera, Piper (ca. 2000 species), Peperomia (ca. 1500-1700 species), and Aristolochia s. l. (ca. 500 species). Sequences of the matK gene and the non-coding trnK group II intron are analysed for a dense set of 105 taxa representing all families (except Hydnoraceae) and all generic segregates (except Euglypha within Aristolochiaceae) of Piperales. A large number of highly informative indels are found in the Piperales trnK/matK dataset. Within a narrow region approximately 500 nt downstream in the matK coding region (CDS), a length variable simple sequence repeat (SSR) expansion segment occurs, in which insertions and deletions have led to short frame-shifts. These are corrected shortly afterwards, resulting in a maximum of six amino acids being affected. Furthermore, additional non-functional matK copies were found in Zippelia begoniifolia, which can easily be discriminated from the functional open reading frame (ORF). The trnK/matK sequence data fully resolve relationships within Peperomia, whereas they are not effective within Piper. The resolution contrast is correlated with the rate heterogeneity between those lineages. Parsimony, Bayesian and likelihood analyses result in virtually the same topology, and converge on the monophyly of Piperaceae and Saururaceae. Lactoris gains high support as sister to Aristolochiaceae subf. Aristolochioideae, but the different tree inference methods yield conflicting results with respect to the relationships of subfam. Asaroideae. In Piperaceae, a clade formed by the monotypic genus Zippelia and the small genus Manekia (=Sarcorhachis) is sister to the two large genera Piper and Peperomia.
Die Gleitfalle als Blumentypus
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