Neolloydia....Who?

Even from a splitter's viewpoint there are occasions when the lumping of genera makes sense. One such "occasion" is probably the amalgamation of Gymnocactus, Rapicactus, Normanbokea, and Turbinicarpus under the Neolloydia prefix.

As an individual I have been convinced of the merits of this action for several years, much to the chagrin of my colleagues, whose derision remains abundant...hence the title of this article!

History

In the beginning (and lo!)... the genus Neolloydia was erected by Britton & Rose in 1922 in order to classify three rather uncomfortable species of cacti formerly placed in Mammillaria - N. conoidea, N. ceratites, and N. grandiflora. A fourth was added by Br.and R in 1923 - N. texensis. But it was not until 1948 that a fifth and final species was added by Backeberg - N. matehualensis.

Since that time, and probably due to the poor definition of generic boundaries, Neolloydia has been a dumping ground for a plethora of "I don't know what to do with it" species from such diverse genera as Mammillaria, Coryphantha, Escobaria, Echinomastus, Cumarinia, and Thelocactus.

It was not until Glass and Foster (1977) showed that there are very few botanical differences between Turbinicarpus and Gymnocactus that the definition of the Neolloydia genus began to take shape. Their work was followed up by John and Riha (1981) and finally by Anderson in 1986. Ultimately it appears that Anderson's work on the Neolloydia complex is now the generally accepted view amongst taxonomists.

From personal observations of Anderson's definitive article however, there is a suggestion that, whilst Gymnocactus and Turbinicarpus are clearly related, Neolloydia in the narrow sense seems equally distinct from these genera as it is from Thelocactus! Marginally so, perhaps, but a point which I am sure has not been lost on the taxonomists.

However, for the need to justify the ramblings of this article (and the very existence of it!) we will presume that all species discussed here are all of the same ilk, at least until we know different...!

Habitat

Neolloydia in the broad sense appear to have a quite specific range in the wild - the dry scrub areas of southern Texas (Big Bend) and the Chihuahua desert of north-eastern Mexico. More specifically they are found growing in limestone or, not uncommonly, gypsum soil, and at a height of between 700 m and 2500 m.

Cultivation

With the exception of the rather dubious N. horripila all species show a distinct intolerance to over watering either by rotting off (Neolloydia in the narrow sense), or by bursting of the epidermis (Turbinicarpus and Gymnocactus). Composts should therefore be very open and gritty. To prevent growing a monster totally out of proportion to their attractive and natural form, one should limit the fertiliser. As for pests, they seem remarkably resilient, particularly the Turbinicarpus and Rapicactus species. The Gymnocactus occasionally suffer from corkiness around the base, and since some of them are rather thin necked, cautious examination should periodically be undertaken to ensure that rot has not set in. Re-rooting is not easy. The most difficult species to grow is arguably N. matehualensis. It usually requires support and seems to suffer from a similar bacterial epidermal problem to that often seen on Ortegocactus macdougallii. N. matehualensis is, for me, quite distinct from other Neolloydia (in the narrow sense) both in terms of cultural demand and its quite unusual glossy grey-green epidermis which on closer inspection appears porous! I have also recorded something about this species which I have never observed elsewhere - last Summer it was scorched by sunlight, yet the bleached areas have largely returned to their former condition.

The flowering habit of most species is similar, a show in the early season and then again in early autumn. The exceptions being Neolloydia (in the narrow sense), N. horripila and N. gielsdorfiana which tend to flower sporadically throughout midsummer, interestingly enough, a trait of Thelocactus!

Species

What follows is a subjective analysis of the recognised plants in the Neolloydia complex (with abbreviated former generic names in parenthesis), and of those whose existence in our collections seems deserved, if not recognised. Please note therefore that taxonomists may not approve!

