<p>Poslednjih godina u balkanskom regionu pa i u Srbiji, zabeleņeno je ńirenje i invazivne korovske vrste Asclepias syriaca. Pretpostavlja se da je u Srbiju dospela iz susedne MaĎarske gde se dugi niz godina gajila kao medonosna biljka. Zahvaljujući alelopatskim osobinama i jakoj kompetitivnosti, ova vrsta ne dozvoljava regeneraciju starih peńčanih polja Najčeńće se javlja u zapuńtenim voćnjacima i vinogradima, na peskovitim terenima pored puteva i ņelezničkih pruga, nasipima, formirajući pri tom čiste sastojine asocijacije ili se javlja sa joń nekoliko korovskih vrsta ali uz apsolutnu dominaciju A. syriaca. Konstatovano je njeno ńirenje sa parloga na obradive povrńine na kojima obrazuje &bdquo;oaze&ldquo;, naročito u usevima pńenice, ječma, suncokreta, soje, u uljanoj repici, voćnjacima i vinogradima. Imajući u vidu sve negativne aspekte nekontrolisanog ńirenja A. syriaca na ruderalnim i poljoprivrednim povrńinama, ciljevi rada bili su praćenje ńirenja i izučavanje njenih biolońkih i ekolońkih karakteristika kao i utvrĎivanje mogućih načina za njeno suzbijanje. Populacija svilenice na teritoriji Vojvodine ima tendenciju ńirenja i ustaljivanja posebno uz puteve i vodene tokove. UtvrĎena je na skoro svim tipovima zemljińta u Vojvodini. Uspeva na zemljińtima od slabo do jako karbonatnih, kao i od slabo do jako humusnih. Podnosi zemljińta čiji je sadrņaj pristupačnih oblika fosfora i kalijuma ispod donje granice optimalne obezbeĎenosti zemljińta, ali i na zemljińtima gde je sadrņaj vrlo visok do ńtetan. Analizom biolońkog spektra sastojina ass. Asclepiadetum syriacae L&aacute;nikov&aacute; in Chytr&yacute; 2009, utvrĎeno je da su one hemikriptofitsko-terofitskog karaktera sa dominacijom hemikriptofita. Godińnja produkcija semena A. syriaca po m2 je od 1.336,53 do 10.109,61, a sa povećanjem gustine biljaka po jedinici povrńine smanjuje se produkcija plodova. Udeo semena svilenice u ispitivanoj zemljińnoj banci semena je vrlo nizak pa A. syriaca ne formira permanentnu banku semena. Dormantnost semena prekida se njegovim izlaganjem niņim temperaturama tokom nekoliko meseci a vlaga igra značajnu ulogu u povećanju klijavosti semena.<br />Alelopatska svojstva svilenice potvrĎena su ispitivanjem uticaja vodenog ekstrakta iz korena na klijavost semena kukuruza, soje i suncokreta u koncentracijama 0,05-0,1g suvemase korena po 1ml destilovane vode. UtvrĎena je i značajna osetljivost gajenog i divljeg sirka na primenjene ekstrakte. Od ekstrakata korena svilenice pripremljenih sa različitim rastvaračima, najveći inhibitorni uticaj na klijavost semena soje imao je butanolni, a na klijanje semena gajenog sirka etil-acetatni i butanolni ekstrakt. Isto je i sa divljim sirkom i ńtirom, dok na klijavost semena kukuruza i soje ni jedan primenjeni ekstrakt nije imao negativan uticaj u značajnijoj meri. Ogledi u poljskim uslovima pokazali su uticaj alelohemikalija iz korena A. syriaca na redukciju prinosa kukurza i sirka. Vodeni ekstrakt korena u koncentraciji primene 0,04g/ml umanjio je prinos kukuruza za 10-15%, krmnog sirka za 12,5% a sirka metlańa za 23%. Ispitivanja mogućnosti hemijskog suzbijanja A.syriaca u poljskim uslovima pokazali su najveću efikasnost herbicida na bazi glifosata, 2,4-D i tembotriona. kao i nakon trogodińnje primene bentazona, bentazona + dikambe, a najmanja efikasnost ustanovnjena je prilikom primene dikambe. Terbutilazin i klomazon u većim preporučenim količinama primene efikasno suzbijaju svilenicu iz semena ako se herbicidi primene pre nicanja, dok imazamoks i oksasulfuron efikasno suzbijajuu tek iznikle biljke svilenice kada se u preporučenim dozama tretiraju biljke u fazi 2-4 lista.</p> / <p>In previous years in the Balkan region, including Serbia, spread of invasive weed species Asclepias syriaca L. was recorded. It supposedly arrived to Serbia from neighbouring Hungary, where it was grown as honey plant. Due to allelopathic properties and strong concurrence, this species does not allow regeneration of old sand fields. It is most common in neglected orchards and vineyards, sandy terrains along roads and railways, embankments, where form pure A. syriaca association, or several additional weed species occur, but with absolute dominance of this weed. From uncultivated areas on which it forms &quot;oasis&quot; its spread was noted especially in wheat, barley, sunflower, soybean, rapeseed crops, in orchards and vineyards. Considering all negative aspects of uncontrolled spread of A. syriaca on ruderal and agricultural areas, the aims of the paper were monitoring of its spread and the study of biological and ecological properties, as well as the establishment of possible ways of its control. On the territory of Vojvodina, common milkweed population has a tendency of spread and colonization, particularly near roads and water flows. It is established on almost all soil types in Vojvodina. It grows on poor to very calcareous soils, as well as on poor to very humic ones. Common milkweed withstands soil with the content of available forms of phosphorous and potassium below the optimum , but also the soils with high to damaging content of these elements. The analysis of the biological spectrum of stands of ass. Asclepiadetum syriacae, L&aacute;nikov&aacute; in Chytr&yacute; 2009, established that it is hemicryptophyte and therophytein, with a domination of hemicryptophytic character.