Morphological, Anatomical, and Phytochemical Studies of Carlina acaulis L. Cypsela.

Carlina acaulis L. has a long tradition of use in folk medicine. The chemical composition of the roots and green parts of the plant is quite well known. There is the lowest amount of data on the cypsela (fruit) of this plant. In this study, the microscopic structures and the chemical composition of the cypsela were investigated. Preliminary cytochemical studies of the structure of the Carlina acaulis L. cypsela showed the presence of substantial amounts of protein and lipophilic substances. The chemical composition of the cypsela was investigated using spectrophotometry, gas chromatography with mass spectrometry, and high-performance liquid chromatography with spectrophotometric and fluorescence detection. The cypsela has been shown to be a rich source of macro- and microelements, vegetable oil (25%), α-tocopherol (approx. 2 g/kg of oil), protein (approx. 36% seed weight), and chlorogenic acids (approx. 22 g/kg seed weight). It also contains a complex set of volatile compounds. The C. acaulis cypsela is, therefore, a valuable source of nutrients and bioactive substances.

Morphoanatomical and phytochemical studies for the quality control of Neurolaena lobata (L.) R.Br. ex Cass. (Asteraceae) / Estudios morfoanatomicos y fitoquimicos para el control de calidad de Neurolaena lobata (L.) R.Br. ex Cass. (Asteraceae)

Neurolaena lobata (L.) R.Br. ex Cass. (Asteraceae)Is a popular folk remedy for in Central America. The plant is of commercial value in Guatemala but so far there is not any monograph to guide regional laboratories on ensuring identity and chemical tests for this species. As identity test we here run macro and micro morphoanatomical studies of the characters of the vegetative organs. We also developed standard chemical tests for quality by both TLC and HPLC for infusions and tinctures of varying alcoholic strength. Their radical scavenging activities in DPPH and NO were also measured. Macro and micro morphoanatomical characters of the vegetative organs present a set of characteristics to facilitate the identification of dry powdered samples of this species. We developed optimal conditions for the TLC and HPLC phytochemical fingerprints of the 4 most common pharmacopoeial liquid herbal preparations from this herbal drug, namely infusion, 70%, 45% and 20% hydroalcoholic tinctures. Our work provides the Latin-American industry with a set of analyses to establish the identity and chemistry of N. lobata samples for quality control purposes.

Neurolaena lobata (L.) R.Br. ex cass. (Asteraceae) es un remedio popular popular en América Central. La planta tiene un valor comercial en Guatemala, pero hasta el momento no existe una monografía que guíe a los laboratorios regionales para garantizar la identidad y las pruebas químicas para esta especie. Como prueba de identidad proponemos estudios macro y micro morfoanatómicos de los caracteres de los órganos vegetativos. También desarrollamos pruebas químicas de calidad mediante CCF y CLAR para infusiones y tinturas de grado alcohólico variable. También se midieron sus actividades de captación de radicales en DPPH y NO. Los caracteres macro y micro morfoanatómicos de los órganos vegetativos presentan un conjunto de características para facilitar la identificación de muestras de polvo seco de esta especie. Desarrollamos condiciones óptimas para las huellas dactilares fitoquímicas de CCF y CLAR de las 4 preparaciones herbales líquidas farmacopéicas más comunes de esta droga herbal, a saber, infusión, 70%, 45% y 20% tinturas hidroalcohólicas. Nuestro trabajo proporciona a la industria latinoamericana un conjunto de análisis base para establecer la identidad y la química de las muestras de N. lobata con fines de control de calidad.

Comparative authentication of three "snow lotus" herbs by macroscopic and microscopic features.

"Snow lotus" is a famous Chinese Materia Medica derived from species of the genus Saussurea (Compositae). To differentiate three representative easily-confused snow lotus herbs, namely, Saussurea involucrata (Kar. et Kir.) Sch.-Bip, Saussurea laniceps Hand.-Mazz., and Saussurea medusa Maxim., macroscopic features of the three herbs were systemically observed, and microscopic features were compared by using ordinary light microscopy, polarized light microscopy and scanning electron microscopy (SEM). The results indicate that, as for macroscopic identification, capitula situation and arrangement, and as for microscopic identification, pollen grains, nonglandular hairs, glandular hairs, and cells of inner surface of the microdiodange can be used to authenticate the three snow lotus herbs. Comprehensive table comparing the characteristics were presented in this study. SEM has been found to provide a number of unique characteristics of pollen grains. Based on the observation of pollen grains, evolution sequence of the three species was speculated. The present method was proven to be efficient, convenient, simple, and reliable, which was successfully applied to the authentication of three snow lotus herbs.

