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Identifying three species of Datura

Identifying three species of Datura



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I have found a large, wild patch, some 200m long, of mainly Datura Stramonium, in our street. I have always been keenly interested and well read on the shamanic, and - very rare - medicinal uses of this species, but scattered amongst them, like family, are two other plants I don't know, but am convinced are also some kind of Datura. I would please like some help on identifying the other two based on the two photos I have of all three.

This is the well known - to me - Stramonium flower, the plant characterized by jagged leaves and the well known spiked seed pod of Jimson Weed seeds.

It looks like user29734's answer is correct, and the above is Morning Glory, but of course subject to confirmation. I now do recognize at least the purple flower as Morning Glory, but from some 30 odd years ago. And it now looks like what I confused with Datura Alba is really just another colour flower of the Morning Glory.

If anyone could please confirm my ID of the Stramonium, and confirm the identification of the second image as morning glory, and suggest what species.

These plants are all around my neighbourhood, with is the northern part of Randburg, South Africa. This city borders on the Northern side of Johannesburg. 1 It's a temperate climate, with very hot and rainy summers, and bitterly cold and dry winters. The soil is reddish and lacks much humus.

It is a semi-rural area, having been rapidly urbanised over the last ten or so years, when it was still just farms and small holdings. Farming is mainly vegetables or lucerne; and lots of maize further out from the city. We are a few km south of the Cradle of Humankind.

1 Strangely enough, there are two cities in the US with the same names, both in California, I believe. I know Johannesburg is.


Okay, so as far as the blue and white flower goes I'm pretty sure it's a cultivar of ipomoea called Flying Saucer which is a hybrid of heavenly blue (I. tricolor) and Pearly gates (also I. tricolor) meaning it's a hybrid and no seed output. As far as the white flowers go it is most likely Datura since it is part of a short shrub and the petals are spiked. However you should determine if it blooms at night because its possible it could be moonflower. It may also be brugsmansia or pearly gates. As far as 'alchemical use' datura contains scopolamine which is potentially lethal if consumed in indiscriminate amounts.


The second flower is, I'm guessing by the valentine shaped leaf and violet tubular flowers, a morning glory vine (ipomoea violacea or ipomea tricolor). These plants both are closely related cultivars of ololioqui and tlitlitzen which had a history of shamanic use in southern Mexico.


The top plant looks like it could easily be D. stramonium. Daturas can be a bit tricky. You should either provide an estimate of the scale or measure the flower, then compare with this simple key of invasive Daturas (in Australia): https://www.samorini.it/doc1/alt_aut/ek/haegi.pdf. It helps to note capsule size and shape. (It is NOT a Brugmansia sp.; those are trees with pendant flowers.) Also, the second flower photo seems to me is clearly a morning glory, possibly as IDed by user29734, and the center leaf seems from the same plant(s) or closely related.


The origin of Datura metel (Solanaceae): genetic and phylogenetic evidence

Using the analysis of nine isozyme systems and cladistic analysis of 32 morphological characters of the Mexican species of the section Dutra of the genus Datura, evidence was sought on the origin of the cultivated D. metel L. The genetic similarity and the phylogenetic relationship suggest that D. metel is related more closely to D. inoxia Mill. than to the other taxa of the section Dutra based upon the small genetic distance between them. The cladistic analysis revealed two main clades: the long-lived, tuberous rooted perennials (D. inoxia, D. lanosa A.S. Barclay ex Bye, D. metel, and D. wrightii Regel) and the tap-rooted annuals (D. discolor Bernh., D. kymatocarpa A.S. Barclay, D. leichhardtii F.V. Muell. ex Benth. ssp. pruinosa (Greenm.) Barclay ex Hammer (syn.: D. pruinosa Greenm.), D. reburra A.S. Barclay). Datura inoxia is the sister taxon of D. metel next to which is D. wrightii while D. lanosa is the basal taxon of this group. The combination of genetic and cladistic data indicates that D. inoxia is most likely the progenitor of D. metel.