<br />Annual seed production of A. syriaca seed per m2 is from 1.336,53 to 10.109,61, and with the increase in plant density per unit area, production of fruits is reduced. The share of milkweed seed in soil seed bank is very low, and therefore A. syriaca does not form permanent seed bank. Seed dormancy is stopped by its exposure to lower temperatures for several months, and moisture has an important role in the<br />increase in seed germination. Allelopathic properties of common milkweed were confirmed by examination of root water extract effects on maize , soybean and sunflower seed germination at concentrations of 0.05-0.1g of dry root weight per 1ml of distilled water. Significant susceptibility of cultivated sorghum and Johnson grass to the applied extract was recorded. From common milkweed root extracts prepared by various solvents, the highest inhibiting effect to soybean seed germination had butanol extract, and to germination of seed of cultivated sorghum, ethyl acetate and butanol extract. It was the same with wild sorghum and redroot pigweed while none of the applied extracts had significantly negative effect on maize and soybean germination. In field trials, allelochemicals from A. syriaca root reduced maize and millet yield. Root water extracts at the applied concentration of 0.04g/ml reduced maize yield by 10-15%, of fodder sorghum by 12.5% and broomcorn by 23%. In studies regarding chemical control of A.syriaca in field conditions,the highest efficiency was recorded for herbicides based on glyphosate, 2.4-D and tembotrione, as well as for bentazone and bentazone + dicamba after tree-years long application.Dicamba was recorded as the least efficient. Terbuthylayine and clomazone herbicides in pre-emergence application at higher label rates efficiently control common milkweed from seed, while imazamox and oxasulfuron applied at lable rates efficiently control young plants of common milkweed at the phase of 2-4 leaves.</p>

Orientador: Jose Roberto Trigo / Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-12T18:54:20Z (GMT). No. of bitstreams: 1 Stanton_MarianaAlves_M.pdf: 682844 bytes, checksum: 36b7ed46fcdc82eacc95256aae48e173 (MD5) Previous issue date: 2008 / Resumo: A produção de defesas contra a herbivoria é essencial para o sucesso reprodutivo das plantas, no entanto pode ser custosa. Esses custos resultam da redução de investimento em outras partes do metabolismo, como o crescimento e a reprodução. Nesse estudo, usamos Asclepias curassavica (Apocynaceae: Asclepiadoideae) como modelo para avaliar como uma planta divide seus recursos entre crescimento, reprodução e defesas. A. curassavica é uma planta anual que possui cardenolidas como defesas. Para verificar como esta responde a herbivoria, simulamos a mesma através de dano artificial (DA) e medimos crescimento (biomassa de folhas e raízes) e reprodução (número de flores, frutos e sementes e biomassa de sementes) em um experimento em longo prazo. Em um experimento em curto prazo, medimos a produção de defesas (concentração de cardenolidas), para avaliar se estas podem estar interferindo no crescimento e reprodução desta espécie. Correlacionamos também, em uma população natural, a concentração de cardenolidas com a percentagem de herbivoria foliar. O hormônio jasmonato de metila (JM) é usado para induzir compostos do metabolismo secundário em plantas, sem o custo adicional da remoção de tecido fotossinteticamente ativo causada por dano artificial ou natural. Usamos esse composto em ambos os desenhos experimentais acima ao invés de remoção da área foliar para avaliar se havia indução de cardenolidas e seu efeito sobre o crescimento e reprodução de A. curassavica. Nossos resultados demonstraram uma redução significativa do número total de frutos, sementes e da massa final de raízes em plantas com tratamento de DA em longo prazo. O crescimento do tratamento DA não diferiu do controle, sugerindo crescimento compensatório das folhas à custa do investimento em raízes e em reprodução. Os custos reprodutivos e de crescimento de raízes detectados no experimento de longo prazo podem resultar da diminuição da capacidade fotossintética em plantas danificadas e investimento simultâneo em crescimento compensatório das folhas. No experimento de curto prazo, não houve alteração da razão de indução de cardenolidas sugerindo que o dano artificial não induz defesas nessa espécie. A ausência de