Morfoanatomia do eixo vegetativo aéreo de Ayapana triplinervis (Vahl) R.M. King & H. Rob. (Asteraceae) / Morphoanatomy of the vegetative aerial axis of Ayapana triplinervis (Vahl) RM King & H. Rob. (Asteraceae)

Ayapana triplinervis (Asteraceae) é uma erva medicinal da Amazônia conhecida como "Japana branca" e "Japana roxa" utilizada como tônico, digestivo, antidiarréico dentre outros. Objetivou-se caracterizar neste estudo a morfoanatomia caulinar e foliar de A. triplinervis, para corroborar se esta espécie apresenta morfotipos, além de fornecer dados para sua identificação. Amostras foram coletadas no Horto de Plantas Medicinais da Universidade Federal Rural da Amazônia e Embrapa Amazônia Oriental, analisadas morfologicamente e segundo técnicas usuais em anatomia vegetal. O caule é cilíndrico branco ou roxo, com epiderme unisseriada persistente e endoderme com bainha amilífera e estrias de Caspary. As folhas são simples, opostas, triplinervadas, com base atenuada ou decorrente. Possuem estômatos anomocíticos, cera em placas ou granular, glândulas peroladas, mesofilo dorsiventral com feixes colaterais e ductos secretores. A. triplinervis apresentou caracteres morfoanatômicos diferentes entre os tipos branca e roxa, o que possibilita a distinção dos dois morfotipos.

Ayapanatriplinervis (Asteraceae), also known as "Japana branca" and "Japana roxa" in Brazil, is a medicinal herb from the Amazon used as tonic, digestive, antidiarrheal, among others. This study aimed to characterize the morphoanatomy of the aerial vegetative axis of A. triplinervis to corroborate that this species has morphotypes, in order to provide data for their identification. Samples were collected in the Garden of Medicinal Plants of the Federal Rural University of Amazônia and Embrapa Amazônia Oriental, and they were analyzed morphologically and by the standardtechniques of plant anatomy. The stem is cylindrical, white or purple, with uniseriate epidermis, trichomes and endoderm with starch sheath and Casparian strips. The leaves are simple, opposite, with attenuate or arising basis; they have anomocytic stomata, epicuticular wax plates or granules, pearl glands, dorsiventral mesophyll with collateral vascular bundles and secretory ducts. Ayapana triplinervis showed different morphoanatomical features betweenthe white and purple types, which enablesthe distinction between the two morphotypes.

[Micro-macroscopical identification of Siegesbeckiae herba].

OBJECTIVE: To comprehend the connections and differences of the three sources of Siegesbeckiae Herba. METHOD: Using traditional Chinese medicine micro-macroscopical identification to identify these three sources of Siegesbeckiae Herba. RESULT: Three sources of Siegesbeckiae Herba are obviously different when using micro-macroscopical identification. CONCLUSION: Micro-macroscopical identification can distinguish three sources of Siegesbeckiae Herba veritably and directly.

[Macroscopic and microscopic identification of Ligularia przewalskii].

OBJECTIVE: To provide macroscopic and microscopic identification basis for Ligularia przewalskii. METHODS: Macroscopic and microscopic identification of roots, stems and leaves of Ligularia przewalskii were carried out with the methods of paraffin section, leaves epidermal section and powder transdermal section. RESULTS: The microscopic characteristics included: Open collateral vascular bundles in stem were not in the same size and arranged in two rings; Lots of fiber bundles scattered in the column parts; There were two vascular bundles in principal vein of leaf; Anticlinal wall of upper epidermis cells was thickened like moniliform, lower epidermis were like waves with irregular; The type of stoma was anomocytic; Calcium oxalate acicular crystal could be seen in the powder. CONCLUSION: These features can provide references for identification of Ligularia przewalskii.

