Table 2 - uploaded by Stefan Wanke
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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 spe...
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Floral and inflorescence structure and ontogeny are described in detail in Peperomia fraseri, an anomalous species of Piperaceae that differs in several respects from other species of Peperomia and other perianthless Piperales (Piperaceae and Saururaceae). Inflorescence structure is atypical in this species, with numerous spikes arranged spirally o...
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... Among those, Aristolochia microstoma BOISS. & SPRUNER, a species endemic to Greece, stands out due to its unique perianth morphology and flower presentation (Wanke, 2006). The limb of the small, purplish-brownish flowers (Figures 1A-C) is reduced to a small beak or missing, and the entrance into the floral tube is reduced to a small pore, responsible for the name of the species (Nardi, 1991). ...
... The limb of the small, purplish-brownish flowers (Figures 1A-C) is reduced to a small beak or missing, and the entrance into the floral tube is reduced to a small pore, responsible for the name of the species (Nardi, 1991). While most Aristolochia species display their often showy flowers above the ground, A. microstoma flowers are presented close to or partly buried in the ground, among leaf litter (Figures 1A,B) or between rocks (Figures 1C,D), often hidden from above (Nardi, 1991;Wanke, 2006). Another unusual feature is the more or less horizontal orientation of the floral tube, which is vertical in other species. ...
... Another unusual feature is the more or less horizontal orientation of the floral tube, which is vertical in other species. Pollinators were hypothesized to be small arthropods living near the ground or in leaf litter (Nardi, 1991;Wanke, 2006). So far, the flower visitors and pollinators, the reproductive biology, and the floral scent of A. microstoma remained unknown. ...
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.
... Most important contributions to the knowledge of species from the Mediterranean were published by Nardi and others ( [31], [32], [33], [34], [35], [36], [37], [38]). [39] provided the first phylogeny of all West Mediterranean Aristolochia species also including few members of Caucasia and the East Mediterranean region. ...
... Rodlets have been found among all Caucasian and East Mediterranean species as well as for most of the Old and New World taxa, indicating that this is likely a plesiomorphic character in East Mediterranean species. Although, A. pistolochia is geographically occurring in the Western Mediterranean area (France, Spain), molecular phylogenetic studies showed this species as sister to all Mediterranean species being more closely allied to A. clematitis ([28], [39]), which has a Eurasian distribution and is not part of the Mediterranean clade neither ( [28], [39]). Also from a morphological point of view these species are separated from other Mediterranean Aristolochia species as they do not possess a rootstock but have either a rhizome or show fleshy roots [39]. ...
... Rodlets have been found among all Caucasian and East Mediterranean species as well as for most of the Old and New World taxa, indicating that this is likely a plesiomorphic character in East Mediterranean species. Although, A. pistolochia is geographically occurring in the Western Mediterranean area (France, Spain), molecular phylogenetic studies showed this species as sister to all Mediterranean species being more closely allied to A. clematitis ([28], [39]), which has a Eurasian distribution and is not part of the Mediterranean clade neither ( [28], [39]). Also from a morphological point of view these species are separated from other Mediterranean Aristolochia species as they do not possess a rootstock but have either a rhizome or show fleshy roots [39]. ...
... The genus Aristolochia (Aristolochiaceae) comprises over 450 species and is distributed in the tropics and in temperate regions (Wagner et al. 2014). In the Mediterranean about 60 species are known, representing a diversity hotspot for the genus (Wanke 2007). Virtually all Aristolochia species exhibit deceptive pollination strategies and it was assumed that flowers mimic the brood sites of their fly pollinators (Vogel 1978;Proctor et al. 1996). ...
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.
... The results also supported the sister group relationship of Endodeca and Isotrema; subgenera ParAristolochia and Aristolochia also showed sister group relationship and were found to be monophyletic. These results were in congruence with the earlier studies [15,16,18,23,24] . The Aristolochia clade was further divided into subclades forming section Diplolobus and Gymnolobus; the section Diplolobus with subsection Podanthemum and subsection Aristolochia. ...
