An expanded description of Lophophora alberto-vojtechii

Šnicer, Bohata and Myšák recently expanded on their original description of the miniature peyote, Lophophora alberto-vojtechii, in the article The Littlest Lophophora – an article that also proposes a new key to the classification of Lophophora.

The hallmark of the new peyote species is its minuscule size, rarely exceeding 2.5 cm (~ 1'') in diameter. The Lilliputian stature is eminently illustrated by the following picture of Vojtěch Myšák (one of the persons for whom the plant is named) photographing a specimen in northern Zacatecas.


Lophophora alberto-vojtechii being photographed by Vojtěch Myšák

The expanded description of Lophophora alberto-vojtechii reads:

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The stem is grayish green with a touch of mauve and is for the most part concealed below ground. The aboveground part (the crown) is flat to slightly convex and 10–50 mm wide in adult plants (average 18 mm, well under an inch). The subterranean part of the stem is cylindrical, extending 25 mm down from the base of the crown (the depth in adult plants increasing with age). The stem is distinctively small and divided into five ribs, or eight in exceptional cases. Trichomes (fine epidermal outgrowths which might be called fuzz or hair) are visible only near the growing point and drop off early, leaving the areoles inconspicuous.

The root is thick and bulbous, up to 30 mm long, and typically 16 mm in diameter where it joins the subterranean stem. It is smooth and a dirty white to creamy yellow color.

The flower is 15–35 mm (average 23 mm) wide. The style is longer than anthers and overtops them, and the petals are long relative to their width, both traits in accord with other members of the Diffusae. Filaments are white, anthers a yellowish orange. The style is white and topped by a white to yellowish or pinkish stigma. Tepals are white to dirty pink, rarely creamy yellowish, and bear a lengthwise stripe in a brownish, salmon, or darker pink shade.

The fruit, dirty white, yellowish, or pinkish, is round to slightly claviform and dries out over time.

Seeds are black, round, 1.15–1.45 mm long × 1.0–1.45 mm wide with the hilum compressed into a broad V–shape. The perimeter of the hilum consists of a pronounced edge. The testa is nodulated, and individual cells of the testa are clearly demarcated.

Type locality: northern San Luis Potosí, Mexico, 1700 meters above sea level. The type specimen was collected by GS Hinton on 1 August 2007 and is kept at the GB Hinton herbarium filed under number 28642.

We have named the plant Lophophora alberto-vojtechii in honor of two people: the late prominent Czech traveler and cactus-hunter Alberto Vojtěch Frič and the living lophophora enthusiast Vojtěch Myšák.

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Lophophora alberto-vojtechii flower visited by an unidentified diptera

Three insects (two kinds of bees, one large and one small, and an unidentified diptera) have been observed visiting flowers of Lophophora alberto-vojtechii. The plant being visited by a fly in the above picture is from the second known locality in San Luis Potosí and has exceptionally numerous tepals.

The mini-peyote grows in flat, alluvial sediments that transform into greasy mud flats during rain periods. L. alberto-vojtechii has a distinctively geophytic lifestyle, in dry periods it withers and withdraws into small cavities below ground where it is covered with dust and dry leaves by the wind, effectively protecting the plant from the harsh environment. When rains arrive, the plants absorb water and swell, pushing the photosynthetic crown back up above ground level.


Zacatecan Lophophora alberto-vojtechii plant hidden by its flower

In Zacatecas L. alberto-vojtechii flowers can on rare occasions reach over 3 cm in diameter, totally obscuring the plant below - specimens of L. alberto-vojtechii from the plains of northern Zacatecas are lighter flowered than those at the type locality.


Lophophora alberto-vojtechii growing in mud flat


All pictures are taken from the article The Littlest Lophophora.

Key to the genus Lophophora sensu Šnicer et al.

In the article The Littlest Lophophora Šnicer, Bohata and Myšák detail the discovery of the new species Lophophora alberto-vojtechii and expand on the description originally given in the June 2008 issue of Cactus & Co.