Morfoanatomía comparativa de dos especies de Acanthospermum (ASTERACEAE) / Comparative morpho-anatomy of two species of Acanthospermum (ASTERACEAE)

RESUMEN: Para el presente trabajo se ha escogido dos especies de las plantas reportadas como antiinflamatorias, dentro del marco del Proyecto "Un aporte para el control de calidad de drogas de origen vegetal: Análisis micrográfico de drogas vegetales empleadas en la medicina popular paraguaya como antiinflamatorias y comercializadas en los mercados de Asunción, Luque, San Lorenzo, Paraguarí y Caacupé": "tapekué", Acanthospermum australe (Loef.) Kuntze y "toro rati", A. hispidum DC. (Asteraceae); con el fin de realizar la caracterización morfoanatómica foliar y caulinar de ambas especies. Los caracteres morfoanatómicos diferenciales encontrados son: A. australe; hierba decumbente o ascendente, hoja levemente pubescente, rómbico-ovada; base atenuada, corto peciolo, fruto aquenio fusiforme cubierto de espinas ganchudas; parénquima en empalizada con dos hileras de células, pelos glandulares que forman una depresión en la superficie foliar . A. hispidum, hierba erecta, hoja ovada o deltoide ovada, base cuneiforme, sésil; fruto aquenio cuneiforme comprimido, cubierto de espinas ganchudas, de las cuales dos sobresalen en forma divergente a manera de cuernos; parénquima en empalizada con una hilera de células. Palabras clave: morfoanatomía, Acanthospermum, planta medicinal.

Microscopical discrimination of the subterranean organs of medicinally used plants of the Cichorieae and their relatives.

CONTEXT: Light microscopy is in most cases a quick method for the identification and discrimination of medicinally used plant drugs; moreover, this technique is very inexpensive. Reliable descriptions of the anatomy of plants and their adulterations are prerequisites for necessary purity controls. OBJECTIVE: The anatomy of the subterranean organs of 18 pharmaceutically useful as well as related but inconsiderable Asteraceae species from nine genera (Taraxacum F. H. Wigg., Leontodon L., Scorzoneroides Moench, Hypochaeris L., Crepis L., Aposeris Neck., Cichorium L., Scorzonera L., and Tragopogon L.; tribe Cichorieae, Asteraceae) is described in detail and graphically illustrated. Features characterizing and discriminating the studied taxa are presented and discussed. MATERIALS AND METHODS: The roots/rhizomes of various species were examined by means of light microscopy. RESULTS: Useful anatomical characters were found for the discrimination between the species, and some of them were examined for the first time. DISCUSSION AND CONCLUSION: Discrimination of most genera and species investigated was possibly based on the anatomy of their underground parts. The identified characters may be effectively used for quality control of commercial drugs and the identification of adulterations.

Identificação de espécies da família Asteraceae, revisão sobre usos e triagem fitoquímica do gênero Eremanthus da Reserva Boqueirão, Ingaí-MG / Asteraceae species identification, use revision and phytochemical screening of Eremanthus genus in Boqueirão Ecological Reserve, Ingaí - Minas Gerais State, Brazil

Com o objetivo de identificar espécies da família Asteraceae, revisar seus usos e realizar triagem fitoquímica preliminar do gênero Eremanthus procederam-se coletas botânicas semanais de espécimes na Reserva Boqueirão, localizada em Ingaí, Minas Gerais. As amostras foram herborizadas e identificadas utilizando-se bibliografia especializada e comparação com espécimes disponíveis no Herbário ESAL, da Universidade Federal de Lavras. A revisão dos usos foi feita através de consulta a obras clássicas e artigos científicos contendo relatos sobre levantamentos etnobotânicos realizados na área de estudo. Para triagem fitoquímica empregaram-se reagentes específicos para cada grupo de metabólito. Foram levantadas 102 espécies da família Asteraceae, sendo 32 delas úteis para o homem. A triagem fitoquímica dos extratos hidroalcoólicos indicaram a presença de açúcares redutores, carboidratos, aminoácidos, taninos, flavonóides, glicosídeos cardiotônicos, carotenóides, esteróides e triterpenóides, depsídeos e depsidonas, derivados de cumarina, saponinas espumídicas, alcalóides, purinas, polissacarídeos e antraquinonas. Não foram detectados ácidos orgânicos, catequinas, lactonas sesquiterpênicas e azulenos.

