‘Cistanthe sp. subsp. subspeciosa,’ a specioid from the Atacama Desert, with comments on the taxonomy, ecology, and evolution of Cistanthe sect. Cistanthe (Montiaceae)

Cistanthe subspeciosa Hershk. (Cistanthe Spach sect. Cistanthe; Montiaceae) here is described as a herbaceous to suffruticose perennial from the vicinity of Copiapó, Chile. Its epithet is juxtaposed with its rank in order to highlight its subspecific aesthetic. In particular, the specioid is infrequent, locally restricted, and lacks unique diagnostic traits. Rather, it is diagnosed by a combination of traits characteristic of different Cistanthe species from the region, none of which can be identified as the referential species to which Cs. subspeciosa might be considered subspecific. The intersection between species nomenclature and species ontology thus is discussed. I hypothesize that Cs. subspeciosa is irrigated primarily by mountain runoff rather than localized precipitation, and that it might be both resistant to and dependent upon high substrate metal concentrations characteristic of the Copiapó region. The ornamental value of this and other Cistanthe species is discussed. Finally, additional historical details pertaining to Calandrinia spectabilis Otto & Dietr. and Cistanthe philhershkovitziana are provided.


Introduction
This tantalizingly-titled paper emerges as a sequel to a previous paper (Hershkovitz, 2019a), which reconstructed the discombobulated taxonomic history of the name Calandrinia speciosa Lehm. (Montiaceae), a mythical species known only from its protolog in an 1831 commercial seed catalog. For over 170 years, this name was applied haphazardly in reference to various species of Cistanthe Spach sect. Cistanthe, a taxon largely endemic to the Chilean Floristic Region (Hershkovitz, 2019b). Among the references to Ca. speciosa Lehm.is Muñoz Schick (1985), who published photos and a description of a so-identified plant growing in the Atacama Desert near Copiapó. No diagnostic basis was provided. The distribution was described as (effectively) 'growing in isolation in the interior of Atacama Province, sometimes with Cistanthe longiscapa (Barnéoud) Carolin ex Hershk.' In 2000, I collected a similar plant from desert flats near Copiapó, ca. 400 m elevation. Based only on the photo in Muñoz Schick (1985), I identified it as "Cistanthe sp., cf. C[alandrinia] speciosa Lehm." (Hershkovitz, 2006). But my knowledge of Cistanthe species diagnostics at that time, was, at best, rudimentary. My prior (though fruitful) research had focused only supraspecific-level diagnostics and phylogeny (Hershkovitz, 1991a(Hershkovitz, , 1993Hershkovitz and Zimmer, 2000;reviewed in Hershkovitz, 2019b). In 2018, I located photos of similar plants, also identified as Ca. speciosa, in an internet blog of an amateur botanist, Joachim, Zora (see below). The locality evidently is near Copiapó, and the plants are similar to Werdermann 405, a 1924 collection from the same general locality. Notwithstanding its low elevation (700 m), Peralta and Ford-Werntz (2008) cited Werdermann 405 as Cistanthe cabrerae (Añon) Peralta, a montane amphi-Andean species whose elevational range they listed as 1800-2500 m.
Reexamination of the evidence led me to conclude that the Copiapó area plants are similar to but distinct from other species of Cistanthe sect. Cistanthe (sensu Hershkovitz, 2006Hershkovitz, , 2018aHershkovitz, , 2019bcontra 1991a, b;1993). For example, besides the 1000+ m elevational discrepancy, they differ from Cs.
The above bases on the description and photos in Muñoz Schick (1985), images of Werdermann 405, blog-published images of live plants, and my notes and recollections. Additional investigation is needed to provide a more thorough description. Some critical data are missing, especially details of the stem geometry in vivo and the ovule/seed number.

Distribution and habitats.
Cistanthe subspeciosa is infrequent in desert flats and low-elevation canyons, 400-700 m elevation, in the vicinity of Copiapó and Tierra Amarilla. Possibly it grows/flowers here only during El Niño years, when this region receives considerable precipitation and, more importantly, mountain runoff. The flowering frequency, however, remains to be confirmed by additional study. I tentatively include in the species a plant I collected in the littoral zone north of Taltal (Antofagasta Region; see below).