correlação entre cardenolidas e porcentagem de dano natural em plantas coletadas em campo pode sugerir dois cenários excludentes: 1. a indução de cardenolidas não seria importante para a defesa de A. curassavica, ou 2. a ausência de correlação, associada a baixa percentagem de herbivoria implicaria em uma defesa constitutiva eficaz contra herbívoros. O tratamento com JM a longo prazo também mostrou uma tendência à redução do crescimento de raízes e redução significativa da biomassa e porcentagem de germinação de sementes. Nas plantas tratadas com JM a curto prazo, houve um aumento significativo de cardenolidas tardio (384 h após tratamento) sugerindo que estas podem contribuir para a redução de aptidão observada no experimento de longo prazo e que existem custos da produção das mesmas. Dano artificial leva à diminuição da aptidão, através de desvio de investimento em raízes para o crescimento compensatório das folhas. No caso do jasmonato, não houve perda de massa fotossinteticamente ativa (folhas) e a redução do crescimento de raízes pode ser resultado de um efeito direto do tratamento de JM ou indireto causado pela indução de outras partes do metabolismo (p.ex. metabolismo secundário) causada por esse fitohormônio. Experimentos futuros devem comparar os presentes resultados com dano real por um dos herbívoros especialistas para avaliar a eficácia do dano artificial em induzir cardenolidas e o papel dessa indução sobre outras partes do metabolismo da mesma / Abstract: Although the investment in defensive traits against herbivory is essential to the reproductive success of plants, it may be costly. These costs result from reduced investment in other metabolic functions such as growth and reproduction. In the present study, we used the milkweed Asclepias curassavica (Apocynaceae: Asclepiadoideae) as a model to study how a plant divides its resources between growth, reproduction and defense. A. curassavica is an annual weed that uses cardenolides as defenses against herbivory. To evaluate how Asclepias curassavica responds to herbivores, we simulated herbivory by artificial damage (AD) and measured growth (leaf and root biomass) and reproduction (number of flowers, fruit and seeds and seed biomass) in a long term experiment. We also measured defensive traits (cardenolide concentration) in a short term experiment to verify whether there is an investment in defense that may interfere with growth and reproduction. We also correlated cardenolide concentration in a natural population with percent leaf damage. As the plant hormone, methyl jasmonate (MJ) is commonly used to induce secondary metabolism in plants without the additional costs of tissue removal, we used this compound in the same experimental designs instead of AD. Our results from the long term AD treatment showed a significant decrease in final root biomass and in total fruit set and seed number. Plants from the AD treatment did not differ from controls in leaf growth, suggesting that there was a compensatory growth in the former at the expense of root growth. The reproductive and growth costs detected in this experiment may result from reduced photosynthetic capacity in damaged plants and concomitant compensatory leaf growth. In the short term, we found no induction of cardenolides compared to controls, suggesting that artificial damage does not induce defenses. The lack of correlation between cardenolides and percentage leaf damage in plants collected from a natural population suggest two self-excluding scenarios: 1. the induction of cardenolides is not important for the defense of A. curassavica, or 2. the lack of correlation, coupled with low herbivore damage, suggests that this plant has an efficient constitutive defense against herbivores. The long term MJ treatment showed a trend in reduced root biomass and significantly reduced seed biomass and percentage germination. In the short term MJ treatment we found a significant increase in cardenolide concentration (after 384 h) suggesting that the production of these defenses is costly and this may have contributed to observed costs in the long term experiment. Apparently, the artificial damage treatment leads to reduced fitness through reduced root growth which is a consequence of compensatory leaf growth. In the methyl jasmonate treatment, there was no removal of photosythetically active tissue (leaves) and the reduced root growth may be a direct effect of this phytohormone or an indirect effect caused by the induction of other metabolic pathways (such as secondary metabolism) caused by this hormone. Future experiments should compare the present results with natural damage by specialist herbivores to evaluate the efficiency of artificial damage in inducing cardenolides and the role of the induction of these substances on other metabolic functions / Mestrado / Mestre em Ecologia

No description available.