The article also gives a key to the species of Lophophora which are divided into two sections Lophophora and Diffusae. These sections were originally introduced by the same authors in the booklet Genus Lophophora Coulter - Kaktusy special 2, 2005 and the division was based primarily on chemical composition but also on other factors like habitat, incompatibility of the species, rib numbers and morphology. Members of the Lophophora section are characterized by having concentrations of the psychotropic (mind-altering) alkaloid mescaline ranging from 15–30% of the total alkaloid content, while members of the Diffusae section contain a maximum of 1.3% mescaline. Šnicer et al. recognize five species of Lophophora.

SECTION LOPHOPHORA
L. williamsii (Salm-Dyck) Coulter

SECTION DIFFUSAE
L. diffusa (Croizat) Bravo
L. fricii Habermann
L. koehresii (J. Ríha) Bohata, Myšák & Šnicer
L. alberto-vojtechii J. Bohata, V. Myšák & J. Šnicer

The following key to the species of Lophophora assumes that L. alberto-vojtechii is allied with the non-mescaline-prevalent species, i.e. it is placed in the Diffusae section.

KEY TO THE SPECIES OF LOPHOPHORA
1	15–30% mescaline in total alkaloids, epidermis tough and thick	Section Lophophora, L. williamsii
1	Maximum 1.3% mescaline in total alkaloids; thin, fine epidermis	Section Diffusae, 2
2	Seed hilum wide (almost circular), testa not nodulated but reticulated	L. koehresii
2	Seed hilum V–shaped, testa nodulated (outer cell walls protruding), individual testa cells clearly demarcated	3
3	Usually five (rarely eight) ribs, stem solitary in nature, miniature (rarely exceeding 25 mm in diameter)	L. alberto-vojtechii
3	Up to 21 ribs, stem solitary or branching in nature	4
4	Flowers dirty white with touch of yellow (rarely pinkish); fruits white to dark pink, usually pale pink; occurring in Queretaro, Mexico	L. diffusa
4	Flowers usually light pink to dark purple-pink, also white; fruits pink to dark purple-pink (when flower is white the fruit is always dark purple-pink); Coahuila, Mexico	L. fricii

Seed morphology is an important factor in the above key. In their original publication on Lophophora alberto-vojtechii Šnicer et al. included SEM images illustrating the differences between the seeds of the various Lophophora species, these images along with their description are included below (Šnicer et al. attribute the SEM images to Gerhard Köhres; the images below are not scanned from the article but based on the ones included in the online French version of the article - just to make sure that credit is given where credit is due ;-)

Lophophora koehresii seed SEM images
L. koehresii has a seed testa structure that is completely different from the other lophophoras. The hilum is wide open, and is almost circular. The testa is not nodulated (the outer cell walls do not protrude above the surface of the seed), but is reticulated [characterized by or having the form of a grid or network], and the cell walls are flat to even. The individual cells of the testa almost merge together. The seed in the pictures is from San Francisco in San Luís Potosí.

Lophophora alberto-vojtechii seed SEM images
L. alberto-vojtechii has round black seeds, 1.15 to 1.45 mm long and 1.0 to 1.45 mm wide. The hilum is compressed into a broad V shape. The perimeter of the hilum consists of a pronounced edge. The testa is nodulated with the outer cell walls protruding. The individual cells of the testa are clearly demarcated. The seed in the pictures is from the type location in San Luís Potosí.

Lophophora fricii seed SEM images
L. fricii has oval seeds, 1.5 mm long and 1.2 mm wide with the hilum compressed into a V shape. The testa is black, relatively coarsely nodulated, with an oval tubercle structure; the tubercles get smaller near the hilum. The individual cells of the testa are clearly demarcated. The seed in the pictures is from El Amparo in Coahuila.

Lophophora diffusa seed SEM images
L. diffusa has seeds which vary greatly in terms of shape and size. The hilum is compressed into a broad V shape. The testa is coarsely nodulated with the outer cell walls protruding. The individual cells of the testa are clearly demarcated. The seed in the pictures is from Niñas Las Palmas in Querétaro.

Lophophora williamsii seed SEM images
L. williamsii has seeds that are 1-1.5 mm long and black. The hilum is compressed into a broad V shape. The testa is nodulated with the outer cell walls protruding. The individual cells of the testa are clearly demarcated. The seed in the pictures is from Mazapil in Zacatecas.