Additional observations. CHILE: Atacama Region. Copiapó Province: road from Paipote to Diego de Almagro ca. 9 km north of Paipote, roadside, sandy soil, 27˚20'S, 70˚11'W, 500 m, 11 September 2000, Hershkovitz 00-91 [Specimen and photos destroyed; habit, flower, and habitat were identical to those in Muñoz Schick's (1985) photo; recorded as having ca., 100 stamens, 15 mm style, and hairy seeds]. ?Antofagasta Region. Antofagasta Province. Taltal Community: beach area along road between Taltal and Paposo ca. 26 km north of Taltal, rocky sandy disturbed flat on east side of road, 24˚10'S, 70˚26'W, 10 m, Hershkovitz 00-51 [Specimen and photos destroyed; Hershkovitz 00-51 appeared superficially similar to am individual evidently of Cistanthe grandiflora growing alongside ; field notes highlighted the contrasts between these individuals, the latter having conspicuously angled stems, ca. 40 stamens, and hairy seeds, while Hershkovitz 00-51 was recorded as having ca. 100 stamens a 15 mm long (exserted) style, and glabrous seeds.] Muñoz Schick (1985: 78),two photos of a plant and flower in a desert flat near Copiapó. The specific locality/elevation are not indicated. (ii) Personal blog of Joachim Zora (http://www.joachim.cl/desierto_florido/html2/flora/calandrinia_speciosa.htm), seven photos of plants and flowers. The locality/elevation are not indicated, but the blog indicates 2017 exploration of Quebrada de Paipote and Quebrada Cerrillos (= Quebrada Carrizalillo). The former is a dry wash originating ca. 10 km E of the Hershkovitz 00-91 locality and extending into the High Andes. Quebrada Carrizalillo originates south of Tierra Amarilla and also extends into the High Andes. Werdermann 405 is from this general region. Exsertion of the style and flowers with (40-)80-100 stamens are evident in Zora's photos. I e-mailed the contact address of Zora's blog requesting precise locality data and permission to reproduce his photos, but received no reply. (iii) Hershkovitz (1991c: Figs. 8, 44, 63, 68, 79, 80) illustrated leaf venation and epidermal morphology of Werdermann 405.

Illustrations. (i)
Etymology. The epithet subspeciosa is derived from appending the prefix sub-(less than) to the feminine form of the adjective speciosa (attractive, showy). Thus, subspeciosa is a valid Latin term meaning "less than attractive/showy." Its applicability to Cistanthe subspeciosa is a matter of opinion. But the epithet was chosen for different aesthetics. Firstly, it refers to one of several forms of Cistanthe historically erroneously referred to as Cistanthe speciosa Lehm. [viz., by Muñoz Schick (1985) and, following this reference, Hershkovitz (2006), Squeo et al. (incl. Muñoz Schick;, and Joachim Zora's blog (see above), which references Muñoz Schick (1985). Hershkovitz (2019a) traced the taxonomic history of the name Ca. speciosa Lehm. and determined that it has not been typified but probably refers to plants of Cs. laxiflora (Phil.) Peralta & D.I. Ford (see below). The epithet subspeciosa also refers aesthetically, but not literally, to the subspecific quality of Cs. subspeciosa, viz., an infrequent form that, while distinct, might be perceived as a subspecies, except that in this case, the referential species is not certain (see below). Finally, the epithet calls attention the millennially and perpetually irresolvable ontology of species and subspecies and the boundaries between them.

1.
Cistanthe grandiflora, Cs. cachinalensis, and Calandrinia taltalensis: Cistanthe subspeciosa shares with Cs. grandiflora (as conceived here) its degree of succulence and glaucous texture, somewhat sloppy habit with large, rubbery ovate leaves crowded at the base and along the lower portion of ascending branches. It differs in having less conspicuously or not angled stems and flowers with 80-100 stamens and long, exserted styles.
As discussed in Hershkovitz (2018aHershkovitz ( , 2019acf. Muñoz Pizarro, 1966), the historically haphazard application of the name Cs. grandiflora has been a major encumbrance in advancing understanding of species taxonomy of Cs. sect. Cistanthe. Much of the problem traces to Reiche (1898a), who lumped all then-named species of the section into a single polymorphic Cs. grandiflora. Taxonomists/floristicians (and, worse, ecologists, biogeographers, and phylogeneticists, including myself) ever since have followed Reiche (1898a) erratically and inconsistently. As a disclaimer, however, my earlier work focused on supraspecific-level diagnostics and relationships. I deferred species identifications to the expertise of numerous contemporary and historical card-carrying species/floristic taxonomists. In the case of Cistanthe, however, nominally "expert" identifications were not much better.
As conceived here, Cs. grandiflora includes essentially all Cistanthe sect. Cistanthe with the conspicuously angled stems, ovate leaves along the lower portion of the inflorescence stems, ca. 50 stamens, and styles not exserted. It is distributed continuously between the Aconcagua and Antofagasta Regions, below 1000 m, especially near the coast. I have seen evidence of possible (but unconfirmed) occurrences in the O'Higgins and Maule Regions, but not south of this (i.e., to Araucania; cf. Peralta and Ford-Werntz, 2008;Rodríguez et al., 2018). Such occurrences are not impossible, though, because the species is weedy and can flower in its first season (Hershkovitz, 2018b;cf. Loudon, 1840). Peralta and Ford-Werntz (2008) listed the distribution of Cs. grandiflora as ranging from Chile's Coquimbo to Araucania Regions, but they included therein Cistanthe discolor (Schrad.) Spach and, implicitly, the southerly-distributed Cistanthe mucronulata (Meyen) Carolin ex. Hershk. (Hershkovitz, 2018a). This taxonomy is untenable (Hershkovitz, 2018a). Peralta and Ford-Werntz (2008) recognized as distinct Cs. cachinalensis (including Ca. taltalensis) in the Antofagasta and Atacama Regions, and in the Antofagasta Region also Cs. lamprosperma. Rodríguez et al. (2018) more or less followed Peralta and Ford-Werntz (2008), but they extended the range of Cs. grandiflora northward to the Antofagasta Region and Cs. cachinalensis southward to the Coquimbo Region. Thus, according to Peralta and Ford-Werntz (2008), the species are parapatric, whereas in Rodríguez et al. (2018), they are sympatric, which begs the question of their distinction.