N. conoidea

(Syn: N. ceratites, N. grandiflora, N. texensis)

Stems slowly clustering to about 15.5 cm tall, usually club shaped. The body is a distinctive yellow/green, with prominent tubercles. Radial spines are white occasionally tipped black. Central spines range from 0-3, even varying on the same plant, black erect and to around 12 mm long. A very variable plant. Two variations to watch out for are the white flowered form of N. texensis, a nice colour break from the uniform magenta common to most forms, and the N. grandiflora form from Guadalcazar which has no centrals, a very pale body, offset by a quite woolly crown for this species.

N. conoidea var. matehualensis

This plant is quite different, or, at least, represents an extreme form of the type above. Its habit is similar to the species but for reasons explained elsewhere it does appear to have distinct merits of its own.

N. subterranea (Rap.)

A compact single stern, club-shaped, with a very long, narrow, neck not often seen on cultivated plants. It is renowned for its very long tuberous root system. Radial spines numerous, about 4 mm long or less. Central spines two, usually darker than the radials, erect, to about 1.5 cm long. There appears to be some conflicting opinions on flower colour, they have been described as yellow-brown by some authorities and magenta by others. I have both this species and its variety zaragozae from documented sources, the type has yellow-brown flowers and is botanically correct in all other details. Subjectively, I suggest that flower colour can vary, since the variety has magenta flowers but it too is botanically correct elsewhere.

N. subterranea var. zaragozae (Rap.)

Similar to the species but without the long neck, slightly larger growing, (and easier), with upward pointing central spine. The crown of this plant appears more densely spined and has more wool.

N. mandragora (Rap.)

Seemingly quite rare in cultivation, and certainly not as well known as its illustrious cousins, despite having been described six years earlier! It is more globular, although similarly spined to N. subterranea. It has a much thicker root system, rather turnip like. Flowers are larger, to 2.5 cm across, greenish-magenta with a reddish mid-vein. Its overall appearance (above ground) clearly shows the relationship between Rapicactus and Gymnocactus.

N. gautii (Gym.)

(Syn.: N. smithii and its fa. senilis, N. beguinii (Gym.))

This species suffers from an identity crisis, but I have elected to refer to it by its generally accepted name. It is larger than many of its cousins, being up to 10 cm high and 6 cm in diameter. Like N. conoidea it is quite wide-ranging in habitat, and several forms are known; e.g. N. smithii (SB308) has no centrals, a globular habit, and brownish radial spines which reflex towards the plant body, whilst N. smithii var. senilis (PZ263) is much whiter spined, more cylindrical in form and usually has 2-3 central spines to 1 cm long. All forms have a magenta flower with lighter margins. This species is prone to rupturing of the epidermis when over-watered.

N. gielsdorfiana (Gym.)

Apparently virtually unknown in habitat, this species makes for a most attractive and striking plant if grown well. Unfortunately its large, angular tubercles often die back at the areoles, spine clusters are easily dislodged and it has a rather slender neck which makes for a rather precarious existence if not supported by a healthy top dressing of grit around it. Like most Gymnocacti it is solitary, with a bluish epidermis. Spines are radial, black with a white base, fading to grey with age, to 2 cm in length. Areoles are woolly, the crown too being quite densely woolly. The flowers which open among the spines, are cream coloured, small, and quite delicate in appearance.

N. horripila (Gym.)

If this species is truly a Neolloydia (in the broad sense), then it is the genus's version of Mammillaria bombycina. Against other members it is almost invasive in its clumping habit, achieving 20 cm or more in diameter with numerous heads. It also grows much further south than its relatives, and unlike any other species, appears susceptible to scorching. Plants have prominent conical tubercles, with star like spine clusters. There is a central spine, usually darker and larger - to 2 cm. Flowers are magenta with a white throat.

N. knuthiana (Gym.)

Contrary to the last species, this is a real joy. Body globular to 6 cm diameter, green with small conical tubercles. Radial spines are numerous to 5 mm, slender usually straw coloured, but there is an attractive white spined version in cultivation. Central spines similar, to 1 cm. Flowers are a delicate shade of pink with a darker mid-stripe, small, to about 1 cm long and 5 mm across. The paler spined version also has white flowers. A slow growing, attractive and unassuming species from Guadalcazar, San Luis Potosi.