... The Aristolochia clade was further divided into subclades forming section Diplolobus and Gymnolobus; the section Diplolobus with subsection Podanthemum and subsection Aristolochia. Podanthemum and Aristolochia showed sister relationship and found monophyletic as in previous studies [17,23] . All the species studied were fully resolved using matK phylogeny. ...
The genus Aristolochia comes under the family Aristolochiaceae, a conspicuous family with distinctive and showy flowers positioned among the basal angiosperms. The taxonomic treatment of the genus Aristolochia is ambiguous. Several generic segregates have been proposed for the genus and most literatures propose four subgenera within two subtribes. The study focused on the relationship of A. indica, A. tagala, A. ringens and A. krisagathra with the different generic segregates. The sequencing was carried out using rbcL, matK, psbA-trnH, ITS and ITS2. However, only rbcL and matK were used to infer the relationship because of the lack of availability of sequences of psbA-trnH, ITS and ITS2 representing the different generic segregates. Both the neighbor joining trees constructed using rbcL and matK placed A. krisagathra as the sister taxon of A. indica with very high bootstrap values. A. krisagathra, A. indica and A. tagala forms a part of the subsection Podanthemum and Diplolobus section along with A. zollingeriana, whereas the exotic A. ringens formed a part of Gymnolobus section.
... The genus Aristolochia consists of between 450 and 600 species growing in temperate and tropical climates worldwide (Wanke, 2007). They are mostly cultivated as ornamentals but most species are also popular medicaments. ...
Aristolochia repens was extracted with 80% ethanol and the extract concentrated in vacuo to obtain the crude ethanolic extract. Phytochemical screening of the crude extract revealed the presence of steroids, terpenoids, flavonoids, anthraquinones, tannins, saponins and reducing sugar. The crude extract showed excellent to moderate inhibitory activity on Pseudomonas aeruginosa, Staphlococcus aureus, Proteus spp , Aspergillus flavus and Candida albican.
... Aristolochiaceae is a large genus of herbs and twining plants, found in the tropical and temperate regions of the world [2]. In the Mediterranean and adjacent Near East 50 Aristolochia species occur, all of them being endemic to a particular area [3] (Figure 1). ...
The genus Aristolochia (Aristolochiaceae) consists of approximately 500 species, most of which are found in tropical, subtropical, and Mediterranean regions. Plants of the genus Aristolochia have been shown to exhibit interesting biological activities including cytotoxic, apoptosis-induced, antimicrobial, anti-inflammatory, antioxidant, antiallergic activities, among other.
... The genus Aristolochia (Aristolochiaceae) consists of about 500 species, growing mostly in tropical and subtropical areas (Neinhuis et al., 2005). The Mediterranean region, including Turkey, Caucasus and Near East, comprises more than 60 species of Aristolochia, which are monophyletic based on molecular phylogeny (de Groot, et al., 2006;Wanke, 2006). They occur in different type of habitats, particularly in subalpine humid or semidry grasslands. ...
... Mediterranean climate as influenced also in differentiation of morphological characters and adaptation of Aristolochia species to substrate, mostly in limestone. Most of species are geophyte, which allow them to pass the dried summer period in dormancy whereas the growth including flowering, in spring time (Wanke, 2006;Costa, 2008). The west Mediterranean Aristolochia species, based on chromosomal numbers, according to Ball (1964) and Mayer & Greuter (1985) are grouped in 4-species complex: A. rotunda, A. sempervirens, A. fontanessii and A. pallida complex. ...