Butterfly weed (Asclepias tuberosa) is a primary larval food source for the monarch butterfly (Danaus plexippus). Planting more butterfly weed may stimulate declining monarch populations. To that end, a habitat suitability map was created for Mississippi in GIS using soil pH, soil texture, and land cover. Herbarium data were derived from the Southeast Regional Network of Expertise and Collections (SERNEC) database. Environmental data were from the USDA National Resource Conservation Service geospatial data gateway. Frequency analysis was used to assign scores to environmental variables of SERNEC occurrences using a suitability index. Global positioning systems (GPS) locations of butterfly weed were collected to validate the model. The most suitable model with 78.9% of GPS points in medium to high suitability was a weighted sum overlay with land cover 50%, soil pH 25%, and soil texture 25%. The suitability map may enable conservationists to identify suitable sites for butterfly weed in Mississippi.

The eastern migratory population of monarch butterflies (Danaus plexippus plexippus) has declined by >80% within the last two decades. One possible cause of this decline is the loss of >= 1.3 billion stems of milkweed (Asclepias spp.), which monarchs require for reproduction. In an effort to restore monarchs to a population goal established by the US Fish and Wildlife Service and adopted by Mexico, Canada, and the US, we developed scenarios for amending the Midwestern US landscape with milkweed. Scenarios for milkweed restoration were developed for protected area grasslands, Conservation Reserve Program land, powerline, rail and roadside rights of way, urban/suburban lands, and land in agricultural production. Agricultural land was further divided into productive and marginal cropland. We elicited expert opinion as to the biological potential (in stems per acre) for lands in these individual sectors to support milkweed restoration and the likely adoption (probability) of management practices necessary for affecting restoration. Sixteen of 218 scenarios we developed for restoring milkweed to the Midwestern US were at levels (>1.3 billion new stems) necessary to reach the monarch population goal. One of these scenarios would convert all marginal agriculture to conserved status. The other 15 scenarios converted half of marginal agriculture (730 million stems), with remaining stems contributed by other societal sectors. Scenarios without substantive agricultural participation were insufficient for attaining the population goal. Agricultural lands are essential to reaching restoration targets because they occupy 77% of all potential monarch habitat. Barring fundamental changes to policy, innovative application of economic tools such as habitat exchanges may provide sufficient resources to tip the balance of the agro-ecological landscape toward a setting conducive to both robust agricultural production and reduced imperilment of the migratory monarch butterfly.

O mimetismo é uma adaptação onde duas ou mais espécies, que podem pertencer a grupos taxonômicos distintos, assemelham-se no padrão geral de cor e/ou forma e que devido a isso são confundidas por um outro organismo. Os tipos clássicos de mimetismo, Batesiano e Mülleriano, nem sempre se ajustam a estudos com flores; esta limitação, todavia, não afeta o chamado Sistema Tripartido. O Sistema Tripartido foi proposto originalmente por Wolfgang Wickler (1968) e aperfeiçoado por Richard I. Vane Right (1976); o sistema é formado por três elementos constantes: modelo (S1), o mímico (S2) e operador (O – o qual Wickler originalmente representou como R – receptor de sinal). O presente estudo concentra-se em três plantas que coexistem na restinga do Parque Estadual de Itapeva, Torres, as quais mostram um padrão convergente na coloração de suas flores. As espécies são Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), ambas produtoras de néctar, e Epidendrum fulgens (Orchidaceae [Epidendroideae]), aparentemente não produtora de néctar e considerado o suposto mímico do sistema. O trabalho de campo foi realizado entre a primavera de 2004 e verão de 2006. Para tanto, foram avaliados aspectos concernentes ao número e identificação dos polinizadores potenciais (especialmente borboletas – excepcionalmente, uma ou duas espécies de mariposas foram encontradas, porém mostrando baixa contribuição); às fenofases de floração e frutificação das três espécies. Também foram realizados