Although I have vacillated on this point (Hershkovitz, 2018a), I am inclined (today) to include Cs. cachinalensis in Cs. grandiflora, because I cannot discriminate the corresponding specioids (cf. Rosas and Estévez, 2010). Philippi (1893) did not characterize the stem geometry of Cs. cachinalensis. Numerous internet photos identify individuals from the Antofagasta Region as Cs. cachinalensis, most of which appear within the range of variation of Cs. grandiflora in the Coquimbo and Atacama Regions.
Possibly there has been a tendency to consider by default all Antofagasta plants to be Cs. cachinalensis or Ca., taltalensis and not Cs. grandifora (e.g., Finger and Tellier, 2010;Orrego et al., 2013). Watson (2019) includes photos of inflorescence termini of plants identified as Cs. cachinalensis (Fig. 57, from the Antofagasta Region and Cs. grandiflora ( Fig.  60) from the Coquimbo Region. The former appears to be coarser, but otherwise I see no difference. Watson (2019) also includes a photo of a whole plant identified as Cs. grandiflora ( Fig. 50) from the Atacama Region, but this plant is a common non-pachycaulescent form (or stage) of Cs. crassifolia (Phil.) Carolin ex Hershk. However, as I will consider in a later work, this form might have experienced gene exchange with Cs. grandiflora. My interpretation at present is that the coarser Cs. cachinalensis forms represent plastic variability corresponding to a stressful habitat and/or developmental mutants ("apospecies;" Hershkovitz, 2019b) likely to interbreed with "normal" Cs. grandiflora in the same zone.
The taxonomy of the specioid Ca. taltalensis remains problematic (cf. Finger and Teillier, 2010;Rosas and Estévez, 2010). Johnston (1929) did not describe the stem geometry and stamen number. He discriminated Ca. taltalensis from Cs. grandiflora mainly on the basis of its coarser inflorescence bracts and short and not reflexed pedicels. I am inclined to believe that these traits are both developmentally plastic in response to environmental conditions and/or genetically easily mutable. In particular, a defunct blog created by Chileans Juan Acosta and Florencia Senoret contains a photo of a plant from Cuesta Paposo they identified as Ca. taltalensis. It appears to be a developmental mutant of Cs. grandiflora with an unelongated but fully mature, flowering and fruiting "capitate" inflorescence (http://blog.spiniflores.com/archives/tag/desierto-florido). Again, my effort to communicate with the authors was unsuccessful.
The Hershkovitz 00-51 tentative Cs. subspeciosa collection from the Taltal area also might be considered a candidate for Ca. taltalensis, but the former has 100 stamens and glabrous seeds and the latter hairy seeds (Johnston, 1929). It seems unlikely that Ca. taltalensis had 100 stamens, or Johnston likely would have noted this. Analysis of Johnston's specimen should resolve this question.
A propos Cs. subspeciosa, had I conceived in 2000 what I conceive currently, I would have recognized Hershkovitz 00-61 as Cs. grandiflora and Hershkovitz 00-51 as a distinct species. Similarly, while I recognized that Hershkovitz 00-91 from near Copiapó was distinct from Cs. grandiflora, I did not appreciate that true Cs. grandiflora grew nearby, in particular Hershkovitz 00-114, collected on sandy flats ca. 30 km SSW. Hershkovitz (2006) identified this only as "Cistanthe sp." My field notes indicate "plants 80 cm, stems angular, petals 3 cm, bright fuchsia [obs.: more accurately rose-magenta], yellow at claw, stamens ca. 40, filaments red, yellow toward base, anthers red, style 2 mm, pink, stigma green, seeds hairy." Thus, this plant differs from Cs. subspeciosa in all of the diagnostic traits. The observation demonstrates that Cs. subspeciosa and Cs. grandiflora s. str. occur geographically proximally. It is not clear whether they may occur "side-by-side," especially at higher elevations in Tierra Amarilla Province.