In this paper A. lutea Desf., and Aristolochia elongata (Duchartre) Nardi, as new species from the family Aristolochiaceae is presented. Other aspects related to several subalpine areas of Albania are described, too. Morphological characters of A. lutea were discussion and compared with it’s closely relatives Aristolochia elongata (Duchartre) Nardi reported on several habitats of South Albania. As the other species like Aristolochia merxmuelleri Greuter & Mayer, A. lutea is a member of the Aristolochia pallida aggregate. Ecological data in growing habitat and area of distribution overall the country is given and mapped. Along with A. lutea, in its distribution habitat are recorded several important species for the Albanian flora like: Campanula hawkinsiana Hausskn. et Heldr., Alkanna graeca Boiss. & Spruner, Alkanna scardica Griseb. Viola acrocerauniensis Erben, Viola raunsiensis W.Becker & Košanin, Fritillaria thessala subsp. jonica (Halacsy) Kamari, Fritillaria orentalis Adams, Orchis pauciflora x Orchis quadripunctata etc.
Keywords: Aristolochia lutea, A. elongata, new species, subalpine ecosystems, important species
... The genus Aristolochia (Aristolochiaceae) consists between 450 and 600 species growing in temperate and tropical climates worldwide (Wanke, 2007). Typically species of Aristolochia, including A. consimilis, are woody vines of tropical areas. ...
Aristolochic acids (AA) are characteristic compounds of the Aristolochia genus and are
known to be nephrotoxic, carcinogenic and mutagenic. Aristolochia consimilis is one of
the most frequently used medicinal plants among Surinamese migrants. Despite the
regulations, these products are still available and continue to be used. Surinamese
consumers might be at risk of potential exposure to aristolochic acid. It is therefore,
essential to investigate A.consimilis and the derived medicinal products for the presence
of AA. In this study, several quantitative and qualitative methods have been used. A
reversed-phase high-performance liquid chromatographic (HPLC) method has been
developed and validated. Separation was accomplished on a Luna C18 column with 0.1%
methanol gradient elution. Crude methanol extracts of A.consimilis and several herbal
teas and alcoholic aphrodisiacs were examined. Quantitative determination of AAI
revealed inter batch variations ranging from not detected to 274.9 ug/g DW, whereas no
detectable amounts were found in the derived medicinal products. Despite these findings,
the results do not guarantee safe use of bitter tonics. Based on the cumulative impact of
aristolochic acids, consumption of these plants on a regular basis is not recommended.
... The genus Aristolochia (Aristolochiaceae) consists of approximately 500 species, most of which are found in tropical, subtropical, and Mediterranean regions (Neinhuis et al., 2005;Wanke 2006;Wanke et al., 2006Wanke et al., , 2007. Aristolochia flowers are highly derived, functioning as a trap for arthropods in order to ensure pollination (Knoll, 1929). ...
... In the Mediterranean and adjacent Near East $50 Aristolochia species occur, all of them being endemic to a particular area (Wanke, 2006). The only exception is Aristolochia clematitis L., which is probably not native to this region (Wanke, 2006) and for which several observations on pollination biology have been made clearly outside its natural range or in cultivation (Daumann, 1971;Havelka, 1978;Ule, 1898). ...
... In the Mediterranean and adjacent Near East $50 Aristolochia species occur, all of them being endemic to a particular area (Wanke, 2006). The only exception is Aristolochia clematitis L., which is probably not native to this region (Wanke, 2006) and for which several observations on pollination biology have been made clearly outside its natural range or in cultivation (Daumann, 1971;Havelka, 1978;Ule, 1898). The most recent studies on the pollination biology of Mediterranean Aristolochia species date back to the 19th century (Delpino, 1868(Delpino, , 1869, dealing with Aristolochia pallida Willd., Aristolochia rotunda L. and Aristolochia sempervirens L. ( ¼ A. altissima Desf.) from Italy. ...
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.
... 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 (1984Nardi ( , 1988Nardi ( , 1991 and Davis & Khan (1961, 1964, 1982, all of these studies were based on morphological characters only. ...
... 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. ...
... China 45 species, 33 of them endemic (Brach & Song 2006, Huang et al. 2009). The highest rate of endemism coupled with species richness in the northern hemisphere with a rate of 100% endemism is confined to the extended Mediterranean region (Wanke 2007). Recent molecular based phylogenies recovered the Mediterranean Aristolochia species as monophyletic , deGroot et al. 2006, Wanke 2007. ...
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.