experimentos de cruzamento em E. fulgens (autopolinização espontânea e induzida, geitonogamia, xenogamia); remoção de polinários de E. fulgens em relação à distância da fonte de néctar (L. camara), bem como uma descrição da anatomia do nectário de E. fulgens para verificar a existência de tecido produtor de néctar. Dentre as 28 espécies de lepidópteros diurnos que foram registradas, destacam-se Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae) e Urbanus simplicius (Hesperiidae), as quais visitam as três espécies de plantas. Não houve sincronia de floração entre as fenofases, ou seja, não houve diferença estatística entre A. curassavica, Epidendrum fulgens e Lantana camara. Os experimentos de cruzamento mostraram que E. fulgens é polinizador-dependente. A remoção de polinários de E. fulgens foi baixa, talvez porque a oferta de flores seja alta. Os testes sobre a anatomia do nectário de E. fulgens sugerem que não há produção de néctar (Anexo 2). Estes resultados sugerem que A. curassavica, Epidendrum fulgens e Lantana camara, no cenário ecológico do Parque Estadual de Itapeva, não constituem um complexo mimético. Porém, isto não invalida a possibilidade de que estas plantas realmente constituam um sistema mimético, considerando-se que o presente estudo foi realizado dentro de um curto espaço de tempo, enquanto que estudos a respeito de processos evolutivos requerem maior tempo de observações. / Mimicry is an adaptation where two or more species which can belong to different taxonomic groups, are similar in color pattern and/or behavior, and that for this reason are not discriminated by a third species. Classical Batesian and Müllerian mimicry are not generally useful to describe examples of plant mimicry; however, the so-called tripartite mimicry system does not suffer this restriction. The Tripartite System was first envisaged by Wolfgang Wickler (1968) and latter elaborated and expanded by Richard I. Vane-Wright (1976); the system is based in three constant elements, the model (S1), the mimic (S2) and the operator (O – which Wickler originally represented as R – signal receiver). The present study concentrates on three plants that coexist at Parque Estadual de Itapeva, Torres, RS, and that show a convergence in the pattern of flower color. The species are Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), both nectar producers and Epidendrum fulgens (Orchidaceae [Epidendroideae]) which does not produce nectar and that is considered the mimic in the system. Field work has been done between the spring of 2004 and the summer of 2006. The variables studied were the number and the identity of the potential pollinators (specifically butterflies – exceptionally one or two moths were found but with little contribution), flower and fruit phenology in the three species. Experiments in the field were conducted in E. fulgens to ascertain the mating system of this plant (spontaneous and induced self-pollination, geitonogamy and xenogamy). Moreover, to complement the test of hypothesis of mimicry of E. fulgens in relation to L. camara, the frequency of removal of pollinaria in the orchid were estimated according to the distance from the nectar source. A description of the anatomy of the floral nectary of E. fulgens was also made, to test for nectar production. In the whole, 28 species of diurnal Lepidoptera were observed visiting the flowers of at least two of the three species of plants; among them, Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae), and Urbanus simplicius (Hesperiidae) did so for the three species of plants. As for the synchrony in flower phenology, there was no statistical significance for any combination of plants. The experiments for mating system of E. fulgens showed that this orchid is pollinator-dependent. Pollinaria removal was very low, which can be related to the extremely large number of inflorescences available in the area. The study of the anatomy of the nectary in E. fulgens suggested that the orchid does not produce nectar in detectable proportions (Appendix 2). Given all the results here reported, there is no indication that at the ecological scenario of the Parque Estadual de Itapeva, L. camara, E. fulgens and A. curassavica are members of a mimetic system. This does not invalidate the possibility that they really are a mimetic system, since the time spent in the study was short and that the evolution of adaptations demand a larger amount of time to be detected.