Acharagma roseana – flowers and fruits

To be honest, Acharagma roseana plants are not particularly ingratiating – the flowers are not breathtaking nor is the spination stunning. To me, the attraction of A. roseana lies in its close relationship with Lophophora and Obregonia, making it an obvious candidate for hybridization experiments.


Acharagma roseana flower with pale yellowish tepals with brownish pink midveins

Gene sequence studies by Butterworth et al. (2002) suggest that Acharagma belongs in a well-supported taxonomic clade with Lophophora and Obregonia. They sum up their results as follows:

LOPHOPHORA CLADE. Although there is strong support for this clade (87% bootstrap, 5 decay steps), few morphological features unite this clade. All members have napiform or carrotlike tap-root systems, although these features are also found in other members of the tribe.

The two species of Acharagma have been a source of taxonomic confusion. […] The rpl16 intron data suggest the removal of these two species from Escobaria, placing them in a well-supported (bootstrap 87%, decay 5) clade containing Obregonia and Lophophora, the latter shown to be polyphyletic based on this topology.

(the closing remark on Lophophora being polyphyletic is an interesting statement in its own right, but that’s a whole other story)


Acharagma roseana flower with pink tepals with darker red-tinged midstripe

In The New Cactus Lexicon, Hunt describes this species as Acharagma roseanum - I’m not sufficiently well-versed in the Latin language to say which form of the name is correct, but according to this site on botanical Latin the specific epithet must match the genus name in gender. As Acharagma is from the Greek a, without, charagma, groove (the gender of which, according to Hunt, is neuter) this would warrant the “-um” ending. I’m a firm believer in applied laziness so I’ll stick to the (apparently erroneous) “-a” ending for a wee bit longer and avoid having to re-label my plants ;-) Hunt accepts two subspecies: ssp. roseanum and ssp. galeanense, the latter having more spines and slightly larger, cylindrical bodies, compared to the egg-shaped body of the typical form.


Comparison of the two different color forms of Acharagma roseana flowers

The Acharagma roseana plants featured in this post were started from seed in 2005 and flowered freely last summer. As mentioned I would like to use these plants for hybridization experiments with Lophophora but I still need to come up with a good protocol for avoiding self-pollination. The locality information for these specific plants is: LX 578; Ramon Arizpe, Coahuila - “Ramon” should probably read “Ramos” but I'll stick to the information from the vendor’s seed list.


Acharagma roseana with fruits

Like most other parts of Acharagma roseana the fruits are not showy either. They are an inconspicuous yellowish green color, but have a surprising, not unpleasant, tart taste reminiscent of gooseberries.

The plants grow in my unheated greenhouse and I hope they survive the month long cold snap that is just now loosening its grip of Denmark.

Lophophora alberto-vojtechii, a new miniature species in the genus Lophophora

In the June 2008 issue of the Cactus & Co magazine Jaroslav Šnicer, Jaroslav Bohata, and Vojtěch Myšák described a new Lophophora species, Lophophora alberto-vojtechii, characterized by its small size, adult plants typically reaching just about 2 cm (less than one inch) in diameter.


Flowering Lophophora alberto-vojtechii at the type locality. Scale is indicated by the coin that is 23 mm in diameter

The full description of the lilliputian species reads:

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The stem is a greyish green with a touch of mauve (greenish, greenish grey to mauve), for the most part concealed in the ground. The aerial part (crown) is flat to slightly convex, 18 mm wide (10 to 50 mm in adult plants); the subterranean part of the stem is cylindrical, extending 25 mm down from the base of the crown (the depth in adult plants increases with age). The stem is distinctively small and divided into five ribs (up to eight ribs in exceptional cases). The trichomes are visible only near the apical meristem; they drop off early, leaving the areoles inconspicuous.

The root is thick and bulbous, 30 mm long and 16 mm in diameter where it joins the subterranean stem, smooth, and a dirty white to creamy yellow colour.

The flower is 23 mm wide (15 mm to 35 mm). The tepals and style are long, the same as in other members of the Diffusae section. The filaments are white, the anthers a yellowish orange. The style is white, while the stigma is yellowish (white to yellowish or pinkish). The tepals are light pink with a distinctive darker stripe (white to dirty pink, rarely creamy yellowish with a stripe in a brownish, salmon or darker pink shade) on the adaxial surface.

