The Caryophyllales are an order of flowering plants that include:

Cacti Amaranths Ice Plants

Carnations Beets Carnivorous Plants

- Many members have fleshy stems or leaves.

Synapomorphies:
Peripheral embryo around a central nutritive perisperm, instead of an endosperm (reflected by the ordinal connotation Centrospermae Betalains pigments (Red/yellow) A proteinaceous plastid distinct similarities in chloroplast DNA (cpDNA)

These characteristic traits do not occur in any other angiosperm clade and in recent decades has formed a solid case for recognizing the Caryophllales, and thus Cactacea as core members, as a monophyletic clade.

Cacti (or cactus) are easily recognized amongst other members of the plant kingdom due to distinct morphological characteristics.

Cacti are commonly known for unique lack of leaves, instead photosynthesis takes place in the stems. Although they are typically recognized by this feature, not all cacti are leafless. In fact, leaves are actually an ancestral trait, and their loss occurred after the break from the rest of the terrestrial plant lineage.

Another distinguishing feature of cacti is the presence of intimidating defensive spines. The spines emerge from structures called Areoles, which are unique to the family. Although variable, they typically appear as woolly or hairy areas on the stems. Spines are distinct from thorns: Spines are modified leaves Thorns are modified branches. Spines provide strong evidence that the loss of leaves occurred after the split. Spines are present even in cacti with leaves, such as Pereskia, Pereskiopsis and Maihuenia, so they clearly evolved before complete leaflessness.

Similar to spines, flowers are also produced from the areoles and are variable.

Typically, the ovary is surrounded by material derived from stem or receptacle tissue, which forms a floral tube. Although strictly speaking, the outside often has areoles that produce wool and spines, thus only the part furthest from the base is floral in origin. The tube also has small scale-like bracts, which gradually change into sepal-like and then petal-like structures, so the sepals and petals cannot be clearly differentiated (tepals).

 Cactus flowers usually have many stamens, but only a single style, which may branch at the end into more than one stigma. The stamens usually arise from all over the inner surface of the upper part of the floral tube The flower as a whole is usually radially symmetrical (actinomorphic), but may be bilaterally symmetrical (zygomorphic) in some species. Flower colors range from white through yellow and red to magenta.

Adaptations to Dehydration

Cacti have a thick, hard-walled, succulent stem, which are often fluted or ribbed. This allows them to swell and store water when it rains, because inside of the stem (cortex) is either spongy or hollow. A thick, waxy coating keeps the water inside the cactus from evaporating. Cacti have a particular increased cortex width in which the photosynthetic outer cortex is farther away from the relatively narrow pith vascular that contains the vascular bundles of the stem (Stele).

Adaptations to Dehydration
Life sustaining nutrients are transported through the xylem and phloem. In large barrel cacti the epidermis and chlorenchyma are often several centimeters away (as much as 40 cm in species). Diffusion of water and nutrients is extremely slow across this distance. Even with a thick covering of cuticle and wax, the epidermis will lose some water to arid desert air. Water must be replaced rapidly if the epidermis and outer cortex tissues are to remain alive.

Adaptations to Dehydration

Rapid translocation throughout the cortex of many cacti is made possible by the presence of a specific system of collateral cortical bundles.

Adaptations to Dehydration
Biology textbooks often highlight the familys Crassulacean-acid metabolism (CAM), an alternative to traditional photosynthesis. The stomata in the leaves remain shut during the day to reduce evapotranspiration, but open at night to collect carbon dioxide (CO2). The CO2 is stored as the four-carbon acid malate, and then used during photosynthesis during the day. The pre-collected CO2 is concentrated around the enzyme RuBisCO, increasing photosynthetic efficiency.

 Cactus (or cacti) are easily recognized amongst other members of the plant kingdom due to distinct morphological characteristics. Besides their spines and thick fleshy stems, the family is known to possess a remarkable ability to survive in extremely dry habitats.

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Habitat:
With the exception of one species, all cacti are native to the New World; and their resilience to harsh conditions has allowed them to inhabit various climate types as they spread across the Americas.