O mimetismo é uma adaptação onde duas ou mais espécies, que podem pertencer a grupos taxonômicos distintos, assemelham-se no padrão geral de cor e/ou forma e que devido a isso são confundidas por um outro organismo. Os tipos clássicos de mimetismo, Batesiano e Mülleriano, nem sempre se ajustam a estudos com flores; esta limitação, todavia, não afeta o chamado Sistema Tripartido. O Sistema Tripartido foi proposto originalmente por Wolfgang Wickler (1968) e aperfeiçoado por Richard I. Vane Right (1976); o sistema é formado por três elementos constantes: modelo (S1), o mímico (S2) e operador (O – o qual Wickler originalmente representou como R – receptor de sinal). O presente estudo concentra-se em três plantas que coexistem na restinga do Parque Estadual de Itapeva, Torres, as quais mostram um padrão convergente na coloração de suas flores. As espécies são Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), ambas produtoras de néctar, e Epidendrum fulgens (Orchidaceae [Epidendroideae]), aparentemente não produtora de néctar e considerado o suposto mímico do sistema. O trabalho de campo foi realizado entre a primavera de 2004 e verão de 2006. Para tanto, foram avaliados aspectos concernentes ao número e identificação dos polinizadores potenciais (especialmente borboletas – excepcionalmente, uma ou duas espécies de mariposas foram encontradas, porém mostrando baixa contribuição); às fenofases de floração e frutificação das três espécies. Também foram realizados experimentos de cruzamento em E. fulgens (autopolinização espontânea e induzida, geitonogamia, xenogamia); remoção de polinários de E. fulgens em relação à distância da fonte de néctar (L. camara), bem como uma descrição da anatomia do nectário de E. fulgens para verificar a existência de tecido produtor de néctar. Dentre as 28 espécies de lepidópteros diurnos que foram registradas, destacam-se Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae) e Urbanus simplicius (Hesperiidae), as quais visitam as três espécies de plantas. Não houve sincronia de floração entre as fenofases, ou seja, não houve diferença estatística entre A. curassavica, Epidendrum fulgens e Lantana camara. Os experimentos de cruzamento mostraram que E. fulgens é polinizador-dependente. A remoção de polinários de E. fulgens foi baixa, talvez porque a oferta de flores seja alta. Os testes sobre a anatomia do nectário de E. fulgens sugerem que não há produção de néctar (Anexo 2). Estes resultados sugerem que A. curassavica, Epidendrum fulgens e Lantana camara, no cenário ecológico do Parque Estadual de Itapeva, não constituem um complexo mimético. Porém, isto não invalida a possibilidade de que estas plantas realmente constituam um sistema mimético, considerando-se que o presente estudo foi realizado dentro de um curto espaço de tempo, enquanto que estudos a respeito de processos evolutivos requerem maior tempo de observações. / Mimicry is an adaptation where two or more species which can belong to different taxonomic groups, are similar in color pattern and/or behavior, and that for this reason are not discriminated by a third species. Classical Batesian and Müllerian mimicry are not generally useful to describe examples of plant mimicry; however, the so-called tripartite mimicry system does not suffer this restriction. The Tripartite System was first envisaged by Wolfgang Wickler (1968) and latter elaborated and expanded by Richard I. Vane-Wright (1976); the system is based in three constant elements, the model (S1), the mimic (S2) and the operator (O – which Wickler originally represented as R – signal receiver). The present study concentrates on three plants that coexist at Parque Estadual de Itapeva, Torres, RS, and that show a convergence in the pattern of flower color. The species are Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), both nectar producers and Epidendrum fulgens (Orchidaceae [Epidendroideae]) which does not produce nectar and that is considered the mimic in the system. Field work has been done between the spring of 2004 and the summer of 2006. The variables studied were the number and the identity of the potential pollinators (specifically butterflies – exceptionally one or two moths were found but with little contribution), flower and fruit phenology in the three species. Experiments in the field were conducted in E. fulgens to ascertain the mating system of this plant (spontaneous and induced self-pollination, geitonogamy and xenogamy). Moreover, to complement the test of hypothesis of mimicry of E. fulgens in relation to L. camara, the frequency of removal of pollinaria in the orchid were estimated according to the distance from the nectar source. A description of the anatomy of the floral nectary of E. fulgens was also made, to test for nectar production. In the whole, 28 species of diurnal Lepidoptera were observed visiting the flowers of at least two of the three species of plants; among them, Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae), and Urbanus simplicius (Hesperiidae) did so for the three species of plants. As for the synchrony in flower phenology, there was no statistical significance for any combination of plants. The experiments for mating system of E. fulgens showed that this orchid is pollinator-dependent. Pollinaria removal was very low, which can be related to the extremely large number of inflorescences available in the area. The study of the anatomy of the nectary in E. fulgens suggested that the orchid does not produce nectar in detectable proportions (Appendix 2). Given all the results here reported, there is no indication that at the ecological scenario of the Parque Estadual de Itapeva, L. camara, E. fulgens and A. curassavica are members of a mimetic system. This does not invalidate the possibility that they really are a mimetic system, since the time spent in the study was short and that the evolution of adaptations demand a larger amount of time to be detected.