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Identifying three species of Datura - Biology

© 2021 Institute for Systematic Botany | Data last modified: 6/24/2021

Wunderlin, R. P., B. F. Hansen, A. R. Franck, and F. B. Essig. 2021. Atlas of Florida Plants (http://florida.plantatlas.usf.edu/). [S. M. Landry and K. N. Campbell (application development), USF Water Institute.] Institute for Systematic Botany, University of South Florida, Tampa.

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Solanaceae: Characters, Distribution and Types

Plants herbs, shurbs rarely trees leaves alternate, flowers solitary or in cymes axillary or terminal flowers pentamerous, actinomorphic, hypogynous, hermaphrodite, calyx persistent, gamosepalous, corolla gamopetalous, campanulate stamens epipetalous gynoecium bicarpellary, syncarpous, ovary obliquely placed, axile placentation swollen placentae ovules many in each locules fruit capsule or berry.

A. Vegetative characters:

Mostly herbs (Petunia, Withania), shrubs and trees.

A branched tap root system.

Aerial, erect, climbing (Solanum jasminoides), herbaceous, or woody, cylindrical, branched, solid or hollow, hairy, or glabrous, underground stem in Solanum tuberosum.

Cauline, ramal, exstipulate, petiolate or sessile, alternate sometimes opposite, simple, entire pinnatisect in Lycopersicurn, unicostate reticulate venation.

Solitary axillary, umbellate cyme, or helicoid cyme in Solanum.

Bracteate or ebracteate, pedicellate, complete, hermaphrodite, actinomorphic, pentamerous, hypogynous.

Sepals 5, gamosepalous, tubular or campanulate, valvate or imbricate, persistent, green or coloured, hairy, inferior.

Petals 5, gamopetalous, tubular or infundibuliform, valvate or imbricate aestivation, scale or hair-like outgrowth may arise from the throat of the corolla tube, coloured, inferior.

Stamens 5, epipetalous, polyandrous, alternipetalous, filaments inserted deep in the corolla tube, anthers dithecous, usually basifixed or dorsifixed, introrse, inferior.

Bicarpellary, syncarpous, ovary superior, bilocular, unilocular in Henoonia, axile placentation placentae swollen, many ovules in each loculus, ovary obliquely placed in some cases nectariferous disc is present style simple stigma bifid or capitate.

Distribution of Solanaceae:

The family is commonly called ‘Potato family’. It is a large family well distributed in tropics and sub-tropics, though a few members are found in temperate zone. The family includes 2,000 species belonging to 90 genera. In India it is represented by 70 species of 21 genera.

Several members are cultivated through out the world for their great economic importance among them are Solarium tuberosum (Potato), Solarium melongena (Bringal), Lycopersicurn esculentum (Tomato) etc.

Economic Importance of Solanaceae:

The family is of great economic importance.

Many members viz., Solanum tuberosum (Potato), Solanum melongena (Brinjal), Lycopersicurn esculentum (Tomato), Capsicum (H. Mirch) etc. are used as vegetables. Physalis peruviana (H. Rasbhari) produces edible berries.

Atropa belladona contains alkaloid Atropine this is used in Belladona plaster. Atropine is used in eye testing. Nicotiana tabacum (tobacco) yields Nicotine. Hyoscyamus niger, Solanum nigrum, Datura (H. Dhatura), Withania somnifera (Ashwagandha) are used medicinally.

Tobacco is obtained from leaves of Nicotiana tabacum and variously used in cigars, bidi, chewing, jarda etc.

Petunia, Cestrum, Lycium, Salpiglossis, Schizanthus are cultivated in gardens for ornamentals.

Primitive characters:

1. Shrubs, trees and perennial climbers.

2. Leaves simple and alternate.

3. Inflorescence solitary axillary or terminal.

4. Flowers actinomorphic, hermaphrodite and hypogynous.

6. Ovules numerous in each loculus.

Advanced characters:

1. Most of the plants are herbaceous and many are annuals.

2. Leaves exstipulate, in some finely divided.

3. Calyx and corolla are gamosepalous and gamopetalous.

5. Reduction in the number of carpels to two.

6. Gynoecium is syncarpous.

Affinities of Solanaceae:

Hallier regarded Solanaceae as a primitive member of the tubiflorae together with the Scrophulariaceae and both have arisen very likely from the Linaceae. Wettstein placed the family in the Tubiflorae along with Convolvulaceae. Rendle placed the family in the Tubiflorae, assigning a separate position for the Convolvulaceae under the order Convolvulales.