N. knuthiana var. ysabelae (Gym.)

This species, from Tule, Tamaulipas, is tentatively placed here on account of its similarity in habit to the type. This is the opinion of Backeberg (1961). However, this relatively unknown plant may be "similar" but it differs markedly on two counts. Firstly, its dwarf size, I have a four year old specimen which is only 3 cm in diameter, so, it is also much slower growing. Secondly, on account of its large taproot; it is this latter feature which suggests the link with Turbinicarpus. Flowers are paler than the type. Whatever its status, this is a most desirable plant, and certainly commands respect for its difficulty of cultivation.

N. saueri (Gym.)

Known from only a small area of Tamaulipas, this small species, to 5 cm diameter, slightly flattened, grey, and with copious amounts of white wool in the crown, is still uncommon in collections even though it was discovered as early as 1928! The radial spines are white, barely covering the angular, flattened and largish tubercles. The single central spine is black, to 2 cm long, and flexible. Flowers are small, with a reddish mid-vein. It is extremely slow growing but not difficult in cultivation.

N. viereckii (Gym.)

Not as common as its variety, this species is recorded as "gently" clustering to around 10 cm - each head being about 5 cm in diameter. The body is blue green with white radial spines to 1 cm in length and generally without centrals. Mature specimens produce copious amounts of white wool in the crown, often enough to obscure the spination! Flowers are magenta with a white throat or all white, small to 1 cm long and 5 mm or less wide. The collection HK1377 seems particularly attractive with much darker flowers and almost cephalium-like wool. From the Jaumave valley, Tamaulipas.

N. viereckii var. major (Gym.)

Despite the name, the variety is only marginally larger than the type, and perversely, is usually solitary. Its two most obvious differences are the larger creamy-white flower and prominent, stout, black central spine to 1.5 cm long. Species of Gymnocactus with the collection number L730 are referable here. Fairly widespread in habitat, it too produces the woolly crown but not to the extent of that produced by the species.

N. pseudopectinata and var. rubriflora (Norm.)

An attractive species which often appears to be a juvenile Pelecyphora aselliformis at first glance. This dwarf plant grows to about 3 cm tall and slightly cylindrical. Spines are aptly described as pectinate and are minute. There are no central spines. Flowers appear in the depressed crown and can be quite striking, being brownish on the outside and a nice bright white on the inner: to about 2.5 cm, across. In 1978 a red flowered variety was discovered and has tentatively been known as a variety ever since. Subjectively, it is probably deserving of that status. The flowers are indeed reddish, with a paler mid-stripe, they are also much larger than the species. But it is its general appearance which merits another look; the reddish factor is also apparent in the overall hue of the spines, rather like the better known rubrispina form of Echinocereus rigidissimus. Susceptible to over-watering.

N. valdeziana and var. albiflora (Norm.)

This quite extraordinary looking plant has been affectionately described as "looking like a bird's dropping". A very accurate description. The plant grows very slowly, arguably slower than an Ariocarpus! It takes on an encrusted rock-like habit. The spines are white, quite plumose and minute enough not to be readily discernible. It grows to around 2.5 cm tall, and possesses a large taproot out of all proportion to the plant above ground. The flowers are magenta, almost as large as the plant. The flower buds often appear in the autumn, only to over-winter before opening in March the following year. There is a white flowered form which when in bloom gives the plant quite a bleached look, not unlike its habitat, I suspect! But unlike the previous species there appears little argument to support varietal status.