O mimetismo é uma adaptação onde duas ou mais espécies, que podem pertencer a grupos taxonômicos distintos, assemelham-se no padrão geral de cor e/ou forma e que devido a isso são confundidas por um outro organismo. Os tipos clássicos de mimetismo, Batesiano e Mülleriano, nem sempre se ajustam a estudos com flores; esta limitação, todavia, não afeta o chamado Sistema Tripartido. O Sistema Tripartido foi proposto originalmente por Wolfgang Wickler (1968) e aperfeiçoado por Richard I. Vane Right (1976); o sistema é formado por três elementos constantes: modelo (S1), o mímico (S2) e operador (O – o qual Wickler originalmente representou como R – receptor de sinal). O presente estudo concentra-se em três plantas que coexistem na restinga do Parque Estadual de Itapeva, Torres, as quais mostram um padrão convergente na coloração de suas flores. As espécies são Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), ambas produtoras de néctar, e Epidendrum fulgens (Orchidaceae [Epidendroideae]), aparentemente não produtora de néctar e considerado o suposto mímico do sistema. O trabalho de campo foi realizado entre a primavera de 2004 e verão de 2006. Para tanto, foram avaliados aspectos concernentes ao número e identificação dos polinizadores potenciais (especialmente borboletas – excepcionalmente, uma ou duas espécies de mariposas foram encontradas, porém mostrando baixa contribuição); às fenofases de floração e frutificação das três espécies. Também foram realizados experimentos de cruzamento em E. fulgens (autopolinização espontânea e induzida, geitonogamia, xenogamia); remoção de polinários de E. fulgens em relação à distância da fonte de néctar (L. camara), bem como uma descrição da anatomia do nectário de E. fulgens para verificar a existência de tecido produtor de néctar. Dentre as 28 espécies de lepidópteros diurnos que foram registradas, destacam-se Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae) e Urbanus simplicius (Hesperiidae), as quais visitam as três espécies de plantas. Não houve sincronia de floração entre as fenofases, ou seja, não houve diferença estatística entre A. curassavica, Epidendrum fulgens e Lantana camara. Os experimentos de cruzamento mostraram que E. fulgens é polinizador-dependente. A remoção de polinários de E. fulgens foi baixa, talvez porque a oferta de flores seja alta. Os testes sobre a anatomia do nectário de E. fulgens sugerem que não há produção de néctar (Anexo 2). Estes resultados sugerem que A. curassavica, Epidendrum fulgens e Lantana camara, no cenário ecológico do Parque Estadual de Itapeva, não constituem um complexo mimético. Porém, isto não invalida a possibilidade de que estas plantas realmente constituam um sistema mimético, considerando-se que o presente estudo foi realizado dentro de um curto espaço de tempo, enquanto que estudos a respeito de processos evolutivos requerem maior tempo de observações. / Mimicry is an adaptation where two or more species which can belong to different taxonomic groups, are similar in color pattern and/or behavior, and that for this reason are not discriminated by a third species. Classical Batesian and Müllerian mimicry are not generally useful to describe examples of plant mimicry; however, the so-called tripartite mimicry system does not suffer this restriction. The Tripartite System was first envisaged by Wolfgang Wickler (1968) and latter elaborated and expanded by Richard I. Vane-Wright (1976); the system is based in three constant elements, the model (S1), the mimic (S2) and the operator (O – which Wickler originally represented as R – signal receiver). The present study concentrates on three plants that coexist at Parque Estadual de Itapeva, Torres, RS, and that show a convergence in the pattern of flower color. The species are Asclepias curassavica (Apocynaceae [Asclepiadoideae]), Lantana camara (Verbenaceae), both nectar producers and Epidendrum fulgens (Orchidaceae [Epidendroideae]) which does not produce nectar and that is considered the mimic in the system. Field work has been done between the spring of 2004 and the summer of 2006. The variables studied were the number and the identity of the potential pollinators (specifically butterflies – exceptionally one or two moths were found but with little contribution), flower and fruit phenology in the three species. Experiments in the field were conducted in E. fulgens to ascertain the mating system of this plant (spontaneous and induced self-pollination, geitonogamy and xenogamy). Moreover, to complement the test of hypothesis of mimicry of E. fulgens in relation to L. camara, the frequency of removal of pollinaria in the orchid were estimated according to the distance from the nectar source. A description of the anatomy of the floral nectary of E. fulgens was also made, to test for nectar production. In the whole, 28 species of diurnal Lepidoptera were observed visiting the flowers of at least two of the three species of plants; among them, Agraulis vanillae maculosa, Dryas iulia alcionea, Heliconius erato phyllis, Tegosa claudina (Nymphalidae), and Urbanus simplicius (Hesperiidae) did so for the three species of plants. As for the synchrony in flower phenology, there was no statistical significance for any combination of plants. The experiments for mating system of E. fulgens showed that this orchid is pollinator-dependent. Pollinaria removal was very low, which can be related to the extremely large number of inflorescences available in the area. The study of the anatomy of the nectary in E. fulgens suggested that the orchid does not produce nectar in detectable proportions (Appendix 2). Given all the results here reported, there is no indication that at the ecological scenario of the Parque Estadual de Itapeva, L. camara, E. fulgens and A. curassavica are members of a mimetic system. This does not invalidate the possibility that they really are a mimetic system, since the time spent in the study was short and that the evolution of adaptations demand a larger amount of time to be detected.