Solanaceae bears a close relationship to the Boraginaceae in alternate leaves, regular flowers and five stamens. It is related to Convolvulaceae in the presence of persistent calyx, twisted corolla and false septum. It is allied to Scrophulariceae but the latter differs from it in actinomorphic flowers and obliquely placed carpels.

Common plants of the family:

An ornamental genus characterised by the presence of 4 didynamous stamens.

2. Cestrum nocturnum (H. Rat Ki Rani or Queen of the night):

A garden plant with flowers emitting sweet smell at night.

3. Datura metal (H. Dhatura or Thorn apple):

A herb with highly poisonous fruits and seeds.

5. Lycoperscium esculentum:

A tall herb with red globose pulpy fruits.

Produces edible reddish berries that are enclosed in a bladdery and persistent calyx.

7. Solanum nigrum (H. Mako):

A roadside herb S. xanthocarpum a spinous xerophytic herb.

8. Withania somnifera (H. Asgandh, Sanskrit – Ashwangandha):

Division of the family and chief genera:

Von Wettstein divided the family in to five tribes and two groups:

A. Embryo curved like a semicircle,.stamens 5, filaments equal.

Gynoecium bicarpellary with 3-5 locules e.g. Nicandra.

Ovary bilocular e.g. Solanum, Capsicum.

Ovary 4-celled, septum dividing placenta equally e.g. Datura.

B. All stamens or 2-4 stamens fertile embryo slightly or almost straight.

All 5 fertile stamens seeds not flat e.g. Cestrum.

Tribe 5. Salpiglossideae:

Stamens 2-4 fertile flowers zygomorphic e.g. Browallia, Salpiglossis.

Important Types of Solanaceae:

1. Solanum nigrum (Fig. 78.1):

Erect, aerial, woody below and herbaceous above, cylindrical with distinct ribs, solid, branched, green.

Alternate, simple, exstipulate, petiolate, ovate, repand, acute, glabrous, unicostate reticulate venation.

Extra-axillary helicoid cymes. Extra axillary position is due to fusion.

Ebracteate pedicellate, complete, hermaphrodite, actinomorphic, pentamerous, hypogynous, small and white.

Sepals 5, gamosepalous, pentafid, valvate, persistent, green, hairy, inferior.

Petals 5, gamopetalous, rotate, valvate, five lobed, white, inferior.

Stamens 5, polyandrous epipetalous, alternipetalous, filaments shorts, equal in length, anthers long and conniving, basifixed, dithecous, and dehiscence by apical pores.

Bicarpellary, syncarpous, ovary superior, bilocular, axile placentation, placentae swollen, ovules many in each loculus, ovary obliquely placed style simple, hairy stigma bilobed.

2. Withania somnifera (Fig. 78.2):

Erect, aerial, cylindrical, solid, woody below and herbaceous above, branched, covered with hair.

Cauline, ramal, opposite, simple, exstipulate, petiolate ovate, entire, acute, covered with glandular branched hair, unicostate reticulate venation.

Ebracteate, sub-sessile, complete, hermaphrodite, actinomorphic, pentamerous, hypogynous.

Sepals 5, gamosepalous, campanulate, pentafid, valvate, hairy, green, inferior.

Petals 5, gamopetalous, pentafid, campanulate, valvate aestivation, corolla lobes inflexed, greenish yellow.

Stamens 5, polyandrous, epipetalous, alternipetalous inserted near the base of corolla tube, filaments equal in length basifixed, dithecous, introrse.

Bicarpellary, syncarpous, ovary superior, bilocular, ovules many, axile placentation, placentae swollen, ovary obliquely placed hairy, style simple stigma bifid.

A berry enveloped by persistent calyx.

Herbaceous, erect, cylindrical, branched, slightly woody and slightly fistular, hairy.