Justification

Before going into the former Turbinicarpus species it is probably worth pausing to note why they have been lumped with Gymnocactus et al. Glass and Foster (1977) noted that, "..it is difficult to separate these two genera either on the basis of distribution, habit and appearance, or flower, fruit, dehiscence or seed". Anderson (1986) commented that it may be that Turbinicarpus "may represent a group of organisms demonstrating Neoteny..." That is, they are capable of all adult functions whilst remaining in a physically juvenile condition. Indeed, in comparison of one year old seedlings it is difficult in some instances to distinguish Gymnocactus from Turbinicarpus. A good example of the neoteny phenomenon is Sulcorebutia steinbachii var. horrida and its juvenile version gracilior.

N. hoferi (Turb.)

This is a recently described species which grows in the gypsum soils of the fabled Geohintonia territory. From photographs it appears fairly typical of the genus in form, although it does not appear to produce the central spines so typical of other members of the genus (with exceptions). Flowers are again fairly typical, whitish to about 1 cm long and slightly less across. It does seem to be developing a reputation for being difficult to grow, and after an initial wave of availability from some nurseries seems to have become very scarce again. It appears to be distinct enough to warrant species status.

N. swobodae (Turb.)

There seems to be even less information about this one than N. hoferi! I have no locality details, although San Luis Potosi / Nuevo Leon would seem a reasonable guess. [See BCSJ Vol.12 (4) 1994 Newsletter page xxviii Progress report on Conservation Projects. Ed.] My two year old seedling has quite prominent, angular tubercles with small star-like clusters of off-white/grey radial spines. New growth suggests that there may be a darker, longer central in the offing. Flowers are unknown but probably typical of the genus.

N. laui (Turb.)

A popular species from San Luis Potosi which seems to have been around longer than the intervening years from its actual discovery in 1975. It is quite different from the norm for Turbinicarpus being much flatter, to 4 cm diameter and 1.5 cm tall. Tubercles are prominent, angular, and surmounted by, usually, straw coloured radial spines up to 2 cm long which stand quite erect. Flowers are a nice colour break, for this genus, being a shell-pink.

N. roseiflorus (Turb.)

I am reluctant to include this species. Primarily, because having possessed at varying times, four specimens (from different sources), no two have been quite alike. I believe that most plants have derived from continental nurseries, and that some authorities believe it to be of hybrid origin. Subjectively, even if it is of habitat origin there is some suggestion that it, and some of the N. schmiedickeanus forms may hybridize naturally.

N. lophophoroides (Turb.)

At 5 cm diameter, this is the largest of the Turbinicarpi or would be if it survived long enough in cultivation. As its name implies, with the exception of a few token, small, radial, grey spines, this species does look remarkably like its namesake, Lophophora. Its blue-grey body contrasts well with the fresh white woolly crown, through which appear the occasional delicate pale pink, and quite large flowers; up to 2 cm across. As lamented above, this species seems prone to rotting, and mature specimens are uncommon in cultivation. It also appears to be difficult to locate in habitat and probably has a very limited range, reportedly near Las Tablas, San Luis Potosi.

N. krainziana and var. minima (Turb.)

Taxonomically these names may not be accurate. N. krainziana is not known in habitat (according to information at hand), whilst the variety minima, which is certainly of habitat origin, caused a sensation when it was discovered a few years ago. Both forms are reportedly from Queretaro and Hidalgo, where their probable parent species N. pseudomacrochele has been collected. N. krainziana will readily form a small clump of dark green bodied, spidery-spined plants. Flowers are typical of the genus - small and off-white, often with a greenish hue. The variety minima is quite extreme, being very thin and columnar - 6 cm tall yet barely 5 mm in diameter. Flowers are minute, quite green and open star-like over the crown of the plant. Spines are different, being quite rigid, occasional darker centrals, radials are white and about 2 mm long, interlacing to form a dose covering over the body, which is difficult to see because of the density of spination.