To identify Asteraceae species, review the utilization and perform a preliminary phytochemical screening of some species of Eremanthus genus, plants were weekly collected in Boqueirão Ecological Reserve, located in Ingaí, Minas Gerais State, Brazil. The samples were herborized and identified by using a specialized bibliography and comparison with the species available in the Herbarium ESAL of the Federal University of Lavras. The utilization review was carried out by means of bibliographical research and ethnobotanical surveys in the sampling area. Specific reagents for each group of compounds were used for phytochemical screening. From the 102 Asteraceae species investigated, 32 were reported to be of use to humans. The phytochemical screening of the hydroalcoholic extracts indicated the presence of reducing sugars, carbohydrates, amino acids, tannins, flavonoids, glycosides cardiotonics, carotenoids, steroids and triterpenoids, depsides and depsidones, coumarin derivatives, soapy saponins, alkaloids, purines, polysaccharides and anthraquinones. On the other hand, organic acids, catechins, sesquiterpene lactones and azulenes were not detected.

Eocene Patagonia fossils of the daisy family.

Fossil capitula and pollen grains of Asteraceae from the Eocene of Patagonia, southern Argentina, exhibit morphological features recognized today in taxa, such as Mutisioideae and Carduoideae, that are phylogenetically close to the root of the asteracean tree. This fossil supports the hypothesis of a South American origin of Asteraceae and an Eocene age of divergence and suggests that an ancestral stock of Asteraceae may have formed part of a geoflora developed in southern Gondwana before the establishment of effective dispersal barriers within this landmass.

Effects of aqueous extract of Aspilia africana on reproductive functions of female Wistar rats.

Effects of Aspilia africana leaf extract on oestrous cycle and ovulation were studied in adult female Wistar strain rats. Cyclic female rats weighing 150 to 200 g were divided into two study groups: the oestrous study and ovulation study group. For the oestrous study, the experimental group received 500 mg kg(-1) b. wt. of the extract for 14 days while the control group received distilled water for the same period. In both groups, vaginal lavage was taken daily from the 5th day to monitor the oestrous cycle. For the ovulation study, there was a control group and two experimental groups. The control group received distilled water while group 1 and 2 received 500 and 1000 mg kg(-1) b.wt. of Aspilia africana leaf extract for 16 days, respectively. The animals were sacrificed on the estrous following the treatment. The results showed a significant decrease in the body weight of the treated rats (p = 0.01) and the oestrous cycle was altered after the commencement of extract. This was indicated by the prolonged proestrous and a reduced dioestrus and estrus. There was a dose-dependent reduction in the ovulation s shown by the reduced number of ova observed in the oviduct from the treated rats compared with control (p<0.05). The extract caused inflammation of the fallopian tube, degeneration in the ovarian cortex in the stroma cells of the ovary and disruption of the endometrium of the uterus. Results suggest that aqueous extract of Aspilia africana leaf has antifertility effect by altering oestrous cycle and causing a dose dependent adverse effect on ovulation in Wistar strain rats.

[Pharmacognostic identification of Centipeda minima].

OBJECTIVE: To conduct pharmacognostic identification of Centipeda minima. METHODS: To conduct characteristic identification, micro-identification and TLC identification of Centipeda minima. RESULTS: We established a systematic pharmacognostic identification method including characteristic identification, micro-identification and TLC identification. CONCLUSION: Brevilin is used as chemical reference firstly in TLC identification of Centipeda minima, and combined with characteristic identification and micro-identification, it will raise the rapidity and accuracy of the identification of Centipeda minima.

Authenticating and distinguishing the eight species of traditional Tibetan medicine "Meiduoluomi" by microscopic technique.

Because of the morphological and macroscopic similarity, many species of Erigeron and Aster (Asteraceae) are confusable and usually used under the same name "Meiduoluomi" in traditional Tibetan medicine (TTM). To find an easy, quick, and reliable method to authenticate and distinguish the eight main medicinal plants of these species, the light microscope was used to reveal the morphoanatomic details. The fixed, sectioned, and stained plant materials and epidermis materials were studied by microscopic techniques. The results of the microscopic features are systematically described and illustrated, and comparison parameters are presented. Furthermore, a key to the eight species of "Meiduoluomi" was constructed. Microscopy can be unambiguously used to authenticate and distinguish the eight main species of TTM "Meiduoluomi."