Alternate, opposite in floral region, simple, exstipulate, petiolate, ovate-lanceolate, acuminate or acute apex, unequal at base, sinuate-toothed or repand margin, glabrous on both sides, unicostate reticulate venation.

Shortly pedicellate, bracteate, hermaphrodite, actinomorphic, complete, hypogynous, pentamerous.

Sepals 5, gamosepalous, tubular, angulate, 5-toothed, teeth triangular lanceolate, acuminate, green, persistent, twisted aestivation, inferior.

Petals 5, gamosepalous, campanulate, 5 or 6 cuspidate acute angles, violet outside, twisted aestivation, inferior.

Stamens 5, epipetalous, filaments long, anthers basifixed, introrse, dithecous.

Bicarpellary, syncarpous, ovary superior, obliquely placed, four celled due to the formation of false septum, axile placentation, many ovules in each loculus, style long, stigma bilobed.

4. Petunia alba (Fig. 78.3):

Herbaceous, erect, cylindrical, solid, branched, hairy, green.

Cauline, simple opposite decussate, exstipulate, ovate, unicostate reticulate venation.

Axillary or terminal condensed cymes.

Pedicellate, bracteate, hermaphrodite, complete, actinomorphic, pentamerous, hypogynous.

Sepals 5, gamosepalous, campanulate, sepals free above and fused below, green, hairy, imbricate aestivation, inferior.

Petals 5, gamopetalous, campanulate, tube hairy, twisted aestivation, inferior.

Stamens 5, polyandrous, epipetalous, alternating with the petals, filaments long, anthers basifixed, dithecous, introrse.

Bicarpellary, syncarpous, ovary superior, obliquely placed, bilocular swollen axile placentation ovules many style long, slightly, twisted, stigma capitate, bilobed.


Health Effects of Alkaloids from African Medicinal Plants

21.5.1 Atropine

Atropine (12) is a naturally occurring tropane alkaloid extracted in several plants of the Solanaceae including Atropa belladonna, Datura inoxia, Datura metel, Datura stramonium, Brugmansia spp. and Hyoscyamus spp. Plants such as Datura metel L., Datura stramonium naturally occur in tropical West Africa [81] . In tropical West Africa, Datura spp. are used in palm wine to add a narcotic and stupefying effects [81] . Datura stramonium also occurs in South Africa where it used frequently as an antiasthmatic treatment [82] . Compound 12 is an anticholinergic drug acting as a competitive antagonist for the muscarinic acetylcholine receptor types M1, M2, M3, M4, and M5 [83] . Topical application of 12 is used as a cycloplegic, to temporarily paralyze the accommodation reflex, and as a mydriatic, to dilate the pupils. Due to its slow degradation, compound 12 is generally used as a therapeutic mydriatic.


Phylogenetics of Datureae (Solanaceae), including description of the new genus Trompettia and re–circumscription of the tribe

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Abstract

Datureae is a tribe in the Solanaceae known for its charismatic large–flowered species (jimsonweeds and angel trumpets). The monophyly of the tribe is well established, but the recent finding that a species previously described in Iochroma (also Solanaceae) belongs in Datureae calls for a reassessment of the tribe's circumscription. Here we estimated the phylogeny of Datureae, including all of its 18 species, using three nuclear regions, and incorporated fossil information to estimate divergence times. Based on this phylogeny, we reconstructed the evolution of key aspects of reproductive morphology and life history to identify diagnostic features. Our molecular phylogenetic analyses suggest that the diversification of Datureae began roughly ca. 35 Ma, around the beginning of the Andean uplift. Within the tribe, Datura and Brugmansia are monophyletic sister taxa and the misplaced species of Iochroma is sister to the remaining species. Based on our morphological analysis, we describe the latter as a new monotypic genus Trompettia. Ancestral state reconstructions identify diagnostic features for each of the three genera and show a large suite of changes along the Datura branch, including the evolution of erect flowers, capsular fruit, and annual life history. Using these features, we formally re–circumscribe Datureae to include all genera and their species and provide a taxonomic key for the tribe.