Somewhat further away is Zöllner 4079 [CHILE. Atacama Region. Tierra Amarilla Province, Valle Jorquera, (obs.: probably 2000-2800 m), 11 January 1979 (L!, NA!)]. From the specimens, I cannot diagnose the floral morphology. I presume they represent Cs. cabrerae because of the elevation. The Copiapó area plants of Cs. subspeciosa are only ca. 70 km WNW of the Zöllner collection, but the elevational difference is >1000 m and the ecology correspondingly drastically distinct. Equally importantly, I have not located Cs. sect. Cistanthe collections in the intervening elevational zone, 1000-2000 m, of the Atacama Region. Peralta (1993) listed the Chilean distribution of Cs. cabrerae as Antofagasta and Atacama Provinces, 1700-2000 m. Peralta and Ford-Werntz (2008) listed its Chile-Argentina elevational range as 1800-2500 m. Inexplicably, they cited Werdermann 405, from 700 m, as the representative Chilean specimen. The distribution in the Antofagasta Region evidently owes to an unidentified locality indicated on a map in Peralta (1993), ca. 550 km NNW (along the Arid Diagonal) of bona fide Cs. cabrerae localities in Atacama Province. The indicated locality is perhaps 25 km NNE of the city of Antofagasta. An elevation above 1700 m is possible in that region, but, in light of the present work, the locality and identification of the specimen beg Coquimbo Region. I am not aware of inclusion of Cs. speciosa Lehm. in other contemporary local floristic listings, and Marticorena and Quezada's (1985) national list did not mention the name at all. This might render the impression that Cs. speciosa Lehm. is endemic to the Atacama Region. Ironically, the specioid itself is possibly endemic. It simply had been misidentified as Cs. speciosa, which is part of the reason I chose the new epithet subspeciosa.

5.
Calandrinia corymbosa: Walpers (1843a;cf. Walpers, 1843b) described this species as originating in Copiapó, evidently what Meyen (1834: 400) referred to as "Thalinum (glauca?)" and described only as a plant with thick, succulent leaves. Walpers' (1843a, b) description was more elaborate, but still did not serve to distinguish Ca. corymbosa from other species of Cistanthe sect. Cistanthe. The only trait that was supposed to distinguish Ca. corymbosa from Cistanthe glauca (Schrad) Lilja (= Cs. grandiflora; Hershkovitz, 2018a;2019a, b) was green rather than glaucous leaves. This trait does not seem to have taxonomic value, as I have found that essentially all species of Cs. sect. Cistanthe vary from more green to more glaucous depending upon conditions.
Other than that, Walpers' (1843a, b) description applies to Cs. cabrerae, Cs. grandiflora, and Cs. subspeciosa. The diagnostic trait of Cs. subspeciosa is the high stamen number and long style. But Walpers (1843a, b) indicated that the flower of Ca. corymbosa was unknown. Paradoxically, Walpers (1843a) indicated that its (northern hemisphere) flowering time was May. This and the diagnostic trait (green versus glaucous) suggests that the species was described from a cultivated plant. But, why would the flower be then "unknown"? Barnéoud (1846) accepted Ca. corymbosa without critical comment, but it appears that he had not seen a specimen. Philippi (1893) included with a question mark the name Ca. corymbosa in his list of segregates of Cs. grandiflora, not describing/discussing it further. Reiche (1898) listed it among "problematic" species of Calandrinia s. l., there reproducing Walpers' (1843a, b) description. The only subsequent mention of Ca. corymbosa that I have located is Squeo et al. (incl. Muñoz Schick;, which lists the species as possibly extinct in Copiapó Province. As noted above, this work listed Ca. speciosa Lehm. as, effectively, endemic to Copiapó and Tierra Amarilla Provinces. The authors apparently did not notice that the plants identified as Ca. speciosa Lehm. conformed better to the description of C, corymbosa. For that matter, they did not consider whether Ca. corymbosa might refer to Cs. cabrerae, since the descriptions agree and Squeo et al. (2008) listed Cs. cabrerae for Copiapó Province. Peralta and Ford-Werntz (2008) listed Cs. corymbosa as a doubtful name, and Rodríguez et al. (2018) did not mention this name.
Meyen (1834) did not indicate precisely the locality or the elevation of his plant. The general locality is an unnamed canyon from "Ramillas," a place name I cannot locate, to Mina Checo. The latter might refer to any of many mines in the historical "Checo mining district," these spanning considerable elevational range and distances from Copiapó (https://www.geovirtual2.cl/minas/Atacama-Checo-de-Cobre-minas-01.htm). Comparison of Meyen's (1834) chronicle with a later English summary (Royal Geographical Society, 1836) implicates an elevation below 1000 m somewhere between Nantuco and Los Hornitos (CHILE. Atacama Region. Tierra Amarilla Province). This approximates the inferred localities of Werdermann 405 and Joachim Zora's plant (see above). But the presence of Cs. grandiflora in this range cannot be ruled out. Meyen evidently continued on to rather high elevations, where Cs. cabrerae might be found.
Yet, there seems to be a better case for accepting M. solanoides than Ca. corymbosa. In the latter case, there is not even a type photo. The description is not Meyen's (1834) original, but Walpers' (1843a) second hand, and possibly based on a cultivated specimen. Description of the diagnostic androecial and gynoecial traits is missing. If the origin indeed is from > 1000 m, it might be Cs. grandifora. If it is from higher up, it might be Cs. cabrerae. Thus, here I reject the name Ca. corymbosa.  (Hershkovitz. 2018b). In addition, I now can identify as Cs. grandiflora most samples in Hershkovitz (2006) identified as "C. sect. Cistanthe sp. [##-##]."