The black sheep of this family is the epiphytic genus Rhipsalis; which has likely spread naturally by birds, to tropical Africa and Madagascar as well as to Sri Lanka and southern India The true benefit of these modifications is expressed in desert landscapes where cacti are the dominant species; here many varieties are able to exist in conditions that are too severe for many plants or animals.

Late 1800s - 1905

The German botanist Alwin Berger (18711931), was known for his pioneering work of succulent nomenclature.

He was one of the first to evaluate the phylogenetic relationships among cacti.
In 1905 he proposed dividing the genus Cereus, which generically included all columnar cacti, into several new subgenera. Bergers peers widely accepted most of these decisions, thus establishing a foundation for the systematic classification of columnar. A couple of years later, Vincent Riccobono elevated Trichocereus to the genus level. However, he only included two species in the transition (T. macrogonus and T. spachianus).

Britton and Rose (1920)

Added an additional 17 species, bringing the species count of the tribe to 19.

Curiously though, they chose not support this decision with any discussion or even speculation on phylogeny. It is reasonable to infer such a drastic change would be highly criticized. However they complimented their work an additional identification guide based on the distinguishing characteristics of each species they observed. This morphological key was the first of its kind, and therefore revolutionary for future identification and classification efforts. Aside from a few additional species, their definition remained the primary accepted model, and the status of the genus remained stable for a large number of years. Unfortunately, because many of their classifications were ambiguous and ignored evolutionary history, their system has had minimal influence on modern hierarchies.

Curt Backeberg (1949)

German horticulturalist and cacti enthusiast, who made many field expeditions to collect data on species location, habitat, morphology, and the occasional photograph.

In 1949, as part of his reworking of the entire Cactacea family, he cleaved the Trichocereus genus in two. This revision split the species by their floral development morphology. Species with white petaled flowers that open at night would remain within Trichocereus While those with red, orange, or yellow colored and day-opening flowers would be transferred into a new genus Helianthocereus.

Backeberg (Cont.)
For more than 30 years he published an enormous stack of literaturewhich described his theories on cactus evolution, organization, identification, and nomenclature. Throughout these texts, he named or renamed well over 220 genera and nearly 3000 species. This new classification system was weighted heavily by his belief; the current geographic distribution reflected a bipolar division in evolutionary history radiating from an ancestral origin of Central America or the Caribbean. However, most of his work was eventually nullified, as current evidence contrarily predicts a South American birthplace. Sadly, he also neglected to submit a herbarium specimen for any species he ever described; essentially forfeiting any possibility of redressing his phylogeny. The botanist David Hunt is quoted as saying that he "named 78 more genera and named or re-named 1200 species without, so far as I know, ever making (or citing) an herbarium specimen. He left a six-volume monograph of the family running to 4000 pages and a trail of nomenclatural chaos that will probably vex cactus taxonomists for centuries."

Ironically, although much of his material is acknowledged as being discordant and inaccurate, it is still widely referenced in todays publications. likely due to sheer magnitude of information Backeberg published on the subject, and the lack of better material.

Franz Buxbaum (1958)

Franz Buxbaum, a skilled plant morphologist, published his ideas on Cactacea phylogeny. Similar to Britton and Rose, Buxbaums redesign included splitting the three subfamilies, that were known at the time, into nine tribes. One of the tribes he proposed was the Trichocereeae, which he designated Trichocereus as the type genus. He further split the tribes into sub-tribes, which were finally divided into lines. Therefore according to this schema, the Trichocereus line was now member within Trichocereinae, a sub-tribe of Trichocereeae. Buxbaum describes the sub-tribe Trichocereinae as being characterized by large and columnar stems and rarely globular; flowers radiate, campanulate to funnel-form; perianth large, mostly white or whitish, sometimes brightly colored; nectar chamber lacking or present; stamen insertion beginning at base of the receptacle or above a nectar chamber.

Buxbaum (Cont.)
Interestingly, in Buxbaums consideration of Trichocereus, he deviates from Backeberg and returns the genus Helianthocereus into the clade.