Myths

“Smoking datura destroys the dangerous tropane alkaloids”

This commonly held assumption is highly suspect. [63] [64] After all, the effectiveness of smoking remedies like Asthmador cigarettes relied on these alkaloids surviving heat. [56] The plant has also long been smoked or burned in traditional ritual contexts for its deliriant and hallucinogenic effects. [2] . It’s safe to assume, based on the historical context of smoking Datura, that burning it does not destroy the tropane alkaloids.

Can it be detected in a drug test?

Only specialist laboratory tests like gas chromatography-mass spectrometry (GC-MS) can reliably detect the tropane alkaloids, but they’re not routinely used for drug screening.

Can Datura cause psychological trauma?

While under the influence of datura, hallucinations and delusions are widely perceived to be real. It may take a week or more to fully recover from the experience. Datura is also thought to carry a greater long-term risk of psychosis than other hallucinogens, but the research is limited. [3]

Are there risks?

There are many health risks associated with datura, including the very real threat of death. The variability of alkaloid levels makes it extremely easy to overdose. People with existing heart conditions should be especially cautious. Pregnant women should also avoid datura, due to the potential impact of excessive acetylcholine levels on the fetus.

Some other risks include physical injury and legal issues. See “Risks” for more information.

Is it legal to grow at home?

It’s usually legal to grow Datura at home. Just be aware of the high risk of poisoning to pets and small children. Sometimes just handling the plant is enough to cause toxic effects. [18]

What are the differences between datura species?

Potency and appearance can vary between the species, but the effects are largely the same. Datura metel has the highest scopolamine content, and usually the highest percentage of alkaloids overall, but the difference isn’t hugely significant.

Unlike most species, the seeds and flowers of Datura wrightii and the flowers of Datura discolor are non-psychoactive. [2]

What is the safest way to take datura?

There are several ways to ingest Datura plants, including Datura Stramonium. The plants can be smoked, brewed into a tea, or converted into a skin ointment. Since all parts of the plant contain variable amounts of psychedelic compounds, some people have been known to chew the seeds as well.

Smoking Datura may be somewhat less dangerous than oral ingestion, but it’s by no means “safe.” All methods have been linked to bad trips, hospitalization, and death. However you choose to take it, the dried plant material should be finely ground to ensure an even distribution of the alkaloids. If an initially small dose is ineffective, a slightly higher dose can be taken a couple of weeks later and so on until the desired effects are reached. Because of the high variability between and often within plants, this process should be repeated for each new batch even if it’s from the same plant. [3]

Despite any preparation you take, Datura is still a dangerous substance. There is no way to tell how much of the psychoactive alkaloids are in different parts of the plant, making the effects of ingestion highly unpredictable. Use extreme caution before ingesting Datura in any way.

Can I microdose datura?

Anecdotal reports suggest that microdosing Datura helps to induce sleep and lucid dreaming.[65][66][67] However, regular dosing—even at a fraction of the hallucinogenic threshold—could lead to a gradual build-up of dangerous toxicity over time.

Does it produce tolerance?

Yes. It takes roughly two weeks to return to baseline sensitivity, and there’s a cross-tolerance effect with other deliriants, including diphenhydramine (Benadryl) and myristicin (from nutmeg). [3]

Can I mix it with other drugs?

Datura is sometimes combined with other drugs, including cannabis, but for safety reasons this isn’t recommended. MAOIs, stimulants, sedatives, antidepressants, antihistamines, and other medications are all contraindicated. Aspirins can be especially problematic. [4]

It’s a good idea to abstain from all other substances for at least a couple of weeks before and after taking Datura.