Molecular diagnostics of
Unfortunately, although nominally interspecific-level DNA markers were employed, sequence divergence among Cistanthe sect. Cistanthe species was extremely low and revealed no significant phylogenetic or taxonomic structure. The relationships illustrated graphically might be consequent to homoplasy, lineage sorting, or gene flow. The two samples here purported to represent Cs. subspeciosa (Hershkovitz 00-51 & 00-91) indeed shared both nuclear and plastid marker genotypes, but these were shared also by a sample here identified as Cs. grandiflora (Hershkovitz 00-214) from a ruderal locality ca. 30 km E of La Serena in the Coquimbo Region. Hershkovitz (2019b) noted that Cs. grandiflora is an especially weedy species, usually found in disturbed sites throughout its extensive range, < 1000 m between at least 24 -33S. Hershkovitz (2006) demonstrates that the genotypes of samples Cs. grandiflora (when correctly identified) are highly variable and shared variously with other species.
Ontology of Cs. subspeciosa. I conceive of Cs. subspeciosa typologically, i.e., as a typereferenced specioid that comprises all individuals considered to be conspecific to Werdermann 405 and to the type of no other named species. As articulated in Hershkovitz (2019b), I do not presume that this or any specioid conforms to any of the numerous ontological "species concepts," least of all the "phylogenetic species concept" (PSC). The PSC appeals to an affirmation that "species are…clades" (i.e., terminal branches of phylogenetic trees; David Baum, U. Wisconsin, written comm., 25 December 2015). The PSC became and possibly remains popular among systematic biology's institutional socioeconomic/political elite. Yet, prior species theory/data (e.g., for gene flow, lineage sorting) already betrayed the PSC idealization as a concept dead-on-arrival. Its subsequent and persistent popularity possibly owes to the operational/mathematical convenience of reducing species to single dimensions for phylogenetic zealots challenged to count any higher.
A type-referenced specioid is non-dimensional (cf. Brooks and M cClennan, 1999), but there is no limit to the number of dimensions in which its existence might be corroborated. Cistanthe subspeciosa here is distinguished from an apospecific variant in its morphological and spatiotemporal geographic dimensions. In particular, the morphologically distinctive plants have been recorded in the Copiapó vicinity on at least four occasions spanning nearly a century: 1924, some few years before 1985, 2000, and 2017. The putative Taltal-area observation is not germane here. The observations suggest that the specioid is locally transitionally stable.
The Copiapó vicinity geography itself explains the infrequency of collection. Significant precipitation arrives to the Atacama Region central valley only during El Niño years, often 10+ years apart. This spawns Chile's "desierto florido" phenomenon, in which perhaps 5000 km 2 of normally barren earth rapidly transforms into a carpet of flowers. This is when botanists flock to the desert to collect plants. But even during such years, rain concentrates in the southerly Huasco Province and decreases precipitously towards and north of Copiapó. Naturally, botanical collection concentrates in the vegetated region, which distracts attention from the fewer and isolated individuals of specioids like Cs. subspeciosa in the otherwise barren margins. Moreover, the more floriferous the desert, the more likely rare and distinct individuals will fall through the collection net. This is especially true in the case of Cistanthe sect. Cistanthe, given the 120 year-old conventional wisdom that they all are the same species (Hershkovitz, 2019a, b). In this context, the four Copiapó-area records of Cs. subspeciosa spread over a century are significant.
Available evidence offers little insight into the history of this specioid. But the impulse to interpret Cs. subspeciosa or any specioid as a static or statistical entity must be resisted. In terms of the nonlinear wave model of species (Hershkovitz, 2019b), it represents a transitionally stable entity, but its present state might be explained by innumerable histories, e.g., a recently-evolved offshoot, a hybrid arising one more times, or a vestige of a once widespread form. It might have migrated/dispersed from elsewhere, or it might be a relict. In terms of the Principle of Evolutionary Idiosyncraticity (Hershkovitz, 2019b), its history and future are determined not extrinsically, but intrinsically and nonlinearly owing to the capacity of living beings to enact their own history via self-organized metabolism, growth, development, and reproduction structurally coupled to a dynamic hierarchically-organized physical milieu (cf. Maturana and Varela;Varela et al., 1992;Salthe and Matsuno, 1995;Mpodozis and Maturana, 2000;Hershkovitz, 2019b). Evolutionary history in a nutshell.
For formal taxonomic purposes, as intimated above, Cs. subspeciosa projects a "subspecific" aesthetic. While unique, it lacks unique diagnostic characteristics, its range is relatively small, and its frequency is low. Its collectively distinctive character emerges from its combination of traits individually shared by other geographically proximal and more broadly distributed species of Cs. sect. Cistanthe. But a subspecific aesthetic also emerges from its taxonomic history. In particular, identification notwithstanding, Muñoz Schick (1985) and myself (Hershkovitz, 2006) indeed recognized that the morphology of the Copiapó plants was distinct from that of other (geopolitically) Chilean-endemic species of Cistanthe sect. Cistanthe. Peralta and Ford-Werntz (2008) identified at least the Tierra Amarilla plants as Cs. cabrerae, while listing Ca. speciosa as a synonym of Cs. grandiflora. In this perspective, the Copiapó-area plants lost their taxonomic distinction. This history is typical of taxonomically subspecific specioids.