Plants are under selection to simultaneously attract pollinators while deterring herbivores. This dilemma can lead to tradeoffs in floral traits, which are traditionally thought to be optimized for pollinators. My dissertation addresses the ecological costs and putative functional significance of nectar secondary metabolites, a paradoxical but widespread phenomenon in the angiosperms. I investigate this issue from the pollinator’s perspective using a series of controlled laboratory investigations focused primarily on the bumble bee Bombus impatiens and the nectar alkaloid gelsemine, from Gelsemium sempervirens. I begin by demonstrating that nectar enriched with the alkaloid gelsemine significantly deters visits from bumble bees at a range of natural alkaloid concentrations. However, this aversion can be mitigated by increasing the sucrose concentration such that the alkaloid-rich nectar is more rewarding than its alkaloid-free counterpart. I then demonstrate that the consumption of gelsemine-rich nectar can inhibit oocyte development and protein utilization in bees, but that this effect is limited to bees of suboptimal condition. Continuous consumption of the nectar alkaloid gelsemine also leads to a reduction in the pathogen load of bumble bees infected with Crithidia bombi, but direct interactions between the pathogen and the alkaloid have no impact on infection intensity. Gelsemine also fails to inhibit floral yeast growth, suggesting that nectar alkaloids may not be universally antimicrobial. Finally, I demonstrate that gross nectar cardenolides from the genus Asclepias are strongly correlated with gross leaf cardenolides and that the majority of individual cardenolides found in nectar are a subset of those identified in leaves. This pattern suggests that nectar cardenolides are a consequence of defense for Asclepias; however, they may not be a costly corollary because bumble bees show an overall preference for nectar cardenolides at mean concentrations. Altogether, my dissertation provides a new perspective on the role of chemical defenses against herbivores in plant-pollinator interactions.

Plants are under selection to simultaneously attract pollinators while deterring herbivores. This dilemma can lead to tradeoffs in floral traits, which are traditionally thought to be optimized for pollinators. My dissertation addresses the ecological costs and putative functional significance of nectar secondary metabolites, a paradoxical but widespread phenomenon in the angiosperms. I investigate this issue from the pollinator’s perspective using a series of controlled laboratory investigations focused primarily on the bumble bee Bombus impatiens and the nectar alkaloid gelsemine, from Gelsemium sempervirens. I begin by demonstrating that nectar enriched with the alkaloid gelsemine significantly deters visits from bumble bees at a range of natural alkaloid concentrations. However, this aversion can be mitigated by increasing the sucrose concentration such that the alkaloid-rich nectar is more rewarding than its alkaloid-free counterpart. I then demonstrate that the consumption of gelsemine-rich nectar can inhibit oocyte development and protein utilization in bees, but that this effect is limited to bees of suboptimal condition. Continuous consumption of the nectar alkaloid gelsemine also leads to a reduction in the pathogen load of bumble bees infected with Crithidia bombi, but direct interactions between the pathogen and the alkaloid have no impact on infection intensity. Gelsemine also fails to inhibit floral yeast growth, suggesting that nectar alkaloids may not be universally antimicrobial. Finally, I demonstrate that gross nectar cardenolides from the genus Asclepias are strongly correlated with gross leaf cardenolides and that the majority of individual cardenolides found in nectar are a subset of those identified in leaves. This pattern suggests that nectar cardenolides are a consequence of defense for Asclepias; however, they may not be a costly corollary because bumble bees show an overall preference for nectar cardenolides at mean concentrations. Altogether, my dissertation provides a new perspective on the role of chemical defenses against herbivores in plant-pollinator interactions.