Materials and Methods

Study species and sampled populations

Datura inoxia (Solanaceae) is a summer annual, self-compatible plant that inhabits arid and semi-arid lands in Mexico and Southern USA (i.e., The Chihuahuan desert) (Barclay, 1959 Lockwood, 1973). High daily fluctuation of ambient temperature is a characteristic of these ecosystems. The onset of flowering of D. inoxia is in July and lasts through September, similar to other Datura species (Bronstein et al., 2009). Datura inoxia produces large, funnel-shaped, and nectar producing hermaphroditic flowers that open at dusk and remain receptive for only one night. During the flowering period, individuals can display from a few up to tens of flowers each night. Flower traits related to mating system such as herkogamy and flower size, vary widely within and among populations (Jiménez-Lobato & Núñez-Farfán, 2012). Further, comparisons between genetic differentiation at neutral loci (FST) and phenotypic differentiation (QST) in floral traits suggest adaptive evolution under diverging selection (Jiménez-Lobato & Núñez-Farfán, 2012). Flowers are commonly visited by honeybees, which collect pollen, and hawkmoths (Manduca sexta, M. quinquemaculata and Hyles lineata) which forage for nectar (Barclay, 1959 Lockwood, 1973 Grant, 1983 McCall et al., 2018 V. Jiménez-Lobato, personal observation).

Two populations of D. inoxia were selected to assess the relationships between herkogamy, selfing rate, and inbreeding. A previous report (Jiménez-Lobato & Núñez-Farfán, 2012) indicated substantial within-population variation in herkogamy and flower size. The Cañada de Moreno population (CM) is located in the State of Querétaro (21° 17′ 43″ N 100° 31′ 00″ W) in the Mexican Bajío at 1,933 m a.s.l. During the flowering period of D. inoxia (July–September), this locality has an average temperature of 18.8 °C, with a daily range from 7.4 °C to 30.8 °C, and a three-month total precipitation of 314 mm. The Mapimí population (Map) is located at 1,157 m a.s.l. in the Mexican Plateau in the States of Coahuila and Durango (26° 41′ 11″ N 103° 44′ 49″ W). This is a more xeric environment, with a quarterly average temperature of 22.8 °C (range: 12.6–32.9 °C) and a total three-month precipitation of 253 mm (Meteorological Service of Mexico: http://smn.cna.gob.mx/). Collection of seed material for experimental analyses was made under the permission SGPA-DGGFS-712-1596-17 (Subsecretaría de Gestión para la Protección Ambiental, Secretaría de Medio ambiente y Recursos Naturales, Mexico).

Variation in the level of herkogamy within and among populations

Thirty reproductive plants were randomly selected and tagged for sampling in a 1 ha area within each population. For each individual plant, 4–6 open flowers were randomly selected to measure herkogamy. Herkogamy was calculated as the difference between pistil and stamen length. Approach herkogamy was defined as the pistil surpassing the stamens in length (henceforth “positive herkogamy”), whereas reverse herkogamy was the opposite trend (henceforth “negative herkogamy”). Absence of herkogamy occurred when pistil and stamens had equal lengths.

Mating system parameters

To estimate mating system parameters from each marked plant in the field, five mature fruits derived from natural pollination were collected, labelled and bagged. In the laboratory, seeds of each fruit were separated and germinated in a greenhouse, and seeds of each fruit within each family (maternal plant) were sown in separate pots. Germination per fruit per family was recorded for 30 days. To obtain an average estimate of germination rate per fruit, we recorded the final number of seeds germinated in each pot. Germination percentage was ≥90% (se = 3.18) for all plants. Once seedlings emerged, we collected leaf tissue from young plants, bagged, labelled, and stored in an ultra-freezer at −97 °C. Finally, we analyzed mating system parameters in 20 seedlings from each of 30 maternal families per population (N = 600).

DNA was extracted from seedlings following the Miniprep protocol (Doyle & Doyle, 1987). Five microsatellite nuclear loci developed for D. stramonium (Andraca, 2009) were amplified for each seedling. Further, we standardized one additional microsatellite locus for D. inoxia (F8: Rw: 5′ -GGACAACATCTTTGCGACCC- 3′), in order to obtain a total of six polymorphic microsatellite loci per individual plant. Primers were labelled with PET, VIC, 6-FAM, and NED dyes (Applied Biosystems, Foster City, CA, USA) (see Supplemental Information: PCR protocols).