The fruit is round to slightly claviform, and dries out over time. The fruit is a dirty white colour, ranging to yellowish or pinkish.

The seed is black, round, 1.45 mm long (1.15 to 1.45 mm) and 1.45 mm wide (1.0 to 1.45 mm). The hilum is compressed into a broad V-shape. The perimeter of the hilum consists of a pronounced edge. The testa is nodulated with protruding outer cell walls. The individual cells of the testa are clearly demarcated.

Type locality: northern San Luís Potosí, Mexico, 1700 metres above sea level.

The type specimen was collected by G. B. Hinton on 1st August 2007 and is kept at the G. B. Hinton herbarium filed under number 28642.

We are naming the plant alberto-vojtechii in honour of two persons, the late prominent Czech traveler and cactus-hunter Alberto Vojtěch Frič and the current Lophophora enthusiast Vojtěch Myšák.

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Lophophora alberto-vojtechii just finished flowering. At this size plants flower regularly

The authors place L. alberto-vojtechii within the section Diffusae (see the Kaktusy 2005, Lophophora Special) along with L. diffusa, L. fricii, and L. koehresii. Based on similarities in habitat (mud flats), growth form, body, fruit, and flower morphology, and loss of trichomes in older areoles, Šnicer et al. conclude that L. alberto-vojtechii is closest related to L. koehresii. However the two species differ in size, the color of the epidermis, seed morphology, and the number of ribs.


Flowering Lophophora alberto-vojtechii at the site in Zacatecas

Šnicer et al. argue that even though the small Lophophora alberto-vojtechii plants might look immature at a first glance they are not, as is evident by old growth epidermis at the ground level and the subterranean part of the shoot.


Lophophora alberto-vojtechii (a) Flat, photosynthetic crown (b) Subterranean part of stem (c) The root

The habitat comprises flat, alluvial sediments, as in the case of Lophophora koehresii. Lophophora alberto-vojtechii has a distinctively geophytic lifestyle, withdrawing into small cavities below ground during dry periods. Here the plants are covered with dust and dry leaves, protected from the sun until the next rains.

The type locality of Lophophora alberto-vojtechii is in the northern part of San Luís Potosí, but it has also been found growing in the states of Zacatecas and Coahuila. The flowers of the Zacatecas plants are more white than those of the plants at the type locality (see picture above).


Flowering Lophophora alberto-vojtechii at the type locality

All in all the description is credible and well argued. Adding to the credibility is the fact that George Hinton collected the holotype and stores it in his herbarium, I doubt he would do that if he wasn’t convinced that this new taxon is for real.


The flower buds are visible very early, as they are not hidden in the wool.

References
Jaroslav Šnicer, Jaroslav Bohata, and Vojtěch Myšák (2008), “Lophophora alberto-vojtechii - an exquisite new miniature from the genus Lophophora”, Cactus & Co 12 (2), 105-117

Curtis’s Botanical Magazine – plate 4296 (vol. 73, 1847), Lophophora williamsii

Since my early days as a “Lophophora aficionado” I’ve heard and read that the first illustration of Lophophora williamsii appeared in Curtis’s Botanical Magazine in 1847 (plate 4296). This fact seems to be very well known – probably because of Anderson’s book Peyote, The Divine Cactus – nevertheless it’s been next to impossible to find a good reproduction of the illustration.

These days are over. The Botanicus Digital Library has scanned the material and made it available online. Below you’ll find copies of the scanned illustration as well as the accompanying description; the description is also reproduced in textual form.

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TAB. 4296.

ECHINOCACTUS WILLIAMSII.

Mr. Williams' Echinocactus.
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Nat. Ord. CACTEAE.- ICOSANDRIA MONOGYNIA.

Gen. Char. (Vide supra, TAB. 4190.)
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ECHINOCACTUS Williamsii ; humilis caespitosus turbinatus inferne teres transversim cicatricatus cinereo-fuscus superne umbilicato-depressus glaucus 6-8-costatus, costis latis convexis parce tuberculatis inermibus pulvilligeris, pulvillis e pilis fasciculatis densis erectis formatis, floribus parvulis subsolitariis albo-roseis.