My initial sensibility was to describe Cs. subspeciosa as a subspecies of Cs. cabrerae, no, wait, maybe Cs. grandiflora, but then again, etc. I then mused over the formally invalid name "Cistanthe sp. subsp. subspeciosa." After all, anyone familiar with the taxonomic, floristic, and broader scientific literature would appreciate that "Cistanthe sp." is by far the most common and broadly distributed species of this genus. The epithet "subspeciosa" has been published validly twice in the botanical realm: Rhynchospora consanguinea f. subspeciosa Kük (Cyperaceae) and Dicranopteris linearis (Burm. f.) Underw. var. subspeciosa Holtt. (Gleicheniaceae). But, in both cases, the epithet has been applied to a subspecific entity. The present instance is the only time the epithet has been applied at and in juxtaposition to the nominal species level. Further investigation might change this status. The name might be reduced to synonymy, or indeed it might be recombined at the subspecific level.
It is intriguing in this context to recall that, while the botanical nomenclatural Code prohibits "species-free" taxa, the International Code of Nomenclature for Cultivated Plants (Brickell et al., 2016) does not. It formally recognizes, e.g., "Genus 'Cultigen,'" which does not imply conspecificity with the type species of that genus. The cultivated Code, however, does defer to the botanical Code as to the legitimacy of application all botanical names in cultivated plant taxonomy. But the relation of the botanical and cultivated Codes is not hierarchical ontologically. They merely intersect operationally. The botanical code is agnostic towards the ontology of biological species, nevertheless obligates species as the universal fundamental biological taxonomic unit. The cultivated Code permits recognition of specioids exclusive of taxonomic species. Paradoxically, per the wave model, the cultivated Code is correct biologically (viz., "not all water pertains to a wave;" Hershkovitz, 2019b). In truth, however, there is no philosophical conflict to the degree that the botanical Code obligates species only nomenclaturally and not ontologically. In no other sense does it specify species ontological equivalency.
Ecology of Cs. subspeciosa. As noted above, Muñoz Schick (1985) indicated that Cs. subspeciosa (as Ca. speciosa Lehm.) grew isolated or together with Cistanthe longiscapa (Barnéoud) Carolin ex Hershk. When I encountered a single individual near Paipote in early September 2000, it was isolated, so strikingly so that I do not recall whether any plants at all grew nearby. 2000 was an El Niño year that yielded an impressive desierto florido in the Huasco Province interior, but the effect gradually decreased northwards. By perhaps 50 km S of Copiapó, the interior desert was fairly barren, the plants, mainly perennials/geophytes and unworldly annual Philippiamra spp., few and far between. Meteorological records (Gobierno de Chile, 2019) corroborate my observations. Precipitation decreased from 73 mm to a mere 37 mm between Vallenar and Copiapó.
However, precipitation is only part of the story of Chile's desert flora, and, in the case of Copiapó-area perennials like Cs. subspeciosa, not the greater part. The greater part is runoff from the Andes, where, with increasing elevation, precipitation becomes several-fold higher (see, e.g., AGRIMED, 2019, and note that rainfall averages combine El Niño, La Niña, and "normal" year totals). Given the height and steepness of the Andes, even a 10 mm sprinkling in Vallenar or Copiapó raises the specter of catastrophic flooding. Thus, if it seems fantastic that a large, succulent individual of Cs. subspeciosa can thrive on less than 40 mm of precipitation, it is because indeed it is fantastic. The proportion of irrigation that a Cs. subspeciosa individual receives from a one-meter radius of rainfall is probably very small. Of the larger remaining proportion, I will not speculate how much represents surface versus aquifer irrigation. For the purposes of this paper, I have not researched literature on the contribution of broader area geological hydrology to the historical origin and episodic maintenance of Chile's desierto florido. I put this on my "to do" list.
Water, however, is not everything, The central and southern Andes are known for their rich deposits of metals, copper being the most abundant in Chile, but also gold, silver, among others, and the Atacama Desert region is said to harbor the world's largest deposits of lithium. Presumably the last helps plants in this challenging environment overcome at least their emotional stress. Rocks of the high Andes east of the Vallenar-Copiapó valley are exceptionally rich in gold, its commercial exploitation politically highly contentious. In fact, Cs. cabrerae is endemic to montane zones immediately beneath the gold-rich peaks on both sides of the Andes. For the purposes of this paper, I have not researched literature on the degree to which the distributions of Cs. subspeciosa and Cs. cabrerae might relate to the quantity/quality of metals filtering in runoff/aquifers. The specioids might be especially resistant to or, via genetic assimilation, dependent upon localized mineral concentrations. This relation might be direct or indirect, i.e., via mineral metabolism of co-localized microbes.