Multilocus outcrossing (tm) and selfing (s = 1 − tm) rates, primary selfing rate (r) and inbreeding coefficient (F) were estimated for each maternal family for each population. Mating system parameters (tm and s) at the family level were estimated with MLTR 3.2 (Ritland, 2002) using the Expectation-Maximization method, which allows missing data and undetected null alleles (Ritland & Jain, 1981). Standard errors and standard deviations were estimated by bootstrapping, with 1,000 replicates and re-sampling individuals at the family level. The frequency of null alleles per locus, per population, was assessed using Micro-Checker v.2.2.3 (Cock et al., 2004). One locus (G8) did not amplify for any of the plants from CM, so analyses were carried out with five loci in that population, and six loci in Map. Selfing rates (s) obtained from molecular markers, after fertilization and germination, might not be completely independent from inbreeding depression and thus may underestimate its true value (Lande, Schemske & Schultz, 1994). Primary selfing rate (r) is a better predictor of mating system because it assesses separately the magnitude of inbreeding depression. Hence, r refers to the proportion of selfed progeny at the time of fertilization (Lande, Schemske & Schultz, 1994). The primary selfing rate (r) was calculated for each maternal family as: r = s/[1 − δ + sδ], where s is the selfing rate obtained from microsatellite loci, and δ is the cumulative inbreeding depression obtained for each population (see below).

The inbreeding coefficient (F) may include components of inbreeding other than just self- or cross- fertilization, such as biparental inbreeding or population substructure. Here, we used F as a proxy of the inbreeding history of each maternal family (i.e., adult cohort) (Latta & Ritland, 1994). Inbreeding coefficient (F) values were inferred for each lineage from the microsatellite loci amplified previously, with GenePop v.4.2 (Rousset & Raymond, 1995 Rousset, 2008). To investigate whether inbreeding depression was associated with the selfing rate and herkogamy, we calculated inbreeding coefficients at equilibrium (Fe) (i.e., progeny cohort) for each lineage, assuming that adult F and tm are constant among generations (Ritland, 1990). We then related these differences with selfing rate and herkogamy. Since Fe increases in relation to F due to self-fertilization, larger differences between F and Fe at each maternal lineage should indicate the presence of higher inbreeding depression. Once selection against inbred progeny has occurred, Fe and F will be equal (Ritland, 1990 Shirk & Hamrick, 2014). This approach yields an estimation of potentially late-acting inbreeding depression in each lineage. Fe was calculated as Fe = (1 − tm)/(1 + tm), where tm is the outcrossing rate calculated from MLTR for each maternal family (Allard, Jain & Workman, 1968 Ritland, 1990).

Inbreeding depression

To estimate inbreeding depression (δ), we collected 150 fruits from different individual plants in each population. These plants did not include the maternal families previously analyzed. From each fruit, we sowed ten seeds on separate pots under greenhouse conditions. Average (±S.E.) environmental variables in the greenhouse (March–September), measured by HOBO sensors (Onset Computer Corporation, Bourne, MA, USA) were: temperature 27.10 ± 0.06 °C relative humidity 42.93% ± 0.09% and light intensity 3,133.59 ± 18.73 Lux (Camargo, 2009). When seeds germinated, one seedling per fruit was randomly chosen and grown under controlled conditions until flowering. For each population, 100 individuals were randomly chosen to act as pollen receptors (mothers) and 50 individuals as pollen donors (fathers). Two manual pollination treatments were applied to each maternal plant: (1) cross-pollination (o), where two flowers were emasculated before anthesis and hand-pollinated with pollen from one donor randomly chosen from the same population (2) self-pollination treatment (s), where two flowers of each receptor plant were fertilized with self-pollen. After pollination, flowers in both treatments were bagged individually with a fine nylon mesh. Since many mother plants did not produce the four flowers needed for pollination treatment application, the final sample included mother plants that produced at least one fruit per treatment (CM: N = 77 Map: N = 41). Two fitness components per pollination treatment were evaluated in each population: seed-set mean (i.e., number of seeds/ number of ovules) and seed mass. Seed mass was obtained from a random sample of 30 seeds per fruit using an analytical balance (Adventurer OHAUS). Allocating more resources to seeds can increase quality of seeds, increasing the likelihood of successful seedlings’ establishment and mothers’ fitness (Stanton & Young, 1994 Byers & Waller, 1999). In D. inoxia, as in D. stramonium, under greenhouse experiments conditions, inbreeding depression on seed mass has been detected even when maternal plants have been fertilized with a single pollen donor (Sosenski, 2004 Jiménez-Lobato, 2013). Likely seed mass may influence germination rate and seedling establishment in Datura species.