ECHINOCACTUS Williamsii. “Lemaire, ex Salm-Dyck in Otto et Dietr. Allgem. Gartenzeit, xiii. p. 385.” Walp. Repert. V. 5. p. 816.
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A neatly-formed species, which has a very pretty appearance when its starry blossoms are expanded. We received several plants of it at the Royal Gardens of Kew, through the favour of the Real del Monte Company, from the rocky hills of their district of mines in Mexico, with many other treasures. It flowers in the summer months.

DESCR. Our largest plants do not much exceed the size represented. They grow in a tufted manner and are often proliferous, as in the instance here shown: the parent plant being, as it were, stifled or subdued by its offspring. Each individual is turbinate: from the base to the crown, or summit, terete, of an ashy brown colour, and scarred with close transverse lines, occasioned, it would appear, by the progressive withering and contraction of the tubercles: the summit is broadly convex, but with a deep depression in the centre, glaucous, traversed from the centre outwards by 6-8 furrows, and thus divided into as many convex ridges, and these again, transversely, but more or less deeply, into rather large, rounded, more or less confluent unarmed tubercles, each of which has a dense tuft or short pencil of compact erect hairs:--no aculei. Flowers proceed from a young tubercle, near the centre of the crown. The base of the calyx is downy. The petals lanceolate, rather numerous, white, externally tipped with pale green, and having a rose-coloured line down the centre. Stamens yellow. Stigma of four spreading rays.

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The scans are courtesy of the Botanicus Digital Library, Missouri Botanical Garden and are free for non-commercial use, as long as attribution is provided.

High-resolution copies of the scans can be found in the files section of the Lophophora Google group.

Back to online articles.

The genus Lophophora – Kaktusy Special 2, 2005: A Review

I just received the new Kaktusy Special monograph on Lophophora. It comprehensively describes the distribution, characteristics and classification of the genus. In an attempt to resolve some of the taxonomic confusion surrounding Lophophora the authors propose a division of the genus into two sections and a change of rank.

Based on factors like chemical composition, habitats, incompatibility of the species, rib numbers and morphology, etc, a division of the genus into the two sections Lophophora and Diffusae is proposed.

Section Lophophora comprises the various forms of the species L. williamsii including the type species (hence the autonym for the section).

Section Diffusae includes the three species related to L. diffusa, i.e. L. diffusa itself, L. fricii, and L. koehresii (aka L. diffusa v. koehresii, aka L. viridescens).

The authors argue convincingly for the taxonomic revision but it would have been interesting if the revision was supported by DNA sequencing results (like Butterworth et al. who confirmed that L. diffusa and L. williamsii are indeed distinct species).

The description for each species includes a detailed distribution map showing the range of the species. The booklet is packed with excellent habitat photos showing the plants natural growth forms (actually some of the very best habitat photos I’ve seen are included in this work – see examples below).


Left – Lophophora williamsii, Sierra de la Paila, Coahuila
Right – Lophophora diffusa, Peña Miller, Queretaro

The booklet is rounded of with a few notes on the cultivation of Lophophora.

The genus Lophophora is a comprehensive and long needed review of the genus. As mentioned it would have been great with DNA sequencing results supporting the change of rank for L. fricii and L. koehresii. Also an index and a list of literature references would have been helpful. That being said, The genus Lophophora must be recommended to anyone interested in these fascinating plants.

The booklet is written by Jaroslav Bohata, Vojtĕch Myšák, and Jaroslav Šnicer; it comprises 48 pages, contains 89 color photos, 3 black and white photos, and 6 drawings. It is available from the Society of Czech and Slovak Cactus and Succulent Growers. Apart from the English edition I believe German and Czech editions are published as well.

Update - March 21, 2006
I have to mention the review of the Genus Lophophora published in the latest edition of CactusWorld (the journal of the British Cactus and Succulent Society). According to the reviewer this work “boldly goes where no taxonomist has gone before” in the attempt at clarifying the systematics of Lophophora ;-)

Köhres's Lophophora information page

I just noticed that Köhres has updated his Lophophora information page.

A lot of Lophophora photos ordered by state, location and species can be found by selecting "Bilder am Standort" (photos from the growing locations) and then "Los geht´s". This takes you to a map of Mexico where you can select a specific state. From the state page you can select a location (from the "Standorte" list). If more than one species grow in a given state, e.g. Coahuila or San Luis Potosi, the locations for the species are differentiated using color.