Buxbaum surmised there is a general trend in cactus evolution, a consistent pattern of increased specialization and morphological reduction in the evolution of various characters of the stems, flowers, fruits, and seeds. He was later supported by Kiesling (1978), who published an illustrated key describing the Trichocereus species of Argentina, and through his observations found Backebergs Helianthocereus to be synonymous.
Buxbaums notion of a directionality, or trend, in cactus evolution has been influential on subsequent considerations of phylogeny. Within the past two decades, a lightly revised and more modernized version Buxbauxs system has gained favor

Heimo Friedrich (1974)

Up until this time, pervious revisions of the clade arose from a desire to split it into smaller, less dissimilar clusters. Contradicting this trend, in 1974, Friedrich preferred instead to compress Trichocereus and several related taxa into a single sister genus Echinopsis. Surely influenced by Buxbaum, he demoted Trichocereus to a subgenus yet again. He reflected this change by renaming the previous sub-tribe and tribe Trichocereinae/Trichocereeae to Echinopsidinae/Echinopsideae.

The unification of several genus required Friedrich to adopt new nomenclature for some species; to avoid homonymous names for distinctly different species.
Still in the pre-DNA era, Friedrich justified the new sub-tribe Echinopsis based on two attributes, hairs in the axils of the scales and disposition of the stamens in two groups. Oddly, he makes this conclusion solely on related floral morphologies, which were known at the time to be observed in other tribe genera or polymorphic in some of the species. Similarly, he related the Echinopsidinae simply by the nectar chamber, but failed to assert which characteristics. What species was type genus of new echino?

Gordon Rowley (1974)

President of the Cactus and Succulent Society Journal Rowley (1974), a proponent Friedrichs idea, increased the audience and accessibility of his Friedrichs work by republishing it in the IOS bulletin

Rowley also provides very brief discourse, in praise of the merger, and condoning the decision based on the previous inability to identify individual groups based on floral characteristics: the undeniable truth is that the only way you can decide whether a plant belongs to Trichocereus or Echinopsis today is by applying a tape measure to its stem and hoping a short stem is not merely a measurement of juvenility. The flowers and fruit show no constant and recognizable differentia. Further, we know species that bridge the gap so there is no sharp discontinuity between tall (cereoid) and dwarf (cactoid) growth forms. Additionally, he amended Friedrichs original work with an enormous list of species that once stripped of their original genus, were lumped into the genus Echinopsis Rowley (1974).

*For those who were wondering; yes, this is the same Gordon Douglas Rowley who attempted to saddle peyote with the very odd purportedly common name of the L.S.D. cactus.

Friedrich Ritter
During the early 1980s he published successive the South American cacti, accumulating some 5,500 pages of notes and data that filled 74 volumes

However, only a minute fraction related to the sub-genus Trichocereus

His only recommendations for the group are integrating a couple separate species into a single species with known variants and the addition of a few new species.
Sadly, like many of the Cactacea pioneers before him, Ritter failed to identify proper identification keys and herbarium submissions that are plagued by absence, loss, inaccuracies or insufficiencies

Considering his focus on Trichocereus was almost negligible, the incompetence of his herbarium specimen is grossly evident.
The type specimen (FR 228a) is mentioned (Ritter: iii., 1979, 1980b) as having been deposited in the

National Herbarium of the Netherlands. According to Eggli et al. (1996), the holotype is lost. (Ritter 1908) erroneously corrected of the spelling name T. chiloensis to T. chilensis.

According to Article 33.3 of the International Code of Botanical Nomenclature (McNeill et al. 2006), the combination T. chiloensis var. eburneus (K. Schum.) F. Ritter is invalid, as Ritter did not provide basionym information: name, place and date of publication. (tricho2012)

Friedrich and Glaetzle (1983)

Nearing the boarder between morphological and molecular analysis, Friedrich and Glaetzle slightly updated the classification. Taking a closer look, they used a scanning electron microscopy (SEM) to study the ultra-structure of the seed testa of Echinopsis sensu lato (broad sense). This revision proposed nine groups, where the first four correspond to species of the subgenus Trichocereus, and in another they also included species of Soehrensia.