N. pseudomacrochele (Turb.)

The previous species and variety may be more correctly placed here not least on the basis for the current craving of geographical variation in single populations. The plant usually remains solitary in habitat, but will clump slowly in cultivation. Stems to 4 cm tall and 3 cm diameter, dark green with a prominent woolly crown. Spines to 2 cm or more, very straggly, thin and yellow-brown in colour. Flowers are the largest of the Turbinicarpi being 2.5 cm long and 3 cm in diameter. Colours are slightly variable, the best being a pink form with a magenta mid-rib. If grown true to type this makes for a very attractive plant. From a locality further south than most.

N. schmiedickeana and varieties: klinkeriana, macrochele, schwarzii, gracilis, dickisoniae and flaviflora.

If N. horripila's party trick is its impression of Mammillaria bombycina then N. schmiedickeana's is Sulcorebutia steinbachii in all its myriad forms. Indeed, as I write this, there are new forms of Turbinicarpus being discovered, most of which are clearly related to this species. Certainly if one considers the current treatment of Rebutia as the way forward in taxonomy, then those rules should easily apply here. All species originate from the Nuevo Leon / San Luis Potosi area of Mexico.

N. schmiedickeana grows to 2.5 cm tall and will offset to form a small cluster. It has a dull-green body with dark spines to 1 cm long, usually curved towards the apex of the plant. Like several of its varieties, the spines have a spongy texture. The flowers are small, to 1 cm diameter, white with a darker mid-rib. There is a more squat form in circulation, usually labelled miquihuanensis.

N. klinkeriana is quite similar to the type but is geographically isolated, (according to current knowledge). It tends to grow somewhat larger - to 5 cm and has a shorter, singular central spine. Flowers are typical for the species.

N. macrochele is quite different. The stems are blue-green, globular with poorly defined tubercles. Spines number 3-6, and up to 3 cm long, flexible and quite tangled, particularly in the crown of the plant, they are whitish to straw coloured. The flowers are slightly larger than the species, more translucent white. A quite distinguished plant, growing to 5 cm high and globular in form.

N. schwarzii is similar to the former variety but more squat and usually only with a single recurved spine to 1 cm long. The flowers are 1.5 cm in diameter and creamy-white. It is particularly prone to bursting of the epidermis. The misnomer polaskii is referrable here.

N. gracilis is a real treat. The body is virtually obscured by a dense covering of long thin Leuchtenbergiain tubercles, each surmounted by one or two long, to 3 cm, flat, papery spines. There are some token radial spines. The woolly crown yields a flower typical of the species. The physique of this plant can justifiably give mealy bugs a real sense of security from probing cotton-wool buds.

N. dickisoniae apparently grows little more than a stone's throw from the previous variety and superficially resembles it. The plant does not have, however, the prominent tubercles and the long papery central spines, usually only one central spine, rounded as opposed to fiat. The flowers are similar to the species.

N. flaviflora is to many the favoured Turbinicarpus to grow. It has a very distinctive form, described by some as almost pagoda-like. The body, cylindrical to 5 cm tall, is grey in colour and coated in a whitish bloom. The spines are spongy, 1 cm long, recurved upwards. The flowers, from which it gets its name are an unusual greenish-yellow and contrast well with the body. A striking species.

Conclusion (Phew!)

Well, subjectively and all, that seems to be the Neolloydia complex as we know it. I am sure that sooner or later, the burning issue of the Neolloydia species (in the narrow sense) will be resolved and we may yet find that we can call them all by some other name, (conceivably a ploy, paid for by the label manufacturers!). Since this is strictly an amateurish stab at defining (I use the word very loosely) a popular group of plants, I welcome any correspondence on the matter in the hope that we may all understand the inhabitants of our greenhouses a little better.

References

• Anderson, E.F. 1986. A revision of the genus Neolloydia. Bradleya 4, pp 1-28.
• Britton and Rose The Cactaceae Vol.3 and Vol.4 1923.
• Glass and Foster A Revision of the Genus Turbinicarpus Cact. & Succ. Jour. USA 52 (4) pp 161-176. 1977.
• John and Riha Unkombinationen aus Gymnocactus in de gattung Turbinicarpus. Kaktusy 17: 15-18, 19:22.

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