Estudo anatômico de folha e caule de Elephantopus mollis Kunth (Asteraceae) / Anatomical study of the leaf and stem of Elephantopus mollis Kunth (Asteraceae)

Conhecida comumente como erva-de-colégio, erva-grossa e sussuaiá, Elephantopus mollis Kunth (Asteraceae) é uma espécie herbácea perene, de base sublenhosa, ramos curtos e flores arroxeadas. Suas folhas são empregadas como emoliente, resolutivo, sudorífico e no tratamento de bronquite, tosse e gripe na medicina popular. O objetivo deste trabalho foi estudar caracteres anatômicos foliares e caulinares, a fim de fornecer subsídios aplicáveis à identificação da planta medicinal. Folhas adultas e fragmentos de caules foram fixados, seccionados e corados com azul de astra/fucsina básica ou com azul de toluidina. Testes microquímicos e análise ultra-estrutural de superfície foram também executados. A folha possui epiderme uniestratificada, recoberta por cutícula estriada e, em vista frontal, tem células com contorno ondulado. Ocorrem tricomas tectores e glandulares. Os primeiros são eretos, pluricelulares e unisseriados. Os tricomas glandulares são capitados e bisseriados. O mesofilo é dorsiventral e a nervura central é percorrida por feixes vasculares colaterais dispostos em arco aberto. O caule, em estrutura secundária inicial, possui epiderme unisseriada com cutícula estriada e tricomas semelhantes aos da folha. No córtex, observam-se colênquima anelar alternado com clorênquima e endoderme com estrias de Caspary. Calotas esclerenquimáticas apõem-se ao floema. Este é formado pelo câmbio vascular em direção à periferia e o xilema é gerado de modo centrípeto. A medula é parenquimática. Pequenos cristais de oxalato de cálcio estão presentes na folha e no caule.

Commonly known as erva-de-colégio in Portuguese, Elephantopus mollis Kunth (Asteraceae) is a perennial herb, with slightly lignified base, short branches and purple flowers. Its leaves are used as emollient, vulnerary and diaphoretic, and to treat bronchitis, coughs and influenza in folk medicine. The aim of this work was to study leaf and stem anatomical characters, in order to expand the knowledge for this medicinal plant identification. Adult leaves and stem fragments were fixed, sectioned and stained either with astra blue/basic fuchsine or toluidine blue. Microchemical essays and scanning electron analysis were also carried out. The leaf has wavy-contoured cells in surface view, forming a single layer and coated with striate cuticle. Non-glandular and glandular trichomes occur. The former is erect, multicellular and uniseriate. The glandular ones are capitate and biseriate. The mesophyll is dorsiventral and the midrib is traversed by collateral vascular bundles arranged in open arc. The stem, in initial secondary growth, has uniseriate epidermis coated with striate cuticle and trichomes similar to the leaf. In the cortex, annular collenchyma alternates with chlorenchyma and an endodermis with Casparian strips is found. Sclerenchymatic caps adjoin the phloem, which is formed outwards by the vascular cambia, while the xylem is produced inwards. The pith is parenchymatic. Small calcium oxalate crystals are present in the leaf and stem.

Style morphological diversity of some Asteraceae species from Argentina: systematic and functional implications.

Stylar micromorphological diversity of 42 Asteraceae species from Argentina was analysed considering species phylogenetic membership and some floral reproductive functions (pollen presentation and pollen reception). In particular, the morphology and organisation of pollen presenter (sweeping hairs) and pollen receptive structures (stigmatic papillae) were described. Results showed that style morphology of the studied species is far more diverse than the categories previously established for Asteraceae, and that it is problematic to relate the sweeping-hair arrangement of species to the only three modes of pollen presentation described for the family, indicating that the hypothesised relationship could be more complex than was formerly thought. For all species with di- or trimorphic florets, the styles of female florets were more slender and without or with more reduced sweeping hairs than the styles of hermaphrodite florets, and divergences of sweeping hair arrangements and morphology were higher among phylogenetically related species. These results suggest that functional aspects of floral morphology seem to be more important than phylogenetic constraints as selective forces determining stylar pollen presentation structures. In contrast, stigmatic-area organisation as well as the morphology of stigmatic papillae remain identical between female and hermaphrodite florets and among phylogenetically related species. Thus, stigmatic papilla morphology seems to be a phylogenetically constrained character in the studied species.

[The flower and blossom morphology of Asteraceae correlates with composition of their pollinators]. / Sviaz' mezhdu stroeniem tsvetkov i sotsvetii slozhnotsvetnykh (Asteraceae) i sostavom ikh opylitelei.