It seems clear that Cs. subspeciosa hyperarid Atacama perennials depend upon hydrological more than direct climatological irrigation. In this context, Hershkovitz (2019b) criticized a slough of "Big Data" research papers published in "high impact" journals purporting to explain ecological, evolutionary, and biogeographical phenomena in terms of correlations with fine-scale global rainfall data. Much of my criticism was epistemological, but, epistemology aside, in situ natural history betrays the farce of this research program. For the benefit of some of (nominal) plant physiological ecologists, Hershkovitz (2019b) explained his admittedly rudimentary understanding that "mean annual precipitation" and "moisture availability" are not the same thing. The popularity of this trendy approach seems to owe less to science than to the ease with which downloaded Big Data can be analyzed with downloaded Big Software, facilitated by artificial intelligence tools that aid researchers deficient in the real thing. Expectedly, the approach [de-]generates [into] numerous "high impact" publications, the Bitcoins of modern institutional science. The ecology of Cs. subspeciosa and the lengthy periodicity of Atacama rains provide just two of innumerable examples that demonstrate the absurdity of mean annual precipitationbased hypotheses.
Ornamental value of Cs. subspeciosa. As noted, the epithet subspeciosa literally means "less than attractive/showy." This hardly applies to the flowers of Cs. subspeciosa, which are among the largest in the genus. With its 80+ yellow anthers contrasting the pink-mauve petals, it is more appealing than the occasionally cultivated "true" Cs. grandiflora. Presumably, Cs. subspeciosa is not hardy, but, like other Cs. sect. Cistanthe spp., it probably can be grown as an annual (cf. Loudon, 1840) or, alternatively, indoors. But the sloppy vegetative architecture shared by Cs. subspeciosa and Cs. grandiflora indeed demands a mother's love. On the balance, Cs. subspeciosa offers a conspicuous accent for lowmaintenance water-conserving wildflower gardens. Its form might be better controlled in a container planting. Without the flowers, however, it might be better suited for greening up landfills, homeless camps, and artillery ranges.
The most widely cultivated species of Cs. sect. Cistanthe is Cs. laxiflora, though this species usually has been distributed incorrectly as Cs. grandiflora or Ca. spectabilis Otto & Dietr. (Hershkovitz, 2019a). Within the past 20 years, cultivation of this species has gone viral especially in southern California, where possibly it has become adventive. Underexploited horticulturally is Cs. discolor (Schrad.) Peralta & D.I. Ford, which has neat rosettes of succulent coriaceous leaves deep green and white-mottled adaxially and literally beet-red (rarely green) abaxially. Given its relatively high-altitude native range in central Chile (1000-2000 m), this species presumably is hardy but, in any case, also can be grown as an annual (Loudon, 1840). One would suppose that the montane Cs. cabrerae also is hardy.
Much of the early taxonomic literature on Cs. sect. Cistanthe was oriented not towards botanical science, but horticulture. This likely contributed to the taxonomic ambiguity and confusion that persists to the present day, as persistence of popularized Latin names diverges from the evolving botanical taxonomy. As noted in Hershkovitz (2019a), the taxa Ca. speciosa Lehm. and Ca. spectabilis were published in horticultural literature, as was Loudon's (1840) early taxonomic summary of Montiaceae species. But even the taxa described/illustrated in nominally botanical literature (Lindley, 1828(Lindley, , 1833(Lindley, , 1839Hooker, 1834aHooker, , b, 1835 emphasized cultivation instructions and not natural history. Like Lehman (1831) and Otto and Dietrich (1833), most of the descriptions were based not on field-collected specimens, but on plants cultivated from acquired seeds. As Loudon (1840) noted, some of the botanical publications did not state even the country of origin. Possibly the emphasis on horticulture owes to economics. Botanical research in this period was financed partially by the sale of ornamental plants and literature to an emerging petit bourgeois that would not have found appeal in the minutia that excites taxonomists.

Addenda to Hershkovitz, 2018c, 2019a
Otto and Dietrich (1833) noted that the plant they described as Ca. spectabilis originated from seed of a plant collected by Carlo Bertero. Based on the protolog and Bertero's itinerary in Chile (Delprete et al., 2002), Hershkovitz (2019a) surmised that Ca. spectabilis might refer to Cistanthe grandiflora s. str., Cs. laxiflora, or Cs. mucronulata. Since posting Hershkovitz (2019a), I located two additional Bertero collections pertaining to Cs. sect. Cistanthe. All bear variations of the identification "Talinum crassifolium." One is Bertero 684 evidently from mountains along the Río Claro in the Rancagua area [CHILE. O'Higgins Region. Cachapoal Province [P: P04583007 (Fig. 2); P04583008]. P04583007 indicates an 1828 date, and this accords with Bertero's itinerary. P04583008 indicates a November, 1829 date, consistent with a notion that dates/specimens (understandably) may have been mixed/corrupted by recipients of these collections. In any case, this collection would seem to solve the riddle of the "missing" Bertero 684 on the "Bertero 684 [and] 1349" (NY: 02065851) sheet of Cistanthe philhershkovitziana (Hershkovitz, 2018c). Bertero 684 appears to be Cs. mucronulata, confirming my speculation. But it possibly is Cs. grandiflora. Its identity must be confirmed by physical examination and by surveying plants near its locality.