Cumulative inbreeding depression coefficient (δ) was calculated for each population as: δ = 1 − w ´ s w ´ o , where w ´ s and w ´ o are the mean fitness of progenies derived from self- or cross-pollination, respectively. Average fitness of self- and out-cross progenies was calculated as the product of seed-set and seed mass (Schemske & Lande, 1985), and it was used to estimate the primary selfing rate (r) at each maternal lineage. This approach yields an estimate of early-acting inbreeding depression

Statistical analyses

Statistical analyses were implemented in R software version 4.0.2 (R Development Core Team, 2020). To estimate phenotypic variation in herkogamy within each population, we quantified the variance components within and between individuals. Using the nlme package (Pinheiro et al., 2017), we fitted a linear mixed model for each population where maternal family and plants nested within families were considered as random factors.

Statistical differences in primary selfing rates (r), inbreeding in adult cohort (F), inbreeding at equilibrium (Fe) and an estimate of inbreeding depression (FeF) between populations were tested by Analysis of Covariance in multcomp package (Hothorn, Bretz & Westfall, 2008). Population was considered as fixed factor and herkogamy as the quantitative variable.

Multilocus outcrossing (tm) and primary selfing rates (r) were strongly negatively correlated (CM: estimate = −1.00, p = 0.000, d.f. = 25 Map: estimate = −0.969, p = 0.000, d.f. = 27), hence we present here only the analyses for primary selfing rates (r). Since r is a proportion, its relationship with herkogamy was analyzed with a beta regression using the betareg package (Cribari-Neto & Zeileis, 2010 Douma & Weedon, 2019). This regression has been proposed for modeling continuous data limited to a specific interval (0, 1) (Ferrari & Cribari-Neto, 2004). We tested for cloglog, logit and log link functions and based on Akaike’s criterion we selected the model that best fit to the data. Estimates of β and ∅ were obtained by maximum likelihood estimation. F, Fe and their differences were associated to herkogamy with a generalized linear model with Gaussian error distribution (Crawly, 2013). It must be noted that Fe is a theoretical prediction based on the selfing rate (r), and these quantities are therefore correlated by construction. Likewise, their correlation with other variables would be very similar. Thus, to avoid redundancy, we present the correlations of selfing rate with other variables.


Phylogenetics of Datureae (Solanaceae), including description of the new genus Trompettia and re–circumscription of the tribe

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1800 Colorado Ave, 80309 Colorado, Boulder, U.S.A.

Abstract

Datureae is a tribe in the Solanaceae known for its charismatic large–flowered species (jimsonweeds and angel trumpets). The monophyly of the tribe is well established, but the recent finding that a species previously described in Iochroma (also Solanaceae) belongs in Datureae calls for a reassessment of the tribe's circumscription. Here we estimated the phylogeny of Datureae, including all of its 18 species, using three nuclear regions, and incorporated fossil information to estimate divergence times. Based on this phylogeny, we reconstructed the evolution of key aspects of reproductive morphology and life history to identify diagnostic features. Our molecular phylogenetic analyses suggest that the diversification of Datureae began roughly ca. 35 Ma, around the beginning of the Andean uplift. Within the tribe, Datura and Brugmansia are monophyletic sister taxa and the misplaced species of Iochroma is sister to the remaining species. Based on our morphological analysis, we describe the latter as a new monotypic genus Trompettia. Ancestral state reconstructions identify diagnostic features for each of the three genera and show a large suite of changes along the Datura branch, including the evolution of erect flowers, capsular fruit, and annual life history. Using these features, we formally re–circumscribe Datureae to include all genera and their species and provide a taxonomic key for the tribe.


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