The correlation between flower morphology and share of different insect groups visiting them was studied for 15 Asteraceae species. We measured length and width of corolla tube of 100 flowers of each plant species and determined proportions of main groups of anthophilous insects during all blooming period. According to corolla length species under study ranged more or less uniformly from 2.16 mm (Tripleurospermum inodorum) up to 21.06 mm (Cirsium heterophyllum). The correlation between share of long-tongued bees (mainly bumblebees) among all visitors of inflorescens and corolla length was positive (r = 0.737, P < 0.01) while for short-tongued flies (Syrphidae, Muscidae, Calliphoridae) it was negative (r = -0.869, P < 0.01). It is interesting, that the point of crossing of regression lines (12 mm) approximately coincides with change in inflorescences coloration. Plants with corolla length less than 10 mm have yellow or white inflorescences that are visited primarily by flies, while the plants with longer corolla have violet or dark blue inflorescences, by bumblebees. The dependence of proportion of short-tongued solitary bees (Andrenidae, Halictidae) on a corolla length was non-linear. It increased with increase in corolla length in an interval of 2.16-6.26 mm (r = 0.930, P < 0.1), but decreased for longer corollas (r = -0.680, P < 0.05). The correlation between corolla length and proportions of beetles and butterflies were insignificant.

Molecular phylogeny of Subtribe Artemisiinae (Asteraceae), including Artemisia and its allied and segregate genera.

BACKGROUND: Subtribe Artemisiinae of Tribe Anthemideae (Asteraceae) is composed of 18 largely Asian genera that include the sagebrushes and mugworts. The subtribe includes the large cosmopolitan, wind-pollinated genus Artemisia, as well as several smaller genera and Seriphidium, that altogether comprise the Artemisia-group. Circumscription and taxonomic boundaries of Artemisia and the placements of these small segregate genera is currently unresolved. RESULTS: We constructed a molecular phylogeny for the subtribe using the internal transcribed spacers (ITS) of nuclear ribosomal DNA analyzed with parsimony, likelihood, and Bayesian criteria. The resulting tree is comprised of three major clades that correspond to the radiate genera (e.g., Arctanthemum and Dendranthema), and two clades of Artemisia species. All three clades have allied and segregate genera embedded within each. CONCLUSIONS: The data support a broad concept of Artemisia s.l. that includes Neopallasia, Crossostephium, Filifolium, Seriphidium, and Sphaeromeria. However, the phylogeny excludes Elachanthemum, Kaschgaria, and Stilnolepis from the Artemisia-group. Additionally, the monophyly of the four subgenera of Artemisia is also not supported, with the exception of subg. Dracunculus. Homogamous, discoid capitula appear to have arisen in parallel four to seven times, with the loss of ray florets. Thus capitular morphology is not a reliable taxonomic character, which traditionally has been one of the defining characters.

Down the tube: pollinators, predators, and the evolution of flower shape in the alpine skypilot, Polemonium viscosum.

We address how a conflict between pollinator attraction and avoidance of flower predation influences the evolution of flower shape in Polemonium viscosum. Flower shape in P. viscosum is the product of an isometric relationship between genetically correlated (rA = 0.70) corolla flare and length. Bumblebee pollinators preferentially visit flowers that are more flared and have longer tubes, selecting for a funnel-shaped corolla. However, flower shape also influences nectar-foraging ants that sever the style at its point of attachment to the ovary. Surveys of ant damage show that plants having flowers with flared, short corollas are most vulnerable to ant predation. Consistent with this result, the ratio of corolla length to flare is significantly greater in a krummholz (high predation risk) population than in a tundra (low predation risk) population. To explicitly test whether the evolution of a better defended flower would exact a cost in pollination, we created tubular flowers by constricting the corolla during development. Performance of tubular flowers and natural controls was compared for defensive and attractive functions. In choice trials, ants entered control flowers significantly more often than tubular ones, confirming that the evolution of tubular flowers would reduce the risk of predation. However, in a bumblebee-pollinated population, tubular flowers received significantly less pollen and set fewer seeds than controls. A fitness model incorporating these data predicts that in the absence of the genetic correlation between corolla length and flare, intermittent selection for defense could allow tubular flowers to spread in the krummholz population. However, in the tundra, where bumblebees account for nearly all pollination, the model predicts that tubular flowers should always confer a fitness disadvantage.