Molecular
Molecular evidence currently suggests that there are two primary clades of columnar cacti that arose from the South American ancestral populations, each having inferred common ancestries. The first clade consists of three former tribes, Browningieae, Cereeae, and Trichocereeae, which all share a unique 300 bp deletion in cpDNA and strongly suggests a close ancestor. Because of the limited data for molecular phylogenetic resolution, this group has been generically named the BCT clade until further relations are determined. Interestingly, Buxbaum (1958) originally proposed that these groups are related to one another and constituted a single major radiation in South American cacti. Arakaki et. al. 2011 indicates that Trichocereinae has a relatively recent evolutionarily origin, about 7.5-6.5 Mya, and that the genetic divergence between members is far lower than the difference shown by morphological and floral characters. Nyffeler & Eggli (2010), also point out that within the sub-tribe Trichocereinae, the most difficult group to interpret is the macro-genus Echinopsis. With most sources agreeing that despite being polyphyletic, all taxa within tribe Echinopsis are intimately related

Past 100 Years?

Since the turn of the new millennium, several molecular studies concerning Cactacea phylogeny have been published; many of which have sampled at least one member of the tribe Trichocereeae or the genus Echinopsis (s.l.) These studies often use specific markers in cpDNA regions.
Nyffeler (2002) in his study of the Cactaceae sequenced the chloroplast genes of 70 species of 48 genera. His results showed that the tribe Trichocereeae was paraphyletic in nine species sampled. In addition, Echinopsis (s.l.) represented by three species did not form a monophyletic clade.

Hernndez et al. (2011) compared the Cactacea on a broader scope, sampling 224 species of 108 genera, and analyzing multiple regions of cpDNA as well as an additional nuclear marker.
They found paraphyly in both the. Trichocereeae (represented by 18 genera and 33 species) and Echinopsis (seven species). More interestingly, they uncovered the three species studied of the subgenus Trichocereus as monophyletic.

Brcenas et al. (2011), used the same gene region as Nyffeler, but utilized a significantly greater range of 532 species
Found that the tribe Trichocereeae was monophyletic, but not the genus Echinopsis (s.l).

Sadly, despite the promise of as better measuring stick, even the current molecular techniques are still unable to absolve the conflicts of this war torn family.

 Species within the clade display a tremendous diversity in vegetative characteristics such as growth habit, size, and habitat preferences, as well as extreme variations in floral morphology and pollinator species, including insect, bat, hawkmoth, and hummingbird syndromes Some argue this clearly demonstrates that, flower characters and pollination syndromes are inconsistent and evolutionarily unpredictable, and therefore the presence or absence of a certain syndrome does not accurately measure phylogenetic relation between two lineages. In this sense, distinctions based on the different floral syndromes, such as those used by Backeberg (1966) to separate genera, now seem devoid of meaning.

1. 6. 3.
7. 2. 4. 5.

Echinopsis sp. 2, the labeled arrow indicates the brush-like section of the dorsal stamen but cluster, and the unlabeled arrow, a Helianthocereus, the dorsal stamen clusterais somewhat similar to that T. candicans, a stamens discrete nectar chamber is not Trichocereus candicans, arrow indicating point about halfway up theof nectar chamber. the of the throat are sheet of parallel sticky filaments orientedin longitudinally. Notice the position of the stigma: it has sunk down into the throat developed, a not unusual circumstance Helianthocereus grouped into a broad but loose dorsal stamen cluster which runs a simple course through the throat, the stamens more and against the brush-like part of the cluster. 4throat complexly developed and totally unlike those of the other trichocerei. Trichocereus spachianus. the stamens of the or less straight for most of their lengths then all distally curving too one side. Echinopsis oxygona, frontal section, which divides the flower and its dorsal stamen cluster into a top half and a bottom Trichocereus thelegonus. dorsal stamen cluster is somewhat similar to that of T. candicans. the black arrow indicates a the massive asymmetric funnel-like structure mentioned in the text. halfNote the style has retracted far into the throat. Trichocereus imperialis the dorsal stamen cluster is somewhat similar to that of T. candicans.

Shaman

Shaman