A very impressive site.

32. Lophophora

J. M. Coulter, Contr. U.S. Natl. Herb. 3: 131. 1894.

Peyote [Greek lophos, crest, and phoreus, a bearer, in reference to tufts of hairs in areoles]

Allan D. Zimmerman & Bruce D. Parfitt

Plants erect, commonly unbranched, becoming many branched and moundlike in old age, deep-seated in substrate. Roots taproots. Stems unsegmented, usually gray-green or blue-green to dark green [yellow-green in L. diffusa of Mexico], usually flat-topped and cryptic in soil surface, sometimes protruding above it with recessed apex, ribbed or weakly tuberculate, 2-7.5 × (4-)5-12 cm, softly fleshy, soft skinned, dull, glabrous; ribs 5, 8, or 13 (rarely 21), low, broadly rounded, straight, vertical or less often helically curved around stem; areoles 3-15+ mm apart along ribs or at apices of low, humplike tubercles, circular, copiously hairy, hairs usually in compact, erect tufts to 7-10 mm; areolar glands absent; cortex and pith not mucilaginous. Spines absent. Flowers diurnal, deeply nestled in copious areolar hairs at stem apex, arising from adaxial edges of areoles, campanulate, 1-3 × (1-)1.5-2.5 cm; outer tepals whitish to greenish pink, midrib greenish, margins entire or minutely fringed or ciliate distally; inner tepals usually white to pink [rarely yellowish white or magenta to reddish violet, at least in Coahuila, Mexico], 8-14(-22) × (1-)2.5-5 mm, margins ciliate or entire; ovary smooth, scales, hairs, and spines absent; stigma lobes (3-)4-8, white or pinkish, 1-3 mm. Fruits indehiscent, white to pinkish [to purple], clavate to nearly cylindric, 11-25 × (2-)4-5 mm, weakly succulent, quickly drying and contracting after ripening, upon drying becoming translucent and brownish white or whitish, spines and scales absent; pulp colorless; floral remnant weakly persistent or tardily deciduous. Seeds black, somewhat pyriform, cylindric, or obovoid, 1-1.5 × 1-1.2 mm, not glossy, with large, flat hilum; testa cells strongly convex. x = 11.

Species 2 (1 in the flora): arid regions, sw United States, Mexico.

1. Lophophora williamsii (Lemaire ex Salm-Dyck) J. M. Coulter, Contr. U.S. Natl. Herb. 3: 131. 1894.

Mescal buttons, divine cactus

Echinocactus williamsii Lemaire ex Salm-Dyck, Allg. Gartenzeitung 13: 385. 1845; Anhalonium williamsii (Lemaire ex Salm-Dyck) Lemaire

Plants 0-50-branched, nearly flush with soil surface. Roots fleshy, broadly carrot-shaped, 6-12 cm. Stems flat or domelike with deeply depressed center; ribs to 25 mm broad. Flowers: outer tepals narrowly elliptic to oblanceolate, apex acute, mucronate; inner tepals elliptic, margins white to greenish pink, midstripes darker, apex mucronate or attenuate; ovary 3-4.5 mm; styles white, 5-14 mm. 2n = 22.

Flowering Mar-May(-Sep). Chihuahuan desert scrub, Tamaulipan thorn scrub, usually on or near limestone hills; 100-1500[-1900] m; Tex.; Mexico (Chihuahua, Coahuila, Nuevo León, San Luis Potosí, Tamaulipas, Zacatecas).

Long used for religious and medicinal purposes by native peoples, Lophophora williamsii is famous for its psychoactive alkaloids, primarily mescaline, which are concentrated in the small, photosynthetic, above-ground portion of the stem. Experienced peyoteros harvest only the top few millimeters of the stem, leaving the deeply recessed apical meristem, thus allowing regeneration. The plants live for many decades and grow very slowly. Anhalonium, a later name for Ariocarpus, has been applied to species of Lophophora, and was widely used in older pharmaceutical literature.

Fruits abruptly ripen up to one year (rarely longer) after flowering.

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Back to online articles.
The Flora of North America web site.