Another collection is Bertero 1349 from Quillota [CHILE. Aconcagua Region. Quillota Province [P: P04583004 (Fig. 3), P04583005 (Fig. 4); see also below]. The first indicates an 1829 collection date, again agreeing with Bertero's itinerary. The second includes an individual of Cs. philhershkovitziana. The large plants appear to me to resemble Cs. grandiflora s. str. rather than Cs. laxiflora, and indeed they cannot be the latter littoral zone species unless sea levels were much higher in those days, or at least there had been one hell of a tidal wave.
But these Bertero 1349 sheets are problematic, because this is the same collection number as the NY sheet of Cs. philhershkovitziana! This suggests that the various curators of the martyred Bertero's Chilean haul did not appreciate how strikingly different was the herbaceous Cs. philhershkovitziana from its large, suffruticose congeners, hence mounted them on the same sheet believing them to be the same. Evidence supporting this hypothesis includes the two mixed collections at K (Hershkovitz, 2018c), the misidentification of Werdermann 39 (E: E00033182) by noted Cistanthe expert I. E. Peralta (MERL), and the nearly two-century oversight of this conspicuous and conspicuously-distributed seaside species by all other botanists, Chilean and non-Chilean alike. All of this history involving so many distinguished botanists, and all these years I believed that it was me that was the crappy Cistanthe taxonomist! However, another of the Bertero sheets is perplexing, P04583006 (Fig. 5). Like NY: 02065851, it is labeled as Bertero 684 [and] 1349, and it is a mixed collection that includes Cs. philhershkovitziana. Also like NY: 02065851, a date of 1830 is indicated. But the collection locality is given as Valparaiso, and indeed Bertero was in Valparaiso in 1830. Superficially, this at least hints that a Cs. laxiflora collection from Valparaiso was possible. Another twist is a secondary dual identification, "Calandrinia speciosa Lehm." ´presumably for the larger plant and "[Calandrinia] picta Gill." presumably for the smaller individual of Cs. philhershkovitziana. This manifests curator recognition that the two plants were distinct. Whether it implicates the original source of Ca. speciosa Lehm. is another question.
However, to me, the large plant of P04583006 also resembles Cs. grandiflora more than Cs. laxiflora, though physical examination is desirable. I am inclined to believe that the label is not Bertero's, and that it is erroneous. Hershkovitz (2018c) noted that Bertero 683 indicates Rancagua and Bertero 1348 and 1350 both indicate Quillota, so it is unlikely that Bertero 684 or 1349 could be from Valparaiso. Also, Bertero departed Chile in September, 1830 (Delprete et al. 2002), whereas Cs. laxiflora is a comparatively belated bloomer, not busting blossoms in the Valparaiso region until latest October and continuing through summer.
But the P04583006 sheet raises another possibility. In particular, there seems to be a missing collection number, since Bertero 684 now is accounted for, while there are two species sharing Bertero 1349. Since P04583006 has two plants and two numbers (Bertero 684 and Bertero 1349), possibly the larger plant is Bertero 684 from Rancagua and the smaller Cs. philhershkovitziana is Bertero 1349 from Quillota. However, there are two sheets with only large plants labeled Bertero 1349. But given the manifest mix-ups in the curation of these collections, no hypothesis can be discarded. Again, forensic physical examination is needed to resolve these questions.
The upshot is that there is positive evidence that the Bertero collection spawning the name Ca. spectabilis could be Cs. grandiflora or Cs. mucronulata, but only circumstantial evidence that it could be Cs. laxiflora. Nonetheless, in horticulture, the name Ca. spectabilis historically has been applied commonly to Cs. laxiflora (Hershkovitz, 2019a). Such things happen in horticultural taxonomic history. I have seen herbarium specimens of cultivated plants of Cs. grandiflora labeled as Calandrinia elegans, whereas no species of this genus ever has been named thusly formally. The only elegans validly published in Calandrinia is Ca. elegans Spach, a synonym of the small annual California red maids, Calandrinia menziesii (Hook.) Torrey & A. Gray (Hershkovitz, 2019a). And the name Calandrinia speciosa, referring to Ca. speciosa Lindl. non Lehm., was applied popularly to Calandrinia menziesii (Hook.) Torrey & A. Gray well into the 20 th century. This is even though the illegitimacy of Lindley's name was recognized by the mid-19 th century (e.g., Walpers, 1846). Thus, effectively, in horticultural usage, the genera corresponding to the epithets elegans and speciosa were swapped. Vernacular transfer of epithets from one to another species of the same genus is all the more common. The identity of Ca. spectabilis thus remains unresolved.      (1830) probably are erroneous. The plant on the left probably is Cistanthe grandiflora from either the Rancagua or Quillota areas, and on the right is Cistanthe philhershkovitziana from Quillota.